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Structure and Composition of Vegetation Along an Elevational Gradient in Puerto Rico - 563

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Structure and Composition of Vegetation Along an Elevational Gradient in Puerto Rico - 563 Journal of Vegetation Science 17: 563-574, 2006 © IAVS; Opulus Press Uppsala. - Structure and composition of vegetation along an elevational gradient in Puerto Rico - 563 Structure and composition of vegetation along an elevational gradient in Puerto Rico Gould, W.A.*; González, G. & Carrero Rivera, G. International Institute of Tropical Forestry, USDA Forest Service, Río Piedras, PR 00926-1119, USA; *Corresponding author; Fax +1 7877666302; E-mail [email protected] Abstract Introduction Question: What are the composition, conservation status, and structural and environmental characteristics of eight mature Vegetation composition in Puerto Rico is controlled tropical forest plant communities that occur along an elevational by four key factors: climate, substrate, topography and gradient. human and natural disturbance (Britton & Wilson 1924; Location: Northeastern Puerto Rico. Methods: We quantified the species composition, diversity, Dansereau 1966; Weaver 1991; Lugo 2005). Climatic conservation status, and ecological attributes of eight mature gradients are related to an elevational rise from sea level tropical forest plant communities in replicated plots located to up to 1340 m on the central cordillera of the island, the sample representative components of important forest types northeasterly trade winds, and the rain shadow effect of occurring along an elevational gradient. A suite of environ- the mountains. Diverse geologic substrates include vol- mental and vegetation characteristics were sampled at each canic, limestone, and serpentine bedrock and colluvial, plot and summarized to characterize communities and analyse alluvial, and marine quaternary deposits. Within these trends along the elevational gradient. broad landscape features community composition is Results: The set of communities included 374 species; 92% controlled by topographic effects on soil moisture and were native, 14% endemic, and 4% critical elements (locally endangered) to the island. All communities, occurring within a development (slope position). Disturbance regimes (e.g. wide range of patch sizes and degree of conservation protec- land use, flooding, fire, landslides, and hurricanes) in- tion, showed a high percentage of native species (> 89% per duce effects on the subsequent succession of secondary plot). The lowland moist forest communities, occurring within vegetation. The resulting landscape in Puerto Rico is a a matrix of urbanization, agriculture, and disturbance, had the mosaic of (1) primary forests that have not experienced highest degree of invasion by exotics. Community descrip- deforestation – according to Wadsworth (1951) and tions were nested within a variety of hierarchies to facilitate Birdsey & Weaver (1982) < 1% of the landscape sup- extrapolation of community characteristics to larger ecosys- ported climax vegetation by 1950; (2) mature secondary tem units. Basal area, above-ground biomass, canopy heights, forests (< 10% cover of the island); (3) young secondary and mean species richness peaked at mid elevations. Conclusions: It is significant that all of these forest communi- forests and shrublands (> 30%); (4) agriculture, grass- ties continue to be dominated by native species while existing lands and pastures, (> 30%); and (5) developed areas in a matrix of human and natural disturbance, species inva- and urban forests, (> 15%). The mature upland forests sion, and forest regeneration from widespread agriculture. The are primarily found in the upland forest reserves which lowland moist and dry forest types represent a minority of the occupy ca. 4% of the island. These protected areas protected forested areas in Puerto Rico, serve as unique ge- harbour our greatest wealth of native, endemic, and netic reservoirs, and should be protected. endangered plant species (Figueroa & Woodbury 1996). There are over 2800 species of native and exotic flower- Keywords: Detrended Correspondence Analysis; Exotics; ing plants in Puerto Rico and 9% of these are endemic to Gradient analysis; Land management; Native species; Phyto- the island (Liogier & Martorrell 1999). Understanding sociology; Puerto Rico. the floristic composition of mature forest communities is important for biodiversity conservation and land man- agement planning as these serve as both reservoirs of Nomenclature: Liogier & Martorrell (1999). species and provide important ecosystem functions. Community-level descriptions of plant associations Abbreviations: AB = above-ground; BA = Basal area; form the basis of hierarchical phytosociological classifi- DCA = Detrended Correspondence Analysis; CE = Critical cation (Braun-Blanquet 1964; Mueller-Dombois & element; I = Introduced; MAP = Mean annual precipitation; Ellenberg 2002). This approach has been more preva- MAT = Mean annual temperature; N = Native. lent in temperate than tropical vegetation studies. A few 564 Gould, W.A. et al. recent studies place tropical plant communities within the compositional differences between communities. The Braun-Blanquet framework (Borhidi 1996; Bussmann plot size, while smaller than typically used to character- 2001; Sánchez-Sánchez & Islebe 2002). While many of ize tropical forest alliances, served well to characterize the major forest formations and alliances in Puerto Rico homogeneous stands of these plant communities. Addi- have been described floristically and structurally (Birdsey tionally, we tested whether plot sizes of 10 m × 10 m, 10 & Weaver 1987; Scatena et al. 1993; Chinea 2002; m × 20 m, or 10 m × 30 m added a significant number of Thompson et al. 2002; Lugo 2005), the description and species and found that not to be the case at our sites. The classification of plant community composition within high degree of β-diversity between the eight forest types these alliances has not been well documented. and the a priori site selection based on compositional This study has three objectives: (1) describe the homogeneity within each forest type resulted in the composition of eight tropical forest plant communities selection of repeatable occurrences of these communi- within a hierarchy related to the primary controls on ties as replicates. vegetation, i.e. climate, substrate, topographic position, Environmental and vegetation characteristics were and disturbance; (2) evaluate the conservation status sampled at each plot including location, elevation, slope, species within these communities; and (3) relate the and aspect. Six soil volumes were taken from 0-10 cm variation in diversity, structure, and site environmental depths at each plot and combined to form a composite characteristics of these communities to gradients of cli- sample, which was ground with a Foss Tecator Cyclotec mate and soil characteristics; which are correlated to (model 1093) mill through a 1 mm stainless steel sieve. elevation. Total N, C and S were determined using the dry com- bustion method by means of a LECO CNS-200 analyzer (St. Joseph, MI, USA). Elemental P, Ca, K, Al, Fe, Mn, Methods and Mg were determined from digestions using a Beckman plasma emission spectrometer (Spectra V) Study sites (Luh Huang & Schulte 1985). Mean annual precipita- tion was sampled bimonthly from July 2001 to July This study was developed in Northeastern Puerto 2004 and mean annual air temperature was determined Rico and focused on plant communities within eight from shaded stations placed at 1 m height. All plant older forest types (> 60 years old) that are representative species were identified following Liogier and Martorrell of Lowland subtropical dry, Lowland subtropical moist, (1999). Species cover was estimated using the Braun- Subtropical wet, Subtropical rain, Lower montane wet, Blanquet cover scale (Westhoff & van der Maarel 1978). and Lower montane rain forest life zones in Puerto Rico Percent cover and height estimates were made for each (Ewel & Whitmore 1973). The sites occur along an vegetation layer (canopy trees, understorey trees, shrub elevational gradient from sea level to over 1000 m. Four layer, and herbaceous layer). Percent cover estimates of the forest types are located in the Luquillo Experi- were made for non vegetation layers including bare soil, mental Forest (LEF) and include Elfin woodland, Sierra root, rock, litter, and water. Tree diameters were meas- palm, Palo colorado, and Tabonuco forest alliances ured at breast height (DBH) for all trees over 2 cm. (Wadsworth 1951; Weaver 1994). Four additional for- est types include Lowland moist and dry forests, and Data analyses flooded Pterocarpus and mangrove forests. The latter four are located on federal, state, and private managed All species were classified as either native or non and unmanaged lands. All sites are on non calcareous native (Little & Wadsworth 1964). Native species were material derived from volcanic bedrock (montane and classified as endemic and/or critical elements using the lowland sites) or Quaternary deposits (coastal sites). classification developed by the Puerto Rico Department of Natural and Environmental Resources for conserva- Field sampling tion purposes (Quevedo 1999). Species were identified by growth form as trees, shrubs, lianas, forbs, graminoids, We selected 24 sampling sites throughout northeast- or epiphytes. For trees, we determined the number of ern Puerto Rico, three of each of the eight forest types. stems per species per plot and forest type and the basal Within these sites, 10 m × 10 m plots were sampled for area per species per plot. We calculated biomass per a total of nine replicates per forest type. Sampling
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