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Quantifying the Deciduousness of Tropical Forest Canopies Under Varying Climates Author(S): Richard Condit, Kristina Watts, Stephanie A

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Quantifying the Deciduousness of Tropical Forest Canopies Under Varying Climates Author(S): Richard Condit, Kristina Watts, Stephanie A International Association for Vegetation Science Quantifying the Deciduousness of Tropical Forest Canopies under Varying Climates Author(s): Richard Condit, Kristina Watts, Stephanie A. Bohlman, Rolando Pérez, Robin B. Foster and Stephen P. Hubbell Reviewed work(s): Source: Journal of Vegetation Science, Vol. 11, No. 5 (Oct., 2000), pp. 649-658 Published by: Wiley Stable URL: http://www.jstor.org/stable/3236572 . Accessed: 10/12/2012 14:51 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Wiley and International Association for Vegetation Science are collaborating with JSTOR to digitize, preserve and extend access to Journal of Vegetation Science. http://www.jstor.org This content downloaded by the authorized user from 192.168.52.67 on Mon, 10 Dec 2012 14:51:47 PM All use subject to JSTOR Terms and Conditions Journal of VegetationScience 11: 649-658, 2000 ? IAVS;Opulus Press Uppsala. Printed in Sweden 649 Quantifying the deciduousness of tropical forest canopies under varying climates Condit, Richardl*; Watts, Kristina2; Bohlman, Stephanie A.3; Perez, Rolando1; Foster, Robin B.4 & Hubbell, Stephen P.5 1CenterforTropical Forest Science, SmithsonianTropical Research Institute, Unit 0948, APO AA 34002-0948, USA; 2SouthamptonCollege (currentaddress: Schoolfor Public and EnvironmentalAffairs, Indiana University,Bloomington IN 47401, USA); 3College of Forest Resources, Universityof Washington,Seattle WA98192, USA;4Botany Depart- ment,The Field Museum,Roosevelt Rd. at LakeShore Dr., Chicago,IL 60605, USA;5Department of Botany, University of Georgia, Athens, GA 30602, USA; *Correspondingauthor; Fax +5072325978;E-mail [email protected] Abstract. Deciduousness is an importantfunctional attribute Introduction of tropical trees, reflecting climatic conditions. Precisely and deciduousness in forests quantifying mapping tropical Deciduousness is a key functional trait both for indi- will be for remote necessary calibrating sensing images vidual trees and for forest communities. Partial or full which attempt to assess canopy properties such as carbon leaf loss during part of the year is an eco- cycling, productivity,or chlorophyll content. We thus set out important since it indicates that forest to assess the degree of canopy deciduousness in three moist, system trait, productivity semi-deciduous tropical forests in central Panama.One site is seasonally reduced, and deciduousness may correlate is a 6-ha research plot near the Atlantic coast of Panama, with important ecosystem-level characteristics, such as where rainfall is 2830 mm/yr. The second site is a 50-ha plot tree height, biomass, and diversity. Since deciduousness on Barro Colorado Island, near the center of the isthmus of changes canopy reflectance, it can be detected in satel- where rainfall is 2570 and the final Panama, mm/yr, site is a lite images (Reed et al. 1994; Bohlman et al. 1998), and 4-ha near the Pacific coast of Panama,where rainfall is plot thus could potentially be mapped over wide areas. Re- 2060 mm/yr. At each site, a randomsample of trees from all flecting its importance in tropical ecosystems, deciduous- canopy species (those with individuals > 30 cm DBH) were ness often provides a main axis for vegetation classifica- visited and scored for deciduousness three times during the tion Hall & Swaine Box 1997 dry season. The estimated peak fraction of deciduous (Beard 1944; 1981; 1996): individuals in the canopy at the wetter site was 4.8%, at the tropical dry forests are deciduous, wet forests are ever- intermediate site, 6.3%, and at the drier site, 24.3%. The green, and forests in between are called semi-deciduous. estimated fraction of crown area deciduous peaked at 3.6%, Deciduousness is thus an important and easy-to-meas- 9.7%, and 19.1% at the wet, medium, and dry sites respec- ure indicator, and models of how tropical forests re- The of tively. percentage canopy species that was deciduous spond to future climatic changes (Steffen et al. 1996) - 14%,28%, and 41 %- was much thanthe higher percentage might be calibrated by assessing how well they predict of deciduous because not all individuals of de- individuals, deciduous behavior under different climates. ciduous species were deciduous. During the 1999 dry sea- This requires calibration sites, where deciduousness son, every individual of all the deciduous species was visited and climate are at the two driersites, and the total numberof deciduous trees carefully measured, and despite the observed closely matched the estimated numbers based on importance of this single character in forest classifica- the smaller 1997 samples. tion and ecosystem function, there are very few quanti- tative estimates of deciduousness in tropical forests. The only indication of the degree of deciduousness typically Keywords: Climate gradient; Deciduousness; Phenology; reported is the fraction of species (e.g. Frankie et al. 1974; Tropicalforest. Reich 1995). Quantitative studies on the fraction of canopy cover lost during seasons of leaf fall are rare. We thus embarked on a straightforward effort to map and Abbreviation: BCI = BarroColorado Island. quantify deciduousness in three forests in central Panama. We sampled every canopy species for deciduous behavior, and then made a complete census of all deciduous species. The three sites, though close together, have different precipitation patterns, so we were able to quan- tify the relationship between climate and deciduousness. This content downloaded by the authorized user from 192.168.52.67 on Mon, 10 Dec 2012 14:51:47 PM All use subject to JSTOR Terms and Conditions 650 Condit,R. et al. Study sites on the Smithsonian Tropical Research Institute's re- search reserve on Barro Colorado Island was set up in The study was carriedout in lowland forests (< 200 1981 (Hubbell& Foster 1983, 1986, 1992; Conditet al. m elevation) adjacent to the Panama Canal. Annual 1995, 1996a, c). Here we use datafrom the 1995 census rainfall even in the driest sites in central Panama ex- of the plot. A 6-ha plot near the Atlantic coast at Fort ceeds potential evapotranspirationand is ample to sus- Shermanwas establishedin 1996, and a 4-ha plot near tain tall, moist, high-biomassforest. Thereis a long dry the Pacific, near the Cocoli River, was established in season, though,running from mid-Decemberuntil mid- 1994. The latter two sites are on former US military April, during which many trees lose all their leaves installations. (Leigh et al. 1982), and there is considerablevariation across the isthmus in the length of the period when potentialevapotranspiration exceeds precipitation(Fig. Methods 1). At the Pacific side of the Canal, near PanamaCity, total annualrainfall is 2060 mm and the dry season lasts All stems > 1 cm diameter-at-breast-height(DBH) 129 days; at BarroColorado in the centerof the isthmus, were identified to species, measured at breast height, rainfallis 2570 mm and the dry season 118 days; while and mapped,throughout all threeplots. Condit (1998b) nearthe Atlantic,rainfall is 2830 mm andthe dry season describes in detail the methods used. We are primarily is 106 days (Fig. 1). Condit (1998a) reportedslightly interestedin deciduousness of trees forming the upper differentestimates for dry season duration,but based on canopy, since this is where leaf-loss would have the an approximationfor potentialevapotranspiration (100 greatestimpact on ecosystem function, and where it is mm/month);here we use directmeasurements of poten- visible remotely. In addition, understory species are tial evapotranspiration(Fig. 1). All three sites have seldom deciduous(Frankie et al. 1974; Wright 1991): to typical lowland-tropical temperatureregimes, with a our knowledge, just one understorytree at BarroColo- mean close to 27 ?C throughoutthe year. rado - Erythrina costaricana - is (Condit et al. 1996b). We have establishedthree forest census plots across Thus, we only surveyed species which reached a DBH the isthmus,spanning this rainfallgradient. A 50-ha plot of 30 cm; we refer to these as canopy species. At Barro 500 - --PET - Sherman BCI Cocoli 400 300 -C E 200 100 0 0 5 10 15 20 25 30 35 40 45 50 week of year Fig. 1. Precipitation and potential evapotranspiration (PET) at the three plot sites. Both were calculated starting with weekly means over 12 - 25 yr, taking a 5-week running mean at each point, then adjusting to monthly by multiplying the weekly mean by 4.35 (the number of weeks in a month). One station is 6 km from the Sherman plot, the second is 1 km from the BCI plot, and the last is 4 km from the Cocoli plot. PET data are from Barro Colorado Island only. Dry season length was estimated as the interval each precipitation curve spent below the PET curve; for example, at Cocoli, precipitation rises above PET at week 15.8 and falls below at week 49.5, which works out to 129 days below PET. The difference between dry seasons is probably even more extreme than this indicates, because there is undoubtedly greater solar radiation and thus higher PET at the drier sites. (BCI data courtesy of the Smithsonian Environmental Sciences Program; Sherman and Cocoli data courtesy of the Panama Canal Commission.) This content downloaded by the authorized user from 192.168.52.67 on Mon, 10 Dec 2012 14:51:47 PM All use subject to JSTOR Terms and Conditions - Quantifyingthe deciduousnessof tropicalforest canopies undervarying climates - 651 Colorado, 139 of the 144 canopy species in the 50-ha (nearlyevery individualwas scored as 3 or 4). plot were censused; at Sherman, 64 of the 66 canopy We tallied the fraction of trees in each category, 0 species were censused; and at Cocoli, 34 of 39 species through4, but we simplify most of our presentationby were censused.
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