Forest Structure, Function and Dynamics in Western Amazonia E-Book
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- Herausgeber: John Wiley & Sons
- Kategorie: Wissenschaft und neue Technologien
- Sprache: Englisch
The Amazon Basin contains the largest and most diverse tropical rainforest in the world. Besides the Andes and the Atlantic Ocean, the rainforest is bounded to the north by the Guiana crystalline shield and to the south by the Brazilian crystalline shield, marked at their edges by cataracts in the rivers and often dominated by grasslands. This book is motivated not just by the Amazon's scientific interest but also by its role in many ecosystem functions critical to life on Earth. These ecosystems are characterized both by their complexity and their interactive, higher-order linkages among both abiotic and biotic components. Within Amazonia, the Western Amazon (west of 65° latitude) is the most pristine and, perhaps, the most complex within the Amazon Basin. This Western Amazon may be broadly divided into non-flooded forests (e.g. terra firme, white sand, palm) and forests flooded with white water (generally referred to as várzea) and with black water (generally referred to as igapó). Here, for the first time, is a book devoted entirely to Western Amazonia, containing chapters by scientists at the forefront of their own areas of expertise. It should be a valuable resource for all future researchers and scholars who venture into Western Amazonia, as it continues to be one of the most beautiful, mysterious, remote and important ecosystems on Earth.
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Table of Contents
Cover
Title Page
Copyright
Dedication
List of Contributors
Prologue
Chapter 1: Introduction
1.1 The Amazon
1.2 The Western Amazon
1.3 About this book
Acknowledgements
References
Chapter 2: A Floristic Assessment of Ecuador's Amazon Tree Flora
2.1 Introduction
2.2 Methods
2.3 Study area
2.4 Herbarium collections
2.5 Floristic inventories
2.6 Data analysis
2.7 Results
2.8 Aguarico-Putumayo watershed
2.9 Napo-Curaray basin
2.10 Pastaza basin region
2.11 Cordillera del Cóndor lowlands
2.12 What factors drive gradients in alpha and beta diversity in Ecuador's Amazon forests?
2.13 The role of geomorphology and soils in patterns of floristic change in Ecuadorian Amazonia
2.14 Potential evolutionary processes determining differences in tree alpha and beta diversity in Ecuadorian Amazonia
2.15 Future directions
References
Chapter 3: Geographical Context of Western Amazonian Forest Use
3.1 Introduction
3.2 Conditions set by the physical geography
3.3 Pre-Colonial human development
3.4 Colonial era
3.5 Liberation and forming of nations
3.6 World market integration and changing political regimes
3.7 Characteristics of the present forest use
3.8 Present population and regional integration
References
Chapter 4: Forest Structure, Fruit Production and Frugivore Communities in Terra firme and Várzea Forests of the Médio Juruá
4.1 Introduction
4.2 Methods
4.3 Results and discussion
4.4 Conclusion
References
Chapter 5: Palm Diversity and Abundance in the Colombian Amazon
5.1 Introduction
5.2 Study area
5.3 Methods
5.4 Results
5.5 Discussion
Acknowledgements
References
Chapter 6: Why Rivers Make the Difference: A Review on the Phytogeography of Forested Floodplains in the Amazon Basin
6.1 Introduction
6.2 The geological history of flood-pulsing wetlands in the Amazon Basin
6.3 Floodplain environments: why rivers make the difference
6.4 Conclusions
References
Chapter 7: A Diversity of Biogeographies in an Extreme Amazonian Wetland Habitat
7.1 Introduction
7.2 Methods
7.3 Construction of a biogeographic framework
7.4 Results
7.5 Discussion
Acknowledgements
References
Chapter 8: Forest Composition and Spatial Patterns across a Western Amazonian River Basin: The Influence of Plant–Animal Interactions
8.1 Introduction
8.2 Methods
8.3 Analysis
8.4 Results
8.5 Discussion
References
Chapter 9: Bird Assemblages in the Terra Firme Forest at Yasuní National Park
9.1 Introduction
9.2 Methods
9.3 Results and discussion
References
Chapter 10: Conclusions, Synthesis and Future Directions
10.1 Conclusions
10.2 Synthesis
10.3 Future directions
References
Index
End User License Agreement
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Guide
Cover
Table of Contents
Begin Reading
List of Illustrations
Chapter 1: Introduction
Figure 1.1 Photograph of the Amazon river near where it “officially” begins in Iquitos, Peru.
Figure 1.2 Map of the Amazon river and some of the other rivers that drain into it, including the Japurá river, with an outline of the Amazon Basin.
Chapter 2: A Floristic Assessment of Ecuador's Amazon Tree Flora
Figure 2.1 Map of the tree plot network established in the Ecuadorian Amazon.
Figure 2.2 Patterns of diversity at the family level in the tree flora of the Ecuadorian Amazon.
Figure 2.3 Species area curve and the estimated number of tree species expected for Ecuadorian Amazonia. Gray shaded area represents the 95% upper and lower bounds for the expected number of species with respect to area sampled. Data are based on a plot network of 70 one-hectare plots established across Ecuadorian Amazonia.
Figure 2.5 Tree alpha diversity (measured as Fisher's α) as a function of (a) longitude and (b) latitude. Loess regression is fitted to the data. The highest values of tree alpha diversity are found in the YNP region close to the Andean foothills and are probably related to climate and soil conditions. The gray bars highlight the Yasuni Park area.
Figure 2.4 Non Metric Multidimensional ordination based on the species-level compositional dissimilarity matrix for 70 1-h plots in
terra firme
forests of the Ecuadorian Amazon. Ellipses represent the 95% confidence interval in grouping plots as part of a particular cluster of similar floristic units. Four floristically distinct regions are defined: triangle ellipse = forests in the lowlands of the Cordillera del Cóndor region; black ellipse = forests in the Aguarico-Putumayo watershed; square ellipse = forests in the hyperdiverse Napo-Curaray watershed; and diamond ellipse = forests in the Pastaza watershed.
Figure 2.6 Floristic changes are correlated with shifts in climate along a longitudinal gradient in Ecuadorian Amazon forests. (a) Scores of DCA axis 1 against longitude. (b) Climatic distance against longitude; climatic differences were measured as score values of a PCA (principal component analysis) on the basis of monthly mean precipitation values and maximum, minimum and mean temperature values (see Ministerio del Ambiente del Ecuador 2013b). (c) Change in tree species composition as a function of climatic distance. (d) Spatial variation in precipitation across Ecuador. Interpolation of precipitation values was done using the kriging method on the basis of monthly mean precipitation values at 1 km
2
resolution. (e) Spatial variation in temperature across Ecuador. Interpolation of precipitation values were done using the kriging method on the basis of mean, maximum and mean temperature values at 1 km
2
resolution. A longitudinal gradient for temperature and precipitation at landscape scales is clear for (d) and (e).
Chapter 3: Geographical Context of Western Amazonian Forest Use
Figure 3.1 Large-scale geographical patterns in Western Amazonia. (a) Physiography. (b) Climate – variations in mean monthly temperature and precipitation (based on: http://www.worldclimate.com/worldclimate/index.htm). (c) Geological formations and dynamics (after Räsänen
et al.
1987). (d) Vegetation.
Figure 3.2 A general outline of the civil and economic histories in northwest South America with special reference to the Western Amazon Region.
Figure 3.3 Political boundaries in northern South America. (a) The early colonial period
ca
. 1650. (b) The late colonial period
ca
. 1800. (c) After the liberation
ca
. 1830 (drawn according to Lombardi
et al
. 1983).
Figure 3.4 The dissolution of independent Gran Colombia in 1830 (based on Lombardi
et al
. 1983).
Figure 3.5 (a) Boundary disputes between Colombia, Ecuador and Peru (1830–1942). (b) Brazilian territorial expansion (based on Lombardi
et al.
1983).
Figure 3.6 Development in Ecuadorian Amazonia in the second half of the 20th century (Eastwood and Pollard 1992).
Figure 3.7 (a) Deforestation in Western Amazonia (courtesy of http://wwf.panda.org/wwf_news/?208511/Keeping-an-eye-on-deforestation). (b) Aerial view showing deforestation patterns near Iquitos, Peru (numbers explained in the text) [(b) image from http://www.esri.com/landing-pages/software/landsat/unlock-earthssecrets].
Figure 3.8 Territorial reservations in Western Amazonia. (a) Indigenous territories. (b) Nature protection areas. (c) Hydrocarbon production areas in different stages of accomplishment (dark grey: leased; light grey: not yet leased) (data from https://raisg.socioambiental.org/).
Figure 3.9 Population in Western Amazonia. (a) Major population centers (with >10 000 inhabitants) in the Amazonian departments/states/provinces (see Table 3.2). (b) Population density in the Amazonian departments.
Figure 3.10 Major transport geographical settings in Western Amazonia. (a) Major river corridors providing fluvial interconnectivity. (b) Roads (source: http://raisg.socioambiental.org/mapa. (c) Airports and connections from Western Amazonian cities (source: http://www.flightconnections.com/).
Chapter 4: Forest Structure, Fruit Production and Frugivore Communities in Terra firme and Várzea Forests of the Médio Juruá
Figure 4.1 Map of the Médio Juruá region of western Brazilian Amazonia, showing the distribution of forest types within the two study reserves. Colors indicate terrain elevation, which corresponds approximately with the boundary between
terra firme
and
várzea
forests more clearly shown by the dashed lines.
Figure 4.2 Comparative views of
terra firme
and várzea forests in the Médio Juruá region of western Brazilian Amazonia, and corresponding field methods in each forest type.
Chapter 5: Palm Diversity and Abundance in the Colombian Amazon
Figure 5.1 Study area in the eastern Colombian Amazon where 71 transects were placed along Río Guaviare (between the town of Inírida and the upstream settlement of Baranco Picure, 200 km to the west), Río Caquetá (from 42 km west to 11 km east of the village/military camp of La Pedrera), Medio Caqutá (225 km west of La Pedrera) and Río Amazonas (at Puerto Nariño, 68 km northwest of Leticia).
Figure 5.2 Some common canopy (a, b, c) and understory (d, e, f) palm species in the
terra firme
forest of eastern Colombian Amazon. (a)
Astrocaryum chambira
. (b)
O. bataua
. (c)
I. deltoidea
. (d)
I. setigera
. (e)
Geonoma maxima
var maxima. (f)
Bactris simplicifrons
.
Figure 5.3 Abundances (ind./ha) of the 15 most abundant palm species in eastern Colombian Amazon, recorded in 71 transects (17.25 ha) in
terra firme
and floodplain forest.
Figure 5.4 Numbers of species of palms in eastern Colombian Amazon in each growth form, as defined in Balslev
et al
. (2011) and in each habitat (
terra firme
/floodplain and terraces).
Figure 5.5 Numbers of species of palms in eastern Colombian Amazon in each architectural form and in each habitat (
terra firme
/floodplain and terraces).
Figure 5.6 Some common canopy (a, b, c, d) and understory (e, f, g) palm species from the floodplain and terrace forests of eastern Colombian Amazon. (a)
Mauriatia flexuosa
. (b)
Astrocaryum jauri
. (c)
A. butyracea
. (d)
E. precatoria
. (e)
Bactris major var major
. (f)
Desmoncus polyacanthos
. (g)
Manicaria saccifera
.
Figure 5.7 Rarefaction curve for palm species richness in 71 transects in eastern Colombian Amazon, total and divided by habitat (
terra firme
/floodplain) and for the three groups of transects made at the Guaviare, Caquetá and Amazonas rivers.
Chapter 7: A Diversity of Biogeographies in an Extreme Amazonian Wetland Habitat
Figure 7.1 The distribution of sampled wetlands along a 250-km stretch of the Madre de Dios river, Peru. The inset shows the locations of the sampled wetlands in relation to the distribution of the Gentry plot network used to assess distributional patterns of families and genera along the elevation gradient. The Gentry sites are classified into two groups differentiated by site elevation, where upwards and downwards pointing triangles correspond to sites at greater than and less than 1,000 m a.s.l., respectively.
Figure 7.2 A comparison of the variability of biogeographic relations among co-occurring individuals (FD
Q
) between the sampled wetland sites and 6 Gentry sites representative of lowland (<500 m a.s.l.) tropical forests. Sites are distributed along a horizontal axis of richness based on a random sub-sample of 365 individual stems (the least number of stems at a single site). Dashed horizontal and vertical lines indicate group means along both axes. While extreme wetland assemblies tend to be less species rich, their constituent taxa are highly variable in regards to their biogeographic patterns.
Figure 7.3 Scatterplots showing the change in biogeographic variability (FD
Q
) in local plant assemblies along the main physiognomic gradient sampled in wetlands (a) and the taxa with the highest relative influence on our measure of biogeographic variability (b).
Chapter 8: Forest Composition and Spatial Patterns across a Western Amazonian River Basin: The Influence of Plant–Animal Interactions
Figure 8.1 Map of MDDB, Peru, showing locations of four long-term forest dynamics plots used in this study and their faunal status. Original base map created by Nelson Gutiérrez, Amazon Conservation Association (2007).
Figure 8.2 Layout and design of each of four study sites, showing 4-ha (200 m × 200 m) forest dynamics plot and centrally located seed fall monitoring grid (104 m × 104 m) and 1-ha (100 m × 100 m) recruitment monitoring area.
Figure 8.3 Mean percentage of all stems comprised by the 10 most abundant families of trees (stems ≥10 cm diameter) in four 4-ha forest dynamics plots spread across the MDDB, Peru: pooled across sites (±SD), and within each site for first 9 families.
Figure 8.4 Mean percentage of all stems comprised by the 10 most abundant species of trees (stems ≥10 cm diameter) in four 4-ha forest dynamics plots spread across the MDDB, Peru: pooled across sites (±SD), and within each site for first 9 species
Figure 8.5 Mean percentage of all stems comprised by the 10 most abundant families of understory woody taxa (stems ≥1 m height, <10 cm diameter) in the central hectare of 4 forest dynamics plots spread across the MDDB, Peru: pooled across sites (±SD), and within each site for first 9 families.
Figure 8.6 Mean percentage of all stems comprised by the 10 most abundant species of understory woody taxa (stems ≥1 m height, <10 cm diameter) in the central hectare of 4 forest dynamics plots spread across the MDDB, Peru: pooled across sites (±SD), and within each site for first 9 species.
Figure 8.7 Median NND (nearest neighbor distance, m), i.e. distance between a stem and its nearest conspecific neighbor within a cohort, averaged across species for three distinct size-age stem cohorts at each of four long-term forest dynamics plots spread across the MDDB, Peru, and pooled across sites.
Figure 8.8 Proportions of species whose observed median NND (nearest neighbor distance of conspecific stems) confirmed to a “clumped”, “random”, or “dissociated” intra-cohort spatial pattern based on its percentile rank compared to 1,000 iterations of randomly sampled heterospecific stems of the same cohort. Results shown separately for each cohort and pooled across cohorts, in each case pooled across all four sites.
Figure 8.9 Median recruitment distance (RD, m), i.e. distance between a focal sapling and its nearest conspecific adult tree, averaged across species at each of four long-term forest dynamics plots spread across the MDDB, Peru, and pooled across sites.
Figure 8.10 Proportions of species whose observed median recruitment distance confirmed to a “clumped”, “random” or “dissociated” inter-cohort spatial pattern based on its percentile rank compared to 1,000 iterations of randomly sampled sapling and adult tree stems. Results shown separately for each site and pooled across all four sites.
Chapter 9: Bird Assemblages in the Terra Firme Forest at Yasuní National Park
Figure 9.1 Study area with the location of the canopy MSFs studied.
Figure 9.2 Composition of each canopy MSF (Family).
Figure 9.3 Territory size of canopy and understory mixed species flocks. Comparison between six studies. Blue circle: canopy flocks; Green: understory flocks. MSF canopy Yasuní: data presented in this study. MSF canopy Cocha-Cashu: data of Munn (1985) in Manu, Peru. MSF understory Yasuní PE: data from English (1998). MSF understory Yasuní MT/LB: data from Baquero (2003) and Tobar (2006). MSF understory GB: data from Buitrón–Jurado (2005) in Yasuní. MSF understory Cocha Cashu: data from Terborgh
et al.
(1990) in Manu, Peru.
Chapter 10: Conclusions, Synthesis and Future Directions
Figure 10.1 La cantina “The Amazons” on the corner of SW 59th Street and south May Avenue in Oklahoma City, Oklahoma (photo courtesy of Nicole Bankert).
Figure 10.2 A plant-centered view of ecosystems illustrating how important ecosystem components, such as Carbon (C), Nitrogen (N), Phosphorus (P) and Potassium (K), move in and out of the total plant phyto-mass.
List of Tables
Chapter 2: A Floristic Assessment of Ecuador's Amazon Tree Flora
Table 2.1 Estimates of tree species diversity, total number of trees and levels of endemism in Ecuador's Amazonian tree communities. Both values for Fisher's alpha and Chao's 2 metrics are shown
Table 2.2 Generalized Least Squares analysis (GLS) comparing monthly mean precipitation values, maximum, minimum and mean temperature values condensed in the axis 1 of a Principal Components analysis (PC1), with respect Fisher's alpha values, Longitude and Latitude. Model selection to obtain the best model possible was performed applying a backward selection and a likelihood ratio test. Significance of the variables interactions for each model is coded as: * = significant at the 5% level; ** = significant at the 1% level
Table 2.3 Generalized Additive Model analysis (GAM) comparing climatic variables condensed in the axis 1 values of a Principal Components analysis (PC1), with Fisher's alpha values, Longitude and Latitude, to detect patterns of tree diversity in Ecuadorian Amazon forests. Significance of the variables interactions for each model is coded as: * = significant at the 5% level; ** = significant at the 1% level
Chapter 3: Geographical Context of Western Amazonian Forest Use
Table 3.1 Primary wood product trade export statistics in 1999 and 2013 at country level. (source: ITTO Annual review statistics database http://www.itto.int/annual_review_output/?mode=searchdata)
Table 3.2 Administrative units and population in Western Amazonia
Chapter 5: Palm Diversity and Abundance in the Colombian Amazon
Annex 5.1 Localities for the 71 transects in which palms were inventoried. Transect number were given in the field and used in the transect database kept at Aarhus University. Habitat is given as FP = floodplain or TF =
terra firme
. Alt is elevation above sea level. X and Y are geographic coordinates recorded with GPS in the field at the beginning of each transect
Table 5.1 Diversity and abundance of palms in eastern Colombian Amazon, studied in 71 transects along the Guaviare, Caquetá and Amazonas rivers
Table 5.2 Number of palm species in 71 transects in the eastern Colombian Amazon with pinnate, palmate, and costapalmate leaves, respectively
Table 5.3 Growth forms of American palms defined by overall size of stems and leaves, whether caulescent or acaulescent, and whether climbers or not (Table based on Balslev
et al
. 2011)
Annex 5.2 Palm species found in 61 transects of 5 × 500 m (15.25 ha) and 10 transects of 4 × 500 m (2 ha) in eastern Colombian Amazon arranged according to their total abundance in this study (column 3; Total ind. all transects). Vouchers were collected in the number series of Rodrigo Bernal (RB) and are deposited in the herbarium (COL) of Instituto de Ciencias, Universidad Nacional de Colombia. Columns 4 and 5 give the total number of individuals collected in
terra firme
and floodplain forest transects, respectively. Palm architecture refers to whether the palms were solitary (sol), cespitose (ces) or colonial (col). Growth form follows the classification proposed by Balslev
et al
. 2011 (1 = Large Tall-Stemmed Palms, 2 = Large-Leaved Medium-short Stemmed Palms, 3 = Medium-sized Palms, 4 = Medium/Small Palms with Stout Stems, 5 = Small Palms, 6 = Large Acaullescent Palms, 7 = Small Acaulescent Palms, and 8 = Climbing Palms). Leaf shape is given as pinnate (pin), palmate (pal) or costa-palmate cos (cop)
Chapter 7: A Diversity of Biogeographies in an Extreme Amazonian Wetland Habitat
Table 7.1 Loadings of the 8 physiognomic variables recorded in sample wetlands on the primary PCA axis. This axis describes a transition from taller forested sites with reduced herbaceous layers (shady understory) to shrubby sites with increasing density of smaller-stemmed, low-statured individuals and a well-developed herbaceous layer dominated by light-loving Cyperaceae and Poaceae
Chapter 8: Forest Composition and Spatial Patterns across a Western Amazonian River Basin: The Influence of Plant–Animal Interactions
Table 8.1 Summary data of four forest dynamics plots used in this study
Forest structure, function and dynamics in Western Amazonia
Edited by
Randall W. Myster
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Library of Congress Cataloging-in-Publication Data
Title: Forest structure, function and dynamics in Western Amazonia / edited by Randall W. Myster.
Description: Chichester, West Sussex : John Wiley & Sons, Inc., 2017. Includes bibliographical references and index.
Identifiers: LCCN 2016042844| ISBN 9781119090663 (cloth) | ISBN 9781119090694 (epub).
Subjects: LCSH: Rain forests – Amazon River Region. | Rain forest ecology – Amazon River Region. Biodiversity – Amazon River Region. Geography – Amazon River Region. | Plants – Amazon River Region. | Amazon River Region.
Classification: LCC SD160. F67 2017 | DDC 577.340985/44 – dc23 LC record available at https://lccn.loc.gov/2016042844.
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Cover image: © Sebastián Crespo Photography / Getty Images
Cover Design: Wiley
Dedicated to the memory of my beloved cat, Shaman.
“Das Schöne ist eine Manifestation geheimer Naturgesetze, die uns ohne dessen Erscheinung ewig wären verborgen geblieben.” (Beauty is a manifestation of secret natural laws, which otherwise would have been hidden from us forever).
J. W. von Goethe
Source: Goethe, Maximen und Reflexionen. Aphorismen und Aufzeichnungen. Nach den Handschriften des Goethe- und Schiller-Archivs hg. von Max Hecker, Verlag der Goethe-Gesellschaft, Weimar 1907. Aus Kunst und Altertum, 4. Bandes 2. Heft, 1823.
List of Contributors
Balslev, H.
Ecoinformatics & Biodiversity, Bioscience
Aarhus University
Aarhus, Denmark
Email:
Bernal, R.
Instituto de Ciencias Naturales
Universidad Nacional de Colombia
Bogotá, Colombia
Email:
Berrio, J. C.
Department of Geography
University of Leicester
Leicester, UK
Email:
Ceron, C.
Universidad Central
Escuela de Biología Herbario Alfredo Paredes
Quito, Ecuador
Email:
Copete, J. C.
Programa de Biología Con Énfasis En Recursos Naturales
Universidad Tecnológica del Chocó
Colombia
Email:
de Vries, T.
Escuela de Ciencias Biológicas
Facultad de Ciencias Exactas y Naturales
Pontificia Universidad Católica del Ecuador
Quito, Ecuador
Email:
Duque, A.
Departamento de Ciencias Forestales
Universidad Nacional de Colombia
Email:
Galean, G.
Instituto de Ciencias Naturales
Universidad Nacional de Colombia
Bogotá, Colombia
Email:
Guevara, J. E.
Department of Integrative Biology
University of California, Berkeley
Berkeley, CA. USA
Email:
Hawes, J. E.
Animal & Environmental Research Group
Department of Life Sciences
Anglia Ruskin University
Cambridge, UK
Email:
Householder, E.
Department of Wetland Ecology
Institute for Geography and Geoecology
Karlsruhe Institute of Technology
Josefstr. 1, Rastatt, Germany
Email:
Iglesias-Balarezo, A.
Rither
Bolivia, Quito
Email:
Janovec, J.
Facultad de Ciencias Ambientales
Universidad Cientifica del Sur
Lima, Peru
Email:
Kalliola, R.
Department of Geography and Geology
University of Turku
Turku, Finland
Email:
Mäki, S.
Department of Geography and Geology
University of Turku
Turku, Finland
Email:
Mogollon, H.
Endangered Species Coalition
Silver Springs, CO, USA
Email:
Myster, R. W.
Biology Department
Oklahoma State University
Oklahoma City, OK, USA
Email:
Neill, D. A.
Universidad Técnica del Norte
Herbario Nacional del Ecuador
Quito, Ecuador
Email:
Palacios, W. A.
Universidad Estatal Amazónica
Puyo, Ecuador
Email:
Pedersen, D.
Ecoinformatics & Biodiversity, Bioscience
Aarhus, Denmark
Email:
Peres, C. A.
School of Environmental Sciences
University of East Anglia
Norwich, UK
Email:
Pitman, N. C. A.
Keller Science Action Center
The Field Museum
Chicago, IL, USA
Email:
Sanchéz, M.
Departamento de Ciencias Forestales
Universidad Nacional de Colombia
Email:
Swamy, V.
San Diego Zoo Institute for Conservation Research
Escondido, California, USA
Email:
Tobler, M. W.
San Diego Zoo Global Institute for Conservation Research
Escondido, California USA
Email:
Toscano-Montero, G.
Escuela de Ciencias Biológicas
Faculta de Ciencias Exactas y Naturales
Pontificia Universidad Católica del Ecuador
Quito, Ecuador
Wittmann, F.
Department of Wetland Ecology
Institute of Geography and Geoecology
Karlsruhe Institute for Technology
Josefstr. 1, Rastatt, Germany
Email:
Prologue
My first experience in the Amazon occurred in 1995 when I went on a “canned” ecotourism trip to the Rio Napo in eastern Ecuador. Although we saw Anaconda, various monkey species, and a troop of Coatimundi, it was what happened during our return that has stayed with me the longest. Because our plane was departing quite early, we had to leave at four in the morning. We piled into the boat and all was well until we got stuck on a sandbar. At that point, all the men were ordered to disrobe and get into the dark water to push. As I was jumping in, I remembered all the movies and documentaries I had ever seen about the Amazon. I wondered: Would I be attacked from below by a mysterious species unknown to Science, Would the bottom be littered with the corpses of “Indians” murdered by the Conquistadors? or, Would I be swept away, my body melting into a mystical union with the Amazon for all eternity?
I survived to tell the tale and as I worked in the Amazon over the next two decades she dazzled in her beauty, complexity and raw wildness, but not always in the most pleasant way. Perhaps my worst experience was an infected insect bite that landed me in the hospital for two weeks on an IV. During my stay I was told that my leg might have to come off! – but luckily the infection had not reached the bone. Alternatively, I can relate the sense of wonder I felt when, on a clear night, I gazed into the southern sky and saw those stars for the first time, or when I looked into the Yoda-like face of a Uakari monkey from only a few feet away, and waited for it to speak.
And so, the Amazon has been both cruel and deeply satisfying. I have learned to give myself over to her, like riding a horse high in the mountains; trusting her, to take me where I want to go.
R.W.M.
Chapter 1Introduction
Randall W. Myster
Abstract
This introduction presents an overview of the key concepts discussed in the subsequent chapters of this book. The book is motivated not just by the Amazon's scientific interest but also by its role in these various ecosystem functions critical to life on Earth. It highlights several of its interactive and higher-order linkages among both abiotic and biotic ecosystem components. The book provides summaries of the author's research in Western Amazonia over the last two decades, in both non-flooded forests and forests flooded with white water and with black water. The Amazonian rainforest is located in the equatorial regions of the South American countries of Brazil, Colombia, Ecuador, Bolivia and Peru. In addition, the Amazonian rainforest influences the entire world's precipitation and weather patterns and, over the longer term, the world's climate. Flooding differs within the Amazon landscape in frequency, timing, duration, water quality, and maximum water depth and height.
KeywordsAmazon landscape; Amazon's scientific interest; biotic ecosystem; black-water floodplain forests; equatorial regions; non-flooded forests; South American countries; world's climate
The Amazon Basin contains the largest and most diverse tropical rainforest in the world. In particular, the Western Amazon basin is the most pristine and, perhaps, the most complex within the Amazon Basin. This book is motivated not just by the Amazon's scientific interest but also by its role in these various ecosystem functions critical to life on Earth. In this introductory chapter, I first describe that complexity and highlight several of its interactive and higher-order linkages among both abiotic and biotic ecosystem components. Then I include summaries of my own research in Western Amazonia over the last two decades, in both non-flooded forests (e.g. terra firme, white sand, palm) and forests flooded with white water (generally referred to as várzea) and with black water (generally referred to as igapó). Finally, I outline the chapters to come.
1.1 The Amazon
When the first Europeans visited the middle of South America in the 1500s, they saw women warriors and named the area after those same figures in Greek Mythology, the Amazons. The two Iberian powers, Spain and Portugal, then fought for control of the Amazon. The Spanish were mainly interested in the wealth of the Incas and so approached the Amazon from the west, but because the Amazon is so flat the Portuguese could colonize (from the east) a much larger part of it. All this was finally resolved by the Treaty of Madrid in 1750, establishing the general boundaries we see today between Brazil (colonized by Portugal) and Bolivia, Peru and Ecuador (colonized by Spain: Hecht 2014).
One may speak of the river itself (the Amazon), the large depression/watershed which surrounds the Amazon and the smaller rivers and streams which drain into it (the Amazon Basin) or the forest that grows in that basin (the Amazonian rainforest). The Amazon river has the greatest discharge of fresh water in the world and is also its second longest river (∼6,400 km). It originates in the foothills of the Andean Mountains of South America (Figure 1.1) and runs east into the Atlantic Ocean. It is not constant, however, and has changed over time. For example, climate change during the Pleisotcene (2,588,000 to 11,700 years ago) lead to a drop in sea level which changed its course, and also a rise in sea level that filled its connecting rivers with sediments, creating large floodplains. The Amazon Basin predates the separation of South America from Africa some 110 million years ago (Junk et al. 2010) and is generally found below 200 m above sea level (a.s.l.), covering over 8 million km2 (Hoorn and Wesselingh 2010).
Figure 1.1 Photograph of the Amazon river near where it “officially” begins in Iquitos, Peru.
The Amazonian rainforest is located in the equatorial regions of the South American countries of Brazil, Colombia, Ecuador, Bolivia and Peru. Besides the Andes and the Atlantic Ocean, the rainforest is bounded to the north by the Guiana crystalline shield and to the south by the Brazilian crystalline shield (Pires and Prance 1985), marked at their edges by cataracts in the rivers and often dominated by grasslands (Myster 2012a). This rainforest is the world's largest tropical rainforest and it is the largest continuous forest of any kind, encompassing over 6 million km2 (Holdridge 1967; Junk et al. 2010; Walter 1973). It produces approximately 20% of the world's oxygen and approximately 10% of the net primary productivity of the entire terrestrial biosphere. Its biodiversity is legendary (present at least since the Pleisotcene), having at least 11,200 tree species (Fabaceae the most common family: Daly and Prance 1989; Hoorn and Wesselingh 2010) and more than 10% of all the species on the planet (Pires and Prance 1985).
Perhaps most importantly for the future of humans, the rainforest interacts intimately with the Earth's carbon (C) cycle, acting both as a carbon “sink”, by taking in large amounts of CO2 through photosynthesis (∼15% of the world's total), and as a carbon “source” as, for example, when its plants decay or burn (Rice et al. 2004). The Amazonian rainforest contains 20–25% of the world's terrestrial C, with one-third below ground in the soil and two-thirds in the above-ground vegetation (McClain et al. 2001). The Amazon rainforest will continue to be a major C player in the future by both contributing to, and suffering the effects of, global warming (Shukla et al. 1990).
In addition, the Amazonian rainforest influences the entire world's precipitation and weather patterns and, over the longer term, the world's climate (Keller et al. 2004). Evaporation and condensation over Amazonia are engines of global atmospheric circulation (Malhi et al. 2008) and this rainforest may even control how much rainfall it itself receives (Pires and Prance 1985). Daily fluctuations in temperature are greater than seasonal fluctuations, that is 27.9°C in the dry season and 25.8°C in the rainy season. Humidity also varies little seasonally, 77% in the dry season and 88% in the rainy season (Prance and Lovejoy 1985) and its prevailing winds come from the east. Because the rainforest is located on the Equator. it has a day length which varies little during the year, as does solar energy input of 767–885 calories per cm2 per day. Up to 6 m of rain falls every year on the Amazon Basin with a pronounced rainy season that begins in the south. One study showed that 25.6% of that precipitation was intercepted by plants and returned to the atmosphere by evaporation, 45.5% was taken up by plants and transpired, and the rest was absorbed into the soil and/or ran off (through the litter and the soil) into rivers and streams (Salati 1985).
White-water runoff from the Andes appears white because of the high concentrations of dissolved solids, mainly alkali-earth metals and carbonates. This white water has the highest concentration of total electrolytes, phosphorus (P), potassium (K) and other trace elements, of all the Amazonian flood waters, with a pH of around 7 (neutral). Conversely, black water is transparent due to its low amount of suspended matter and has the lowest concentrations of these ions, with high amounts of humic acid (resulting in a pH of between 4 and 5) and clear water is between these two in amounts of sediments and pH. Productivity follows these nutrient trends (Junk and Furch 1985). The resulting Amazonian floodplain forests are the most diverse flooded forests in the world with at least 1,000 tree species, existing since the early Cretaceous (145–66 million years ago: Junk et al. 2010). In general, forest in black- and clear-water areas are less diverse, with less litter production, smaller trees and less herbaceous growth than forests in white-water areas (Junk et al. 2010). Clear-water rivers come from areas where erosion is less intensive but the more sandy a soil is, the more likely it is to give up its humic substances and create black water. Most leaching occurs during the rainy season and when human activity increases erosion.
The large rainfall results in low fertility soil (and has for millions of years: Jordan 1985), which leads to most of the nutrients being stored in the plant biomass with a fast, efficient closed system of nutrient cycling. P availability mostly limits productivity in terra firme forests (McClain et al. 2001). The large rainfall also leaches out significant nutrients, such as nitrogen (N), calcium (Ca), P and K out of the leaves and stems of trees and shrubs, adding soluble inorganic and organic substances which allow epiphytic plants to live without a root system. This makes the water cycle a significant interactive link between the soil and the biota. Trees adaptations (e.g. dense root mats at the soil surface) must then be “fine-tuned” (niche-packing) in order to take up and store nutrients efficiently, and this may be one of the reasons for the large biodiversity found in the Amazon. Other reasons may include the relatively constant wet and warm climate (including a predictable flood pulse) and a heterogenous edaphic substrate. Other common adaptations found in Amazon trees include:
tough, leathery, long-lived leaves;
supporting colonies of algae and lichens which help recover leached nutrients; and
sprouting roots from branches and leaves.
Human activities (logging with or without burning and agriculture) can affect many different aspects of the ecosystem – making it another significant interactive linkage – by leading to large losses of biomass (up to 90% C loss compared to the primary forest), with nutrients leaching out of the necromass and into the rivers and streams. Large species diversity may also lead to a large diversity of plant community types, many as yet unknown (Myster 2009, 2012b).
Within the Amazonian rainforest is the dominant, unflooded terra firme forest. The terra firme has the same physiogomy and many of the same structural characteristics of similar unflooded rainforests throughout the rest of the Neotropics (Everham et al. 1996; Kalliola et al. 1991; Lopez and Kursar 1999; Whitmore 1989). For example, the Amazonian terra firme forest also contains a large amount of above-ground biomass (AGB) and a complex strata of emergent trees, canopy trees, understory trees, palms, understory shrubs, climbing vines, saplings, seedlings, epiphytes, hemi-epiphytic stranglers, lianas, herbs and ferns. Along with this vertical structure, it also has an extensive horizontal structure of various-sized “gaps”, light flecks, micro-topological relief and patchy soil nutrient availability in acid, clayey-loamy shallow soils with extensive organic matter in its upper layers. As is true for trees elsewhere in the Neotropics, many of the terra firme trees have large buttresses and shallow roots, complex growth/sprouting architectures, and large epiphyte and liana communities (Janzen 1984). Within the broad classification of terra firme forest are different types of forest which differ in soil characteristics, for example terra firme proper on clay or loam soils, white-sand forests on soils with large amounts of quartz, and palm or swamp forests on standing water (Tuomisto et al. 2003).
Flooding within the Amazon Basin generates floodplains and flooded forests, which cover approximately 3 million km2 (Junk 1989; Parolin et al. 2004). Most of this water is the nutrient rich “white” water from the Andes, which creates forests generally called várzea (Junk 1984) and the rest is “black” and “clear” water, which is nutrient poor forest runoff and creates forests generally called igapó (Junk 1989; Prance 1979; Sioli 1984). There are also forests created from a mixing of the water types (Myster 2009). Differences in nutrient availability in the water may thus be as important in determining the structure, function and dynamics of flooded forests in the Amazon as is the nutrient availability in the soil for unflooded terra firme forests. The more studied várzea has light levels on the forest floor similar to terra firme (1–3% of ambient: Wittmann et al. 2010a,b), but flooding creates oxygen deficiency, reduced photosynthesis and low water conductance, so that flooding may be a greater source of mortality than desiccation. In addition, high nutrient levels within these várzea forests can lead to trees with rapid growth rates and low wood densities (Parolin et al. 2010). Importantly trees within these forests must time their reproduction cycles in relation to the flooding, where some species grow mainly during the flooded times of the year and reproduce when the waters subside, while other species merely “endure” flooding and reproduce only during the dry times of the year (Junk et al. 2010). White-water areas are used more than black- or clear-water areas for agriculture – because of more nutrients and the predictability of the flood pulse (Junk et al. 2010) – but effects of human land use (Myster 2007a) in white-water areas may be reduced due to flooding (Pinedo-Vasquez et al. 2011).
Flooding differs within the Amazon landscape in frequency, timing, duration, water quality, and maximum water depth and height (Junk et al. 2010; Myster 2009). Such variation within the flooding gradient (Myster 2001) greatly affects the distribution and abundance of plant species (Ferreira and Stohlgren 1999; Junk 1989; Lamotte 1990), leading to inundated vegetative associations created by the rise of the water table on a regular, seasonal basis. In general, flooded forest types, vegetation formations and plant communities lie on a continuum defined by:
the duration of the aquatic and terrestrial phases of the annual cycles; and
the physical stability of the habitat influenced by sedimentation and erosion processes (Junk
et al.
2010).
In general, the soils in the floodplain are less acid than those of the terra firme, but may have a greater concentration of exchangeable cations such as Mg+2 and Na+2 (reviewed in Honorio 2006).
Because the Amazon and its tributaries are very dynamic – often changing their routes within a time span of a few decades (Junk 1989; Kalliola et al. 1991; Pires and Prance 1985) – it may well be that forests that are unflooded today were flooded in the past and vice versa. It is not surprising that many terra firme species establish ecotypes (Myster and Fetcher 2005) in the flooded forest (Wittmann et al. 2004; 2010a,b). For example:
the
terra firme
species
Guazuma ulmifolia
and
Spondias lutea
have developed flood-resistant ecotypes now found in
várzea
;
várzea
species such as
Ceiba pentandra
and
Pseudobombax munguba
occur in
terra firme
; and
several species of the genus
Maquira
occur in both unflooded and flooded forests.
This ecotropic dynamic and the high species richness of the surrounding Amazonian terra firme rainforest, which disperse seeds into flooded forests – when combined with flooding and its associated environmental heterogeneity – suggests that flooded forests will have a unique biology and ecology (Kalliola et al. 1991). Furthermore, it is expected that flooding creates specific tree species zonational distributions (Whitmann et al. 2010a), largely determined by the submergence tolerance of their seedlings (Parolin et al. 2004). Finally the predictability of the flood “pulse” (Junk et al. 2010) – both past and present – facilitates adaptation and thus, along with differences in the surrounding biota and a variety of soil types (Honorio 2006; Junk 1989), may help create complex and diverse forest associations throughout the Amazon Basin (Myster 2009).
1.2 The Western Amazon
Studies of tree endemism in the Amazon (ter Steege et al. 2006; Whitmann et al. 2013) show that there is a natural division between the Western Amazon rainforest (Myster 2009) and the Central Amazon rainforest (Junk et al. 2010), where the Japurá river joins the Amazon river (Figure 1.2). In this book, I will accept that evidence and define Western Amazonia as everything in the Amazon Basin west of latitude 65oW, including the Mamiraua Reserve in Tefé, Brazil.
Figure 1.2 Map of the Amazon river and some of the other rivers that drain into it, including the Japurá river, with an outline of the Amazon Basin.
This western part of the Amazon Basin is composed of young and relatively fertile sediment, which makes the flora of more recent origin than that of Eastern Amazonia (reviewed in Dumont et al. 1990; Terborgh and Andresen 1998). Furthermore, the higher fertility of these soils may lead to more forest turnover and evolution, resulting in the high biodiversity sampled here compared to Central Amazonia (Bierregaard et al. 2001; Gentry 1993). Indeed the Western Amazon rainforest contains some of the most diverse areas or “hotspots” on Earth (Myers et al. 2000; Myster 2007b) due, in part, because it is largely intact, pristine and unaffected by human activity (<5% of its total area is in secondary forest: Gorchov et al. 2004; Neff et al. 2006; Soares-Filho et al. 2006) and so suffers the smallest loss of biodiversity and forest fragmentation (Bierregaard et al. 2001) in all of the Amazon. The large biodiversity of the Western Amazon may also be a consequence of the large areas within it, which have been surrounded by only rainforest for millions of years, without border effects or edge effects. Because the Western Amazon rainforest has extensive variation in edaphic, climatic, topographic and geological conditions, a large number of distinct plant communities are expected (Myster 2009, 2012b). Indeed, sampling has shown that up to 59% of recorded species are found in only one plot among the many sampled (Duivenvoorden 1995), and shared species between wetland and upland forests is very low (Dumont et al. 1990).
All tropical rainforests are hot and humid with large biodiversity. However, in addition to the global and ecosystem issues raised before, the Amazon in general, and the Western Amazon in particular, are also unique because of their high level of complexity – the many ways that different components of the Amazon ecosystem interact – and often at a higher-order than other tropical forests. Some of those interactions are outlined above, and in coming chapters authors will discuss many more Amazonian interactions within their own area of interest. I also point to these recent studies that have highlighted important Amazon ecosystem interactions among:
soil type, land use and tree growth (Moran
et al
. 2000);
water availability and forest biodiversity (Paine
et al
. 2009);
deforestation, fire and drought (Davidson
et al
. 2010);
recruitment and scatter-hoarding (Russo
et al
. 2005);
recruitment and seed dispersal by fish (Anderson
et al
. 2009);
hunting of large primates, richness and density (Nunez-Iturri and Howe 2007);
soils, geography distances and floristics (Ruokolainen
et al
. 2007);
flooding duration and the availability of phosphorus, iron and aluminum (Chacon
et al
. 2008);
soils, herbivory and plant defenses (Fine
et al
. 2004);
forest responses, drought and the C cycle (Phillips
et al
. 2009; Saleska
et al
. 2007); and
climate change and flooding patterns (Langerwisch
et al
. 2012).
With this ecological background in mind, and as background to the chapters that follow, it may be useful to review the ideas and different approaches used in the past to investigate Neotropical forests. First, it has certainly been a positive development over the last few decades of Neotropical forest research that several permanent forest plots has been set up and sampled on a regular basis for a considerable time. These have included large – usually 25–50 ha – plots in terra firme forest (i.e. Barro Colorado Island in Panama, Yasuní National Park (YNP) in Ecuador, La Planada in Colombia, Luquillo Experimental Forest in Puerto Rico: Brokaw et al. 2012) and smaller plots/transects in various Neotropical forests (Balslev et al. 1987; Myster 2007b, 2010; Parolin et al. 2004; Worbes et al. 1992; author unpub. data). The data collected has advanced our understanding of the structure of these forests and how that structure changes through time (Brokaw et al. 2012; Junk et al. 2010; Losos and Leigh 2004). I applaud those efforts and hope they will continue. In addition to plot sampling, approaches have included tabulation of common, or otherwise defined important, species traits (ecophysiological: Parolin et al. 2004; Poorter 1999; phenology: Parolin et al. 2010; mycorrhizal association: Falster et al. 2015; Meyer et al. 2010; Myster et al. 2013), measurement and estimation of different aspects of the C cycle (Baker et al. 2007; Schongart et al. 2010; Townsend-Small et al. 2005) and other biogeochemical cycles (Calle-Rendon et al. 2011; Kern et al. 2010), remote sensing used to discover large-scale vegetation patterns and tree associations (Melack and Hess 2010; Peixoto et al. 2009) and seed/seedling mechanistic experiments (e.g. Kubitzki and Ziburski 1994; Paine and Beck 2007; Wittmann et al. 2010b).
While these plots and other research approaches have had success, field experimentation and modeling approaches tried in Neotropic forests over the last 30–40 years (Carson and Schnitzer 2008) continue to be a challenge, especially for those trying to discover what controls forest dynamics: in particular, those approaches that include hypotheses about static, unchanging patterns of biodiversity (Connell 1971; Janzen 1970), which then need to be “maintained” over time (McDade et al. 1994; Zimmerman et al. 2008), and co-existence of species (Dalling and John 2008; Mabberley 1992). Alternatively, the authors in this book will progress with better methods of investigation into forest structure, function and dynamics, which I will put together into a new and better synthesis of Neotropic rainforest investigation, in the concluding chapter.
1.2.1 Case study: Sabalillo Forest Reserve
I now review my own studies in the Western Amazon. My first study site was Sabalillo Forest Reserve (SFR: 3o20′3″S, 72o18′6″ W: Frederickson et. al
