Allpahuayo: Floristics, Structure, and Dynamics of a High-Diversity Forest in Amazonian Peru Author(S): Rodolfo Vasquez Martinez and Oliver L
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Allpahuayo: Floristics, Structure, and Dynamics of a High-Diversity Forest in Amazonian Peru Author(s): Rodolfo Vasquez Martinez and Oliver L. Phillips Reviewed work(s): Source: Annals of the Missouri Botanical Garden, Vol. 87, No. 4 (Autumn, 2000), pp. 499-527 Published by: Missouri Botanical Garden Press Stable URL: http://www.jstor.org/stable/2666143 . Accessed: 12/07/2012 11:15 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]. Missouri Botanical Garden Press is collaborating with JSTOR to digitize, preserve and extend access to Annals of the Missouri Botanical Garden. http://www.jstor.org ALLPAHUAYO: FLORISTICS, Rodolfo Vdsquez Martznez2and STRUCTURE, AND Oliver L. Phillips3 DYNAMICS OF A HIGH-DIVERSITY FOREST IN AMAZONIAN PERU' ABSTRACT This paper describes the results of a floristicinventory at the Allpahuayo Reserve, near Iquitos in Amazonian Peru. Two long-termone-hectare plots were established using a pre-determinedsampling grid, with each individual tree and liana over 10 cm diameter collected at least once, except forpalms. The plots were re-censused after5 years to quantify forestdynamics. Floristic analysis shows that the Allpahuayo forestis among the most diverse site yet inventoried,with 281 to 311 species per hectare, and at least 466 species and 61 families in the 1277-stem two-hectare sample, confirmingthat upper Amazonia is a world center of tree biodiversity.The ecologically most dominant and speciose family in the plots is Fabaceae sensu lato, with 231 stems and 89 species; no other family represents more than 7% of the species or 10% of the stems. In contrast to the exceptional floristicdiversity, both the structureand the dynamics of the Allpahuayo forest are similar to those recorded fromother old-growthneotropical forests. Many tree and liana canopy species were previously unknown to both the Iquitos area and to Amazonian Peru, which demonstrates the significance of Amazon ecological studies to systematic botany. RESUMEN Este documento describe el estado actual de los resultados de un inventario floristicoa largo plazo, en la Reserva Biol6gica de Allpahuayo, cerca de Iquitos, en la Amazonfa Peruana. Alli se establecieron dos parcelas de una hect. cada una, usando un muestreo pre-determinado, cada Arbol, palmera o liana, mayores o iguales que 10 cm de diAim. fu6 marcado y colectado, excepto las palmeras que fueron colectadas solo una vez por especie. Las parcelas fueron re-censadas despues de 5 afios para cuantificar el proceso de la dinamica del bosque. El analisis de los resultados nos muestra que el bosque de Allpahuayo, esta entre los bosques mas diversos hasta ahora inventariados, con 281 a 311 especies por hect., y al menos 466 especies y 61 familias en 1277 individuos en dos hect. muestreadas; 6stos datos apoyan la idea de que el oeste de la Amazonia es un centro mundial de diversidad de arboles. En las parcelas la familia Fabaceae s.l., es la mas dominante ecologicamente y en numero de especies, contiene 231 tallos y 89 especies; ninguna otra familia representa mas que el 7 por ciento de las especies o el 10 por ciento de los tallos. En contraste a la excepcional diversidad floristica, tanto la estructura del bosque como la tasa de mortalidad en Allpahuayo, estan dentro los rangos de otros sitios neotropicales. El valor de los estudios ecol6gicos para la botanica sistematica, se demuestra en tanto que, antes de que empezaramos a tomar datos ecol6gicos e instalar parcelas permanentes, muchas especies de arboles y lianas eran desconocidas tanto en el area de Iquitos, como en la Amazonia Peruana. Key words: Amazonia, floristics,inventoiy, Neotropics, species richness, species turnover. ' We gratefullyacknowledge the help of institutionalsupport fromInstituto de Investigaciones de la Amazonia Peruana (IIAP) for permission to work in Allpahuayo Reserve and for logistical support forour work; the John D. and Catherine T. MacArthur Foundation for long-termsupport of the Missouri Botanical Garden's floristicand ecological research at Allpahuayo and elsewhere in Peruvian Amazonia; the National Geographic Society for support for forest dynamics research in Amazonian Peru (grant #5472-95), and the U.K. Natural Environment Research Council for a Research Fellowship (OP). Melchor Aguilar, C6sar Grandez, Rosa Ortiz de Genthy,Nestor Jaramillo, Dennis Milanowski, JohnJ. Pipoly III, Peggy Stern, Henk van der Werff,and Arturo Vasquez Martinez assisted in establishing, inventoiying,and re-censusing the Allpahuayo plots. Herbarium determinationswere provided by numerous systematistsat MO and other herbaria, and we are especially grateful to: C. C. Berg (BG); T. Pennington, Sir G. Prance (K); A. Chanderbali, R. Gereau, R. Liesner, J. J. Pipoly III, G. Schatz, C. Taylor, M. Timana, H. van der Werff,and the late A. H. Gentry(MO); D. Daly, S. Mori, and M. Stern (NY). The maps were drawn by A. Manson at the School of Geography Graphics Unit, University of Leeds. We thank V. Hollowell, Sir G. Prance, and H. van der Werfffor their constructive suggestions to improve our manuscript. 2 Missouri Botanical Garden, Apartado 020, Ja6n, Cajamarca, Peru. 3Centre for Biodiversity and Conservation, School of Geography, Universityof Leeds, Leeds LS2 9JT, U.K. ANN. MISSOURI BOT. GARD. 87: 499-527. 2000. 500 Annals of the MissouriBotanical Garden -00 ~~~~~~~~~~~~~~~73015'W IQUITOS 100% PERU 500K 3045'S 800W 700W A 10Km B1 Figure 1. Map showinglocation of studyregion in relationto Iquitos and Peru. -A. Peru. -B. Iquitos region, showinglocation of Allpahuayo.Modified from Kalliola et al. (1998). Biodiversityis at riskin muchof the tropics. Yet floristicunderstanding by generatinglarge numbers biologicalresearch and conservationinitiatives are ofnew collectionsthat are associatedwith site-spe- hampered by inadequate baseline information. cific ecological information.Botanical institutions Thus, sufficientdata on the numbers,kinds, and play a vital role in initiatingand supportingthis abundanceof most major plant and animaltaxa are work,and contributeessential expertise for ensur- not available. Moreover,knowledge of species as- ing accuratevoucher determination. Equally, eco- sociationsand biogeographicdistributions of taxa logical inventoriesgenerate much biogeographical remainmeager (McNeely et al., 1990; NationalSci- data ofrelevance to monographicand phylogenetic ence Board, 1990; National Research Council, studies. Such eco-floristicresearch has recently 1992; Phillips& Raven, 1996). been initiatedin severaltropical countries, includ- In theNeotropics, the extraordinary species rich- ing Madagascar(Lowry et al., 1997; Rakotomalaza ness, ourlimited taxonomic understanding, and the & Messmer,1999), Colombia (Rudas, 1996), and physicalinaccessibility of mostareas are formida- Peru (Gentry,1988a, b). This paper reportsthe re- ble barriersto phytogeographicinvestigation. Tra- sults of a similarresearch program at the Allpa- ditionalbiological inventory efforts that rely mostly huayoReserve, near Iquitosin northernPeru. on ad hoc collectionefforts and lengthymonograph studies cannot address the knowledgegap alone. STUDY SITE Alternativemethods are criticalto improveour un- derstandingof the factors determining species com- The AllpahuayoReserve (3O57'S, 73026'W) lies positionand theecological dynamics of tropical for- southwestof Iquitos in AmazonianPeru, between est ecosystems.In recent years there has been theblackwater Rio Nanayon thenorthwest and the growingscientific interest in moreecological, plot- Iquitos-Nautaroad to the southeast(Fig. 1). This based workas a means of understandingtropical 2750-ha reserveis administeredby the Peruvian forests(e.g., Gentry,1988a, b; Phillips & Gentry, Institutefor Amazonian Research (IIAP). The cli- 1994; Phillipset al., 1994; Dallmeier& Comiskey, mateis humidand hot (withthe mean annual pre- 1998a, b). This approachcan contributegreatly to cipitationabout 3000 mmand an averagetemper- 0 0 ~~0m500m 25 B20 24...... 24 I 23 ~~~~~~~~23m I5 229....Z 22 2m c 21 ~~~~~~21 213 C 21 21~~~~~~~~~~~~1 20 20~~~~~~~~~~~~~~< ................ .......~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~C .....................~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~- ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~~CC 19 19 ............~~~~~~~~~~~~~~~~~ ........... ...............~ ~ ~ ~ ~ ~ ~ ~ ~ 4-lO~~~~m-~~~ 500x 300m x mm e-1O ......... Fiur 2 Dara sowngte rrngmetofpltswthnerannt507 50 00m amliggrd t lpauao.Th21mplt7(1"an octe t heede ofthgid Adiioalprale(rasets"E.ad.U,.haedinth.fgue).Frcopaisn.e tmpray amle f 00X mwee stblsed10 minfrm heede "".aels.so th rlaiv ostinso adiioal50 2 mpot tatinopoat tasets"E ad"U a ohr its n eu CucoA aznio.ndScuar/AER)bu.nt.tAlpauao 502 Annals of the Missouri Botanical Garden atureof 260C; Marengo,1998). AllpahuayoReserve posited in two Peruvian herbaria(AMAZ, USM) lies between110 m and 180 m above sea level. and at MO, withpartial collections existing at HAP Edaphicconditions are variable,representing a mo- and the UniversidadAgraria La Molina (MOL), as saic of patches rangingin texturefrom clayey to well as distributionsto familyspecialists