Flora and dynamics of an upland and a floodplain forest in Peña Roja, Colombian Amazonia

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1 Flora and dynamics of an upland and a floodplain forest in Peña Roja, Colombian Amazonia Flora y dinámica de bosques de tierra firme y de várzea en Peña Roja, Amazonia colombiana

2 Londoño, AC 2011 Flora and dynamics of an upland and a floodplain forest in Peña Roja, Colombian Amazonia. PhD dissertation, Universiteit van Amsterdam. The work presented in this Thesis was mainly funded by Tropenbos - Colombia, the Netherlands Organisation for International Cooperation in Higher Education (NUFFIC) and COLCIENCIAS. Research was conducted at the Institute for Biodiversity and Ecosystem Dynamics (IBED), Universiteit van Amsterdam. ISBN: Lay-out: Ana Catalina Londoño Vega. Cover: Canopy of the upland plot (left); young leaves of Pseudomonotes tropenbosii (centre); canopy of the floodplain plot (right). No part of this dissertation may be reproduced in any format, by print, photocopy, microfilm or by any other means without the written permission from the author.

3 Flora and dynamics of an upland and a floodplain forest in Peña Roja, Colombian Amazonia Flora y dinámica de bosques de tierra firme y de várzea en Peña Roja, Amazonia colombiana ACADEMISCH PROEFSCHRIFT ter verkrijging van de graad van doctor aan de Universiteit van Amsterdam op gezag van de Rector Magnificus prof. dr. D.C. van den Boom ten overstaan van een door het college voor promoties ingestelde commissie, in het openbaar te verdedigen in de Agnietenkapel op dinsdag 21 juni 2011, te uur door Ana Catalina Londoño Vega geboren te Medellín, Colombia

4 Promotiecommissie Promotores: Co-promotor: Prof. dr. A.M. Cleef Prof. dr. H. Hooghiemstra Dr. J.F. Duivenvoorden Overige leden: Prof. dr. P. Baas Prof. dr. P.J.M. Maas Prof. dr. R.G.A. Boot Prof. dr. J.H.D. Wolf Dr. ir. H.F.M. Vester Faculteit der Natuurwetenschappen, Wiskunde en Informatica

5 A mi Padre A mi Esposo Sólo lo que cambia, permanece Heráclito

6

7 Contents 1. Introduction 9 2. Site description of the upland and floodplain plots A new genus and species of Dipterocarpaceae from the Neotropics. I. Introduction, taxonomy, ecology, and distribution Arquitectura de Iryanthera tricornis, Osteophloeum platyspermum y Virola pavonis (Myristicaceae) Composición florística de dos bosques (tierra firme y várzea) en la región de Araracuara, Amazonia colombiana Forest dynamics of an upland and a floodplain plot near Peña Roja, Colombian Amazonia Synthesis 141 Literature 149 Summary, Resumen, Samenvatting 179 Appendices 191 Agradecimientos 235 Curriculum vitae 239

8

9 1 Introduction Ana Catalina Londoño Vega

10 1.1 The importance of Amazonian forests The Amazon Basin, which encompasses the catchments of the Amazon River and the rivers of the Guiana Shield that drain into the Atlantic Ocean, and a substantial part of the basin of Orinoco River, is covered by the largest continuous area of tropical forests in the world. Amazonian forests represent the habitat for about one-tenth of all species of the world. As such, the Amazon Basin has a fundamental role in the origin and conservation of genetic resources worldwide. However, Amazonian forests, just as most of the tropical forests in Africa and Asia, are severely threatened by mankind. Deforestation rates of humid tropical forests vary around 142,000 km 2 per year (FAO 2001; Achard et al. 2002; Fearnside and Laurance 2003). In the past two decades increasing attention has been paid to the important role of the Amazon Basin in Global Change, especially regarding climate change, the cycling of water and carbon, and the emission of greenhouse gasses (Junk and Furch 1985; Phillips et al. 1998; Canadell et al. 2000; Watson et al. 2000; Holmgren et al. 2001; Houghton et al. 2001; Clark et al. 2003; Malhi and Phillips 2004). Changes in atmospheric CO 2 concentrations probably influence the composition, diversity and dynamics of the forests (Clark et al. 2003; Laurance et al. 2004). It is in this context that studies about the biomass and carbon fixation of tropical forests are being intensified in recent years (Brown 1997; Silver et al. 2000; Brown 2002). Most of these studies depend on measurements of forest productivity in permanent plots, which is often estimated on the basis of yearly tree growth by diameter increments and tree mortality. 1.2 Permanent plots to study long-term forest dynamics The adequate conservation of tropical forests and wise use of their natural resources depend to a large extent on our knowledge of the variation in these forests, both in space and in time. Long-term studies in permanent forest plots consisting of repeated surveys in well-demarcated and georeferenced plots, allow the quantification of the temporal variation of forest resources (Campbell et al. 2002). Non-permanent plots to describe Amazonian forests along environmental gradients have been used successfully in exploratory surveys (Gentry 1988b; Duivenvoorden and 10

11 Lips 1993; ter Steege et al. 2003; Duque 2004; Benavides 2010). Yet, such plots have the disadvantage that they, similar to a photograph, only provide a limited view at a certain time. In contrast, permanent plots not only admit a detailed description of the habitat at a particular site, but also allow detecting temporal changes in the forest. Such information is indispensable to predict future changes in the diversity and distribution of species (Losos and Leigh 2004). Beyond the initial inventory of the vegetation in permanent plots it is necessary to measure forest changes in the long run, through continuous monitoring of the floristic composition, structure, growth, mortality and survival of species, preferably at more than one site (Comiskey et al. 1999). Permanent plots have been used extensively to study long-term changes of vegetation and the natural processes that allow the coexistence of species in relation to the environment (Bakker et al. 1996; Herben 1996; Rees et al. 2001; Hubbell 2004). Early studies conducted in tropical forests primarily aimed at measuring the diameter growth for silvicultural management and timber harvesting (Bell 1971). Recent studies include other objectives such as quantifying carbon stocks and their relation to global flows (Dallmeier 1992; Phillips et al. 1998; Orrego et al. 2003). The oldest plots in tropical forests stem from the first part of the twentieth century, and were established in Philippine dipterocarp forests (Richards 1952), on the islands of Trinidad and Tobago (Bell 1971), in Peninsular Malaysia (Manokaran and Swaine 1994) and in Uganda, Africa (Sheil et al. 2000). The largest plots (25 to 50 ha) are now part of the CTFS network (Center for Tropical Forest Science) of the Smithsonian Tropical Research Institute (STRI) (Losos and Leigh 2004), which manages, for example, the well-known plot on Barro Colorado Island (Panama). Another network is that by the Amazon Forest Inventory Network (RAINFOR) (Malhi et al. 2002; Malhi and Phillips 2004), which mostly supports 1-ha plots. In Colombia, permanent forest plots were initially installed to monitor growth of timber species like Cupressus lusitanica (Tschinkel 1972; del Valle 1979), Cordia alliodora, Tectona grandis, Eucalyptus saligna, and Cariniana pyriformis. In the 1980's permanent forest plots were started in the Chocó region (western Colombia) (González et al. 1991; González 1995; del Valle 1996a, 1998ab, 1999) and Amazonia (Londoño 1993, this dissertation). In 2000 about 70 permanent plots were being maintained 11

12 (Vallejo et al. 2005; Álvarez et al. 2008). By mid 2006, these plots covered at least 100 ha. Overall, the two plots described in this dissertation have particular significance as they are among the oldest permanent forest plots in Colombian Amazonia, and in Colombia as a whole. 1.3 Floristic and ecological research in the permanent forest plots at Peña Roja Between 1986 and 2000 the Tropenbos-Colombia Programme supported forest ecological research in the mid catchment of the Caquetá River, with its epicenter in Araracuara (Fig. 1-1). The programme started with a land unit survey of a large area (10,000 km 2 ), east of Araracuara. Based on the ecological maps (Duivenvoorden and Lips 1993, 1995) in late 1988 several sites were selected as representative for the principal land units, in order to develop long-term monitoring studies of forest ecosystem functioning. The following selection criteria were applied: a) Representation: the land unit should have a wide cover in the Middle Caquetá area and in the Colombian Amazon as a whole, b) Accessibility: the site should be well accessible by river from Araracuara to allow efficient transport of personnel and equipment, c) Control, surveillance and security: sites should offer some protection for permanently installed research equipment (Duivenvoorden and Lips 1990). The sites selected were located in the area of Nonuya community of Peña Roja. One site belonged to the land unit classified as Tertiary Sedimentary Plain and the other site was located on a rarely inundated flood plain of the Caquetá River (Duivenvoorden and Lips 1990). With all parties involved it was agreed to develop a long-term research on the flora, structure and dynamics of the forest vegetation at these two monitoring sites. At each site a permanent plot of 1.8 ha was established for the long-term study of the forest. Uniform criteria were used for the establishment, i.e. the sites were chosen in such a way that very large (multiple tree gaps by wind throw) canopy gaps were avoided. Also, the physiography, forest (Duivenvoorden et al. 1988) and soils were as homogeneous as possible. From that moment on, in and outside these plots, a series of ecological studies have been successfully carried out by students from Colombia and other nationalities, for example those regarding plant-animal relationships 12

13 (van Dulmen 2001; Parrado 2005), hydrology (Téllez 2003) and nutrient cycling (Tobón 1999; Tobón et al. 2004ab). Fig Location of Araracuara in Colombian Amazonia. The shaded area around Araracuara and Peña Roja is given in more detail in Fig There is no doubt that the activities of Tropenbos-Colombia triggered the intensification of the botanical explorations in Colombian Amazonia. In spite of considerable progress in the past years (Sánchez 1997; Rudas and Prieto 2005) it is still common that about one fifth of the species encountered in Amazonian forest surveys remains unidentified. Also, most identifications rely on sterile specimens. Therefore, the contribution of taxonomy to describe and categorize the species diversity of Amazonian forests is of utmost importance. Obtaining fertile botanical specimens of tree species is a challenge in tropical forests, because many species are sterile during a large part of the year. More importantly, many tree species are hard to encounter during botanical surveys because of their scattered occurrence in small populations. In this regard, the discovery of a new species of the family of Dipterocarpaceae, in one of the permanent plots established in the Peña Roja community was an extraordinary event. Before this discovery, the family of Dipterocarpaceae, which belongs to 13

14 one of the most dominant, species rich, and economically important tree families in tropical forests of Asia (Whitmore 1975; Ashton 1982), was only represented by one single species in the Neotropics (Pakaraimaea dipterocarpacea Maguire & Ashton recorded in Guyana). The record of Pseudomonotes tropenbosii Londoño, Álvarez and Forero in the upland plot near Peña Roja along the Caquetá river emphasizes the phytogeographical links of Colombian Amazonia with the Guiana Shield region (Duivenvoorden and Lips 1995) and even with Africa and Madagascar (via the subfamily Monotoideae; Morton 1995), and underscores the unique position and value of this permanent plot. The architecture of a plant is the morphological expression of its genome in a given time, and a result of the balance between endogenous growth processes and exogenous constraints exerted by the environment (Barthélémy et al. 1991; Hallé 1995). The objective of architectural analysis is to identify the endogenous processes that control growth and shape of the whole plant, by means of observation. Three key concepts have been developed: the architectural model (Hallé and Oldeman 1970), the architectural unit (Édelin 1977) and the reiteration (Oldeman 1974). The concepts of model and reiteration unit have provided a valuable tool for studying the structure and form of plants. Plant architecture further determines how resources are allocated in plants, for example, regarding the capture of light, water transport, mechanical stability, and resistance to wind (Vester 1997; Poorter and Werger 1999). Vester (1997) analyzed successive architectural development of tree taxa in successional forests near Araracuara. He showed that on the basis of detailed observations, architectural analysis yields understanding of the mechanisms of succession in secondary forest or in old-growth forests. Architectural analysis thus has the potential to supply significant information regarding forest dynamics in permanent plots, to complement the demographic information of recruitment, growth and mortality. Arguably, two principal results emerged from the ecological surveys of the Middle Caquetá area (Duivenvoorden and Lips 1993, 1995; Urrego 1997; Quiñones 2002; Duque 2004; Sánchez 2005; Benavides 2010). The first was the clear recognition of the high tree species diversity per unit of area (Valencia et al. 1994; Duivenvoorden 1996), which ranks among the highest level for tropical forests worldwide. Several explanations for the 14

15 high upper Amazonian tree species diversity have been put forward (Duivenvoorden 1995; ter Steege et al. 2006). In most of these, the unique properties of the geological setting (speciation triggered by the unique configuration and the highly dynamic environment in the Tertiary; Hoorn and Wesselingh 2010) and the climatic history of northwestern Amazonia (continuous high humidity throughout the Pleistocene favouring species survival; Lips and Duivenvoorden 1994; Mayle et al. 2004) play a key role. The second result was the evidence and documentation that forest composition and forest diversity systematically changed across land units, which differed regarding flooding, drainage and soil nutrient concentrations. High levels of forest diversity were consistently recorded in the well-drained uplands (Duivenvoorden 1996; Duivenvoorden and Duque 2010). On the other hand, poor drainage, seasonal flooding and extremely low levels of soil nutrient availability were always associated with low levels of tree species diversity (Urrego 1997; Duivenvoorden and Duque 2010). Furthermore, litterfall measurements in combination with studies of the standing stock of litter on the forest floor indicated that the aboveground decomposition of organic matter (a proxy of net primary above-ground forest productivity; Vogt et al. 1998) was substantially higher in floodplains of the Caquetá than in well-drained uplands (Lips and Duivenvoorden 1996). Summarizing, in view of the physiographic setting of the two permanent forest plots at Peña Roja, a general picture emerged from the ecological survey that soil fertility, forest disturbances, and above-ground forest productivity in floodplains of the Caquetá River were higher than in the well-drained uplands. However, because these survey results were not based on permanent plots, information about forest dynamics could not be incorporated. In more recent studies in Amazonia at continental scales, forest dynamics have been correlated with forest diversity. Wide-scale correlations between tree mortality (Phillips et al. 2004), wood specific gravity (Baker et al. 2004), above-ground biomass (Baker et al. 2004; Mahli et al. 2006), above-ground wood productivity (Malhi et al. 2006) and tree species diversity (ter Steege et al. 2006) have been reported along geological gradients from eastern Amazonia to upper Amazonian watersheds. In short, low diversity forests were associated to low levels of forest dynamics (low mortality, low productivity) and low levels of soil fertility, whereas high diversity forests showed a relatively 15

16 high mortality (and high wood productivity) and were found on relatively rich soils. Is this general scheme, obtained from Amazonia as a whole, useful to predict forest dynamics and forest diversity in a spatially more limited area like that of Colombian Amazonia? 1.4 Aims and outline of this dissertation The principal aim of this dissertation is to provide basic knowledge about the structure, species composition, and forest dynamics of two permanent forest plots established in contrasting land units (upland and floodplain), in the central part of Colombian Amazonia. In Chapter 2 a description of the two plot sites is given, including details of the plot design and set-up. This information serves as background to the next chapters, but is also essential to warrant a sound continuation of the monitoring activities in the coming decades. Chapter 3 reports on the discovery and description of the new genus and species of the family Dipterocarpaceae that appeared as one of the more dominant species in the upland plot. Chapter 4 gives a treatment of the architecture of three subcanopy species of the nutmeg family (Myristicaceae), a pantropical family of mostly tree species. It describes the development of these species from seedlings in the undergrowth to senile trees. The main purpose of this chapter was to evaluate the extra value of architectural analysis for studies regarding forest dynamics in permanent plots. The following two chapters give accounts on the species composition (Chapter 5) and forest dynamics (Chapter 6) in the two permanent plots. Which species, genera and families characterize these plots? How is the distribution of taxa among families and genera? What is the species composition relative to the habits or growth forms? How are the modes of death? Do mortality and recruitment differ between the plots? Do the patterns in tree species composition and forest dynamics in the plots correspond to predictions based on the general model of Amazonian forest diversity as function of geology? Finally, in Chapter 7 a synthesis is given, followed by the summaries in English, Spanish and Dutch and the appendices. 16

17 2 Site description of the upland and floodplain plots Ana Catalina Londoño Vega

18 2.1 Location and accessibility The permanent plots are located about 30 km east (and 70 km downstreams) of Araracuara (Fig. 2-1), in the basin of the Caquetá River, Colombian Amazonia. This area is part of the community of Peña Roja (Nonuya ethnicity), which is under the jurisdiction of the so-called Corregimiento de Puerto Santander, Departamento del Amazonas. The access to Araracuara from Bogotá is by air. Reaching the upland plot requires about 2 km walking from the maloca in Peña Roja. The floodplain plot can be reached by river, navigating upstream along the Caquetá river from the maloca in Peña Roja for about 2.5 km and then walking ca. 0.5 km. 2.2 Climate The information about the current climate in the study area is based on two meteorological stations: one in Araracuara and one in Peña Roja. At Araracuara daily records were available from (station code located at 160 m altitude above sea level; Duivenvoorden and Lips 1995). Climate data from Peña Roja were automatically recorded every 20 minutes between November 1992 and August 1997 (Tobón 1999) at a station located at 0 39' S and 72 5' W at 150 m above sea level, at 2.5 km distance from the upland and floodplain plots. Disregarding slight differences between the records from both climate stations, which can be attributed to the different length of the registration period, the climate of this part of the Middle Caquetá Basin is characterized by an annual rainfall of ca mm, an average yearly temperature of 25 C and a relative humidity of 87% (Fig. 2-2). This climate is classified as Afi, equatorial superhumid with no dry season (Köppen 1936), i.e. more than 60 mm rainfall in all months and a temperature difference of less than 5 C between the warmest and the coldest month. According to the life zone system of Holdridge (1982; Holdridge et al. 1971) the Middle Caquetá area is classified as humid tropical forest (bh-t). 18

19 Figure 2-1. Location of the two permanent plots (M1 = upland plot; M2 = floodplain plot) at Peña Roja, in the middle part of the Caquetá Basin, Colombian Amazonia. The map is derived from Duivenvoorden and Lips (1995). 19

20 Figure 2-2. Climate diagram of Araracuara, Colombian Amazonia. Taken from Duivenvoorden and Lips (1995). Both sources (Duivenvoorden and Lips 1995; Tobón 1999) agree that the daily fluctuations in temperature are higher than the yearly fluctuations, and that the range of day and night temperatures is larger in the dry period than in the wet period. High temperatures occur in January and February and low temperatures in June. The latter are associated with a so-called "friaje" or "cold spell" which is a local phenomenon caused by the movement of cold air masses coming from the south of the continent through the Amazon Basin, and whose appearance has also been reported in Brazil (Salati 1985; Tobón 1999). The average monthly maximum temperature at Araracuara ranged from 29.5 C to 32.1 C and the average monthly minimum values from 21.2 C to 22.6 C (Duivenvoorden and Lips 1995). At Peña Roja the maximum temperature rarely exceeded 35 C and the minimum did not fall below 19 C (Tobón 1999). Despite the similarity between the mean annual precipitation at Araracuara and Peña Roja, there were a few differences in the yearly distribution pattern. At Araracuara, the average annual rainfall was 3059 mm and showed a unimodal distribution with high values around May, and low values around January (Fig. 2-2; Duivenvoorden and Lips 1995). At Peña Roja the annual average rainfall was 3420 mm and showed a bimodal pattern with a slight decrease in June (Tobón 1999) and between 20

21 December and February. The wettest month was September not May, as in Araracuara. The distribution of rainfall through the year is determined by the trade winds (Domínguez 1987; Botero 1999). Although Araracuara belongs to the Southern Hemisphere its rainfall pattern is typical for that of the Northern Hemisphere, and is furthermore characterized by the absence of a pronounced dry season (Salati 1985). In Colombian Amazonia the pattern of rainfall is controlled by the north-south movement of the belt of intertropical convergence. Most of the rainfall occurs in the afternoon and evening (Tobón 1999; Téllez 2003). At Peña Roja, the wettest year was 1994, and the driest year was On average it rained on 197 days per year, corresponding to 616 hours of rainfall per year (Tobón 1999; Téllez 2003). 2.3 Hydrology Peña Roja is located in the catchment of the Caquetá River. This river originates in the Andes and, with a length of 2200 km, it is the largest river in Colombian Amazonia. It is a so-called white-water river, which implies that its water tends to have a whitish color due to suspended clay. The river water has a neutral ph (Duivenvoorden and Lips 1993). The water level of the Caquetá River varies annually according to four seasons: low water, rising water, high water and falling water (Rodríguez 1999). The variation in river water level recorded at several stations along the river ranges from 6.5 m to 8.5 m, with peaks during July-August and low water levels between December-February (Duivenvoorden and Lips 1993; Urrego 1997). In addition to water levels associated to the annual flood cycles, occasionally (once every 3 to 11 years) the river water level rises to exceptional heights. The origin of this phenomenon is unknown, but it has been linked to global climate change, periods of large sunspot activity, or to the influence of El Niño events (van der Hammen and Cleef 1992; Botero 1999). The floodplain of the Caquetá River is called várzea (Prance 1980; Junk 1984, 1990; Padoch et al. 1999). It is built up by fine sediments which are deposited during overflows (Eden et al. 1982; Hoorn 1990; Duivenvoorden and Lips 1993, 1995). Currently there is no evidence of 21

22 gravel transport. The Caquetá River is actively eroding sediments of its fluvial terraces, especially along its southern river bank (Eden et al. 1982). 2.4 Land units The upland plot (also denominated monitoring plot 1, or M1) is located on a land unit, which was classified as Tertiary Sedimentary Plain by Duivenvoorden and Lips (1993, 1995). This land unit covers between 85-90% of the Colombian Amazon (PRORADAM 1979; Botero 1999). It can be seen as upland or "tierra firme" because it is never flooded by river water (Botero 1999; Duivenvoorden et al. 2001). Locally, uplands are also known as "monte firme" (Spanish) or "baj+hó (Muinane). The permanent plot is located along the upper slope of a valley (Fig. 2-3) developed in the Tertiary Sedimentary Plain along the left side (downstream) of the Caquetá River, opposite Sumaeta Island (Fig. 2-1), in the Peña Roja community. The plot coordinates are 0 39'31'' S, 72 4'38'' W. Its altitude is approximately 210 m above sea level. The soils in the upland plot were classified as Ultisols: Kanhapludults in stable positions and Paleudults in positions with more active erosion (SSS 1987; Alarcón 1990). The soils are deep and show a ABtC profile. In some sectors gravelly sandy materials are found at the top, sometimes with abundant charcoal, which may well be a result of ancient human activity. Textures are sandy loam to clay in the upper horizons, sandy loamy clay in the middle part, and clay or sandy clay in the lower part of the pedon. A detailed soil profile description of a soil pit located just outside the permanent plot (20 m from the northwestern border) was presented by Duivenvoorden and Lips (1993, 1995; plot 125) and is in Appendix 1 (profile 125). At this same location (plot 125) Lips and Duivenvoorden (1996) measured a yearly above-ground fine litter fall of 680 ± 54 g m -2 y -1 (mean ± one SD) in The Mean Residence Time of the organic material in the ectorganic horizons was 3.3 y (Duivenvoorden and Lips 1995). 22

23 Figure 2-3. Detailed site map of the upland plot (Alarcón 1990; Tropenbos-Colombia 1990). A: relief of the basin showing 2-m isohypses; the values denote the altitude relative to an arbitrary reference point; the total area shown covers 7.2 ha; B: map showing the terrain units and the location of the permanent plot; C: cross-sections showing representative soils along the slopes of the U-shaped valley and the V-shaped valley; D: cover and name of the terrain units. 23

24 The floodplain plot (also denominated monitoring plot 2, or M2) is located on a sporadically inundated floodplain of the Caquetá River (Duivenvoorden and Lips 1993, 1995). This land unit is characteristically built up by so-called river bank complexes (convex-concave systems of up to 2 m high river banks alternating with depressions, which run more or less parallel to the main channel of the Caquetá River) and poorly drained basins. The plot was established along the right bank of the Caquetá River (downstreams) at 250 m distance from the river, and about 2.5 km north of Sumaeta island (Fig. 2-1). The coordinates of the plot are: 0 37' S, 72 10' W. Its altitude is approximately 155 m above sea level. The floodplain of the Caquetá River is locally called "rebalse" or "bajo" (Spanish), and "cajahó (Muinane). In principle, the plot is only flooded when the water level of the Caquetá River is exceptionally high. This occurs every 9-11 years (according to local indigenous informants). Such events are known as "conejeras". Between May and July 1989 the water level reached up to about 2 m above the average surface level of the plot. Each year, between May and July, the concave parts of the plot (Fig. 2-4) become inundated by rain water for about one month. This inundation usually starts in a depression in the eastern part of the plot, which carries water only during the rainiest time of year. For example, during June-July 1990, some parts of the plot were covered by standing water, at a depth of less than 1 m for one month. Due to the low frequency of flooding by the Caquetá River, and the inundation of the lower parts by rainwater, the forest in the floodplain plot can be seen as intermediate between a seasonal várzea and a swamp (sensu Prance 1980). However, in this dissertation the terms várzea and floodplain forest are used, in line with other studies in the area (Urrego 1991). Given its transitional nature and the relatively low influence of flooding by river water, we suggest caution when comparing the results presented here with those of typical Amazonian várzea. The soil forming processes in the floodplain plot are highly influenced by the continuous supply of organic material from the vegetation and the periodic deposition of sediments during river floods. The fluctuations of the water table induces the alternating reduction and oxidation of the soil pedon. Evidence for this are the gray, olive-green and orange mottles, which occur especially in the soils of the basins and at larger depth in the soils of the convex river banks. Soils were classified as Typic Tropaquept, 24

25 Typic and Aquic Dystropept (SSS 1987; Ordóñez 1990). Duivenvoorden and Lips (1993, 1995) described a soil pit in the plot (Appendix 1, profile 126). In the forest directly surrounding this pit they measured a yearly above-ground fine litter fall of 1070 ± 132 g m -2 y -1 (mean ± one SD) in (Lips and Duivenvoorden 1996). The organic material in the ectorganic horizons had a Mean Residence Time of only 1.0 y (Duivenvoorden and Lips 1995). 2.5 Plot location, plot size and sampling set-up The position of the plots within each land unit was selected such as to ensure that the terrain characteristics regarding relief were similar. Because slopes were hardly developed in the floodplain unit, the upland plot was established at the summit and upper slope positions to minimize the slope (Fig. 2-3). Both plots were set up in forests that lacked any sign of recent human activities. Also in the buffer zones of 100 to 300 m around the plots (Phillips and Baker 2002; Vallejo et al. 2005), no signs of human activities were present. The conspicuous presence of charcoal in the soils of the upland plot (Alarcón 1990) and the remains of pottery found in nearby fluvial terraces of the Caquetá River (Mora et al. 1991; Mora 2003) demonstrate the ancient human occupation and associated change of the forest cover at the monitoring sites. However, at the time of the establishment of the plots the forests did not show any evidence of recent human interventions. This was also concluded from the presence in the forests of large trees with highly valuable timber, as Cedrela odorata (Meliaceae, red cedar) in the floodplain forest and Mezilaurus itauba (Lauraceae, Itaúba) in the upland forest. At the start, in 1989 (Table 2-1) one square 1-ha plot was installed at the upland site (part I in Fig. 2-5). Permanent plots of 1 ha have traditionally been the standard unit of sample area in tropical moist forest inventories, facilitating comparison and also yielding an optimal ratio area/perimeter (Synnott 1979, 1991; Jonkers 1987; Alder and Synnott 1992; Dallmeier 1992). However, on the basis of analyses (not shown) of the first data about the forest diversity using Hill's family of curves (Pielou 1975; Magurran 1988; Krebs 1989) the plot size was increased to 1.8 ha (Londoño and Alvarez 1991). To enlarge the original upland plot of 100 x 25

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