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Past Vegetation Changes in Amazon Savannas Determined Using Carbon Isotopes of Soil Organic Matter Author(S): T Past Vegetation Changes in Amazon Savannas Determined Using Carbon Isotopes of Soil Organic Matter Author(s): T. M. Sanaiotti, L. A. Martinelli, R. L. Victoria, S. E. Trumbore, and P. B. Camargo Source: BIOTROPICA, 34(1):2-16. Published By: The Association for Tropical Biology & Conservation DOI: http://dx.doi.org/10.1646/0006-3606(2002)034[0002:PVCIAS]2.0.CO;2 URL: http://www.bioone.org/doi/full/10.1646/0006-3606%282002%29034%5B0002%3APVCIAS %5D2.0.CO%3B2 BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. BIOTROPICA 34(1): 2±16 2002 Past Vegetation Changes in Amazon Savannas Determined Using Carbon Isotopes of Soil Organic Matter1 T. M. Sanaiotti Departamento de Ecologia, Instituto Nacional de Pesquisas da AmazoÃnia, Manaus 69000-000, AM, Brazil L. A. Martinelli 2, R. L. Victoria Centro de Energia Nuclear na Agricultura, Av. CentenaÂrio 303, Piracicaba, 13416-000, SP, Brazil S. E. Trumbore Department of Earth System Science, University of California, Irvine, California 92697-3100, U.S.A. and P. B. Camargo Centro de Energia Nuclear na Agricultura, Av. CentenaÂrio 303, Piracicaba, 13416-000, SP, Brazil ABSTRACT We investigated the variation of stable (d13C) soil carbon isotopes in relation to depth in seven of the most important savanna areas to adjacent contiguous forests in the Amazon region. The d13C of bulk organic matter in all pro®les from forested sites increased with soil depth. In forest pro®les from AmapaÂ, Alter do ChaÄo, and Roraima, the enrich- ment was less than 3.5½ between deeper soil and surface layers, suggesting that C3 plants have remained the dominant vegetation cover. On the other hand, in forest soil pro®les from Humaita and Carolina sites, the d13C enrichment was greater than 3.5½, indicating the in¯uence of past C4 vegetation or a mixture of C3/C4 vegetation (woody savanna). The surface d13C values in the savanna pro®les were 5±13½ greater than the comparable forest pro®les, indicating the in¯uence of C4 vegetation. Two kinds of isotopic distribution were observed in deeper layers. The savanna pro®les at Alter do ChaÄo, Chapada dos Parecis, and RedencËaÄo had relatively constant d13C values throughout the pro®le, suggesting minor past changes in the vegetation composition. In pro®les at AmapaÂ, Roraima, HumaitaÂ, and Carolina, d13C values decreased with depth from the surface and converged with comparable forest values, suggesting more woody savanna in the past than exists currently. RESUMO NoÂs investigamos neste estudo a variacËaÄo em profundidade dos isoÂtopos estaÂveis de carbono (d13C) da mateÂria orgaÃnica do solo (SOM) em sete aÂreas de savanas e ¯orestas da regiaÄo AmazoÃnica. Os valores de d13C da SOM aumentaram com a profundidade do solo. Nos per®s em ¯oresta do AmapaÂ, Alter do ChaÄo e Roraima o enriquecimento isotoÂpico com a profundidade foi menor que 3,5½, sugerindo que plantas do tipo C3 foram sempre o tipo de vegetacËaÄo dominante. Por outro lado, nos per®s em ¯oresta de Humaita e Carolina, o enriquecimento isotoÂpico foi maior que 3,5½, indicando a in¯ueÃncia no passado de uma vegetacËaÄo do tipo C4, ou uma mistura de vegetacËaÄo C3/C4 (savana lenhosa). Os valores de d13C na superfõÂcie do solo em savanas foram cerca de 5 a 13½ maiores que os per®s em ¯oresta, evidenciando a in¯ueÃncia da vegetacËaÄo C4. Dois tipos de distribuicËaÄo isotoÂpica foram observados em camadas mais profundas. Nas savanas de Alter do ChaÄo, Chapada dos Parecis e RedencËaÄo os valores d13C foram constantes ao longo do per®l do solo, sugerindo que naÄo houveram mudancËas signi®cativas na vegetacËaÄo. Nos per®s do AmapaÂ, Roraima, Humaita e Carolina, os valores de d13C diminuiram com a profundidade do solo, aproximando-se aos valores encontrados na ¯oresta, sugerindo a existeÃncia no passado de uma savana mais lenhosa que a actual. Key words: Amazon; Brazil; carbon isotope; radiocarbon; savanna; tropical forest; vegetation change. THE AMAZON REGION IS OFTEN VIEWED AS A CONTIN- savannas are those located at the forest periphery. UOUS STAND of rain forest, but it contains patches Non-isolated savannas include the northern border of savanna vegetation. According to their geograph- of the central Brazilian savannas, the Cerrado, ical location, Amazonian savannas may be classi®ed which is the most extensive example of this vege- as either isolated or non-isolated. Isolated savannas tation type (Sarmiento 1984). Because of its im- are surrounded by rain forest, while non-isolated portance as the second most extensive vegetation type in Amazonia, the origin of savannas has been 1 Received 5 June 2000; revision accepted 17 July 2001. debated for many years (Ducke & Black 1953, 2 Corresponding author; e-mail: [email protected] Egler 1960, Andrade-Lima 1966, Eiten 1972, 2 Past Vegetation Changes in Amazon Savannas 3 Brown & Ab'Saber 1979, Rizzini 1979, Irion 1982, Prance 1982, Kubitzki 1983, Sarmiento 1984, Bigarella and Ferreira 1985, Cole 1986, Sa- naiotti 1996). Several lines of evidence, including paleolim- nological (van der Hammen 1972, Absy & van der Hammen 1976, van der Hammen 1983, Absy et al. 1991), paleofaunal (Rancy 1993), and the car- bon isotopic composition of soil organic matter (Desjardins et al. 1996, Pessenda et al. 1996, Gou- veia et al. 1997), suggest that the dynamics of ex- pansion and contraction of savanna regions result from climatic ¯uctuations during the Quaternary period (Prance 1982, Sarmiento & Monasterio 1975). Several authors have pointed out that there FIGURE 1. Brazilian Amazon basin showing the sat- were drier periods during the Pleistocene and the ellite image location of all study sites. Isolated savannas: (1) AmapaÂ; (2) Alter do ChaÄo; (3) Roraima; and (4) Hu- Holocene periods, when tropical forests were re- maitaÂ. Non-isolated savannas: (5) Chapada dos Parecis; placed by savanna-like vegetation having a predom- (6) RedencËaÄo; and (7) Carolina. inance of grasses (van der Hammen 1974, Absy & van der Hammen 1976, Ab'Saber 1977, Absy 1980, Bigarella & Andrade-Lima 1982, Leyden dominant plants in the savannas) are isotopically 1985, Markgraf 1989, Bush and Colinvaux 1990, distinct, it is possible to detect shifts in tropical Bush et al. 1990, Absy et al. 1991, Markgraf 1991). forest zones to grassland (or vice versa) from the The middle Holocene, from ca 6000 to 4000 years d13C signature of organic matter in soils (Desjar- B.P., was identi®ed as one of such drier periods in dins et al. 1996, Martinelli et al. 1996, Neill et al. several places of South America, including the Am- 1996, Bird & Pousai 1997). azon region (Absy 1980; Markgraf 1989; Servant et al. 1989; Absy et al. 1991; Ledru 1992, 1993; MATERIAL AND METHODS Servant et al. 1993). The presence of charcoal dat- ed from ca 3900 to 1800 years B.P. found in the SOIL SAMPLING. Sampling locations for forest±sa- northern Amazon (Desjardins et al. 1996) suggests vanna comparisons are identi®ed in Figure 1. Four the occurrence of more recent climatic changes in of these sites (AmapaÂ, Alter do ChaÄo, HumaitaÂ, that region as well as in other regions of Amazonia and Roraima) are ``isolated'' savanna pockets that (Servant et al. 1993). are completely surrounded by forest, and three There are seven major savanna areas in the Am- (Chapada dos Parecis, RedencËaÄo, and Carolina) are azon region; in two of them (Roraima and Ron- classi®ed as non-isolated savannas. Soils in the sites doÃnia), the past vegetation dynamics were investi- generally showed a dystrophic character and with gated by using the carbon stable isotopic compo- 1:1 clay type (Table 1). Climatic conditions were sition of soil organic matter (Desjardins et al. 1996, similar at all sites, most of them classi®ed as equa- Gouveia et al. 1997, Pessenda, Gomes et al. 1998, torial hot humid (Nimer 1989). Rainfall varied Pessenda et al. 1998a). In both places, major veg- from 1500 to 1750 mm in Roraima and Carolina etation changes occurred in the past; these studies up to 2750 mm in Amapa and Humaita (Table 1). inferred that savanna areas replaced forested areas The regional vegetation type was very uniform for of the early Holocene period during the middle forest sites, tropical dense forest, following the clas- Holocene. si®cation of RADAMBRASIL (1974), or terra ®r- In this study, we investigated whether or not me forest according to Pires and Prance (1985). similar vegetation changes occurred in other major The regional vegetation types of savanna sites were savanna areas of the Amazon. We compared soil more diverse, varying from savanna park to open carbon isotopic composition and its variation with woody savannas (Table 2). The major difference depth in seven of the most important savanna areas between savanna park and open woody savannas is with nearby contiguous forests in the Amazon re- that in the latter type there is a higher number of gion. Since the main control of the d13C in soil trees (RADAMBRASIL 1974).
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