Articles

Andean Influences on the

Biogeochemistry and Ecology
of the Amazon River
MICHAEL E. M C CLAIN AND ROBERT J. NAIMAN

Although mountains often constitute only a small fraction of river basin area, they can supply the bulk of transported materials and exert strong
regulatory controls on the ecological characteristics of river reaches and floodplains downstream. The Amazon River exemplifies this phenomenon.
Its muddy waters and its expansive and highly productive white-water floodplains are largely the products of forces originating in distant Andean
mountain ranges. The Amazons character has been shaped by these influences for more than 10 million years, and its present form and host of
diverse organisms are adapted to the annual and interannual cycles of Andean inputs. Although the Andes constitute only 13% of the Amazon River
basin, they are the predominant source of sediments and mineral nutrients to the rivers main stem, and Andean tributaries form productive
corridors extending across the vast Amazonian lowlands. Many of the Amazons most important fish species rely on the productivity of Andean
tributaries and main-stem floodplains, and annual fish migrations distribute Andean-dependent energy and nutrient resources to adjacent lowerproductivity aquatic systems. Mountain-lowland linkages are threatened, however, by expanding human activities in the Andean Amazon, with
consequences that are eventually felt thousands of kilometers away.
Keywords: Amazon, Andes, nutrient subsidies, land use, fisheries

he Amazon River exits the Andes mountains more

than 4000 kilometers (km) from its estuary, but along its
essentially flat and serpentine path through the lowlands of
northern Brazil it maintains the character of an Andean river
(figure 1). The indelible imprint of this distant mountain range
on the main-stem channel of the worlds largest river has been
noted by naturalists and researchers for more than a century,
but the multifaceted nature of Andean influences on the hydrology, biogeochemistry, and ecology of the river system have
only come to light during the past two decades. Other fundamental but still obscure linkages remain to be discovered.
Long before scientists took interest in the study of Amazon environmentsin fact, long before Europeans discovered the rivernative peoples of the lowland Amazon
recognized the unique characteristics of Andean tributaries.
Agriculture thrived on the fertile floodplains of these muddy
rivers and gave rise to some of the regions first and most successful chiefdoms (Meggars 1984). Native Amazonians also
capitalized on the rich fish stocks of Andean tributaries.
Alfred Russel Wallace (1853) was perhaps the first naturalist
to write about the white-water, clear-water, and black-water
river types of the Amazon basin and to relate the color of
tributaries to the nature of their drainage basins (figure 2).
Wallace astutely linked the sediment load of white-water
tributaries to erosion in their steep Andean headwaters, and
identified clear-water rivers with the crystalline mountains

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of Brazil (the Guyana and Brazilian shields). He knew that

black-water rivers emerged from lowland sources, and he
correctly attributed their dark coloring to leaching of decaying
leaves, roots, and other vegetable matter (Wallace 1853).
Another naturalist of that time, Henry Bates (1863), marveled
at the transport of volcanic pumice in the main-stem Amazon River and correctly assigned its origin to volcanic ranges
thousands of kilometers away in the Ecuadorian Andes. He
imagined these porous stones as vehicles transporting seeds
and insect eggs downstream and thereby dispersing organisms
far beyond their original ranges. Over the last 50 years,
systematic investigations have further advanced scientists
understanding of the environment and distinct aquatic ecosystems of the lowland Amazon River (summarized in Sioli
[1984], Junk [1997], and McClain et al. [2001]).
Steep terrain and young lithologies make the Andes an
important source of sediments and solutes to the lower
reaches of the Amazon River. The most visible characteristics
of the main-stem Amazon and its Andean tributaries are
high discharge and heavy loads of suspended and bedload
Michael E. McClain (e-mail: mcclainm@fiu.edu) is with the Department of
Environmental Studies at Florida International University in Miami, and
Robert J. Naiman (e-mail: naiman@u.washington.edu) is with the School of
Aquatic and Fishery Sciences at the University of Washington in Seattle. 2008
American Institute of Biological Sciences.

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Figure 1. Nine major rivers flow from the Andes to form fertile corridors across the lowland Amazon (shown in bold). The
Ucayali, Maraon, and Napo rivers drain southern Ecuador and northern and central Peru, converging to form the mainstem Amazonas, which becomes the Solimes River where it crosses into Brazil. The Caquet River flows from Colombia,
becomes the Japur upon entering Brazil, and merges with the main stem at about 65W (west). The Putumayo River
flows from Colombia and Ecuador to become the Ia in Brazil. The Madeira River collects the Andean tributaries flowing
from southern Peru and Bolivia and traverses thousands of kilometers of lowland Amazon rainforest before merging with
the main-stem Amazon at about 59W.
sediment. Associated with this sediment load are abundant
organic matter and nutrients. The ramifications of a high particulate load are also far-reaching in their geomorphological,
biogeochemical, and ecological effects on the lowland river corridor. Large sediment loads and flooding have created broad
floodplains, and associated nutrients support diverse and
productive floodplain forests, macrophyte beds, and lakes of
seasonal importance to the life cycles of organisms in the rivers
and adjoining uplands. In an important ecological feedback,
the products of floodplain primary production eventually return to the main-stem river in floodplain runoff, becoming
important energy sources for heterotrophic communities
living there (Richey et al. 1990, Melack and Forsberg 2001).
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Many fish also migrate annually into Andean tributaries

from low-fertility black-water and clear-water tributaries to
spawn and feed in resource-rich white-water channels and
floodplains. Upon their return, migrating fish transport organic matter and nutrients that subsidize the food webs of
black-water and clear-water rivers.
Many fundamental aspects of the geomorphology, biogeochemistry, and ecology of the main-stem Amazon are
therefore linked to the magnitude and variability of water and
materials supplied from the Andes. In fact, the dominant
downstream trend in biogeochemical and trophic characteristics of the main-stem Amazon and its large Andean tributaries is the progressive dilution of Andean contributions by
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lowland tributary inputs (Devol and Hedges 2001).
Even though we are beginning to understand the
dynamics of Andean-derived materials in the mainstem Amazon River corridor, and the degree to
which lowland ecosystems depend on upstream
inputs, we still know little about the nature and
variability of processes that mobilize these materials from the Andes and modify them during downstream transport and storage in the extensive
floodplains.
In this article, we briefly introduce the geomorphology and ecological zones of Andean headwater regions of the Amazon, as these are poorly
known even among scientists specializing in Amazon ecology. We then examine the multifaceted
ways in which the main-stem Amazon River is
influenced byand depends onAndean inputs.
We conclude by exploring frontiers in research linking Andean and lowland parts of the Amazon, considering the possible impacts of increasing
human-related development and climate change in
the Andean Amazon.

The Andean Amazon

The Andes mountains rise steeply along the western
margin of the Amazon basin and stand 3000 meters
above sea level (masl) in elevation over much of
their length (figure 1). Approximately half of the Andean Amazon lies at elevations between 500 and
2000 masl, while most of the remainder is between
2000 and 4000 masl; about 16% is above 4000 masl
(table 1). The highest point in the Amazon basin is
the Nevado de Huascaran in the Cordillera Blanca
of Peru, at 6768 masl, but several other peaks extend
above 6000 masl. Active volcanoes are prominent
features of the Ecuadorian and Bolivian Andes. The
eastern cordillera of the Altiplano, a high-elevation
endorheic basin containing Lake Titicaca, forms on
one of the widest sections of the Andes, spanning
nearly 300 km near the lake.
Characterization of the precipitation, soils, and
vegetation of the Andean Amazon is fundamental to
understanding Andean influences on the lower
Amazon River (figure 3). Precipitation is greatest on
the lower and mid slopes of the cordillera (500 to
3000 masl) because of orographic controls on air
masses coming from the east. The wettest parts of
the basin lie in the eastern cordillera of Colombia
and near the PeruBolivia border, where annual
Figure 2. The main rivers of the Amazon have long been classified according to the color of their waters, which also reflects
their source. (a) The Ia (Putumayo) River is a characteristic white-water river colored by the high loads of sediments
transported from the Andes. (b) The Negro River is the largest of the black-water rivers, tinted by high levels of dissolved
organic matter leached from low-lying areas of sandy soils. (c) The Rio Tapajos is the most notable of the clear-water rivers
carrying low levels of sediments and organic matter from the crystalline Guyana and Brazilian shields. Photographs:
Margi Moss (http://brasildasaguas.com.br).
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Table 1. Elevation ranges of the Andean Amazon.
Elevation (meters
above sea level)
5001000
10012000
20013000
30014000
40015000
> 5000
Total

Area
(square kilometers)
111,804
170,514
117,018
120,671
100,766
2444
623,217

Area
(percentage)
18
27
19
19
16
<1
100

Source: Compiled from Shuttle Radar Topography Mission

90-meter data.

precipitation may exceed 4 meters (figure 3a). The most

abundant soil order in the Andean Amazon is inceptisol
(61%), a young, mineral-rich soil that occurs at mid elevations.
More developed but less fertile ultisols occupy 16% of the region and occur mostly at lower elevations in Peru. Mollisols,
or grassland soils, are the third most abundant soil order, covering 6% of the region, primarily near the PeruEcuador
border and at higher elevations in southern Peru. Exposed rock
is common at very high elevations (greater than 4000 masl)
in southern Peru.

The major vegetative cover types in the Andean Amazonmapped using Advanced Very High Resolution Radiometer satellite imagery (Eva et al. 1998)are submontane
(700 to 2000 masl) and montane (2000 to 3700 masl) forests,
which together constitute approximately 42% of the region
(figure 3b, table 2). Montane herbaceous vegetation interspersed with shrubland and agriculture is also widespread, covering nearly a quarter of the region. As of 2000, at least 40%
of the region had been converted to human uses or fragmented by these uses (JRC 2000). The most intense human
alteration has historically been at high elevations (> 3000
masl), where high levels of alteration continue today; but
change is increasingly concentrated at mid and lower elevations as colonization continues and roads spread across the
region (Mena et al. 2006).
The modern Amazon River is born in numerous Andean
springs, but cartographers locate the most distant source of
the river at 5300 masl on the northern slope of Nevado
Mismi. From this stream, the Carhuasanta, the main stem of
the Amazon, changes names at least nine times: from Carhuasanta to Lloqueta, Hornillos, Apurimac, Ene, Tambo,
Ucayali, Amazonas, Solimes, and finally Amazon below the
confluence of the Solimes and Negro rivers. The entire
north-south length of the Andean Amazon basin is drained

Figure 3. (a) Areas of higher precipitation are focused on the lower slopes of the Andes, with maximal registered precipitation
in the headwaters of the Madre de Dios River in southwest Peru and the Napo River of central Ecuador. (b) Montane forests
dominate the land cover between 500 and 3000 meters above sea level and transition into natural high-elevation grasslands
above. Compiled from Shuttle Radar Topography Mission 90-meter data and Global Land Cover 2000 data (CJRC 2000).
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Table 2. Land cover of the Andean Amazon basin.
Land-cover class
Forest (includes areas
of fragmented forest)
Grassland and shrubland
(includes pasture)
Wetland
Cropland
Dryland
Water
Ice
Urban
Totals

Area
Area
(square kilometers) (percentage)
329,574

53

215,755

34

231
71,216
6375
1832
1031
394
626,408

<1
11
1
<1
<1
<1
100

Note: The difference in the total area reported in tables 1 and 2 is

due to grid size differences of the initial raster data sets.
Source: Compiled from Global Land Cover 2000 data (JRC 2000).

by eight major riversthe Caquet, Putumayo, Napo,

Maraon, Ucayali, Madre de Dios, Beni, and Mamor (figure
1).

Andean influences on the load

of the Amazon main stem

portional to the areal coverage of the Andes. Although annual

precipitation on the lower slopes of the Andes exceeds the
Amazon average, higher valleys of the Andes are more arid,
and thus the average precipitation for the entire range is not
likely to be greatly different from precipitation for the basin
as a whole. But while Andean contributions of water to the
main-stem Amazon may be proportional to area, contributions of sediments and solutes are disproportionately
greater. Moreover, energy and nutrients carried from the
Andes by the river appear to largely drive main-stem productivity, both directly and indirectly through biophysical
feedbacks with the massive lowland floodplain.

Inorganic sediments and solutes

Four decades ago, Ronald J. Gibbs wrote that the Andean
mountainous environment controls the geochemistry of the
Amazon River (Gibbs 1967). He had sampled the Amazon
main stem and 16 of its major tributaries and had compared
total particulate and solute concentration data for the wet and
dry seasons against nine environmental parameters. On the
basis of strong correlations with the environmental parameter mean relief, Gibbs concluded that the Andes were the
source of 82% of the total suspended solids exported by the
Amazon River. The importance of Andean sources of suspended sediment to the main-stem Amazon River was reaffirmed by the subsequent work of Robert Meade and others,
who concluded that between 90% and 95% of the suspended

The main-stem Amazon River integrates the flow of subbasins containing distinct combinations of geology, soils,
and vegetation. There are four major Andean tributaries to
the main-stem Amazon River: the Solimes, Ia,
Japur, and Madeira (figure 1). (Andean tributaries
to the main stem are defined as those with headwaters above 500 masl in the Andes mountains, assuming that the western limit of the main-stem
Amazon River is set as the BrazilColombia border.)
Where they intersect with the main stem, the combined mean annual flow of these white-water tributaries is approximately 90,000 cubic meters per
second: roughly half of the main-stem Amazon
Rivers mean annual discharge, or five times the
flow of the Mississippi River (Dunne et al. 1998).
The Andes cover only about 13% of the Amazon
basin upstream of bidos, and Andean tributaries
may flow through hundreds to thousands of kilometers of lowlands (below 500 masl) before connecting with the main stem.Yet most measurements
of Andean contributions to the main-stem Amazon have been made at the main-stem confluences
of the four Andean tributaries. Clearly these rivers
have accumulated water, particulates, and solutes
from the lowlands before reaching the main stem, Figure 4. The disproportionate loads of sediments carried by the main
and therefore one must be careful to consider what Andean tributaries are evident when comparing the inflows of (a) water
part of these loads actually derived from the Andes and (b) sediments to the main-stem Amazon river from its major triburather than from the lowlands. In the case of water, taries. Inputs at the top of each diagram represent the contributions of
we noted that the combined flow of the Andean trib- the Amazonas/Solimes River flowing from Peru. Data were compiled by
utaries amounts to approximately half of main- R. H. Meade from water-discharge data listed by Carvalho and da Cunha
stem flow, but the volume of water actually (1998) and from the sediment-discharge data of Dunne and colleagues
originating in the Andes is probably roughly pro- (1998).
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sediment load of the main stem derived from the Andean tributaries (figure 4; Meade 1984, Meade et al. 1985).
Returning to the question of how much of the water and
suspended particles carried by the Amazon River originate
from the Andes mountains, we speculated that less than a
quarter of the water originates in the Andes but that most
suspended sediments could originate in mountain areas.
Loads of suspended and bed sediments measured along the
entire length of the Madeira River, from its Andean headwaters
to its confluence with the main stem, show a sharp decrease
in sediment load (as much as 60%) at the base of the Andes,
a decrease in the mean diameter of suspended particles in the
piedmont region, and a progressive decrease in the mean
diameter of bed sediments (Guyot et al. 1999)all indicators
of a declining energetic capacity to transport materials. These
characteristics indicate that Andean rivers supply more than
enough sediment to account for the total load of sediments
in the lowland sections of the Andean tributaries. Conclusive
evidence of an Andean source is found in the mineralogical
and isotopic composition of the suspended sediments. The
mineral composition of sediments in the main-stem Amazon
correlates well with that of the Ucayali and Maraon rivers
in the Andes (Gibbs 1967). Measurements of neodymium,
strontium, and lead isotopic ratios reaffirm that Andean
sources account for an overwhelming proportion of the
main-stem sediment load (Allegre et al. 1996).
Andean-derived suspended sediments bring a large flux of
minerals into the main-stem Amazon River, but they also bring
other elements and materials. Andean tributaries deliver an
order of magnitude more particulate nitrogen (1170 megagrams [Mg] per year) and phosphorus (806 Mg per year) to
the main stem than their lowland counterparts (119 and 43
Mg per year, respectively; Richey and Victoria 1993). Most particulate nitrogen is likely to be organic, whereas phosphorus
is mainly phosphate strongly adsorbed to iron and aluminum
oxide surfaces (Berner and Rao 1994). The availability of
this phosphorus to main-stem organisms is not known, but
significant amounts of phosphorus are released from Amazon sediments upon entering the estuary and may be available to organisms on the floodplains (Melack and Forsberg
2001). The question of whether particulate nitrogen and
phosphorus actually derive from the Andes or from some
intermediate river section is tied to the origin of the fractions
with which they are associated. The tendency of phosphate
to adsorb to mineral surfaces links this nutrient to the Andean
sources of the mineral sediment, but the organic association
of nitrogen is tied to that of the particulate organic fraction,
which is less well understood.
Two features of the Andes enhance their importance to
the solute geochemistry of the Amazon River and to its ecological characteristics. First, the Andes contain the only significant deposits of evaporites and carbonates in the Amazon
basin (Stallard and Edmond 1983). High fluxes of Ca2+ (calcium), Mg2+ (magnesium), HCO3 (bicarbonate), and SO42
(sulfate) ions occur in rivers draining carbonate deposits,
and high fluxes of Na+ (sodium) and Cl (chloride) ions
330 BioScience April 2008 / Vol. 58 No. 4

occur in rivers draining evaporite deposits. Rivers draining

basins containing carbonates generally have total cation
charges of 450 to 3000 microequivalents (eq) per liter (L),
and rivers draining basins containing evaporites may have
total cation charges of greater than 70,000 eq per L near the
salt sources (Stallard and Edmond 1983). The rich mineral
content of Andean tributaries underpins the ecological
productivity of downstream reaches. Black-water and clearwater tributaries draining lowland portions of the basin, by
contrast, have total cation charges below 300 eq per L and
are characteristically considered to have low ecosystem-scale
productivity. The second distinguishing feature of the Andes
is the intensity of its weathering regime, which increases the
concentration of ions in solution. Among the Amazon tributaries that drain basins dominated by less-weatherable
silicate rocks, Andean rivers have consistently higher total
cation concentrations (Stallard and Edmond 1983).
Few data exist that would allow us to estimate the proportional contribution of major ion fluxes to the main stem
from the Andes. Robert Stallards work demonstrates that
solute concentrations are elevated in Andean rivers, but without measurements of discharge it is not possible to calculate
fluxes. Furthermore, one-time flux measurements are not
representative of annual or interannual contributions to the
main stem. Unfortunately, no suitable data exist for Colombian, Ecuadorian, or Peruvian Andean tributaries, and thus
no estimation can be made regarding the Andean contribution of major ions to flow in the Solimes River from these
countries. We may speculate, however, on the basis of the high
ion concentrations in Andean rivers, that the Andean contribution to the main-stem solute load is dominant, especially
for certain elements found preferentially in Andean lithologies. For the headwaters of the Madeira River in Bolivia, Andean fluxes can be estimated with some confidence, thanks
to a 10-year data set (Guyot et al. 1992). Over the period of
these data, the specific flux of total dissolved solids from Andean basins was 80 Mg per km2 per year, while the specific flux
from lowland Bolivian basins was 7 Mg per km2 per year. The
headwaters of the Madeira River contain few carbonate and
evaporite deposits in comparison with the headwaters of the
Solimes River in Peru. Thus it is likely that the Peruvian Andes contribute an even larger percentage of the major ions delivered to the main stem.

Organic matter
Andean-derived suspended sediments carry a significant
amount of organic matter, 90% of which is made up of particles less than 63 micrometers (m) in diameter (Richey et
al. 1990). Variations in the fluxes of fine particulate organic
carbon (FPOC; particles < 63 m) along the main stem correlate closely with variations in suspended sediment fluxes,
suggesting a close physical association. In fact, the vast majority of FPOC (> 90%) cannot be physically separated from
mineral material and is therefore probably physically bound
to it (Keil et al. 1997). This physical association has been
shown to reduce the rate of organic matter decomposition and
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enhance its preservation. Total organic carbon is approximately
1%, by mass, of suspended sediment in the main stem, constituting a flux of 5 to 14 teragrams (Tg) of carbon per year
to the Atlantic Ocean (Richey et al. 1990).
Measurements show that more than 90% of particulate organic carbon (POC; > 0.5 m) in the main-stem Amazon
River comes from Andean tributaries, but how much actually
originates in the Andes Mountains? POC behaves more or less
conservatively in the main stem, suggesting that it resists
decay and is derived from distant sources (Richey et al. 1990).
Just how refractory and how distant the sources are can be estimated from a suite of molecular, elemental, and isotopic techniques used to characterize the organic matter and to trace it
back to its sources (Hedges et al. 1986, 2000, Aufdenkampe
et al. 2007). Concentrations of total lignin-derived phenols,
carbon-to-nitrogen ratios, and stable carbon isotope ratios
point to terrestrial plants, and more specifically the leaves of
terrestrial plants, as the main source of main-stem organic
matter. Algae and aquatic plants, so abundant on the extensive Amazonian floodplain, are important sources of labile
organic matter, fueling microbial metabolism in the main
stem, but do not persist in the system (Richey et al. 1990). The
depletion of carbohydrates and the increasing abundances
of nonprotein amino acids and diagnostic lignin-derived
phenols confirm that the organic matter is highly degraded,
especially the FPOC fraction. Moreover, these characteristic
signatures extend up the Madeira and Solimes rivers and into
the Andean foothills (Hedges et al. 2000, Aufdenkampe et
al. 2007). Richey and colleagues (2002) estimated that the
main-stem Amazon River transports only 7% of the organic
matter supplied to the river basinwide, supporting the finding that it also transports the most degraded and recalcitrant
materials.
The isotopic data, however, provide the most definitive
information on the age and general source area of particulate
organic matter in the main stem and its Andean tributaries.
For main-stem FPOC to have a true Andean source, much of
it would have to be hundreds to thousands of years old. This
is because little main-stem FPOC (and little of the fine sediment with which it is associated) is transported directly
from the Andes; most is stored for varying periods of time in
point-bar and floodplain sediments (Dunne et al. 1998).
FPOC does, in fact, have the lowest levels of bomb carbon14 (14C) of any organic matter fraction in the main-stem
Amazon (+19 14C per thousand []), suggesting an average turnover time of hundreds of years (Hedges et al. 1986).
Allowing for the dilution of the bomb 14C signal by younger
organic matter, this implies that a significant portion of
main-stem FPOM may be Andean.
The actual proportion of FPOC of Andean origin has been
approximated using delta carbon-13 (13C) stable isotopic
ratios as a fingerprint of its origin. The 13C of plant leaves
is positively correlated with elevation, and ratios in the
Peruvian Andes have been found to range from about 30
at 1000 to 2000 masl to 26 at 4000 masl (TownsendSmall et al. 2005, 2007). The values of leaves from prominent
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floodplain and upland forest trees along the main-stem river

also average 30, indicating that there is no clear isotopic
separation of leaf 13C between lowland forests and Andean
forests below 2000 masl of elevation (approximately 50% of
the Andean Amazon area; table 1). Unlike plant leaves, however, there is a clear separation of FPOC 13C between Andean
and lowland rivers, and this separation can be used to estimate
the relative proportion of each in the main stem. FPOC in
purely lowland rivers has 13C values consistently near 28.5
(Quay et al. 1992). The 13C of FPOC discharged in the
main-stem Amazon River at bidos is 27.4 and thus
indicates a mixture of the Andean and lowland sources. If the
Peruvian value for 13C of FPOC exiting the Andes (approximately 26.5) is taken as the Andean end member and
28.5 is taken as the lowland end member, FPOC at bidos
is a mixture of 50% Andean FPOC and 50% lowland FPOC.
Alternatively, if the Bolivian end member of 25.5 is used,
FPOC at bidos is a mixture of 33% Andean and 67%
lowland FPOC (Quay et al. 1992, Hedges et al. 2000).
Interestingly, the 13C of FPOC in each of the major
Andean tributaries (the Solimes and Madeira rivers) where
they meet the main stem is 26.8. This suggests that these
rivers carry FPOC that is largely of Andean origin and account
for 82% of the FPOC input to the main stem. If only 30% to
50% of FPOC entering the Atlantic Ocean is of Andean origin, then there is a 50% to 70% reduction in Andean-derived
FPOC in the main-stem section of the river. This reduction
probably occurs through sediment exchange with the floodplain and gradual decomposition of Andean organic matter
while in storage. Recent research using a dual-isotope approach
(14C and 13C) estimated the degree of mineralization of
Andean-derived FPOC with transport downstream and
concluded that nearly all Andean FPOC was mineralized in
the river and floodplain system (Mayorga et al. 2005). Taken
together, the Andes largely regulate the particulate load to
the main-stem Amazon River, not simply with respect to its
particulate mineral load but also with respect to associated
nutrients and organic matter.
The four major Andean tributaries contribute approximately 50% of the dissolved organic matter (DOM) input to
the main stem (Richey et al. 1990), but unlike particulate
organic matter, this DOM appears to derive largely from
lowland sources. Neither mass-balance nor chemical-tracer
approaches support important Andean contributions of
DOM to the lowland or main-stem Amazon. DOM accumulates in swampy environments that are common throughout the lowland Amazon, and in rivers and streams that drain
areas of spodosol soils (McClain and Richey 1996). In the
central Brazilian Amazon, fluxes of DOM to groundwater in
the spodosols characteristic of the Rio Negro subbasin are
approximately 20 times greater than those in the oxisols
characteristic of much of the rest of the lowland Amazon
(McClain et al. 1997). In the Rio Negro basin, high groundwater DOM concentrations (approximately 3000 micromoles
of carbon) also appear in surface water draining spodosols,
whereas in oxisol terrains, fringing wetlands appear to be
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important sources of DOM. DOM concentrations are uniformly low in the few studies on Andean rivers (Guyot and
Wasson 1994, Hedges et al. 2000, Saunders et al. 2006). In the
Madeira subbasin, there is a distinct increase in DOC concentrations in rivers below 500 masl, and this additional
DOC appears to derive from floodplains and wetlands such
as those of the Bolivian Llanos de Mojos (Guyot and Wasson
1994).

Andean influences on the productivity

of the main-stem Amazon
The productivity of the main-stem Amazon is tied to the
productivity of its floodplain, a system built of Andeanderived materials and fueled by mineral nutrients from the
Andes (Melack and Forsberg 2001). Over a 2010-km reach of
the Amazon main stem, the mean lateral flux of sediments
(1570 to 2070 Tg per year) between the channel and adjoining floodplain exceeds the downstream flux (1200 Tg per
year), and approximately 500 Tg per year of upstream-derived
sediment and associated nutrients accumulate on the floodplain and in channel bars (Dunne et al. 1998). This process
builds the fertile floodplain soils along Andean tributaries and
the main stem. By contrast, floodplains along non-Andean,
lowland tributaries are far more depleted in mineral nutrients.
The Amazon River maintains year-round lateral exchanges
with its floodplain, and especially with its abundant lakes.
The floodplain is a highly productive system, with an estimated
regional net production of 113 Tg of carbon per year occurring over an area of 67,900 km2, from the BrazilianColombian
border to near the rivers mouth (figure 5; Melack and Forsberg 2001). This translates to 17 Mg carbon per hectare per
year, which exceeds the productivity of upland Amazon
forests by a factor of five; in fact, the Amazonian floodplain
is among the most productive ecosystems on Earth. The

majority of primary productivity is attributed to macrophyte

(65%) and floodplain forest (28%) communities. Subtracting estimates of carbon loss to respiration and burial, about
90 Tg carbon per year are available for export to the mainstem river, where the additional carbon fuels respiration
(Melack and Forsberg 2001, Mayorga et al. 2005).
A portion of the supply of Andean nutrients to the floodplain can eventually be traced back into the main stem not only
as labile organic matter but as part of myriad organisms that
move between the floodplain and channel. Large numbers of
fish move onto the floodplain annually to exploit its productivity and utilize its habitats (Goulding 1993). In fact,
annual movements onto the floodplains of Andean-influenced
white-water rivers are the most common form of migration
among Amazon fishes and are critical to maintaining the
regions fisheries (Goulding et al. 1997). Of the 24 species in
the Brazilian Amazon that are most important to humans (in
nutritional and economic terms), most migrate as part of their
life cycle, and most rely to some extent on the resources
delivered from the Andes (Araujo-Lima and Ruffino 2004).
One of the most sought-after fish is the tambaqui (Colossoma
macropomum). This omnivorous/frugivorous fish occurs
over the length of white-water rivers but only in the lower
reaches of black-water rivers. It feeds in flooded forests
during high water and migrates back into the channel
during low water. Tambaqui, like many other species, spawns
along the margin of white-water rivers, and the larvae are
washed onto floodplains by the rising waters. There they
feed and seek shelter beneath the ubiquitous macrophyte
beds (Araujo-Lima and Goulding 1997). A number of other
characids important to Amazon fisheries (Brycon spp.,
Mylossoma spp., Myleus spp.) also follow this migration
pattern (Araujo-Lima and Ruffino 2004), using the floodplain
for feeding and nursery habitats and for transporting

Figure 5. Nutrients and mineral substrates carried by Andean tributaries and deposited on floodplains fuel the highest
primary productivity rates per hectare in the Amazon basin. This schematic illustrates the balance of organic carbon on
the main-stem Amazon floodplain between 70.5W (west) and 52.5W (refer to figure 1 for extent). This balance indicates
that large quantities (approximately 90 teragrams) of organic matter are returned to the river channel annually to fuel
in-channel respiration. All quantities are for total organic carbon unless otherwise noted. Source: Melack and Forsberg
(2001) and Richey and colleagues (1990). Abbreviations: DOC, dissolved organic carbon; POC, particulate organic carbon.
332 BioScience April 2008 / Vol. 58 No. 4

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resources back to the river as they migrate. Isotopic tracers
have shown that C3 macrophytes, floodplain trees, and
phytoplankton account for 82% to 97% of the carbon in 35
species of adult fishes examined (Forsberg et al. 1993).
Phytoplankton, while accounting for a small proportion of
the total primary productivity on floodplains, represents the
primary source of carbon to characiform fishes (AraujoLima et al. 1986).
Migrations are also important in distributing the enhanced
productivity of Andean-influenced white-water rivers and their
floodplains to less productive black-water and clear-water
environments. Many Amazon fish migrate from black-water
and clear-water rivers to the main stem and other whitewater rivers to spawn. In fact, all commercially important
species appear to spawn only in white waters (Goulding et al.
1997). During times of the year other than the spawning
season, some move back into black-water and clear-water
environments, and in the event of predation or death, the organic matter and nutrients of their bodies serve as subsidies
to these less productive ecosystems. Jaraqui (Semaprochilodus
spp.) is an example of a fish that migrates from black-water
rivers into white-water rivers to spawn (figure 6a). These
predictable migration routes are stalked by larger predators
that congregate at the confluences of black-water and whitewater rivers, such as the Amazon River dolphin, or boto (Inia
geoffrensis).
Many other fish use the main stem and its Andean tributaries as migration corridors, most notably large predatory catfish (Pimelodidae) moving upriver to Andean spawning
areas. Catfish making long-distance migrations are quantitatively the most important predators in the river system, and
they are also the most important species to fisheries along the
rivers length (Barthem and Goulding 1997). The most remarkable of these migrations is that of the dorado, or dourada,
catfish (Brachyplatystoma spp.; figure 6b), which travels as far
as 5000 km in one direction (Goulding et al. 2003). Statistical data on size classes along the entire length of the Amazon
River reveal that dorado spawn in headwater regions (including Andean foothills) and that the young are washed
downstream to nursery areas in the Amazon estuary (Barthem
and Goulding 1997). Preadult dorado move upriver again,
completing the approximately 8000-km migration over several years. Dorado and a number of other migrating catfish
are heavily fished along the river, so their numbers are significantly reduced by the time they reach the rivers of the piedmont and Andean foothills.
In Andean piedmont regions, characins emerge as the
most important fishery species in biomass; the most important among these is Prochilodus nigricans, known as
boquichico in Peru. Boquichico is a fine-particle feeder that ingests detritus and algae, and has a maximum length of less than
40 centimeters. During the low-water season, it lives in floodplain lakes and channels of the Amazon piedmont, but at the
initiation of rising water it leaves the floodplain and migrates
en masse up Andean tributaries to spawn (Diaz-Sarmiento and
Alvarez-Len 2004). Collectively, the fish migrations illustrate
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the critical connections between the Andes and downstream

biotic communities and ecological processes, as well as the importance of maintaining both lateral and longitudinal connectivity throughout the Amazon.
Enormous sediment loads, fluxes of nutrients and refractory organic matter, and ultimately the fertility of the expansive floodplains reflect the many influences of distant
Andean mountain ranges on the main-stem Amazon and
other white-water tributaries (figure 7). The rivers character
has been shaped by these materials for more than 10 million
years, and its present form and host of diverse organisms
are adapted to the annual and interannual cycles of Andean
inputs. It is safe to say that the ecology of the modern Amazon main stem has been built on substrates and nutrients de-

Figure 6. Migrations of many Amazon fish are strongly influenced by the pursuit of resources and habitats tied to
Andean tributaries. (a) The jaraqui (Semaprochilodus
insignis) is an example of species that, as adults, live
mostly in black-water rivers or lakes, but migrate to
white-water rivers to spawn. Juvenile jaraqui also use
white-water floodplains as their nurseries. (b) The
dourada (Portuguese) or dorado (Spanish) catfish
(Brachyplatystoma spp.; B. rousseauxii in photo) are the
farthest-migrating species known in the Amazon. They
hatch in the Andean foothills, use the Amazon estuary as
their nursery, and then migrate thousands of kilometers
up Andean tributaries to spawn. Photographs: Michael
Goulding.
April 2008 / Vol. 58 No. 4 BioScience 333

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rived from the Andes, and that the decoupling of the mainstem Amazon from its mountain headwaters would lead to
dramatic changes in the rivera pattern reflected in many of
the worlds other great rivers.

Andean processes regulating fluxes

to lowlands: A research frontier
The Andes exert strong influences on the main-stem Amazon,
and these influences strengthen as one travels upstream along
the major Andean tributaries. But what processes regulate the
fluxes of Andean derived materials, and how do these processes
vary spatially and temporally in the Andean Amazon? Unfortunately, little research to date addresses these questions,
and obtaining regional numbers is exceedingly difficult.
Nevertheless, current rates of land-use change in the Andean Amazon are among the highest in the Amazon basin;
40% or more of the region already has been significantly
fragmented and otherwise affected by human alterations
(Eva et al. 1998). How will land-use change and possible flow
regulation alter fluxes of particulates and solutes to the lowland Amazon, and what other forms of contamination might
be emitted by growing mountain populations? Research

addressing these human-related questions is still relatively restricted spatially in the Andean Amazon, but such research is
essential for the coming decade if effective regional agreements
are to be forged about the future of the Amazon basin.
Concerning sediment fluxes, it is important to note that instantaneous loads in lowland rivers are largely decoupled
from those in mountain rivers. Where lowland Andean tributaries remain white with high sediment loads year-round,
mountain rivers are generally clear during the dry season
and white only during storm-runoff events (Townsend-Small
et al. 2008). Their sediment fluxes may fluctuate greatly on
daily or weekly timescales in response to individual storm and
landslide events (Guyot et al. 1999), whereas lowland river
fluxes, like their hydrographs, fluctuate according to dampened seasonal cycles. Meandering lowland rivers maintain their
sediment loads by continually resuspending and depositing
materials within their channels (Meade et al. 1985, Dunne et
al. 1998), effectively mining sediments accumulated in the
piedmont over long timescales through discrete depositional
events (Aalto et al. 2003). To understand mountain-lowland
linkages, one therefore needs to consider erosional processes
over a broad range of timescales.

Figure 7. Andean influences on the ecology and biogeochemistry of the Amazon may be grouped into three interacting sets of
processes. Andean exports of water, sediment, nutrients, and organic and biological material exert fundamental control and
produce the white-water characteristics of Andean tributaries and the mainsteam Amazon itself. Floodplain building by
these Andean-derived materials provides the substrate and nutrition fueling productive flooplain forests, macrophyte beds,
and lakes. Fish migrate throughout these systems and along tributaries, capitalizing on the productivity of white-water
river systems and transferring a small quantity of Andean-derived energy and nutrients to nutrient-poor black-water
and clear-water systems.
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At timescales stretching into millions of years, and at the
spatial scale of the entire mountain range, climate seems to
exert a fundamental control on erosion processes in the
Andean Amazon. Montgomery and colleagues (2001) analyzed
the topographic, climatic, and tectonic variability of the entire Andes cordillera and concluded that morphology is more
closely related to climate than to tectonic processes.
Erosion from the mountain range over the past 25 million
years has come predominantly from the northern Amazon
Andes (north of 15 south), where historical rates of erosion
are up to twice as high as in the drier southern portion of the
Amazon Andes (southern Peru and Bolivia). Linked to this
long-term erosional history, a striking and relevant geomorphological characteristic of the high Andes is a shift from
steep-sided, V-shaped valleys to gently sloped, U-shaped
valleys between 3000 and 3500 masl. Although much reduced in size today, glaciers have been important in shaping
high Andean valleys. Moreover, the gentle valley slopes exposed
by glacial retreat result in reduced physical erosion in the
highest portions of the Andes.
At subregional spatial scales and shorter timescales, vegetation may assume a first-order control of erosion rates.
Erosion rates in the Beni and Mamor river basins of Bolivia
range from 521 to 6000 metric tons per km2 per year and from
310 to 2600 metric tons per km2 per year, respectively (Guyot
et al. 1988). Topography, lithology, rainfall, and vegetation all
play roles in explaining differences in erosion between basins,
but vegetation plays the dominant role. Rates of erosion are
greatest in the southernmost basins, where vegetation is
sparse. In the north, where rainfall is greater but subbasins are
heavily forested, erosion rates are considerably lower.
The controlling influence of vegetation on erosion at both
subregional and hillslope scales is significant because land-use
change is the most prolific form of anthropogenic disturbance
in the Amazon (figure 8). Erosion is less intense in densely vegetated parts of the Andes, despite high rainfall on erosionprone slopes. The stabilizing effects of natural vegetation are
lost, however, following deforestation, and land management
practices become important variables in explaining fluxes of
sediments, organic matter, and nutrients from newly created
agricultural fields and pastures. Studies conducted in midelevation (2000 to 2500 masl) valleys of the Peruvian Amazon find increased fluxes of sediments, organic matter, and
nutrients in rivers draining valleys with greater proportions
of agriculture and pastures (Waggoner 2006). Similar trends
have been observed in the Napo River basin of Ecuador,
where clear correlations were found between overall river
health and the level of anthropogenic alterations (Celi 2005).
Continued investigations of land-use impacts on stream and
river sediment loads are one of the most pressing research
needs in the Andean Amazon today. Studies of land-use
impacts on rivers and streams should emphasize riparian
zones, both because they are control points for land-to-river
material transfers (Naiman and Dcamps 1997, Naiman et al.
2005) and because they are favored for agriculture in the
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Andean Amazon as a result of the relative fertility of their soils

(McClain and Cossio 2003).
It was recognized early on that concentrations of major ions
and trace elements in Andean Amazon rivers were linked to
the lithologies of the major subbasins, and subsequent work
has supported this link (Sobieraj et al. 2002). The most focused
impacts that humans have on major ions and trace-element
fluxes from the Andes is through mining, which is widespread at higher elevations. Contamination of soils and vegetation by heavy metals has been documented near mines and
downstream of mining operations (Hudson-Edwards et al.
2001). Accumulations of metals in river invertebrates have even
been measured downstream of the point at which contamination of bottom sediments is no longer detectable (Bervoets
et al. 1998). Mercury contamination from placer gold-mining
operations is a significant concern in many Amazonian areas,
and mercury accumulations in fish and in the hair of riverine people have been linked to gold-mining operations as far
as 150 km upstream in the upper Beni subbasin of Bolivia
(Maurice-Bourgain et al. 1999). Although of considerable
local concern, the current impacts from mining appear to be
limited to river reaches immediately downstream of mining
sites. Expansion of mining activities, however, may eventually
lead to significant changes in the fluxes of heavy and trace
metals to adjoining Amazon lowlands. Quantifying the
composition, magnitude, and ecological consequences of
increased heavy metal fluxes is an important need in the Andean Amazon.
The dependence of lowland river corridors on sediments
and nutrients derived from the Andes requires unobstructed
connectivity between the two regions. No major Andean
tributary to the Amazon is currently dammed, although
Brazil is pursuing plans to build two major dams on the
Madeira River. Hydroelectric installations are common, however, on streams and small rivers close to major mining operations, to urban areas, or to other significant human
settlements. Peru has five significant hydroelectric projects
under way in its Amazon region, and the Peruvian Ministry
of Energy and Mines has identified dozens more potential dam
sites, some on prominent rivers such as the Maraon, Huallaga, Tambo, and Urubamba. Dams trap large volumes of sediment, and could cause major readjustments over the long
term in the geomorphology of downstream river sections
and the eventual sediment starvation of some downstream
reaches. The ill effects of dams on river organisms and riparian
environments are well known (e.g., Dudgeon et al. 2006)
and could be especially destructive in the Andean Amazon,
where biodiversity is high and many fish species migrate annually between mountains and the lowland rivers and floodplains. Far too little is known at this point about the extent
to which riverine organisms and riparian environments rely
on open linkages between mountains and adjacent lowlands
in the western Amazon. It is therefore impossible to predict
what the short- and long-term consequences of widespread
dam building would be. We suspect, on the basis of evidence
presented here and evidence from other regions with
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Figure 8. The Oxapampa Valley in central Peru illustrates a number of the forces threatening the ecological health
of Andean and downstream river reaches, including the deforestation and cultivation of steep slopes and the urban
development of narrow valley bottoms. Future damming of valleys such as this could significantly affect downstream
fluxes of sediments and nutrients. Photograph courtesy of Thomas Saunders.

numerous dams, that eventually the consequences would be

severe, as they have been for other rivers (e.g., the Columbia
River in the United States).
A wild card in all discussions of future scenarios in the
Andean Amazon is the effect of climate change, including the
feedbacks between land use and climate. There is already
strong spatial variability in todays Andean climate, due to the
areas topographic complexity. Even though the response of
Andean environments to El Nio/La Nia events is complicated, the trend is toward heavier than normal rainfall (Kane
2000), resulting in increased landslide intensity. This may
not be the case, however, in the future. Rainfall in the Andean
Amazon is sensitive to the water balance of the lowland Amazon, and this balance is expected to change in predictable ways.
Because rain in the Andean Amazon is ultimately derived from
the Atlantic Ocean, it must be transported across the lowland
Amazon basin in westward-moving air masses. During this
westward movement, moisture cycles between the atmosphere and land surface, and estimations are that roughly
55% of the rain falling in the Amazon basin is derived from
evapotranspiration within the basin (Marengo and Nobre
2001). For the eastern slopes of the Andes, the percentage of
rainfall derived from evapotranspiration is probably higher.
336 BioScience April 2008 / Vol. 58 No. 4

Consequently, continued deforestation should lead to reduced levels of precipitation in the Andean Amazon (Chagnon
and Bras 2005).
Both elevated carbon dioxide (CO2) and the conversion of
forest to managed uses are predicted to reduce evapotranspiration and thus the amount of water moving westward
toward the Andes. Elevated CO2 alone is predicted to reduce
evapotranspiration in the Amazon by about 4% through reductions in stomatal conductance, and this should also reduce
rainfall. Conversion of forest to pasture across the entire
Amazon basin is predicted to reduce evapotranspiration by
as much as 20% (Lean et al. 1996). These changes in the
regional water balance will certainly affect terrestrial and
aquatic ecosystems of the Andean Amazon and thereby fundamentally alter the mountain-to-lowland fluxes discussed
here. As investigations of these questions proceed at a basin
scale, and as confidence in predicted changes increases, Andean policymakers should carefully examine local impacts.
The Amazon River system is unique in many ways
because of its size and orientation along the equator, but the
controls by its Andean headwaters are not unique. In fact,
many of the mountain-lowland linkages we have discussed
should be relevant to other major river systems. Similar
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controls are certainly observed in the adjoining Orinoco
River system (Edmond et al. 1996, Jepson and Winemiller
2007) and are likely to be important in the major rivers
draining the Himalayas, namely the Indus, Ganges, Brahmaputra, and Mekong. The fundamental ecological importance
of these linkages stresses the need to manage even the worlds
largest rivers in a basin context.
Although our knowledge of the nature and magnitude of
mountain-lowland linkages in the Amazon basin can serve to
inform research and management in the Amazon and in
basins around the world, much remains to be learned.
Research in recent decades has illuminated the nature and
magnitude of mountain-lowland linkages along the mainstem Amazon river, but investigations in the Andes lag far
behind. Researchers still know little about the fluxes of sediments and associated nutrients from the Andes on a regional scale, and even less about the spatial and temporal
variability in those fluxes. We know equally little about the degree to which river organisms depend on habitat and other
resources of Andean rivers during annual and multiyear migrations. In the midst of our incomplete ecological knowledge,
the Andes are being rapidly transformed into a managed
landscape where rivers are modified and where montane
forests and high-altitude grasslands are converted to pastures and agricultural fields. Filling these knowledge gaps is
an immediate scientific challenge with important ramifications for the sustainability of the Amazon River basin as a
whole. Brazil, the downstream beneficiary of Andean inputs
from its upstream neighbors, should take special interest in
these issues. Over the long term, the most productive components of the Brazilian Amazon River system are also the
most vulnerable to poor management decisions in the Andes.
Brazils own plans for large-scale hydroelectric development,
new road building, and agricultural intensification should pay
similar consideration to the important hydrological and ecological linkages uniting the larger basin.

Acknowledgments
We wish to acknowledge our colleagues and collaborators in
the Andean Amazon who have informed and influenced our
understanding of Andean-Amazon linkages, especially Jay
Brandes, Remigio Galarraga, Michael Goulding, Jean Loup
Guyot, Carlos Llerena, Jos Efrain Ruiz, Richard Chase Smith,
and Amy Townsend-Small. We thank the Inter-American
Institute for Global Change Research, the US National Science
Foundation, and the Andrew W. Mellon Foundation for supporting our research in the Amazon basin. Daniel Gann and
Anna Boyette provided critical support with graphics. Michael
Goulding, Margi Moss, and Thomas Saunders contributed
photos. This manuscript was improved by the comments of
John Melack and three anonymous reviewers.

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