Kvist&Nebel - 2001 - Peruvian Flood Plain Forests

Forest Ecology and Management 150 (2001) 326
A review of Peruvian ood plain forests: ecosystems,

inhabitants and resource use
Lars Peter Kvist, Gustav Nebel*
Department of Economics and Natural Resources, The Royal Veterinary and Agricultural University,
Unit of Forestry, Rolighedsvej 23, 1958 Frederiksberg C, Denmark
Abstract
The lowland Peruvian Amazon remains sparsely populated and densely forested. Few roads exist and rivers provide much of
the infrastructure. Over 12% of the area is comprised of ood plains inundated by the larger rivers, but due to their easy access
and relatively fertile conditions, they provide a much larger share of the resources extracted in the region, and sustain most of
the rural villages. The largest area of annually ooded land, constituting more than 60,000 km2, surrounds the lower reaches of
the Ucayali and Maranon rivers above their conuence to the Amazon proper, including almost 90% of the 20,600 km2 large
Pacaya-Samiria National Reserve. The entire area is constantly reshaped by erosion and deposition by the two main rivers, but
also by smaller rivers which carry fewer sediments and less nutrients. The vegetation constitutes a complex mosaic of habitats
dened by combinations of hydrological, physical, chemical and biological characteristics. Sixteen habitats including 12
forest formations are classied and described. Most ood plain inhabitants have lost their native identity, but they descend
mostly from Amerindians rather than from recent immigrants, and preserve much knowledge on the ood plain environment
and its habitats and uses. They combine agriculture, shery, hunting and extraction of other forest products, and they market
increasing amounts of these products. Growing populations and an increase in the need for monetary incomes as well as in
external economic interventions, increasingly endanger economically important ood plain plants, animals and shes. This is
even the case within the Pacaya-Samiria National Reserve, making it urgent to develop and implement sound management
systems in the marginal zones of the reserve, and elsewhere in Peruvian ood plains. # 2001 Elsevier Science B.V. All rights
reserved.
Keywords: Wetlands; Amazon; Natural resources; Land-use; Vegetation classication; Pacaya-Samiria National Reserve
1. Introduction
On a global scale wetlands cover some 200530
million ha corresponding to approximately 3% of the
land surface, and about 60% is covered with forest
vegetation by which most is under the inuence of
fresh water. The tropics contains considerably more
wetlands than the temperate and boreal regions, and a
*
Corresponding author.
E-mail address: gne@kvl.dk (G. Nebel).
larger part is forested (Lugo et al., 1990a). Many

wetlands provide economically important services
for shery, agriculture and forestry (e.g. Welcomme,
1985; Bayley, 1989; Bayley and Petrere, 1989; RosTonen, 1993; de Jong, 1995), and at the same time
constitute important habitats for plant and animal life
(e.g. Soini et al., 1996; Junk and da Silva, 1997; Junk
et al., 1997; Petermann, 1997), generally stressing the
need for multiple-purpose management adapted to the
ecosystems and the people interacting with these.
According to Junk (1993) and Irion et al. (1997) the
main rivers of the Amazon basin have fringing ood
0378-1127/01/$ see front matter # 2001 Elsevier Science B.V. All rights reserved.
PII: S 0 3 7 8 - 1 1 2 7 ( 0 0 ) 0 0 6 7 9 - 4
L.P. Kvist, G. Nebel / Forest Ecology and Management 150 (2001) 326
plains covering 300,000 km2. In Peru more than

60,000 km2 of alluvial lands are exposed to annual
ooding by sediment rich rivers originating in the
Andes, and in Brazil a similar or even larger area is
ooded by the sediment rich Amazon proper (Salo and
Rasanen, 1989; Junk, 1997).
The productivity of ood plains inundated by turbid
water is enhanced by the nutrients deposited during
the annual ooding (e.g. Furch, 1997), while long or
irregular inundation periods on the other hand may
reduce plant growth (e.g. Gill, 1970; Junk, 1989; Irion
et al., 1997). Many Amazonian ood plains are attractive to human settlement and activities due to their
relatively fertile conditions combined with predictable
monomodal ooding patterns and easy access, while
the more extensive terra rme areas tend to have
leached soils unsuitable for permanent cultivation,
and are located away from the main rivers making
transport difcult. Consequently, when the Europeans
arrived, ood plains were more densely populated than
terra rme (Meggers, 1971; Denevan, 1976; Hemming, 1987; Roosevelt, 1989; Grenand and Grenand,
1993; Moran, 1993), and the rural population in the
Peruvian Amazon still depends much upon the ood
plain resources (e.g. Hiraoka, 1985a; Padoch, 1988; de
Jong, 1995).
In Peru, many ood plain inhabitants preserve an
intimate knowledge of the ood plain environment
and its resources (e.g. Hiraoka, 1985a; Padoch, 1988).
For instance, they manage crops and trees in systems
adapted to the environment (de Jong, 1995, 2001; de
Jong et al., 2001) and they have selected varieties
performing well in the ood plains (Srensen et al.,
1997). Their knowledge may be used to sustain the
development of adaptive ood plain management
systems in Peru and elsewhere. This applies in particular to the Cocama-Cocamilla and the ShipiboConibo, which are the two largest native ethnic groups
found at Peruvian ood plains. However, the nonindigenous majority of rural ood plain inhabitants,
locally known as riberenos, also preserves much traditional knowledge, reecting that they descend from
indigenous peoples (Padoch, 1988; Lonzoy, 1992; San
Roman, 1994).
Depletion of valuable ood plain resources due to
over-extraction and loss of habitats as a consequence
of changed land-uses are the major treats for populations of ood plain plants and animals. Increasing
amounts of game, sh or valuable timber are commercialised at the markets of the regional towns. Most
of the Peruvian ood plain forests have been subject to
extraction for subsistence as well as commerce,
including large-scale timber logging. Consequently,
the populations of certain animal and plant species
providing commercially valuable products have been
reduced (Tables 1 and 2, respectively). According to
the character of the use this may put a risk on the
genetic diversity of these populations (e.g. FAO,
1993a; Namkoong et al., 1996). The extraction of
some ood plain resources is likely to increase further
as a raising rural population aspiring for better living
standards than previous generations live at or close to
the ood plains. One of their best economic options at
the moment appears to be an intensication of the
extraction of resources from ood plain forests.
Contrasting to the current small-scale and diversied family-based cultivation, a more intensive and
wide-spread agricultural production may be an alternative option for better-off inhabitants and extern
interests with access to production factors. On a global
scale much ood plain forest has already been converted to such agricultural land-uses (Lugo et al.,
1990b), and in the Brazilian Amazon extensive areas
have been cleared along the middle Amazon both
upstream and downstream from the town of Manaus
(Goulding et al., 1996), mainly for grazing of cattle
and water buffaloes (Ohly, 1985). IIAP (1997) called
for an intensied agricultural production in the ood
plains of the Peruvian Loreto Department due to their
superior edaphic conditions as compared to terra
rme.
Both excessive extraction of selected valuable ood
plain forest species and wide-spread forest conversion
to other land-uses may be problematic, as ood plain
forest ecosystems are both ecologically (Welcomme,
1985; Goulding et al., 1996; Junk, 1997) and socioeconomically important (Anderson, 1990; Peters,
1990; de Jong, 1995; Kvist et al., 2001a). Depletion
of certain species can put a risk on their genetic
diversity and socio-economic contribution, and
large-scale conversion of forest ecosystems may have
large and unpredictable consequences on the whole
ood plain environment. However, it is likely that
Peruvian ood plains increasingly become converted
to intensive agriculture, as they contain the most fertile
soils of the Amazon region (Furch, 1997; Junk, 1997).
Table 1
Important faunal resources in Peruvian ood plains in the lower Ro Ucayali and lower Ro Maranon regionsa
Scientific name
Local name
English name
Use/problem
Status
Fishes
Arapaima gigas
Osteoglossum bicirrhosum
Paiche
Arawana
Arapaima
Aruana
Food, pet
Food, pet
Depleted
Locally depleted
Reptiles
Melanosuchus niger
Caiman crocodylus
Geochelone denticulata
Podocnemis sextuberculata
Podocnemis unfilis
Podocnemis expansa
Lagarto negro
Lagarto blanca
Motelo
Cupiso
Taricaya
Charapa
Black caiman
Spectacled caiman
Big headed turtle
Giant Amazon river turtle
Hide, pest
Food, hide
Food
Food
Food
Food
Depleted
Locally depleted
Locally depleted
Locally depleted
Depleted
Endangered
Birds
Ara miacao
Crax mitu
Cairina moschata
Guacamayo rojo
Paujil
Sacha pato
Scarlet macaw
Razor-billed curassow
Muscovy duck
Food, pet
Food
Food
Depleted
Locally depleted
Locally depleted
Mammals
Myrmecophaga tridactyla
Priodontes maximus
Lagothrix lagothricha
Alouatta seniculus
Cacajoa calvus rubicundus
Pithecia monachus
Ateles belzebuth
Ateles paniscus
Cebus apella
Cebus albifrons
Felis pardalis
Panthera onca
Lutra longicaudis
Pteronura brasiliensis
Trichechus inunguis
Tapirus terrestris
Oso hormiguero
Yungunturo
Choro
Coto
Huapo colorado
Huapo negro
Maquisapa ceniza
Maquisapa negro
Mono negro
Mono blanco
Tigrillo
Otorongo
Nutria
Lobo del ro
Vaca marina
Sacha vaca
Giant anteater
Giant armadillo
Common woolly monkey
Red howler monkey
Red uacari monkey
Monk saki monkey
White-bellied spider monkey
Black spider monkey
Brown capuchin monkey
White-fronted capuchin monkey
Ocelot
Jaguar
Southern river otter
Giant otter
Manatee
Brazilian tapir
Food
Food
Food
Food
Food
Food
Food
Food
Food
Food
Pelt, pest
Pelt, pest
Pelt, food
Pelt
Food
Food
Depleted
Depleted
Depleted
Locally depleted
Endangered
Depleted
Endangered
Endangered
Locally depleted
Locally depleted
Locally depleted
Depleted
Locally depleted
Endangered
Endangered
Locally depleted
Species tabulated are those negatively inuenced by human activities. Based on Soini et al. (1996).
Although the forests by nature are adapted to large but

scattered perturbations (Foster et al., 1986; Foster,
1990a,b; Worbes et al., 1992; Salo and Kalliola,
1993; Worbes, 1997), the consequences of widespread conversions are uncertain and should be
avoided as far as possible. Therefore, land-use systems
must be developed which properly conserve the biological diversity of the ood plains and their related
socio-economic values. Yet, such systems demand a
profound understanding of the productive but complex
ood plain environment and the interactions between
the ecosystems and the people using their resources.
The present paper describes some physical and
biological aspects of the dynamic Peruvian ood plain
environment. Furthermore, an impression is given of

how people interact with the ood plain forests, and
patterns of the present use of the natural resources is
described. The work mostly draws upon experiences
from the Jenaro Herrera municipal district located
along the lower Ro Ucayali from the Puinahua channel located further upriver the Ro Ucayali and from
the Ro Maranon near its conuence with the Ro
Samiria tributary. These areas are all located within or
close to the 20,600 km2 large protected PacayaSamiria National Reserve, which is mainly constituted
by ooded forests located between the lower courses
of the Ucayali and Maranon rivers (Rodriguez et al.,
1995).
Table 2
Plant resources of the Peruvian ood plains in the lower Ro Ucayali and lower Ro Maranon regionsa
Scientific name
Family
Local name
Main use
Status
Aniba spp., Ocotea spp.

Brunfelsia grandiflora D. Don
Calophyllum brasiliensis Cambess.
Castilla ulei Warb.
Cedrela odorata L.
Ceiba pentandra (L.) Gaertn.
Euterpe precatoria Mart.
Ficus insipida Willd.
Mauritia flexuosa L. f.
Maytenus macrocarpa (Ruiz and Pav.) Briq.
Minquartia guianensis Aubl.
Phytelephas macrocarpa Ruiz and Pav.
Scheelea spp.
Smilax sp.
Swietenia macrophylla King
Uncaria spp.
Virola spp.
Lauraceae
Solanaceae
Clusiaceae
Moraceae
Meliaceae
Bombacaceae
Arecaceae
Moraceae
Arecaceae
Celastraceae
Olacaceae
Arecaceae
Arecaceae
Smilacaceae
Meliaceae
Rubiaceae
Myristicaceae
Moena
Chiric sanango
Alfaro, Lagarto caspi
Caucho
Cedro
Lupuna
Huasa
Oje
Aguaje
Chuchuhuasi
Huacapu
Yarina
Shapaja
Zarza
Aguano, Caoba
Una de gato
Cumala, Cumala caupuri,
Aguano cumala
Timber
Roots for medicine
Timber
Resin
Timber
Plywood
Palm heart
Resin
Fruit harvest
Cortex for medicine
Beams for construction
Leaves for roofing
Leaves for roofing
Roots for medicine
Timber
Cortex for medicine
Timber, plywood
Depleted
Locally depleted
Locally depleted
Depleted
Depleted
Depleted
Locally depleted
Locally depleted
Locally depleted
Locally depleted
Locally depleted
Locally depleted
Locally depleted
Depleted
Heavily depleted
Locally depleted
Depleted
a
Species used for subsistence purposes are principally locally depleted close to villages, while commercialised species are generally more
widely depleted.
2. Geography and climate
Pasco, Junin, Cusco and Puno Departments. Based

on studies of satellite images Salo and Kalliola (1993)
found that only 0.8% of the Peruvian tropical lowland
Amazon became deforested during 1015 years up to
1993. As also observed by Gentry and Lopez (1980)
and Dourojeanni (1990) the deforestation of the montane forests above 600 m elevation is much more
advanced.
An approximation of the Peruvian lowland Amazon
population is provided by the gures of the three major
departments, showing that only around 5% of the total
Peruvian population of 22,639,443 live in the region
(Table 3). The population density of 0.83.2 inhabitants/km2 was low compared with the national gure
Dourojeanni (1990) stated that approximately

960,000 km2 in Peru drains to the Amazon, by which
around 770,000 km2 belong to the Amazon region,
including 90,000 km2 tropical montane forests located
above 600 m elevation on the eastern slopes of the
Andes. Approximately 680,000 km2 are situated in the
lowlands below 600 m elevation, where the natural
vegetation is tropical lowland rain forests. The Loreto,
Ucayali, and Madre de Dios Departments cover
556,446 km2, corresponding to 82% of the total Peruvian lowland area (Table 3). The remaining area is
located in the Amazonas, San Martin, Huanuco,
Table 3
Geographic and demographic data for the Peruvian tropical lowland Departments of Loreto, Ucayali, and Madre de Diosa
Department
Loreto
Ucayali
Madre de Dios
a
Surface
(km2)
368852
102411
85183
556446
Based on INEI (1997).
Population
1972
1981
1993
Rural
population (%)
1993
409772
130030
25154
564956
516371
178135
35788
730294
736161
331824
69854
1137839
42
35
43
Population density
(habitat/km2)
1993
Population growth (%)

19721981
19811993
2.0
3.2
0.8
2.6
3.5
3.9
3.0
5.3
5.7
of 17.6 inhabitants/km2. Around 60% of the lowland

Amazon population comprise urban people concentrated in the few larger towns: Iquitos, Pucallpa,
Puerto Maldonado and Yurimaguas. The rural inhabitants are mostly settled along the main rivers where
the ood plains are located, implying that extensive
areas of the region remain almost uninhabited. However, the population of the lowland Amazon seems to
have expanded faster than the national averages
(Table 3).
It was estimated that 62,000 km2 in the Peruvian
lowland Amazon are exposed to annual inundation of
sediment rich rivers originating in the Andean mountains, corresponding to 12% of the region (Salo et al.,
1986; Salo and Rasanen, 1989). In addition, extensive
areas are ooded by water from local streams and
rivers carrying much less sediments, or by rainwater.
Adding these latter areas which receive few sediments,
ood plains may constitute up to 20% of the Peruvian
lowland Amazon region, corresponding to the gure

estimated for the entire South American tropical lowland by Junk (1993, 1997). Poorly drained areas are
also found in relatively plain topographical uplands far
from any rivers (Encarnacion, 1993), further increasing the percentage of temporarily or permanently
inundated areas in the Peruvian lowland Amazon.
The ood plains are extensive along the Amazon
proper and along the two largest Peruvian Amazon
tributaries: the Ro Ucayali and the Ro Maranon.
More than 60,000 km2 of ood plain forests, lakes
and swamps is located mostly south of the Maranon
river surrounding the lower reaches of both the
Ucayali and the Huallaga rivers, as well as the smaller
tributaries Ro Maquia, Ro Tapiche, Ro Pacaya, and
Ro Samiria (Rasanen et al., 1992, 1993). This extensive ooded area was designated as the PastazaMaranon sub-valley by Rasanen et al. (1987), and
as the Ucamara depression by Villarejo (1988). It is
Fig. 1. Climate at Jenaro Herrera as recorded on open land at Centro de Investigaciones de la Amazonia Peruana (CIJH). All gures are mean
monthly values calculated on basis of daily measurements. Data from the period 19701980 were used for calculation of all the gures, except
for precipitation values where data from the period 19651999 were used.
the largest of four inundation and deposition subvalleys recognised in the western Amazon basin
(Rasanen et al., 1987, 1990, 1993), the other three
being the Ucayali, Beni and Acre inundation and
deposition sub-valleys.
The mean annual temperatures in the Amazon basin
vary from 23 to 278C, and the variations through the
year are small, although polar air sometimes sweeps
the continent from the south and temperatures drop to
10158C. Precipitation varies from between 1500 mm
per year in the northern and southern parts of the basin
to more than 6000 mm per year on some Andean
slopes, although 20003000 mm is the average (Salati,
1985; Eden, 1990). In the Peruvian lowland Amazon
the northern parts are the most humid, while to the
south the climate gets drier and more seasonal (Kalliola and Puhakka, 1993).
At Jenaro Herrera the climate is typical humid,
lowland tropical (Fig. 1). In the period 19701980
the mean annual temperature was 25.98C with only
small uctuations over the year. During the same
period the monthly average of hours of sun uctuated
between 98 and 171. The mean annual evaporation
was 566 mm with a relatively constant monthly mean
of 47 mm. During the period 19651999 the mean
annual precipitation was 2715 mm, with mean
monthly precipitations ranging between 140 and
309.
3. Flooding patterns and water types
In wetland ecosystems the ooding pattern has
profound environmental impacts and is a factor distinguishing these areas from other ecosystems. The
oodings can take place on a more or less regular
basis, be of different amplitudes, and last for longer or
shorter periods. The inundation of ood plains of
larger rivers often follows a predictable monomodal
pattern with considerable amplitudes, whereas the
ood plains along smaller creeks and rivers as well
as in smaller depressions tend to be inundated by
rather unpredictable polymodal oods. Larger depressions and poorly drained areas may in contrast be
ooded on a regular basis with only small oods
correlating with the uctuations in the precipitation
(Junk, 1997). In view of the importance of the ooding
events, the ood plains should be seen as ecologically
indivisible from and closely connected to the river and

to their catchment area, according to the ood pulse
concept of Junk et al. (1989).
The Amazon river and many of its larger tributaries
are characterised by annual monomodal ood pulses.
Junk (1984, 1989) and Irion et al. (1997) studied the
ooding patterns at different Brazilian sites. Based on
data from Manaus covering a 90-year measurement
period they found a mean ood amplitude of 9.95 m
peaking around June and having the water level minimum around October. They stressed the ecological
implications of the relatively predictable ooding as
regards the time of maximum and minimum water
levels, as well as the fact that several consecutive years
may occur with high as well as low maximum or
minimum water levels.
The monthly average water levels observed through
daily measurements from September 1987 to February
1997 in Iquitos at the Amazon river and at the Ucayali
river close to Jenaro Herrera are shown in Fig. 2 and
maximum and minimum values recorded for each
month during the same period also appear. For both
sites the average peak period was in MarchMay and
the lowest water levels occurred in AugustOctober,
but in certain years maximum and minimum water
levels were recorded earlier or later. The peak level
reached at Jenaro Herrera was on the average higher
than at Iquitos, and it also occurred earlier, which may
be due to the upriver location of Jenaro Herrera as
compared to Iquitos. In both cases the peak of the
water level suggests a correlation with the amount of
local precipitation (Fig. 1), although the uctuations
of the river probably depend more on the rain patterns
at the more distant Andean catchment area. At the
water level peak we estimate that more than 90% of
the ood plain is inundated, while in the low water
season usually less than 10% remains covered with
permanent lakes, swamps and rivers. However, in
some years (e.g. in 1993 and 1994) almost the entire
ood plain inundates, when the water level rises 12 m
above its peak level in average years. Even the highest
ood plain sites normally inundate under such conditions, because they were deposited during previous
ooding peaks, implying that their height correspond
with the maximum water level. During such ooding
events crops and trees susceptible to ooding stress
may be damaged or die and terrestrial ood plain
animals may drown.
Fig. 2. Relative water level in the period from September 1987 to February 1997 in the Ro Ucayali at Jenaro Herrera, and in the Amazon river
at Iquitos. Monthly average with maximum and minimum recordings over the period. Water levels at the Amazon river at Iquitos were
measured daily by the harbour authorities (Marina de Guerra del Peru, Servicio de Hidrograa y Navegacion de la Amazonia), and at the
Ucayali river at Jenaro Herrera twice daily by the Centro de Investigaciones Jenaro Herrera (CIJH).
Gentry and Lopez (1980) postulated an increase in

annual maximum water levels at Iquitos in the period
19621978 as a consequence of deforestation in the
Andes mountains, but according to Nordin and Meade
(1982) their data were poorly interpreted. Fig. 3 shows

the annual maximum and minimum water levels
recorded at Iquitos in the period 19801997. In this
period we found no indication of an increased peak
Fig. 3. Annual maximum and minimum water stages in the Amazon river at Iquitos during the period January 1980 through October 1997.
Water levels were measured daily by the harbour authorities (Marina de Guerra del Peru, Servicio de Hidrograa y Navegacion de la
Amazonia).
10
ood level, although deforestation on the Andean

slopes continued from 1978.
Three main types of Amazonian rivers are distinguished according to their water colours (Sioli, 1968,
1984; Schmidt, 1972; Junk, 1983; Eden, 1990).
``White water'' rivers contain large amounts of suspended sediments as well as a considerable concentration of nutrients, and their reaction is close to
neutral. ``Black water'' rivers are coloured by slowly
degradable phenolic substances, are very nutrient
poor, and their reaction is acid. ``Clear water'' rivers
neither contain signicant concentrations of suspended sediments nor organic matters, and are consequently more transparent than both white and black
rivers, while their reaction may vary considerably.
In addition, some authors recognise ``mixed water''
rivers as a separate category (Encarnacion, 1985,
1993). This term denes black or clear water mixed
with white water.
Clear water rivers and streams come from geologically old and highly leached bedrock areas. They are
relatively rare in Peru while some major Brazilian
tributaries are of the clear water type, e.g. the Ro
Xingu and the Ro Tapajos originating in central
Brazil (Eden, 1990). Black water rivers and streams
are common in Peru, but are not among the largest
rivers. They originate from sandy areas subject to
podzolization, and from poorly drained swampy areas
(Encarnacion, 1993). The Ro Pacaya and the Ro
Samiria, which give the name to the Pacaya-Samiria
National Reserve, and which respectively ow
into the Ro Ucayali and the Ro Maranon, are of
the black water type. The same goes for the Nanay
river discharging into the Amazon close to Iquitos
(Encarnacion, 1993). In contrast, the landscape in the
Peruvian Amazon is dominated by large white water
rivers. The Amazon proper belongs to this category,
and from the north to the south the large Amazonian
tributaries Putumayo, Napo, Pastaza, Maranon,
Huallaga, Ucayali, and Madre de Dios. These rivers
all originate in the Andes mountains, and carry sediments eroded from their slopes, as well as dissolved
alluvium from their lowland river basins (Eden, 1990).
However, the rivers having their entire watersheds in
the lowlands of the Peruvian Amazon are also of the
white water type (Rasanen, 1993), since they mostly
drain geologically young and easily erodable landscapes consisting of sediments deposited by river
aggradation (Rasanen et al., 1987, 1993). This contrasts with the ancient and highly leached landscapes
drained by the clear water rivers of central Brazil and
Guyana.
The mixed category of water is important in the
Peruvian Amazon. Many rivers receive both clear,
black and white water from different streams in their
watershed, and may in fact consist of mixed water
although they in practice are referred to the colour
category with which their water corresponds best. The
term mixed water is instead applied for rivers where
the type of water change over the year, either seasonally or irregularly. For instance many local streams
carry black water during the dry season and white
water during the more rainy months of the year,
reecting that the increasing current during the latter
period also makes them more erosive (Encarnacion,
1993; Rasanen, 1993).
At Jenaro Herrera this change was observed in the
annually ooded forest of Lobillo located at the conuence of the local black-water stream Aucayacu and
the ood plain channel of Lobillo connecting with the
Ucayali river. Local rain storms increase the discharge
from the Aucayacu channel causing ooding of the
Lobillo forest by black water. In contrast, the Lobillo
forest oods with white water when the water level in
the huge Ro Ucayali rises, and considerable amounts
of sediments are deposited.
Within the Pacaya-Samiria National Reserve waters
both mix and alternate in a much larger scale. Most
lakes and swamps in the reserve receive black water
from local rivers and streams, except in periods succeeding rising water levels in the surrounding Ucayali
and Maranon rivers. In these periods white water may
pour into the Reserve for days or weeks, especially
from the Ucayali river, entering through narrow channels and through the usually black water Pacaya river.
The current in the Pacaya river may even change
direction temporarily, and carry white or mixed water
upriver rather than black water down river (Rasanen
et al., 1993). The extensive lakes, swamps and channels within the Pacaya-Samiria National Reserve are
interconnected with both the Ucayali and the Maranon
rivers. Fishermen in canoes pass across the reserve
between the two main rivers, and water may ow
between its numerous water-bodies, pending on uctuations in the two main rivers as well as in local rivers
and streams.
4. Fluvial processes and landscape patterns

Fluvial processes continuously affect the ood plain
landscape on different scales in time and space, which
profoundly inuences the environmental conditions
and creates distinct landscape patterns. The extension
of the ood plain ecosystems involved can be difcult
to determine, but Junk et al. (1989) dened them as
areas subject to periodic inundation, resulting in
adaptations and development of characteristic community structures. The meander belt comprises areas
subject to active river migration along the river channel, whereas ood basins are located further away
from the river channel, and tend to have a topographically lower, poorly drained, and in humid climates
often swampy terrain. Water from the river inundates
the meander belt, while the ood basin is also inundated by water from small local rivers and rainfall
(Reineck and Singh, 1980; Kalliola et al., 1991a;
Kalliola and Puhakka, 1993; Junk, 1997). Most of
the landscape located in the south and eastern part of
the Pastaza-Maranon sub-valley (the Ucamara depression) is shaped by the Ucayali river, and several of the
smaller tributaries follow its relic meander belts, e.g.
the Tapiche, Pacaya, and Samiria rivers (Dumont and
Garca, 1992). The present meander belt of the lower
Ucayali river is 1020 km wide (Kalliola et al., 1992).
In the Jenaro Herrera region it is bordered by nonooded uplands to the west, and extensive ood basins
to the east (Dumont et al., 1990), but upriver from
Requena its meander belt is surrounded by ood
basins to both sides.
The rivers are the driving force in the uvial
dynamic processes and in the Peruvian lowland Amazon the river channels take forms as meandering,
anastomosing and braiding. The Ro Ucayali is an
example of a strongly meandering river distinguished
by a single river channel with a high sinuosity. In
addition to being moderately meandering the Maranon
river is partly anastomosing and even braiding, which
is characterised by several river channels and high and
low sinuosity, respectively. Meandering, which is
probably the most common form of large rivers in
the Peruvian Amazon, is favoured by low slopes,
cohesive bank material, a high ratio of suspended load
to bed load, and a relatively steady discharge (Reineck
and Singh, 1980; Puhakka et al., 1992; Rasanen et al.,
1992; Kalliola and Puhakka, 1993).
11
Erosion and sedimentation are basic processes of

river dynamics. In meandering rivers the erosion predominantly occur at the outer side of meanders,
resulting in lateral river migration. Sudden and dramatic shifts in the course occur when a river channel
migrates. This process known as river avulsion may be
caused by the gradual building up of the terrain in the
meander belt due to sedimentation (Reineck and
Singh, 1980; Rasanen et al., 1992). The effect of
erosion is illustrated by the conditions at the Ucayali
river. South of Requena the river channel migrates up
to 160 m each year (Kalliola et al., 1992), while the
erosion is slower in the Jenaro Herrera region (de
Jong, 1995). River avulsions occur suddenly, for
example half-way on the Ucayali river between
Pucallpa and the conuence with the Maranon river,
where more than 50 km of the channel relocated up to
80 km some 200 years ago (Parssinen et al., 1996).
Likewise, the Puinahua channel probably represent a
recent river avulsion, and still has a narrow and
topographically low meander belt surrounded by
extensive swamps (Kalliola et al., 1991a). In the latter
case much water continues to ow through the ancient
channel of the Ucayali river.
Sedimentation takes place in different places on the
ood plain, which for practical purposes can be
divided into three major groups of deposits: (1) channel deposits, (2) bank deposits, and (3) ood basin
deposits (Reineck and Singh, 1980; Kalliola and
Puhakka, 1993).
Channel deposits are of special interest in meandering rivers, where they create point bars forming
sequences of ridges in meander scrolls. The point bar
deposits develop apically at and parallel with the inner
sides of the meanders as long and narrow formations.
They have proportions according to the river size, and
are characterised by a sequence of coarse material in
the bottom and ner material upwards (Reineck and
Singh, 1980). After inundations point bars emerge
temporarily as beaches, which during a few exposed
months are colonised, initially by annual weeds. At
each inundation the sedimentation raises the height of
the point bars, eventually transforming them to ridges,
which are stabilised by a perennial vegetation of
grasses, shrubs and small trees. The ridges are separated by narrow depressions known as swales, which
are lled up by ner sediments (Reineck and Singh,
1980; Puhakka et al., 1993). In the Ucayali river the
12
ridges are approximately 6 m high and extend 24 km.

They are separated by swales being 25 m lower than
the ridges (Rasanen et al., 1992). In meandering river
systems channel ll deposits also ll up river channels
abandoned due to cut-off processes or avulsions. They
are rapidly sealed near the active river channel, but the
sedimentation in cut-off lakes is a slow process (Reineck and Singh, 1980).
Bank deposits sediment as levees or crevasse splays
on the river banks. Levees are the topographically
highest points in the ood plains, principally located at
the outer side of the meanders or along straight courses
of the river, forming large and plane surfaces. Crevasse splays are formed by sediments depositing along
channels cut through natural levees or banks, especially at the outer side of meanders. In both cases
sediments are deposited at high water levels when the
velocity of the ooding water decreases, as the river
banks are overown. Levees can block the draining of
extensive areas located behind them, resulting in
forest dead and the formation of swamps or swampy
forest types (Reineck and Singh, 1980; Kalliola and
Puhakka, 1993). In the Peruvian lowland Amazon the
highest terrain formations in the ood plains probably
derive from bank deposits.
Flood basin deposits refer to the accumulation of
sediments passing the river banks and depositing
further away from the river course, especially in the
ood basin areas. The sediments are few and nely
grained, since the coarser fractions tend to deposit
closer to the river. Water from local rivers and streams
also enter ood basins, but they also tend to carry few
ne sediments. However, an annual sedimentation of
several ton of ne sediments per hectare has been
reported from black water inundated areas (Walker,
1995), and over time the deposited sediments gradually mask the relief of the ood basins. Flood basins
are therefore modied more slowly than the meander
belts.
The uvial dynamics processes described above
form complex landscape patterns in ood plains and
they act on different scale in time. The various processes act differently in time and space, resulting in a
patchy landscape with different inter-mixed habitats.
Salo and Rasanen (1989) classied the geomorphological processes which inuence the ood plain landscape patterns and its biotic responses. Sedimentation
and the resulting formation of uvial bars, meander
scrolls, levees, crevasse splays and ood channels are

considered to act on a scale of 1100 years, and
supposed to have extensions ranging up to 100 ha.
Over longer periods of 10010,000 years channel
avulsions, drainage reversals, changes in channel patterns and formation of terraces are supposed to inuence larger areas. Formation of basins take place in
periods up to millions of years and cover vast areas.
The biological responses to these processes result in
the formation of different vegetation communities
differentiated according to succession stage and environmental conditions. For example, the present meander belt forests may have established recently as
colonising vegetation at newly deposited levees or
meander scrolls, while mature meander belt forests
may be rare or absent, corresponding to the situation
described by various authors (e.g. Foster et al., 1986;
Foster, 1990a; Gentry and Terborgh, 1990; Worbes
et al., 1992; Kalliola and Puhakka, 1993; Worbes,
1997). At the Ucayali river the site-turnover rate,
which is the average annual percentage of the meander
belt eroding away, was estimated to approximately
0.3% (Kalliola et al., 1992). On the Maranon river the
corresponding value is probably of the same size. For
example, in 1996 the river eroded more than 500 m of
land at its conuence with the Ro Samiria, forcing the
people in San Miguel to relocate their houses repeatedly. Site-turnover rates of the sizes mentioned suggest that large parts of the meander belt vegetation in
the Pacaya-Samiria region erode away within a few
hundred years. Therefore, older meander belt forests
are probably rare or absent.
5. Vegetation
An efcient extraction as well as a proper management demand overview as regards the distribution and
availability of resources. Therefore, it is necessary to
separate different sites according to the composition
and structure of the vegetation, although it implies a
number of difculties (e.g. Greig-Smith, 1983). This is
illustrated by the distinctions between vegetation
types applied by people living from natural resources.
Amazonian ood plains are by no means an exception,
as different habitats of varying sizes, and many transitional zones appear in an inter-mixed manner (e.g.
Worbes et al., 1992; Puhakka and Kalliola, 1993;
Tuomisto, 1993).
According to Klinge et al. (1990) it is useful in the

classication of ood plain vegetation to consider: (1)
hydrological criteria, (2) physical and chemical criteria, and (3) biological criteria. For each of these sets
of criteria several indicators can be identied.
Among the hydrological criteria especially aspects
of the inundation pattern with emphasis on ooding
duration has been referred to as a decisive factor in
determining the vegetation composition and structure
for wetland forests in general (e.g. Bacon, 1990; Lugo
et al., 1990b; Junk, 1993), as well as for Amazonian
ood plain forests (Worbes, 1983, 1986, 1997; Junk,
1989, 1997; Klinge et al., 1990; Worbes et al., 1992;
Kalliola and Puhakka, 1993; Junk and Piedade, 1997).
Studies show that the ooding tolerance for trees differ
with species as well as with the size of individuals, and
that different plant physiological and anatomical adaptations to ooding exist (Gill, 1970; Crawford, 1982;
Hook, 1984; Kozlowski, 1984; Junk, 1989; Schluter
et al., 1993; Armstrong et al., 1994; Worbes, 1997).
Furthermore, reproductive patterns may be adapted to
the ooded environment (Gottsberger, 1978; Goulding, 1980, 1983; Ziburski, 1991; Ayres, 1995;
Kubitzki and Ziburski, 1994; Worbes, 1997). However, rare events such as several consecutive years with
high minimum ooding levels may inuence the
development of the vegetation more strongly than
the average ooding pattern (Junk, 1989; Irion et al.,
1997; Junk and Piedade, 1997). Indicators as amplitude, frequency, and duration of ooding can therefore
help to classify ood plain vegetation.
The type of water inundating the ood plains
strongly inuences the chemical characteristics of
their soils (Furch, 1997), and the erosion and deposition of the river shape the landscape (Salo and Rasanen, 1989). Thus, physical and chemical indicators
related to soil characteristics, water type, and landscape form may serve to classify ood plain vegetation
(Prance, 1979, 1980; Encarnacion, 1985, 1993; Pires
and Prance, 1985; Kubitzki, 1989).
Among the biological criteria the succession stage
of the vegetation may be useful to distinguish ood
plain vegetation. Several authors found that the oristic composition and structure of ood plain forests
was correlated with this factor (Salo et al., 1986;
Foster et al., 1986; Foster, 1990a,b; Worbes et al.,
1992; Worbes, 1997). Furthermore, characteristic
plant families and species may characterise various
13
ood plain forest formations (Prance, 1979, 1980;

Pires and Prance, 1985; Campbell et al., 1986; Balslev
et al., 1987; Worbes et al., 1992; Freitas, 1996;
Worbes, 1997; Nebel et al., 2001a,b).
Based on experiences from Brazilian forests Prance
(1979, 1980) provided the most widely cited classication of ooded and swampy Amazon forest vegetation. He based his terminology on the regularity of
ooding and the types of inundating waters, while the
succession stage of the vegetation was not considered.
His main distinction was between varzea, which is
ooded by white water, and igapo, which is ooded by
black or clear water. Varzea in Brazil may according to
Prance (1979) approximate the forests named
tahuampa in Peru. The igapo forests as found in Brazil
may hardly exist in Peru, because the rivers and soils
rarely are as nutrient-poor in Peru as in much of the
Brazilian Amazon.
Several authors described the vegetation of Amazonian ood plain forests outside Peru (Black et al.,
1950; Takeuchi, 1962; Braga, 1979; Keel and Prance,
1979; Worbes, 1983, 1986, 1997; Pires and Prance,
1985; Campbell et al., 1986; Balslev et al., 1987;
Colonnello, 1990; Klinge et al., 1990; Worbes et al.,
1992; Ayres, 1995; Junk, 1993; Macedo and Anderson, 1993). Most of these studies were conducted in
the central Amazon near Manaus. Junk and Piedade
(1993, 1997) described the herbaceous vegetation of
the region.
As regards ood plain forest vegetation in the
Peruvian Amazon Tuomisto (1993) compared existing
classications. The most cited works are those of
Malleux (1971, 1975 cited in Tuomisto, 1993) and
Encarnacion (1985); the latter recently updated by
Encarnacion (1993). The focus of Malleux was the
timber potential of the various forest types, and his
classication was based on aerial photos. Encarnacion
(1993) instead based his classication on extensive
eld work, and the vegetation types and terminology
recognised and applied by local people. In addition,
several other workers provided valuable information
that may help to dene and describe types of Peruvian
ood plain vegetation, especially Kalliola et al.
(1991a) concerning swampy vegetation, Seidenschwarz (1986) as well as Kalliola et al. (1991b) as
regards young colonising vegetation, and Kahn and
Mejia (1990) concerning forests dominated by palms.
Mature vegetation as well as aspects of succession of
14
ood plain vegetation were described by several

authors (Salo et al., 1986; Gentry, 1988; Foster,
1990a,b; Gentry and Ortiz, 1993; Puhakka and Kalliola, 1993; Puhakka et al., 1993; Freitas, 1996; Nebel
et al., 2001a). Based on studies of maps Lopez and
Freitas (1990) surveyed and categorised forests in the
Jenaro Herrera region, to some extent applying the
classication of Encarnacion (1985). Lamotte (1990)
described the relations between uvial dynamics and
forest vegetation on an island in the Jenaro Herrera
region.
In Table 4 we classify some major habitats at
Peruvian ood plains exemplied by the areas surrounding the lower Maranon and Ucayali rivers, corresponding to the Pastaza-Maranon sub-valley (the
Ucamara depression). The classication is based on
the work of Encarnacion (1985, 1993), and consistency with the terminology used by local people is
attempted. The indicators used in this connection are
inundation period, characteristics of the inundating
water, soil drainage, geographical location according to the main river course, vegetation structure,
vegetation succession stage, and presence of character

plants. These indicators represent the three sets of
criteria mentioned above, and help to identify 16
vegetation formations. It should be noted that distinction between young and later succession stages is
restricted to the meander belt. Here perturbations of
the vegetation by river dynamics processes occur more
often than in ood basin areas, making it necessary to
distinguish young vegetation stages. In addition, the
local population extracts more resources from meander belt habitats (see Table 6), and hence apply a more
detailed terminology to these formations, compared to
the more distant and usually less fertile ood basins.
The limit between vegetation formations is gradual
rather than clear-cut, and much vegetation may have
intermediate characteristic.
Restinga denotes the upper parts of the ood plains,
which only ood in some years or for a shorter period.
We nd that the restinga forests on the average are
ooded up to 3 months each year, and consider it
convenient to distinguish between high and low restinga, being located on sites inundating on an annual
Table 4
Key to ood plain forest vegetation formations at the lower Ro Ucayali and lower Ro Maranon regions
A. Woody vegetation
B. Good drainage
C. Riverine mixed forest, white to mixed water flooded (meander belt)
D. Flooded annually 01 month
Later succession stage (away from river)
Young succession stage (near river)
D. Flooded annually 13 months
D. Flooded annually 36 months
C. Non-riverine mixed forest, mixed to black water flooded (flood basin)
Flooded annually 01 month
Flooded annually 13 months
Flooded annually 36 months
B. Poor drainage to permanently flooded
Mixed broadleaved forest on riverine sites (meander belt)
Mixed broadleaved forest on non-riverine sites (flood basin)
Dominant tree Pseudobombax munguba
Dominant palm Mauritia flexuosa
A. Non-woody vegetation
Permanent swampy and floating vegetation
At least periodically with open water
Annual weedy vegetation on clayey riverine sites
Annual weedy vegetation on sandy riverine sites
1. High restinga
2. Young restinga
3. Low restinga
2. Young restinga
4. Tahuampa
5. Young tahuampa
6. High flood basin restinga
7. Low flood basin restinga
8. Flood basin tahuampa
9. Bajial
10. Flood basin bajial
11. Pungal
12. Aguajal
13.
14.
15.
16.
Pantanal
Lake and river
Barrial
Playa
average of up to 1 month and between 1 and 3 months,

respectively. Restinga forests are high-stature, close
and diverse, and ve categories are distinguished (nos.
1, 2, 3, 6 and 7 in Table 4). High restinga forests (nos. 1
and 6) tend to be located on relatively large and plane
surfaces, since they typically have developed from
bank deposits. In contrast, low restinga (nos. 3 and 7)
often grow on land created by channel deposits shaped
as typical meander scrolls composed of long and
narrow ridges interrupted by swales (known as
bajiales, no. 9). The most recently deposited restingas
in the meander belt are designated as young (no. 2),
and certain valuable pioneer species such as Calycophyllum spruceanum (Bentham) Hooker f. ex Schumann and Ficus insipida Willd. are typically
prevalent. In older succession stages of restinga forests
(nos. 1, 3, 6 and 7), which are characterised by gapphase dynamics (Foster et al., 1986; Worbes et al.,
1992; Worbes, 1997) these species are often absent or
only present with large individuals. Young restinga
forests tend to be relatively similar despite differences
in the length of the inundation periods. They gradually
establish on new land created in uvial processes, and
considerable amounts of bank deposits cause a built up
of the terrain level, implying that the trees must be able
to sustain this deposition. Due to river migration most
forests in the meander belt are young, while forests on
ood basin restingas (nos. 6 and 7), which are located
far from the present courses of the main river, may be
thousands of years old. In the Pacaya-Samiria region
most ood basin restingas were deposited by former
courses of the Ucayali and Maranon rivers. These
areas may be characterised as relic meander belts.
Tahuampa forests (nos. 4, 5 and 8 in Table 4) are
located lower in the terrain than their restinga counterparts. They are ooded for longer periods lasting from
3 to 6 months on an annual average, and the soils are
usually well-drained. The tahuampa forests are oristically distinct from restinga forests, but are also closed,
mixed and high-stature. The presence or absence of
early succession species distinguishes the meander belt
mature tahuampa forests (no. 4) from recently established young tahuampa forests (no. 5). Flood basin
tahuampa forests (no. 8) are located away from the river
channel, and are more nutrient-poor and older than
tahuampa forests in the meander belt (nos. 4 and 5).
Extensive parts of the ood plain environment
covered with woody vegetation are poorly drained
15
(nos. 9, 10, 11 and 12 in Table 4), implying that they

often are waterlogged outside the annual inundation
period. Much fewer tree species thrive in irregularly or
permanently waterlogged areas (Junk, 1989) than in
areas characterised by regular although often extended
annual inundations, i.e. restinga and tahuampa. The
forests of poorly drained areas are often stunted and
species-poor with many large herbs and vines in the
understorey due to light-open conditions. These
swampy forests may be composed by a mix of canopy
species (nos. 9 and 10) or be dominated by a single tree
species (nos. 11 and 12). Bajial forests (no. 9) are
typically located close to the main rivers in the meander belt ooded by white water. The understorey is
often very dense and almost impassable due to dense
populations of large herbs (Calathea, Heliconia and
Ischnosiphon) and many vines, lianas and fallen trees.
Bajiales may be long and narrow depressions (swales)
in scroll-complexes or constitute extended areas cut
off from the rivers by natural levees blocking the
drainage. Flood basin bajial forests (no. 10) have a
oristically mixed canopy and occur in ood basin
areas where they ood with mixed or black water from
the rivers as well as by rainwater. Therefore, the soil
conditions are typically more nutrient-poor and acid
than in bajial forests (no. 9). Aguajal forests (no. 12)
are given their name from the aguaje palm Mauritia
exuosa L. f. which dominates extensive water-logged
areas, especially in the ood basins. They are ooded
by water from mixed or black water rivers, and by
rainwater. In many aguajales the palms Euterpe precatoria Mart. and Oenocarpus mapora H. Karst. are
also major constituents. Pungal forests (no. 11) derive
their name from the dominating punga tree Pseudobombax munguba (C. Martius and Zuccarini) Dugand
belonging to the Bombacaceae family. It often dominates swampy forests, but tend to grow in more
nutrient-rich places than the aguaje palm, in particular
in the interface between other forest types and swamps
surrounding ood plain lakes and rivers, corresponding to pantanal and lake and river (nos. 13 and 14 in
Table 4). Most lakes arise in former river beds of cutoff meanders, and swamp vegetation with rooted
rather than oating vegetation develops as sediments
are deposited. Thereupon the punga tree may invade
the area, resulting in formation of a pungal, which may
later on in the succession be replaced by either
tahuampa, bajial or aguajal, depending on the local
16
conditions. Barrial and playa (nos. 15 and 16 in

Table 4) are located along the main river courses
and result from deposition of clay and sand, respectively. The successions taking place on these sites
result in restinga, tahuampa or bajial forests.
6. Population: history, culture, and demography
The rst Spanish and Portuguese explorers reported
that the ood plains along the Amazon river were
densely settled with numerous and large Amerindian
villages organised in chiefdoms stretching along hundreds of kilometres of river (Meggers, 1971; Denevan,
1976; Moran, 1989; Roosevelt, 1989). However, epidemics introduced from Europe as well as European
slave raiding soon reduced the Amerindian populations, and the survivors were settled at missionary
stations where various ethnic groups often became
mixed (Hemming, 1987; Parker, 1989).
In Brazil the last Amerindian ood plain cultures
lost their native tongue more than two centuries ago,
and their descendants have since then spoken Portuguese (Ross, 1978; Hemming, 1987; Grenand and
Grenand, 1993). They are now known as caboclos,
but still preserve much Amerindian cultural heritage,
especially concerning the management of natural
resources in the ood plain environment (Frechione
et al., 1989; Parker, 1989).
The majority of the present population living in
Peruvian ood plains speaks Spanish, and are designated as riberenos (``riverbank people''). Like the
caboclos in Brazil they descend from Amerindian
people, which to a wide extent have mixed with
immigrants and therefore also are considered as mestizos. Soon after the arrival of the Europeans the
Peruvian Amerindian ood plain populations were
devastated and strongly decimated (Santos, 1992).
Later in the 16th and 17th century Jesuits as well as
Franciscanian monks (Biedma, 1989; Ardito, 1993)
assembled the native people of the region at missionary stations and converted them to Catholicism. They
applied the Quechua language originating from the
Incas in the Andes region as a common church language (Stocks, 1981; Santos, 1992; San Roman,
1994). The Jesuits were expelled from Peru in
1767, but since then Quechua has been used by various
indigenous groups, especially along the Napo, Pastaza
and Tigre rivers. Many Quechua words referring to

natural resources have entered the current Spanish
practised in the Amazon.
During the rubber boom from approximately 1880
to 1915 the ``mestization'' of native people in the
Peruvian Amazon accelerated. New immigrants
entered the region and the notorious ``patrons'' known
as rubber-barons forced Amerindian groups to extract
rubber from the natural populations of rubber trees
(mainly Hevea brasiliensis (Willdenow ex Adr. Jussieu) Mull. Arg.). They often mixed different ethnic
groups and forced them to move to distant areas
abundant in rubber trees (Collier, 1981; Pennano,
1988). Around 1915 the market for rubber collapsed
due to competition from plantations of rubber trees in
Malaysia. The patron system, however, survived with
the owners of the extraction estates and farms still
virtually owing the local inhabitants. During the following decades the extraction of several other forest
products (Padoch, 1988) resulted in long-distance
migrations and the continued mixing of different in
fact enslaved ethnic groups, as described by Padoch
and de Jong (1990) for the Santa Rosa community
located upriver from Jenaro Herrera. In Santa Rosa
the inhabitants descend from ve native groups, but
today the village is a typical ribereno or mestizo
community.
In spite of disruptive forces two large ood plain
indigenous cultures survived to the present in Peru: the
Pano speaking Shipibo-Conibo and the Tupi-Guaran
speaking Cocama-Cocamilla. The former group currently numbers approximately 25,000 (San Roman,
1994) and people live in more than 100 communities
along the central and upper Ucayali river and its
tributaries (Timoteo, 1989), approximately corresponding in geological terms to the Ucayali sub-valley
(Rasanen et al., 1993). The Shipibo-Conibo culture is
known for their beautiful skirts, and their famous
ceramics. The Cocama-Cocamilla often tend to hide
their native identity (Stocks, 1981), and therefore may
appear less ethnically distinct than the ShipiboConibo. However, according to San Roman (1994)
they may be even more numerous numbering 35,000.
They mainly live along the lower reaches of the
Ucayali, Maranon and Huallaga rivers, corresponding
to the Pastaza-Maranon sub-valley (Rasanen et al.,
1993), but also further down-river along the Amazon
proper at least to Iquitos (Villarejo, 1988). The
17
Table 5
An example of the demographic situated in a Peruvian municipal district (Jenaro Herrera) located at the lower Ro Ucayalia
Village
Population size
Upland
Foundation year
Founders
Casa Grande
Cedro Isla
Florida
Iricahua
Jenaro Herrera
Nuevo Aucayacu
Nuevo York
Nuevo San Juan
11. de Agosto
Padre Ginevez
Progreso
Pumacahua
Yanallpa
280
300
150
120
2000
200
100
200
150
100
250
150
400
No
No
No
No
Yes
Yes
No
No
No
No
No
Yes
No
1930
1904
1990
1905
1944
1982
1993
1989
1990
1920
1942
1930
1911
Cocama Indians from nearby Iricahua

From the San Martn Department
From Tamshiyacu near Iquitos and from Pucallpa
Cocamas Indians from the Puinahua channel
Inhabitants from four nearby villages
Evangelists from the San Martn Department
Evangelists from nearby Cedro Isla and P. Ginevez
From Tamshiyacu near Iquitos
From adjacent Yanallpa
From adjacent Cedro Isla
No information
No information
From the Amazon river and the San Martn Department
a
Population sizes for villages are estimates for 1995. Indications of upland signies that people also use terra rme for subsistence
purposes.
Cocamilla subgroup lives along the lower Huallaga

river (Stocks, 1981). In the Puinahua channel of the
Ro Ucayali as well as in the Ro Samiria area of the
Maranon river more than 80% of the population
apparently are ethnic Cocamas, while the corresponding percentage in the Jenaro Herrera region probably
does not exceed 20%.
The dynamic demographic and cultural history of
the Peruvian ood plains is illustrated by the development in the Jenaro Herrera district. The Iricahua and
Casa Grande communities (Table 5) are considered
Amerindian Cocama villages, although their inhabitants are losing their native identity. In contrast Cedro
Isla, which is located at the same riverine island as
Casa Grande, was founded by immigrants from the
San Martn Department of the Andes. Villages located
close to each others can therefore be surprisingly
different according to their histories and ethnic roots,
as also noted by Padoch and de Jong (1990). The
dissemination of various religious convictions in the
Peruvian Amazon region, as described by Neyra
(1992), also results in formation of new villages. Tiny
Nuevo York was established in 1993 by around 10
evangelic Protestant families moving from the Catholic Cedro Isla village. In 1982 the village Nuevo
Aucayacu was also established by evangelic Protestants, in this case from the San Martn Department.
During the past 100 years the number of villages as
well as the population has grown considerably in the
Jenaro Herrera area (Table 5). Cedro Isla, Iricahua and
Yanallpa were the rst villages known to exist early in

the 20th century, and had a few hundred inhabitants.
Today more than 4000 people live in 12 villages, and
the municipal town Jenaro Herrera, which was established in 1944, had more than 2000 inhabitants in
1993. A similar demographic trend is seen elsewhere
along the main rivers where the number of villages
increases, keeping the average size of small villages
stable, whereas large villages have a fast population
growth, developing into small towns.
The Jenaro Herrera district consists of non-ooded
uplands to the east bordered by ood plains to the
west. The three communities of Nuevo Aucayacu,
Pumacahua and Jenaro Herrera are located on the
border between ood plains and terra rme. Most
crops are grown on the terra rme. The remaining
10 communities are located in the ood plains where
they grow most of their crops, and people occasionally
extract resources from the upland forests (Kvist et al.,
2001a). This option hardly exists for communities
located further up the Ucayali river, for instance in
the Puinahua channel, where the closest terra rme
areas are located around 80 km away.
7. Extraction and economic use of ood
plain habitats
The importance of the ood plain habitats identied
in Table 4 for extractive and economic activities is
18
Table 6
Generalised socio-economic importance according to different extraction purposes of Peruvian ood plain habitats in the lower Ro Ucayali
and lower Ro Maranon regionsa
Agriculture
1. High restinga
2. Young restinga
3. Low restinga
4. Mature tahuampa
5. Young tahuampa
6. High flood basin restinga
7. Low flood plain restinga
8. Flood basin tahuampa
9. Bajial
10. Flood basin bajial
11. Pungal
12. Aguajal
13. Pantanal
14. Lake and river
15. Barrial
16. Playa
Annual
Perennial

Fish

Game

Extraction
Food
Construction
Technical
Firewood Medicine Commerce

a
The scores are subjectively added on the basis of socio-economic studies conducted in ood plain villages in the Jenaro Herrera, the Ro
Samiria, and the Ro Pacaya regions. As an artifact of the applied method, the total number of crosses within each activity does not reect its
total socio-economic value. Major importance (); intermediate importance (); minor importance ().
shown in Table 6. The habitats located in the meander

belt obtain scores for almost all types of activities,
while the habitats of the ood basin are less important.
This may reect that the meander belt resources are
most abundant and have a high production potential or
it may be due to a higher population density in the
meander belt where the favourable infrastructure
creates incentives for an intense utilisation of the
resources. A close relationship is often observed
between the size of villages and the extension of
nearby located high restinga constituting the most
important ood plain habitat for production of ooding sensitive fruit trees and stable crops such as
plantain and manioc.
Annual crops as corn, rice, beans and various
cucumbers are predominantly grown on annually
ooded areas located near the main rivers (Table 6).
Several authors (Hiraoka, 1985a,b, 1989, 1992; Bergman, 1990; Chibnik and de Jong, 1992; Padoch and de
Jong, 1992; de Jong, 1995) described the complex but
risky ood plain agriculture in Peru. Annual and
perennial crops are likely to be destroyed by early,
high or extended inundations, and the river may
erode elds away. Most households therefore seek
to minimise the risk of loss by cultivating different

crops, habitats and sites. Recently deposited and more
or less vegetation free barrial and playa habitats are
easily cultivated, as little effort is needed to clear and
maintain elds on these sites. However, the cropping
period is short and sudden raises in the water levels can
cause inundation and destruction of the crop elds.
The fertile and briey inundated restinga habitats are
also widely used for cultivation of annuals, but weed
problems tend to get high in these habitats, demanding
a high labour input or application of adapted farming
systems such as shifting cultivation. Flooding sensitive perennials are mostly grown on the highest restingas. Almost all ood plain households combine
agriculture with shing, hunting and extraction of
forest products. This is a way to minimise the impacts
of agricultural failure and to generate other incomes
than from sale of crops (Kvist et al., 2001a).
Fishing is the second most important economic
activity for people living from the ood plain habitats,
and most households are regularly engaged in this
activity (Kvist et al., 2001a). The close interaction
between sh populations and ood plain forests was
described by various authors (Gottsberger, 1978;
Goulding, 1980, 1983, 1985; Guerra, 1995; Junk et al.,

1997). It is common that many sh species migrate to
the ooded forests to feed during the inundation, and
when the water level falls the sh density in lakes and
rivers is raising. The migration of the shes, which is
instigated by rapidly increasing or decreasing water
levels, creates favourable shing conditions. At low
water levels the sh catches are also good due to the
high density of shes trapped in congested small water
bodies. People sh in almost all ood plain habitats
(Table 6), but shery in the forests is limited to the part
of the year with high water levels, although remnant
pools and muddy depressions in bajial forest habitats
can be rich in shes throughout the low water season.
Fishermen often undertake trips lasting for days or
weeks where the catch is preserved by fuming or
salting and sold to the urban markets (Kvist et al.,
2001a).
For a majority of the ood plain people hunting is
only an occasional or rare activity, although some
specialise in this profession and sell the largest proportion of their catch as fumed meat (Kvist et al.,
2001a). During the inundation terrestrial animals ee
to non-inundated high restingas, and hunters take
advantage of the situation in which the prey may be
killed only using machetes (Table 6). Hunting is
consequently a seasonal activity in the ood plains.
It is mainly realised during months with high water
levels when shery is difcult. Hunters in canoes
paddle through the ooded forest to remote ood
basin restingas, where they may spend several days.
They also hunt at restingas located closer to human
settlements, but on these sites the larger game species
tend to be depleted or extinct (Bodmer, 1995; Soini
et al., 1996). During the low water level season
hunting in aguajal habitats is relatively common, since
many animals gather there to eat palm fruits.
While subsistence hunting often depletes populations of game species (Bodmer, 1995), few if any plant
species have been depleted by subsistence extraction,
at least not in the Jenaro Herrera region (Kvist et al.,
2001b). Table 6 divides extracted ood plain plant
products to six categories according to the purpose of
their extraction: food, construction, technical purposes, rewood, medicine, and commerce. Products
extracted for sale are included in the latter category,
implying that the remaining ve categories only contain products extracted for subsistence purposes. The
19
categories are identical to and dened by Kvist et al.

(2001b), except that rewood appears separately
rather than within the technical category. The extraction of subsistence products from Peruvian ood plain
forests is sparsely described, but Mejia (1988) and
Lopez (1988) provided some information.
The food category is dominated by extraction of
edible fruits, since other parts of wild plants except
palm-hearts are only rarely consumed. Fruits for subsistence are mainly extracted from restingas in the
meander belt, although aguajal and pantanal habitats
also supply fruits (Table 6). Restingas provide a
diversity of various fruits while aguajales mainly
produce fruits of the palm species M. exuosa and
palm-hearts of E. precatoria. Kvist et al. (2001b)
described traditional construction in Peruvian ood
plain villages. The materials are mostly extracted from
meander belt forests located close to the villages
where construction takes place, and only rarely from
more distant areas. Products from restinga forests are
more often used than products from tahuampa forests
(Table 6). Materials extracted for technical purposes,
e.g. tools and handicraft, come from a large selection
of forest formations, reecting that this category of
uses itself is diverse. In addition, relatively small
amounts of plant material are often needed for technical purposes facilitating transport from distant areas.
Medicinal plants are also easy to transport, and a
considerable diversity of species may serve, implying
that medicinal plants are also extracted from a variety
of forest formations, although apparently high and
young restingas are of major importance. In contrast,
rewood is extracted from forests near the villages
(Table 6). Young restingas are important due to the
prevalence of the much preferred C. spruceanum and
their location near the villages in the meander belt,
although other restinga and tahuampa habitats also
provide appreciated rewood species (Kvist et al.,
2001b).
Extracted plant products have been negotiated and
interchanged since aboriginal times, but large-scale
commercial extraction began with the arrival of the
Europeans to the Amazon. Many products have since
been marketed and some species were depleted decades or even centuries ago, e.g. Castilla elastica Sesse,
Aniba roseadora Ducke and Smilax spp., and commercially valuable species generally tend to get overexploited (Kvist et al., 2001b). In the Jenaro Herrera
20
area products extracted for commerce primarily origin

from aguajal and restinga habitats (Table 6), where
particularly fruits, palm-hearts and timber are
extracted. Fruits as well as palm-hearts are mainly
extracted on a smaller scale by ood plain villagers,
and various authors (Padoch, 1988; Peters et al.,
1989a,b; Peters, 1990; Peters and Hammond, 1990)
discussed commercial extraction of fruits from Peruvian ood plain forests. People may also engage in
marketing timber, but this activity is more often
carried out by people specialising in this activity, as
described below.
FAO (1993b) stated that commercial logging in
Amazonian ood plain forests has ``. . . proved an
economic `blessing' for the Amazon forest industries:
in 1973 they provided 80% of Amazonian timber, and
60% in 1981''. Calophyllum brasiliense Cambess.,
Carapa guianensis Aubl., Ceiba pentandra (L.)
Gaertn., Hura crepitans L., Ocotea spp., Platymiscium
spp., and Virola spp. were named for their importance.
It was further stated that timber can now only be
obtained from ood plain forests in extreme upstream
reaches.
In a study of the timber industry in the Brazilian
Amazon Ros-Tonen (1993) similarly found that until
the early 1970s as much as 8090% of the raw material
originated from the ood plains, while this gure was
60% by the end of the decade, and she estimated that
by 1993 it dropped to approximately 15% of the total
round-wood supply in the Brazilian Amazon region.
In the same period the production rose considerably to
around 50 million m3 in 1989. The species mentioned
as important are C. guianensis, C. pentandra, Hevea
guianensis Aubl., H. crepitans, Maquira schlerophylla
(Ducke) C.C. Berg (synonym Olmedioperebea schlerophylla Ducke), and Virola spp. Wood from many of
these species supply the growing plywood industry,
though Virola surinamensis (Rol.) Warb. was the
second most important source of export sawn-wood
in 1988. Ros-Tonen (1993), and White (1978) from
Peru, described how wood extraction from the varzea
mainly is carried out by lumber-men working in often
remote areas with status as public domain. They
typically sell their production via middlemen, who
frequently supply the basic necessities for the operation in a classical Amazon patron system.
Macedo and Anderson (1993) described the intense
extraction of V. surinamensis stands occurring with
extremely high total standing volumes (243 m3/ha) at

a black water river swamp site at the Brazilian Amazon estuary. It turned out that the extraction highly
violated existing limitations on minimum cutting diameters (45 cm), and furthermore destroyed the possible regeneration of stands that could turn out to be
valuable future supplies for the timber industry. The
authors recommended to improve and enforce adequate forest policy measures to secure a sustained use
of the stands. Barros and Uhl (1995) studied the
extraction of wood along the Amazon river and estuary in the Brazilian Amazon State of Para. Several
logging, transport and processing patterns were identied. In the varzea they found that logging is characterised by manual felling and extraction from the
forest, typically carried out by independently operating logging teams consisting of three men, who on a
daily basis extract around 5 m3 of round-wood, which
is often sold to intermediaries transporting logs to the
mills. Advance payment to logging teams is common.
The transport principally takes place in log rafts,
which are carried to small, medium as well as large
processing industries. Prices obtained for varzea logs
are lower than for terra rme logs (approximately
half). However, the transportation costs are considerably lower: the costs of raft transport amounts to only
around 3% of costs connected with truck transport of
logs from terra rme forest.
In the Peruvian Amazon logging from ood plain
forests still provides large amounts of raw material for
the commercial timber industry. Table 7 shows the
total volume of wood extracted and produced in 1996
in the Loreto and Ucayali Departments, and the proportion originating from tree species growing in ood
plain forests. Not all wood of these species is extracted
from the ood plain forests, as many of the species
also grow in terra rme forests. However, the importance of ood plain forests in the Loreto and Ucayali
Departments is underlined by the fact that around 90
and 60% of the extracted wood was supplied by
species growing in the ood plains.
The principal commercial timber species currently
extracted from the Peruvian ood plains principally
are C. spruceanum, C. brasiliense, Cedrela odorata
L., C. pentandra, H. crepitans, Maquira coriacea
(Karsten) C.C. Berg, Swietenia macrophylla King,
Virola spp., and several Lauraceae and Leguminosae
species (see also Lopez and Freitas, 1990; Freitas,
21
Table 7
Extraction and use of commercial timber in 1996 in the Loreto and Ucayali Departmentsa
Loreto
Roundwood
Bolaina (Guazuma spp.)
Caoba (Swietenia macrophylla)
Capinur (Maquira coriacea)
Capirona (Calycophyllum spruceanum)
Carahuasca (Annonaceae)
Catahua (Hura crepitans)
Cedro (Cedrela odorata)
Copaiba (Copaifera paupera)
Cumala (Virola spp.)
Huayruro (Ormosia spp.)
Huimba (Ceiba samauma)
Lagarto caspi (Calophyllum brasiliense)
Lupuna (Ceiba pentandra)
Mari mari (Leguminosae)
Moena (Lauraceae)
Pashaco (Leguminosae)
Pumaquiro (Aspidosperma spp.)
Quinilla (Sapotaceae)
Requia (Guarea spp.)
Ubos (Spondias mombin)
Yacushapana (Terminalia oblonga)
Total
Total flood plain species
Ucayali
Sawnwood
Plywood and
veener
Sawnwood
Plywood and
parquet
37
31868
5031
560
3191
27154
8295
27884
120
4353
60323
338
3031
437
493
68
47
22861
1001
17079
777
21425
2397
49
50
697
55
69
1
1053
24754
473
14654
1428
7
26719
2567
19686
17247
177
1182
3426
9999
1794
333
266
105
281
188930
173183
71171
66508
25807
25807
209976
123447
45066
Figures are volumes (m3) for timber extracted from both terra rme and ood plains, from ood plains only, and according to species
growing in ood plains. The gures for total extraction of ood plain species include volumes actually extracted from terra rme populations,
as some of the tabulated species grow on both zones. Production of plywood and parquet in the Ucayali Department was not specied to
species. Unpublished data from Banco Central de Reserva del Peru, Surcursal Iquitos, based on economic reports from the Ministry of
Agriculture.
1996). Probably many more species are potentially

commercial. Extraction often is by small operators
working for or selling their product through middlemen, as well as by larger scale more planned operations. In some cases oating sawmills may be
involved. The existing forest legislation is technically
more or less well-equipped, but in practice inadequately enforced. For example, it demands preparation
of management plans for larger concessions, which
in practice becomes mere bureaucratic formalities,
because they as a rule are prepared as desk studies
without experimental backing and with no concern for
their implementation. Legal access to Amazonian
forest resources is admitted through different sized
concessions, by which short-term exploitation licenses
issued for areas less than 1000 ha are the most
common and easily obtained (FAO, 1993b). This

implies that forest extractors, even if they possess a
legal permission, are more encouraged to apply a ``cut
and run'' approach. This problem especially prevails
in the ood plain forests where timber can be extracted
by means of simple manual methods minimising
investments in extraction equipment.
Acknowledgements
We are indebted to the many amiable persons met
during the eld work conducted in different communities. Without their acceptance and participation the
eld studies had been impossible. Working facilities
and assistance in eld work was provided by Centro de
22
Investigaciones Jenaro Herrera (CIJH) of the Instituto

de Investigaciones de la Amazonia Peruana (IIAP).
The WWF-DK/AIF Pacaya-Samiria conservation project and staff provided assistance during the eld work
in the Ro Pacaya and the Ro Samiria regions. Marina
de Guerra del Peru, Servicio de Hidrograa y Navegacion de la Amazonia contributed data for the river
water levels at Iquitos. Staff at The Royal Veterinary
and Agricultural University (KVL) and at the Center
for International Forestry Research (CIFOR) commented on earlier versions of this manuscript. Finance
was provided by the Danish International Development Agency (DANIDA) and from the PacayaSamiria conservation project supported by the World
Wide Fund for Nature, Denmark (WWF-DK) and
Arbejdernes Internationale Forum (AIF).
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Forest Ecology and Management 150 (2001) 326

A review of Peruvian ood plain forests: ecosystems,

larger part is forested (Lugo et al., 1990a). Many

plains covering 300,000 km2. In Peru more than

Although the forests by nature are adapted to large but

environment. Furthermore, an impression is given of

Aniba spp., Ocotea spp.

2. Geography and climate

Pasco, Junin, Cusco and Puno Departments. Based

Dourojeanni (1990) stated that approximately

Based on INEI (1997).

Population growth (%)

of 17.6 inhabitants/km2. Around 60% of the lowland

lowland Amazon region, corresponding to the gure

indivisible from and closely connected to the river and

Gentry and Lopez (1980) postulated an increase in

(1982) their data were poorly interpreted. Fig. 3 shows

ood level, although deforestation on the Andean

4. Fluvial processes and landscape patterns

Erosion and sedimentation are basic processes of

ridges are approximately 6 m high and extend 24 km.

scrolls, levees, crevasse splays and ood channels are

According to Klinge et al. (1990) it is useful in the

ood plain forest formations (Prance, 1979, 1980;

ood plain vegetation were described by several

vegetation succession stage, and presence of character

average of up to 1 month and between 1 and 3 months,

(nos. 9, 10, 11 and 12 in Table 4), implying that they

conditions. Barrial and playa (nos. 15 and 16 in

and Tigre rivers. Many Quechua words referring to

Cocama Indians from nearby Iricahua

Cocamilla subgroup lives along the lower Huallaga

Yanallpa were the rst villages known to exist early in

Firewood Medicine Commerce

shown in Table 6. The habitats located in the meander

to minimise the risk of loss by cultivating different

Goulding, 1980, 1983, 1985; Guerra, 1995; Junk et al.,

categories are identical to and dened by Kvist et al.

area products extracted for commerce primarily origin

extremely high total standing volumes (243 m3/ha) at

1996). Probably many more species are potentially

common and easily obtained (FAO, 1993b). This

Investigaciones Jenaro Herrera (CIJH) of the Instituto

Biedma, M., 1989. La conquista franciscana del alto Ucayali. IIAP/

Guerra, H.F., 1995. Estado actual del conocimiento de la pesqueria

Mighty Tropical River and its Basin. Dr. W. Junk Publishers,

Villarejo, A., 1988. As es la selva. CETA, Iquitos, 330 pp.

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