Long-term change within a Neotropical forest: assessing differential functional and floristic responses to disturbance and drought
BRIAN J. ENQUIST,
CAROLYN A. F. ENQUIST,
BRIAN J. ENQUIST
Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, AZ 85721, USA
The Santa Fe Institute, Santa Fe, NM 87501, USA
1Contributed equally.
Search for more papers by this authorCAROLYN A. F. ENQUIST
School of Natural Resources and Environment, University of Arizona, Tucson, AZ 85721, USA
USA National Phenology Network, Tucson, AZ 85719, USA
The Wildlife Society, Bethesda, MD 20814, USA
1Contributed equally.
Search for more papers by this author
BRIAN J. ENQUIST,
CAROLYN A. F. ENQUIST,
BRIAN J. ENQUIST
Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, AZ 85721, USA
The Santa Fe Institute, Santa Fe, NM 87501, USA
1Contributed equally.
Search for more papers by this authorCAROLYN A. F. ENQUIST
School of Natural Resources and Environment, University of Arizona, Tucson, AZ 85721, USA
USA National Phenology Network, Tucson, AZ 85719, USA
The Wildlife Society, Bethesda, MD 20814, USA
1Contributed equally.
Search for more papers by this author
Brian J. Enquist, Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, AZ 85721, USA, e-mail: [email protected]
Abstract
Disentangling the relative roles of biotic and abiotic forces influencing forest structure, function, and local community composition continues to be an important goal in ecology. Here, utilizing two forest surveys 20-year apart from a Central American dry tropical forest, we assess the relative role of past disturbance and local climatic change in the form of increased drought in driving forest dynamics. We observe: (i) a net decrease in the number of trees; (ii) a decrease in total forest biomass by 7.7 Mg ha−1 but when calculated on subquadrat basis the biomass per unit area did not change indicating scale sensitivity of forest biomass measures; (iii) that the decrease in the number of stems occurred mainly in the smallest sizes, and in more moist and evergreen habitats; (iv) that there has been an increase in the proportion of trees that are deciduous, compound leaved and are canopy species, and a concomitant reduction in trees that are evergreen, simple-leaved, and understory species. These changes are opposite to predictions based on recovery from disturbance, and have resulted in (v) a uniform multivariate shift from a more mesic to a more xeric forest. Together, our results show that over relatively short time scales, community composition and the functional dominance may be more responsive to climate change than recovery to past disturbances. Our findings point to the importance of assessing proportional changes in forest composition and not just changes in absolute numbers. Our findings are also consistent with the hypothesis that tropical tree species exhibit differential sensitivity to changes in precipitation. Predicted future decreases in rainfall may result in quick differential shifts in forest function, physiognomy, and species composition. Quantifying proportional functional composition offers a basis for a predictive framework for how the structure, and diversity of tropical forests will respond to global change.
Supporting Information
Additional Supporting Information may be found in the online version of this article:
Appendix S1. Text (Background, Methods, Discussion).
Appendix S2. Graphs.
Appendix S3. Tables.
Appendix S4. Species list of the San Emilio Forest Dynamics Plot.
Appendix S5. Data Summary. List of species inhabiting the San Emilio tropical dry forest.
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References
- Allen CD, Macalady AK, Chenchouni H et al. (2010) A global overview of drought and heat-induced tree mortality reveals emerging climate change risks for forests. Forest Ecology and Management, 259, 660–684.
- Arroyo-Mora JP, Sánchez-Azofeifa GA, Kalacska MER, Rivard B, Janzen DH (2005) Secondary forest detection in a neotropical dry forest landscape using Landsat 7 ETM+ and IKONOS imagery. Biotropica, 37, 497–507.
- Asner GP, Hughes RF, Varga TA, Knapp DE, Kennedy-Bowdoin T (2009) Environmental and biotic controls over aboveground biomass throughout a tropical rain forest. Ecosystems, 12, 261–278.
- Austin MP (1977) Use of ordination and other multivariate descriptive methods to study succession. Vegetatio, 35, 165–175.
- Bakker JP, Olff H, Willems JH, Zobel M (1996) Why do we need permanent plots in the study of long-term vegetation dynamics? Journal of Vegetation Science, 7, 147–156.
- Bazzaz FA, Wayne PM (1994) Coping with environmental heterogeneity: the physiological ecology of tree seedling regeneration across the gap – understory continuum. In: Physiological Ecology: A Series of Monographs, Texts, and Treatises. Exploitation of Environmental Heterogeneity by Plants. Ecophysiological Processes Above- and Belowground (eds MM Caldwell, RW Pearcy), pp. 349–390. Academic Press, San Diego.
- Bell G (2001) Neutral Macroecology. Science, 293, 2413–2418.
- Borchert R (1994) Soil and stem water storage determine phenology and distribution of tropical dry forest trees. Ecology, 75, 1437–1449.
- Borchert R (1998) Responses of tropical trees to rainfall seasonality and its long-term changes. Climatic Change, 39, 381–393.
- Brando PM, Nepstad DC, Davidson EA, Trumbore SE, Ray D, Camargo P (2008) Drought effects on litterfall, wood production and belowground carbon cycling in an Amazon forest: results of a throughfall reduction experiment. Philosophical Transactions of the Royal Society B-Biological Sciences, 363, 1839–1848.
- Brown S (1997) Estimating Biomass and Biomass Change of Tropical Forests: A Primer. Forest Resources Assessment, Food and Agriculture Organization of the United Nations, Rome.
- Brown S, Lugo AE (1984) Biomass of tropical forests: a new estimate based on forest volumes. Science, 223, 1290–1293.
- Budowski G (1970) The distinction between old secondary and climax species in tropical Central American lowland forests. Tropical Ecology, 11, 44–48.
- Bullock SH (1997) Effects of seasonal rainfall on radial growth in two tropical tree species. International Journal of Biometerology, 41, 13–16.
- Chave J, Condit R, Lao S, Caspersen JP, Foster RB, Hubbell SP (2003) Spatial and temporal variation of biomass in a tropical forest: results from a large census plot in Panama. Journal of Ecology, 91, 240–252.
- Chave J, Condit R, Muller-Landau HC et al. (2008) Assessing evidence for a pervasive alteration in tropical tree communities. Plos Biology, 6, 455–462.
- Chazdon RL, Field CB (1987) Determinants of photosynthetic capacity in six rainforest piper species. Oecologia, 73, 222–230.
- Chesson P (2000) General theory of competitive coexistence in spatially-varying environments. Theoretical Population Biology, 58, 211–237.
- Chidumayo EN (2005) Effects of climate on the growth of exotic and indigenous trees in central Zambia. Journal of Biogeography, 32, 111–120.
- Christensen JH, Hewitson B, Busuioc A et al. (2007) Regional climate projections. In: Climate Change 2007: The Physical Science Basis. Contribution of Working Group I to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change (eds S Solomon, D Qin, M Manning, Z Chen, M Marquis, KB Averyt, M Tignor, HL Miller), pp. 848–940. Cambridge University Press, Cambridge, UK.
- Clark DA, Piper SC, Keeling CD, Clark DB (2003) Tropical rain forest tree growth and atmospheric carbon dynamics linked to interannual temperature variation during 1984–2000. Proceedings of the National Academy of Science of the United States of America, 100, 5852–5857.
- Clark DB, Palmer MW, Clark DA (1999) Edaphic factors and the landscape-scale distributions of tropical rain forest trees. Ecology, 80, 2662–2675.
- Clark JS, Mclachlan JS (2003) Stability of forest biodiversity. Nature, 423, 635–638.
- Condit R (1998) Ecological implications of changes in drought patterns: shifts in forest composition in Panama. Climatic Change, 39, 413–427.
- Condit R, Ashton PS, Manokaran N, Lafrankie JV, Hubbell SP, Foster RB (1999) Dynamics of the forest communities at Pasoh and Barro Colorado: comparing two 50-ha plots. Philosophical Transactions of the Royal Society. London B, 354, 1739–1748.
- Condit R, Hubbell SP, Foster RB (1995) Mortality rates of 205 neotropical tree and shrub species and the impact of a severe drought. Ecological Monographs, 65, 419–439.
- Condit R, Hubbell SP, Foster RB (1996a) Assessing the response of plant functional types to climatic change in tropical forests. Journal of Vegetation Science, 7, 405–416.
- Condit R, Hubbell SP, Foster RB (1996b) Changes in tree species abundance in a neotropical forest: impact of climate-change. Journal of Tropical Ecology, 12, 231–256.
- Condit R, Watts K, Bohlman SA, Perez R, Foster RB, Hubbell SP (2000) Quantifying the deciduousness of tropical forest canopies under varying climates. Journal of Vegetation Science, 11, 649–658.
- Daws MI, Mullins CE, Burslem DFRP, Paton SR, Dalling JW (2002) Topographic position affects the water regime in a semideciduous tropical forest in Panama. Plant and Soil, 238, 79–90.
- Diaz S, Cabido M (2001) Vive la difference: plant functional diversity matters to ecosystem processes. Trends in Ecology & Evolution, 16, 646–655.
- Engelbrecht BMJ, Comita LS, Condit R, Kursar TA, Tyree MT, Turner BL, Hubbell SPN (2007) Drought sensitivity shapes species distribution patterns in tropical forests. Nature, 447, 80–83.
- Engelbrecht BMJ, Kursar TA (2003) Comparative drought-resistance of seedlings of 28 species of co-occurring tropical woody plants. Oecologia, 136, 383–393.
- Enquist BJ, Leffler AJ (2001) Long-term tree ring chronologies from sympatric tropical dry-forest trees: individualistic responses to climatic variation. Journal of Tropical Ecology, 17, 41–60.
- Enquist BJ, Sullivan JJ (2001) Vegetative Key and Descriptions of Tree Species of the Tropical Dry Forests of Upland Sector Santa Rosa. Area de Conservación Guanacaste, Costa Rica, 68pp.
- Enquist BJ, West GB, Brown JH (2009) Extensions and evaluations of a general quantitative theory of forest structure and dynamics. Proceedings of the National Academy of Sciences of the United States of America, 106, 7046–7051.
- Enquist BJ, West GB, Charnov EL, Brown JH (1999) Allometric scaling of production and life-history variation in vascular plants. Nature, 401, 907–911.
- Enquist CAF (2002) Predicted regional impacts of climate change on the geographical distribution of diversity of tropical forests in Costa Rica. Journal of Biogeography, 29, 519–534.
- Fleming TH (1986) Secular changes in Costa Rican rainfall: correlations with elevation. Journal of Tropical Ecology, 2, 87–91.
10.1017/S0266467400000638 Google Scholar
- Frankie GW, Baker HG, Opler PA (1974) Comparative phenological studies of trees in tropical wet and dry forests in the lowlands of Costa Rica. Journal of Ecology, 62, 881–919.
- Franklin J, Drake DR, Bolick LA, Smith DS, Motley TJ (1999) Rain forest composition and patterns of secondary succession in the Vava'u Island Group, Tonga. Journal of Vegetation Science, 10, 51–64.
- Fyllas NM, Patiño S, Baker TR et al. (2009) Basin-wide variations in foliar properties of Amazonian forest: phylogeny, soils and climate. Biogeosciences, 6, 2677–2708.
- Gentry AH, Emmons LH (1987) Geographical variation in fertility, phenology, and composition of the understory of neotropical forests. Biotropica, 19, 216–227.
- Gerhardt K, Hytteborn H (1992) Natural dynamics and regeneration methods in tropical dry forests–and introduction. Journal of Vegetation Science, 3, 361–364.
- Gilbert GS, Harms KE, Hamill DN, Hubbell SP (2001) Effects of seedling size, El Niño drought, seedling density, and distance to nearest conspecific adult on 6-year survival of Ocotea whitei seedlings in Panama. Oecologia, 127, 509–516.
- Gillespie TW, Grijalva A, Farris CN (2000) Diversity, composition, and structure of tropical dry forests in Central America. Plant Ecology, 147, 37–47.
- Girvetz EH, Zganjar C, Raber GT, Maurer EP, Kareiva P, Lawler JJ (2009) Applied climate-change analysis: the climate wizard tool. PLoS One, 4, e8320.
- Givinsh TJ (1978) On the adaptive significance of compound leaves, with particular reference to tropical trees. In: Tropical Trees as Living Systems (eds PB Tomlinson, MH Zimmermann), pp. 3251–3380. Cambridge University Press, Cambridge.
- Gonzalez OJ, Zak DR (1994) Geostatistical analysis of soil properties in a secondary tropical dry forest, S. Lucia, West Indies. Plant and Soil, 163, 45–54.
- Harms KE, Condit R, Hubbell SP, Foster RB (2001) Habitat associations of trees and shrubs in a 50-ha neotropical forest plot. Journal of Ecology, 89, 947–959.
- Hartshorn GS (1992) Possible effects of global warming on the biological diversity in tropical forests. In: Global Warming and Biological Diversity (eds RL Peters, TE Lovejoy), pp. 137–146. Yale University Press, New Haven.
- Heimann M, Reichstein M (2008) Terrestrial ecosystem carbon dynamics and climate feedbacks. Nature, 451, 289–292.
- Holbrook NM, Whitbeck JL, Mooney HA (1995) Drought responses of neotropical dry forest trees. In: Seasonally Dry Tropical Forests (eds SH Bullock, HA Mooney, E Medina), pp. 243–270. Cambridge University Press, New York, NY.
10.1017/CBO9780511753398.010 Google Scholar
- Horn H (1971) The Adaptive Geometry of Trees. Princeton University Press, Princeton.
- Hubbell SP (1997) A unified theory of biogeography and relative species abundance and its application to tropical rain forests and coral reefs. Coral Reefs, 16, S9–S21.
- Hubbell SP (2001) The Unified Neutral Theory of Biodiversity and Biogeography. Princeton University Press, Princeton, NJ.
- Hubbell SP, Foster RB (1986) Biology, chance and history and the structure of tropical rain forest tree communities. In: Community Ecology (eds J Diamond, TJ Case), pp. 314–329. Harper and Row, New York.
- IPCC (2007) Intergovernmental Panel on Climate Change (ed. IPCC). The IPCC Data Distribution Center, Geneva.
- Jackson PC, Meinzer FC, Bustamante M et al. (1999) Partitioning of soil water among tree species in a Brazilian Cerrado ecosystem. Tree Physiology, 19, 717–724.
- Janzen DH (1983) Costa Rican Natural History. University Chicago Press, Chicago.
10.7208/chicago/9780226161204.001.0001 Google Scholar
- Janzen DH (1986) Guanacaste National Park: Tropical Education, and Cultural Restoration. Editorial Univ. Estatal a Distancía, San José, Costa Rica.
- Janzen DH (1988) Management of habitat fragments in a tropical dry forest: growth. annals of the Missouri botanical garden, 75, 105–116.
- Janzen DH, Leisner R (1980) Annotated check-list of plants of Lowand Guanacaste Province, Costa Rica, exclusive of grasses and non-vascular cryptogams. Brenesia, 18, 15–90.
- Kalacska M, Sanchez-Azofeifa GA, Calvo-Alvarado JC, Quesada M, Rivard B, Janzen DH (2004) Species composition, similarity and diversity in three successional stages of a seasonally dry tropical forest. Forest Ecology and Management, 200, 227–247.
- Kraft NJB, Valencia R, Ackerly DD (2008) Functional traits and niche-based tree community assembly in an Amazonian forest. Science, 322, 580–582.
- Lal R, Kimble JM, Stewart BA (2000) Global Climate Change and Tropical Ecosystems. CRC Press, New York.
- Lawrence D (2005) Regional-scale variation in litter production and seasonality in tropical dry forests of southern Mexico. Biotropica, 37, 561–570.
- Leffler AJ, Enquist BJ (2002) Carbon isotope composition of tree leaves from Guanacaste, Costa Rica: comparison across tropical forests and tree life history. Journal of Tropical Ecology, 18, 151–159.
- Leigh EGJ, Windsor DM, Rand AS, Foster RB (1990) The impacts of the El Niño drought of 1982–1983 on a Panamanian semideciduous forest. In: Global Ecological Consequences of the 1982–1983 El Niño-Southern Oscillation (ed. PW Glynn), pp. 473–486. Elsevier Press, Amsterdam.
- Leighton M, Wirawan N (1986) Catastrophic drought and fire in Borneo tropical rainforest associated with the 1982–1983 El Niño Southern Oscillation event. In: Tropical Forests and the World Atmosphere, AAAS Selected Symposium 101 (ed. GT Prance, pp. 75–101. Westview Press Boulder, Colorado.
- Letcher SG, Chazdon RL (2009) Lianas and self-supporting plants during tropical forest succession. Forest Ecology and Management, 257, 2150–2156.
- Malhi Y, Wright J (2004) Spatial patterns and recent trends in the climate of tropical forest regions. Philosophical Transactions of the Royal Society B, 359, 311–329.
- Mantua NJ, Hare SR (2002) The Pacific decadal oscillation. Journal of Oceanography, 58, 35–44.
- McDowell N, Pockman WT, Allen CD et al. (2008) Mechanisms of plant survival and mortality during drought: why do some plants survive while others succumb to drought? Tansley Review. New Phytologist, 178, 719–739.
- Meehl GA, Stocker TF, Collins WD et al. (2007) Global Climate Projections. In: Climate Change 2007: The Physical Science Basis. Contribution of Working Group I to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change (eds S Solomon, D Qin, M Manning, Z Chen, M Marquis, KB Averyt, M Tignor, HL Miller), pp. 749–845. Cambridge University Press, Cambridge, UK.
- Meinzer FC, Andrade JL, Goldstein G, Holbrook NM, Cavelier J, Wright SJ (1999) Partitioning of soil water among canopy trees in a seasonally dry tropical forest. Oecologia, 121, 293–301.
- Neelin JD, Munnich M, Su H, Meyerson JE, Holloway CE (2006) Tropical drying trends in global warming models and observations. Proceedings of the National Academy of Sciences USA, 103, 6110–6115.
- Nepstad DC, Carvalho CRD, Davidson EA et al. (1994) The role of deep roots in the hydrological and carbon cycles of Amazonian forests and pastures. Nature, 372, 666–669.
- Nepstad DC, Tohver IM, Ray D, Moutinho P, Cardinot G (2007) Mortality of large trees and lianas following experimental drought in an amazon forest. Ecology, 88, 2259–2269.
- Nepstad DC, Verssimo A, Alencar A et al. (1999) Large-scale impoverishment of Amazonian forests by logging and fire. Nature, 398, 505–508.
- New M, Todd M, Hulme M, Jones P (2001) Precipitation measurements and trends in the twentieth century. International Journal of Climatology, 21, 1899–1922.
- Norberg J, Swaney DP, Dushoff J, Lin J, Casagrandi R, Levin SA (2001) Phenotypic diversity and ecosystem functioning in changing environments: a theoretical framework. Proceedings of the National Academy of Sciences, 98, 11376–11381.
- Oliveira-Filho AT, Curl N, Vilela EA, Carvalho DA (1998) Effects of canopy gaps, topography, and soils on the distribution of woody species in a central Brazilian deciduous dry forest. Biotropica, 30, 362–375.
- Opler PA (1978) Interaction of plant life history components as related to arboreal herbivory. In: The Ecology of Arboreal Folivores (ed. GG Montgomery), pp. 23–31. Smithsonian Institution Press, Washington, DC.
- Opler PA, Frankie GW, Baker HG (1976) Rainfall as a factor in the release, timing, and synchronization of anthesis by tropical trees and shrubs. Journal of Biogeography, 3, 231–236.
10.2307/3038013 Google Scholar
- Overpeck JT, Rind D, Goldberg R (1990) Climate-induced changes in forest disturbance and vegetation. Nature, 343, 51–53.
- Phillips OL (1996) Long-term environmental change in tropical forests: increasing tree turnover. Environmental Conservation, 23, 235–248.
- Phillips OL, Aragao LEOC, Lewis SL et al. (2009) Drought sensitivity of the Amazon rainforest. Science, 323, 1344–1347.
- Phillips OL, Baker TR, Arroyo L et al. (2004) Pattern and process in Amazon tree turnover, 1976–2001. Philosophical Transactions of the Royal Society of London Series B-Biological Sciences, 359, 381–407.
- Phillips OL, Gentry AH (1994) Increasing turnover through time in tropical forests. Science, 263, 954–958.
- Phillips OL, Malhi Y, Higuchi N et al. (1998) Changes in the carbon balance of tropical forests: evidence from long-term plots. Science, 282, 438–442.
- Poorter L (2005) Resource capture and use by tropical forest tree seedlings and their consequences for competition. In: Biotic Interactions in the Topics: Their Role in the Maintenance of Species Diversity (eds D Burslem, M Pinard, S Harley), pp. 35–64. Cambridge University Press, Cambridge, UK.
10.1017/CBO9780511541971.003 Google Scholar
- Pounds AJ, Fogden MPL, Campbell JH (1999) Biological response to climate change on a tropical mountain. Nature, 398, 611–615.
- R Development Core Team (2009) R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna, Austria.
- Reich PB, Borchert R (1984) Water stress and tree phenology in a Tropical Dry forest in the lowlands of Costa Rica. Journal of Ecology, 72, 61–74.
- Saleska SR, Didan K, Huete AR, Da Rocha HR (2007) Amazon forests green-up during 2005 drought. Science, 318, 612.
- Schnitzer SA (2005) A mechanistic explanation for global patterns of liana abundance and distribution. American Naturalist, 166, 262–276.
- Smith TM, Shuggart HH, Woodward FI (1997) Plant Functional Types: Their Relevance to Ecosystem Properties and Global Change. Cambridge University Press, Cambridge, UK.
- Sobrado MA (1986) Aspects of tissue water relations of evergreen and seasonal changes in leaf water potential components of evergreen and deciduous species coexisting in tropical forests. Oecologia, 68, 413–416.
- Sobrado MA, Cuenca G (1979) Aspectos del uso de agua de especies deciduas y siempreverdes en un bosque seco tropical de Venezuela. Acta Cientifica Venezolana, 30, 302–308.
- Sorensen TC (1998) Tropical dry forest regeneration and its influence on three species of Costa Rican monkeys. University of Calgary, Calgary, MA.
- Stephenson NL, Van Mantgem PJ (2005) Forest turnover rates follow global and regional patterns of productivity. Ecology Letters, 8, 524–531.
- Stowe LG, Brown JL (1981) A geographic perspective on the ecology of compound leaves. Evolution, 35, 818–821.
- Swaine MD (1996) Rainfall and soil fertility as factors limiting forest species distributions in Ghana. Journal of Ecology, 84, 419–428.
- Swaine MD, Grace J (2007) Lianas may be favoured by low rainfall: evidence from Ghana. Plant Ecology, 192, 271–276.
- Swenson NG, Enquist BJ (2007) Ecological and evolutionary determinants of a key plant functional trait: wood density and its community wide variation across latitude and elevation. American Journal of Botany, 94, 451–459.
- Swenson NG, Enquist BJ (2009) Opposing assembly mechanisms in a Neotropical dry forest: implications for phylogenetic and functional community ecology. Ecology, 90, 2161–2170.
- Swenson NG, Enquist BJ, Thompson J, Zimmerman JK (2007) The influence of spatial and size scale on phylogenetic relatedness in tropical forest commuities. Ecology, 88, 1770–1780.
- Ter Braak CJF, Similauer P (1998) CANOCO 4 Reference Manual and User's Guide to Canoco for Windows. Centre for Biometry, Wageningen.
- Thompson J, Brokaw N, Zimmerman JK et al. (2002) Land use history, environment, and tree composition in a tropical forest. Ecological Applications, 12, 1344–1363.
- Thompson JD, Weiblen G, Thomson BA, Alfaro S, Legendre P (1996) Untangling multiple factors in spatial distributions: Lilies, gophers, and rocks. Ecology, 77, 1698–1715.
- Townsend AR, Asner GP, Cleveland CC (2008) The biogeochemical heterogeneity of tropical forests. Trends in Ecology & Evolution, 23, 424–431.
- Valencia R, Foster RB, Villa G et al. (2004) Tree species distributions and local habitat variation in the Amazon: large forest plot in eastern Ecuador. Journal of Ecology, 92, 214–229.
- Vargas AB, Trejos VFS (1994) Changes in the general circulation and its influence on precipitation trends in Central America: Costa Rica. AMBIO, 23, 87–90.
- Webb CO, Peart DR (2000) Habitat associations of trees and seedlings in a Bornean rain forest. Journal of Ecology, 88, 464–478.
- West GB, Enquist BJ, Brown JH (2009) A general quantitative theory of forest structure and dynamics. Proceedings of the National Academy of Sciences of the United States of America, 106, 7040–7045.
- Windsor DM, Rand AS (1985) Cambios Climáticos en los registros de lluvia en Panama y Costa Rica. Impretex, Panama, SA.
- Woods P (1989) Effects of logging, drought, and fire on structure and composition of tropical forests in Sabah, Malaysia. Biotropica, 21, 290–298.
- Yáñez-Espinosa L, Terrazas T, López-Mata L (2006) Integrated analysis of tropical trees growth: a multivariate approach. Annals of Botany, 98, 637–645.
- Yavitt JB, Wright SJ (2008) Seedling growth responses to water and nutrient augmentation in the understorey of a lowland moist forest, Panama. Journal of Tropical Ecology, 24, 19–26.
- Zimmerman JK, Everham EMI, Waide RB, Lodge DJ, Talyor CM, Brokaw NVL (1994) Responses of tree species to hurricane winds in subtropical wet forest in Puerto Rico: implications for tropical tree life histories. Journal of Ecology, 82, 911–922.
