Biodiversity and Conservation 8: 551±586, 1999.

Ó 1999 Kluwer Academic Publishers. Printed in the Netherlands.

Riparian ¯ooded forests of the Orinoco

and Amazon basins: a comparative review

JUDITH ROSALES GODOY1;2; , GEOFFREY PETTS2

and JUKKA SALO3
1
Universidad Nacional Experimental de Guayana, Centro de Investigaciones EcoloÂgicas, Av. Las
Americas, Edif. General de Seguros, Piso 3. Puerto Ordaz. Edo., Bolivar, Venezuela; 2 The University
of Birmingham, School of Geography and Environmental Sciences, Edgbaston B152TT, Birmingham,
UK (address for correspondence); 3 University of Turku, Department of Biology, FIN-20014 Turku,
Finland; *Author for correspondence (Fax: +44-121-4145528; E-mail: jdr640@novell5.bham.ac.uk)

Received 18 May 1998; accepted in revised form 11 August 1998

Abstract. This paper compares the non-deltaic, riparian-¯ooded forests of the Orinoco and Am-
azon River basins. Ecological relationships between these forests and their environments that can
be useful in establishing schemes for biodiversity conservation are identi®ed. Adaptations of species
to ¯ow seasonality, ¯ooding intensity, sedimentation pattern and nutrient depletion are described.
The variability and diversity of riparian-¯ooded forests is related to (i) landscape evolution (re-
gional-scale, long-term), (ii) water quality (basin scale, long-term) and (iii) hydrology and geo-
morphology (sector-scale, medium-term). The ¯oristic analysis has produced a preliminary list of
242 tree species common to the riparian-¯ooded forests of both basins. This relatively high number
of species is related to connectivity between the riparian corridors of both basins and the e€ective
operation of dispersal mechanisms. Highly oligotrophic environments add uniqueness at the re-
gional scale through the evolution of endemic species presenting adaptations not only to ¯ooding
but also to nutrient depletion. The process of genetic diversi®cation and the evolution of genotypes
adapted to ¯ooding are suggested to explain longitudinal gradients at tributary junctions and
¯oodplain-upland ecotones where current ¯uvial dynamics are unpredictable over ecological time
scales. The paper presents information that may be used to devise appropriate measures to evaluate
sites for riparian biodiversity conservation and management.

Key words: Amazon River, ¯ooded forests, IgapoÂ, Orinoco River, riparian forests, VaÂrzea

Introduction

Increasing concern about the importance of riparian corridors for conserving

biodiversity and protecting river quality has been noted by a number of au-
thors over the last two decades (see e.g. Petts 1997). Urbanisation, construction
of rail and road networks, dam building, water diversion, channelisation,
mining, disposal of industrial and domestic wastes, use of pesticides and her-
bicides, and inappropriate land use leading to increased sediment production,
have impacted upon the integrity of ¯uvial hydrosystems (Petts and Amoros
1996). Sustainable management and restoration of riparian corridors are
552

important for conserving biodiversity, aesthetic quality, and human health

(Naiman et al. 1992; Shaefer and Brown 1992; Petts 1997). Achievement of
these goals poses a major challenge to governmental and non-governmental
organisations. It is a multidisciplinary task, requiring biological, hydrological
and geomorphological studies to be combined within a landscape perspective
(Frissel et al. 1986; Gregory et al. 1991; Malanson 1993; Petts and Bravard
1996; Wassen and Groothans 1996; Hughes 1997; Ward 1998).
Riparian habitats are recognised as refuges for maintaining species richness
and corridors for species dispersal (Meave et al. 1991; Naiman et al. 1993;
Meave and Kellman 1994; Johansson et al. 1996). Natural levels of ¯ood
disturbance and habitat heterogeneity are important in sustaining the biodi-
versity of riparian ecosystems (Salo et al. 1986; Salo 1988, 1990; Bornette and
Amoros 1996; Ward 1998). However, Mitsch and Gosselink (1998) explain
how riparian zones respond at continental scales to climate and geology; at
river scales to altitude, ¯ood regime and channel dynamics; and at local scales
to slope, moisture gradients, sediment sorting, and biotic processes. Conser-
vation and management of riparian systems must integrate these di€erent
perspectives of time and space.
In the South American Region, many rivers retain their pristine character,
though river regulation for hydroelectric power generation (Petts 1990a) start-
ing in the 1970s, has led to impacts that are still not completely evaluated.
However, high levels of poverty and underdeveloped economies require most of
the countries in this region to use their resources at rates that lead to the
degradation of the very environment that sustains local populations. Besides
the hydroelectric projects, mining, logging, forest ®res and industrial and ag-
ricultural development threaten the integrity of riparian ecosystems (Tundisi
1994; Junk 1995; Fearnside and Barboza 1996). Development of sustainable
management strategies or restoration opportunities therefore requires an im-
proved understanding of the characteristics and functioning of these ecosys-
tems. However, few studies in this region have incorporated a holistic approach
that can be useful in the development of strategies for maintaining the values of
these ecosystems. Only recently have more integrated approaches considered
¯oodplains as complex and unique systems (Salo et al. 1986; Kalliola et al.
1992a, 1992b; Junk 1997a). Junk (1997a) noticed that most of the studies on
riparian systems have been oriented towards either limnological or terrestrial
aspects but few are really focused on the unique properties and processes of the
¯oodplain. The ¯ood pulse (Junk et al. 1989) o€ers an approach for such
integration.
This analysis focuses on two of the most important river basins of South
America, namely the Orinoco and the Amazon, and explores the patterns of
ecological relationships of woody species within riparian (i.e. seasonally-
¯ooded) forest ecosystems. Based mainly on an extensive literature review,
 553

riparian areas of the Orinoco and the Amazon river basins are described in
terms of:
± the riparian environment, including geological histories of both basins,
connectivity between riparian corridors, and the regional- and local-scale
patterns of riparian habitats,
± the riparian forest vegetation, considering ¯oristic similarities, biological and
ecological processes, diversity, and their environmental determinants.

Study area

The Amazon is the largest river in the world in terms of discharge, the Orinoco is
the third largest (Figure 1, Table 1). Both river systems have similar unit dis-
charges (discharge/drainage area) and comparable sediment yields (Meade
1994). High run-o€ occurs from the Guayana Shield Region, which dominates
the ¯ow in the Orinoco, and from the Negro River in the Amazon basin. The
Amazon also receives high discharges from Andean rivers such as the Madeira.
The Andean mountains contribute 85% to 90% of the sediment yield of both
river systems (Martinelli et al. 1989; Meade et al. 1990; Meade 1994). Both the
Orinoco and Amazon rivers have important ¯oodplains (Hamilton and Lewis
1990; Sippel et al. 1994), but in relation to their drainage areas, the Amazonian
¯oodplains are most extensive. Large ¯oodplains within the Orinoco basin in-
clude the 80,000 km2 of the Orinoco Llanos (Junk 1997a) and ¯oodplains along
the Atabapo, Ventuari and Caura tributaries (see Figure 1 and Berry et al. 1995).

Methods

In addition to original data collected by the senior author and collaborators in

di€erent studies along the Orinoco basin, data from di€erent bibliographical
sources of both the Orinoco and Amazon rivers have been reviewed and an-
alysed. In this process, we follow a comparative scheme to ®nd out if the
patterns found for the Amazon have equivalents in the Orinoco.
For the ¯oristic analysis, a preliminary list of tree species common to rip-
arian forests of both the Orinoco and the Amazon river basins has been derived
from a comparison of: (i) the published lists of literature and additional un-
published data (Prance pers. comm.) on riparian forests of the Amazon with
(ii) the botanical collections of the ®rst author and collaborators (Claudia
Knab-Vispo, Elio BricenÄo, Gabriel PicoÂn, Paul Berry) along riparian forests of
black-water tributaries of the Orinoco (Caroni, Caura, Mapire and Atabapo;
Figure 1). Additional information on these and other riparian forests within
the Orinoco basin includes white waters and clear waters of the lower plains or
 554

Figure 1. The Orinoco and Amazon River basins and their main tributaries. A ± Hydrographs: (a) Orinoco River in Ciudad BolõÂ var
gauge station, (b) Amazon River in Obidos gauge station.
 555

Table 1. Some general characteristics of the Orinoco and the Amazon rivers.

Orinoco Amazon

General characteristics
Drainage area (km2) 1,080,000b 6,150,000a
Discharge (m3/sec) 36,000b 200,000a
(runo€ m3/s/km2) (0.033) (0.033)
Total sediment (t/y) 150,000,000d 1200,000,000c
Sediment yield (t/km2/yr) (139) (195)
Fringing ¯oodplain area of the 7,000e 92,000e
main channel (km2)
(percent of the basin) (0.65%) (1.5%)
Floodplains including tributaries Unknown 307.000h
and minor a‚uents (km2)
Tributaries Tributaries

Physico-chemistry White Black White Black

Metab Atabapob Solimoes Negro

pH 7.4 4.4 7.0f 5.2f
Conductivity (lS/cm) 72.9 14.6 58.0f 9.0f
Calcium (mg/l) 8.95 0.07 8.83g 0.39g
Magnesium (mg/l) 1.67 0.03 1.26g 0.13g
Potassium (mg/l) 0.93 0.15 1.40g 0.46g

Sources: a Meade et al. (1991); b Weibezahn (1990); c Sippel et al. (1994); d Hamilton and Lewis
(1990); e Meade (1994); f Junk and Furch (1985); g Konhauser et al. (1994); h Junk (1997a).

Llanos bajos (Smith pers. comm.), white waters of the lower Orinoco ¯ood-
plain, black-water swamps in the high plains or Llanos altos and clear waters
of the Cataniapo river (Bernardi 1957; Gonzalez 1986, 1987; TeraÂn and Duno
1988; Ramirez and Brito 1990; Colonello 1990a, 1990b, 1991; Castillo 1994;
Bevilacqua and Gonzalez 1994; Ponce et al. 1994). Information for the clear or
black-water tributaries of the Upper Orinoco in the Amazon state, was taken
from the TROPICS database of the Missouri Botanical Garden Herbaria and
the Types database of the New York Botanical Garden. They were also used to
verify the authorities and presence of all the species cross-referenced in both the
Orinoco and the Amazon river basins.

Results and discussion

The Riparian environment

Paleoenvironmental context
Since the Cretaceous, the Amazon and Orinoco basins have followed similar
evolutionary paths characterised by three main phases. The ®rst phase was
dominated by the progressive uplift of the Andes and continental deposition.
556

The second phase was dominated by the formation of a large lake, Lake Pebas,
in Western Amazonia (18±11 million years B.P., Hoorn et al. 1995; RaÈsaÈnen
et al. in press), and marine incursions leading to the extension of estuarine
systems (c. 11±c. 8 million years B.P.) including sporadic marine incursions
from the Caribbean into the Western Orinoco basin (RaÈsaÈnen et al. 1992,
1995). During these phases, the Paleo-Orinoco and Paleo-Amazon systems
were probably characterised by two contrasting patterns, with suspended-
sediment dominated systems draining the Andes along a narrow foreland belt
between the mountains and the lacustrine areas, and east to west ¯owing clear/
black-water dominated rivers draining the Shield areas. The predominant ¯ow
direction of the Paleo-Orinoco in that time was northwards. Quartz sands
originating from the Shields were deposited in the valley of the current Negro
River, contributing to its modern oligotrophic character. The third phase was
the ¯uvial period of the last c. 8 million years, giving a tentative age for the
westward drainage of the Amazon and the Orinoco (RaÈsaÈnen et al. 1987,
in press). Consequent upon the uplift of the northern Andean mountains
(Cordillera de Merida), the northern ¯ow of the Palaeo-Orinoco system was
closed, and the parallel eastward drainage of the Orinoco and Amazon rivers
was created. Degameroo (1996) gives evidence for the eastward evolution of
the Orinoco river. Frailey et al. (1988) postulated the presence of a Pleistocene-
Holocene lake with a northern outlet to the Caribbean sea, following the uplift
of the Andes, but this has been refuted by recent research (Tuomisto et al.
1992; RaÈsaÈnen et al. 1995).
Understanding the evolution of the Pebas system is required to reconstruct
the history of modern riparian biota along the Amazon and Orinoco rivers.
Fluvial systems of the Orinoco and the Western Amazon would have experi-
enced repeated fragmentation, reorganisation, ¯ow reversals, and shifts be-
tween black-water systems and suspended-sediment rich environments. The
connection between both river systems and their landscapes has been evident
since the Miocene, and the modern Upper Orinoco-Rio Negro link through the
Canal del Casiquiare (Figure 1) is a remnant of this long evolution. The tec-
tonic history is dominated by the Andean uplift (see Neller et al. 1992), but
rivers draining the Guianan-Brazilian crystalline shields have also experienced
some tectonic control. Tilting (Garner 1966; BricenÄo 1982, Schubert et al.
1986), and an increase in rainfall about 6,000 years ago have determined the
modern course of ancient shield rivers, such as the Caroni and Paragua in
Venezuela. Furthermore, Holocene bank deposits on the alluvial plains of the
Orinoco River above its current day bankfull level are proposed by Carbon
and Schubert (1994) as evidence that the Eastern Plains region has been tec-
tonically uplifted, at least to the end of the Pleistocene and possibly up to the
present.
 557

Evidence for Pleistocene aridity, associated with glacial periods, has been
related to contractions of forests and expansions of savannas in the Amazon
region. This has lead to the formulation of the refuge hypothesis to explain
areas with higher biodiversity (Ha€er 1969; Prance 1982; Whitmore and Prance
1987), and to explain phases of high erosion related to the dominance of open
or less dense vegetation types (Tricart 1975, cited in Baker 1986). However,
some evidence indicates that during glacial periods cooling instead of aridity
determined the distribution of species (Colinvaux 1996) and that extreme
changes in vegetation density did not occur (Irion et al. 1997). E€ects may have
been most signi®cant in the northern Orinoco basin because of its higher lat-
itude. Paleo-dune systems, dated as late Pleistocene, are found in the Lower
Plains (Roa 1979). Evidence for changes in sea level during the Pleistocene is
seen in the morphology of present ¯oodplains in the lowlands of the Amazon
(Van der Hammen et al. 1992; Irion et al. 1997) and possibly along the Orinoco
as well.

Contemporary patterns

a. Regional-scale. Tropical climates with isothermal regimes characterise the

Orinoco and Amazon basins. Seasonal precipitation gives both rivers a char-
acteristic discharge regime (Junk and Furch 1985; Hamilton and Lewis 1990)
with a highly predictable annual ¯ood (see in Figure 1, A ± Hydrographs).
Water levels can ¯uctuate between 10 and 15 m per annum with a maximum
¯ood duration of 6 to 8 months. The Amazon basin is dominated by tropical
rainforest associated with annual precipitation of more than 2,000 mm. In the
Orinoco basin, rainforests are mainly found in the Guayana Region and on the
Andean mountain slopes. An extensive area of this basin has an annual pre-
cipitation below 2,000 mm and is dominated by savannas, where gallery forests
characterise the river corridors.
Di€erences in the composition of sediments in both the Amazon and the
Orinoco Rivers relate to three distinct geological regions: the tectonically active
Andean mountains, the crystalline shields, and the foreland depressions
(Martinelli et al. 1989; Stallard et al. 1990; Meade 1994; Lewis et al. 1995).
White-water rivers, rich in suspended sediments, with high pH and conduc-
tivity, originate in the Andes mountains and the tectonically-controlled An-
dean forelands. Clear-water and humic-rich black-water rivers (oligotrophic,
acidic and with low suspended sediment concentrations) characterise the
tributaries of the shields and the plains in both the Orinoco (Vegas-Villarubia
et al. 1988; Lewis and Saunders 1989, 1990; Weibezahn 1990; Stallard et al.
1990; Paolini 1995; Edmond et al. 1995, 1996) and the Amazon basins (Irion
1984; Sioli 1984; Junk and Furch 1985; Richey et al. 1991; Martinelli et al.
1993; Konhauser et al. 1994; Furch and Junk 1997).
558

b. Sector scale patterns. In both basins, the modern channel patterns re¯ect the
runo€ and sediment regimes superimposed upon geomorphological controls
(slope and valley width) related to the long term evolution of the two rivers in
the di€erent geological regions. High diversity and complexity of channel
patterns along the river network of the Amazon basin is well documented
(Irion 1984; Salo et al. 1986; Salo 1990; Kalliola et al. 1992b; Mertes et al. 1995,
1996), but there are only isolated references for the Orinoco (Garner 1966;
Zinck 1974, 1980, 1982; Schubert et al. 1986; Carbon and Schubert 1994).
Table 2 provides some examples of the predominant channel types found
along the Orinoco river basin. As a broad pattern, river channels have three
general forms which relate to a downstream sequence. Rivers draining the
Andes, with high sediment loads and steep slopes are characterised by actively
braided systems. Downstream, high sediment loads and lower slopes combine
to produce wandering meanders with an intricate scroll-bar topography related
to high rates of meander migration. Further downstream, low gradient reaches
with anastomosing patterns are marked by sinuous but stable channels re-
stricted by levees, with ¯oodplains being dominated by overbank deposition.
This pattern has been clearly described for the Amazon basin by Mertes et al.
(1995, 1996) and Baker (1986). Also it can be seen in the Apure, Meta and
Guaviare tributaries of the Orinoco draining from the Andes, although at a
smaller scale, considering their huge di€erences in discharge in comparison
with the Andean tributaries of the Amazon. Overbank deposition associated
with anastomosing channels is indicated by Hamilton and Lewis (1990) as the
primary mechanism of ¯oodplain aggradation along most of the main Orinoco
River. However from our own observations, the main channel of the Orinoco
River shows either straight, braided or anastomosing patterns in di€erent
reaches. It has narrower fringing ¯oodplains, smaller lakes, and few blocked-
valley lakes in comparison with the Amazon (Hamilton et al. 1992), and an
internal delta occurs on the left margin of the middle reaches between the
con¯uence of the Meta and Apure rivers (Figure 1).
The Shield rivers ± the right tributaries of the Orinoco and the northern and
southern tributaries of the middle-Amazon ± are characterised by con®ned
reaches and bedrock channels. These black-water or clear-water rivers have
low sediment loads and stable channels, and their degree of sinuosity is related
to channel slope. Where the channel ¯ows through unconsolidated beds,
sinuosity can be very high. An anabranching pattern is found in the area
downstream of the con¯uence of the Negro and Branco Rivers, known as
Archipielago das Analvilhanas.
More dynamic reaches are found at some tributary con¯uences characterised
by low channel slope and in¯uenced by the hydrological dynamics of the main
river channel. Backwater e€ects have been clearly demonstrated by Meade
et al. (1991) for several tributaries of the Amazon and by Vargas and Rangel
Table 2. Some examples of river channel patterns found in lower valleys of the Orinoco river basin (based upon Petts 1990b).
Single-thread
Straight Sinuous
Channel planform Wandering Meandering (from single
to complex meanders)
Channel stability High Low to high
Characteristic patches Narrow riparian zone; Low Wide ¯oodplain zone: Silt
to moderate development overbank deposits, sand
of lateral sand beaches levees, abandoned channels,
organic mud backswamps;
Point-bar systems
Dominant processes Low lateral erosion and Overbank sedimentation,
channel accretion Meander cuto€s
Lateral erosion
in rivers with low This is the dominant pattern Con¯uence of tributaries such
sediment loads, along the main channels of as the Caura River in the
the Shield rivers: Caura, Orinoco, or the Mato River
Caroni, Ventuari in the Caura. But also in
valleys with low slopes in the
Caroni (e.g. Kukenan and
Yuruani valleys). Middle
Nichare, Cano Maarik in the
Paragua watershed.
in unconstrained Lower Guaviare
valleys common in Lower Vichada Capanaparo
the Andes and Cinaruco
lowlands of the Lower Apure, inner delta
main channels with shows distributary
high sediment loads characteristics (Baker 1986)
Multi-thread
Straight Sinuous
Braided Anastomosing
Low High
Wide ¯oodplain zone; Sand Wide ¯oodplain zone;
and gravel non-vegetated Point-bar systems;
islands; Abandoned Abandoned channels;
channels Organic mud backswamps;
Vegetated islands
Lateral erosion and channel Overbank sedimentation
accretion Channel avulsion
Channel avulsion
Anastomosing seems to be
occurring in zones
upstream of rapids or
waterfalls. In the Caroni
river they were called
`deranged patterns' by Garner
(1966)
Some reaches of the Upper
Orinoco, Lower Atabapo
With low development in the Lower Orinoco reaches down
Middle Orinoco between stream from the con¯uence
the con¯uence of Apure and of the Caura. Middle
Caura rivers, some sectors Orinoco between the
of the rivers Santo Domingo, con¯uence of the
Caparo and Uribante Atabapo and Arauca
in the high Apure basin. River.
Middle Apure and Meta R. Lower Meta
559
560

(1996) for the Caura tributary of the Orinoco River. Tributary con¯uences
show a greater duration of inundation, a greater rate of sedimentation in the
extreme reaches of the backwater zone, and a greater rate of channel move-
ment than adjacent sectors. In low-sediment load rivers, the reach in¯uenced
by backwater e€ects from the main river is characterised by accretion in both
upstream and downstream locations (Figure 2), and these are both of eco-
logical importance.
At the mouths of clear and black-water tributaries of white-water rivers,
tributary impoundment during high river ¯ows also leads to sedimentation in
downstream and upstream reaches of the impounded channel (Sioli 1984; Ro-
sales 1990; Vegas-Villarubia and Herrera 1993). In the lower reaches of these
tributaries, sedimentary deposits are characteristic of the main river. In the
upper reaches of the impounded area, the coarser sediments, characteristic of the
tributary basin settle out of suspension, but ®ner silt and colloidal particles are
transported toward the central area where they are deposited due to the very

Figure 2. Con¯uence e€ect on soils in the Mapire River con¯uence to the Orinoco River (after
Rosales 1990). P = phosphorous, Ca.Mg/Na.K = calcium magnesium/sodium potassium.
 561

slow-¯owing, impounded waters. Rosales (1990) showed how this process leads
to richer soils in the lower and upper reaches of the con¯uence of the black-water
Mapire River to the white-water Orinoco River (Figure 2). Vegas-Villarubia
and Herrera (1993) found higher total suspended solids in the upper and lower
reaches of that Mapire con¯uence with lower values at middle locations during
highwaters (September in the Orinoco, see Figure 1). The lower Caura, tributary
of the Orinoco, seems to follow a similar pattern (Garcia 1996).
In rivers with high sediment loads, con¯uences are associated with highly
dynamic channels, such as the extensive internal delta at the con¯uence of the
Apure and Orinoco. In such cases, the rich nutrient loads of both the main
channel and the tributary waters create more uniform patterns of productivity.
Studies in oligotrophic environments however need to consider the hydrolog-
ical, geomorphologic and ecological signi®cance of these riparian sectors.

The Riparian forests

Following the di€erences between white, clear and black waters, described by
Sioli (1984), riparian forests of the Amazon River basin have been classi®ed by
Prance (1979) as two major types: VaÂrzea (on white-water rivers) and IgapoÂ
(on clear and black-water rivers) with permanent or seasonal ¯ooding.
For the Amazon basin, these forests have been described in great detail (i.e.:
Rodriguez 1961; Takeuchi 1962; Prance 1979; Keel and Prance 1979; Revilla
1981; Campbell and Frailey 1984; EncarnacioÂn 1985; Pires and Prance 1985;
Campbell et al. 1986, 1992; Balslev et al. 1987; Kubitzki 1989; Dumont et al.
1990; Lamotte 1990; Klinge et al. 1990, 1995; Kalliola 1992a; Worbes et al.
1992; Worbes 1997; Ferreira 1992, 1997a, 1997b; Duivenvoorden and Lips
1993; Duivenvoorden 1995, 1996; do Amaral et al. 1997; Parolin and Ferreira
in press; Ferreira and Stohlgren in press). Worbes (1997) presents a compre-
hensive overview for the Central Amazon.
On the contrary, ¯ooded forests of the Orinoco basin are poorly docu-
mented. Only two pioneering studies can be noted. The ®rst is the Mapire
Project, which has been coordinated by Rafael Herrera from the Instituto
Venezolano de Investigaciones CientõÂ ®cas (IVIC). This is a multidisciplinary
study of the ¯ooded forests in the lower sector of the Mapire River, a black-
water tributary of the lower Orinoco. For this Project, Rosales (1990) identi®ed
a longitudinal gradient based on relationships between ¯oristic and environ-
mental variables, from eutrophic (VaÂrzea, Orinoco-related waters) to oligo-
trophic (IgapoÂ, Mapire-related waters) ¯ooded forests. Colonello (1990a,
1990b, 1991) focused on an eutrophic (VaÂrzea-type) ¯ooded forest in the lower
Orinoco ¯oodplain (Laguna de Mamo). More recently, riparian forests have
been investigated in the Guayanan black-water rivers: the Caroni (Rosales et al.
1993) and Caura rivers (BricenÄo 1995; Huber 1996; Rosales 1996; Knab-Vispo
562

et al. 1997; BricenÄo et al. 1997; Rosales et al. 1997; Nelda Dezzeo pers. comm.).
Other studies in riparian areas of the Orinoco basin are related mainly to
swamps and gallery forests (TeraÂn and Duno 1988; Ramirez and Brito 1990;
Meave et al. 1991; HernaÂndez 1992; Gonzalez 1986, 1987; Kellman and
Tackaberry 1993; Kellman et al. 1994; Ponce et al. 1994; Tackaberry and
Kellman 1996).

Floristic relationships
The preliminary species list (see Appendix) of plant woody species that occur in
the riparian forests of both the Orinoco and the Amazon ¯uvial corridors
includes a relatively high number of species (242), which indicates that con-
nectivity between both corridors have been e€ective in the intermixing of the
taxa. The most important families in terms of number of common species are
the Myrtaceae (23), Mimosaceae (21), Fabaceae (18), Euphorbiaceae (17),
Caesalpiniaceae (12), Arecaceae (11), Clusiaceae (10), Lecythidaceae (10), and
Rubiaceae (10).
The ¯ora of the ¯ooded forests re¯ects the di€erent hydrological, biogeo-
chemical and geomorphological settings described for the Orinoco and Ama-
zon river basins. Prance (1979) wrote one of the ®rst publications that identify
¯oristic elements associated either with permanent or seasonal VaÂrzea or IgapoÂ
forests based on water chemistry. Kubitzki (1989) also concluded that water
chemistry is an important discriminator of plant communities of the Amazon
¯ooded forests. He further argued that the VaÂrzea has a ¯oristic composition
similar to that of the non-inundated Amazonian forests. The ¯ora of IgapoÂ
studied in the region of Central Amazon is related to communities typical of
oligotrophic white sand environments (the old Negro and TapajoÂs Rivers re-
gion). Given the young geological age of the Central and Northern Andes this
author also suggested that the present VaÂrzea systems are younger than the
Igapo systems (Kubitzki 1989), which have evolved since the Upper Creta-
ceous. However, the VaÂrzea systems as limnological entities have been present
in Western South America since the earliest uplift (Miocene), with the begin-
ning of mobilisation of Andean sediments, giving some of the VaÂrzea taxa
sucient evolutionary age for speciation. Higher diversi®cation in IgapoÂ
forests, as stated by Kubitzki (1989) and Worbes (1997), would be a result of
the di€erent histories and ages of the ecosystems following the sequence of
fragmentation previously described for the Pleistocene and the Holocene.
A comparison of the number of species/families made by Worbes (1997)
between IgapoÂ, VaÂrzea, and terra ®rma forests for the Central Amazon showed
a high diversity of Myrtaceae and Euphorbiaceae in ¯ooded forests. This is
supported in this study (see Appendix). We found a high diversi®cation in
Mimosaceae, between which, the genera Inga is shown to be restricted to
neotropical areas with high numbers of riparian species within the Orinoco and
 563

Amazon basins (see Pennington 1997). Furthermore, in the genera Camp-

siandra, which constitutes one important genera in the ¯ooded forests of the
Orinoco and Amazon, Stergios (1996, 1997) has found a high morphological
diversi®cation which allowed the division of the former 2 species and 1 variety
to 23 new taxa, including species and varieties, many of them endemic or of
restricted distribution. In the lower Caura alone he identi®ed the presence of
three di€erent species. The relationship between this diversi®cation and the
history of channel-¯oodplain evolution (the repeated fragmentation, connec-
tion, and disconnection along the Orinoco and Amazon basin riparian corri-
dors) is an issue which deserves further biogeographic investigations.
At the generic level, most of the taxa reported for the riparian forests of the
Amazon are also widely distributed along the Orinoco. Exceptions are some
genera belonging to the Igapo ¯ooded forests, which are con®ned to mostly
sandy ¯oodplains in the region of the Atabapo, Upper Orinoco, Guainia and
Casiquiare-Rio Negro river basins (Huber and AlarcoÂn 1988; Kubitzki 1989;
Huber 1995; Berry et al. 1995). This region is known for its high endemism and
diversity (Kubitzki 1989; Huber 1994; Berry et al. 1995) which appears to be
determined by the extreme conditions of oligotrophism. The possible repeated
fragmentation during the di€erent phases of the Pleistocene and Holocene in
that region can also be taken into consideration to explain this high diversity.
Many of the common species are also present in upland forests of both
basins. They might have evolved genotypes that allow individuals of certain
populations to survive ¯ooding. Species that also occur in upland forests are
found along the ¯oodplain or channel edges in environments with low levels of
¯ood duration or depth. As Rosales et al. (1997) and Ferreira (1997a) state,
infrequently ¯ooded forests or riparian forests with low ¯ooding intensities
resemble the ¯oristic composition of upland communities. However, detailed
studies are needed concerning the species that might also be experiencing un-
predictable ¯ood pulses or predictable ¯ood duration at di€erent recurrence
intervals ± say ®ve, 10 or 50 years ± (Junk 1997b). Genetic diversi®cation can
be expected to occur in the populations present in these edges of the ¯oodplains
(see Keeley 1979; Davy et al. 1990; Ferreira in press).

Biological and ecological processes

a. Flood seasonality and phenological patterns. Junk et al. (1989) propose that the
¯ood pulse is the principal driving force for the existence, productivity, and in-
teractions of the major biota in Central Amazon ¯oodplain systems. The strong
predictability of this pulse along large tropical rivers as the Amazon is associ-
ated with the development of strong seasonal ecological patterns (Junk 1997a).
The presence of annual rings in the wood of certain species tolerating in-
undation in ¯ooded forests of the Amazon and the Orinoco is evidence of an
564

annual dormancy period, which seems to be pulse-related (Worbes 1985, 1997;

Parolin and Ferreira in press; Nelda Dezzeo pers. comm.). In addition, it has
been reported that many of the tree species in the ¯ooded forests shed their
leaves during the submersion phase (Worbes 1985, 1997). However, for most of
the species tolerating inundation in the VaÂrzea forests of the lower Orinoco,
Colonello (1991) found that leaves were continuously renewed throughout the
year.
With regard to fruiting seasonality, the timing of the ¯ood season is an
important factor for e€ective mechanisms of dispersal. For the Amazon
¯ooded forests, Goulding (1980, 1988) and Kubitzki and Ziburski (1994) found
that fruit production occurs at the onset of the high water season. Similar
observations have been made by Colonello (1991) for the Orinoco ¯ooded
forests and have been noted by Conrad Vispo (pers. comm.) in the ¯ooded
forests of the lower Caura. Kubitzki (1985) and Kubitzki and Ziburski (1994)
stressed the evolution of dispersal mechanisms such as hydrochory and
icthyochory, for ¯ooded forests species. Icthyocory has also been reported as
very important for the Amazonian dispersal of ¯oodplain plant species (Gotts-
berger 1978; Goulding 1980, 1983, 1985; Goulding et al. 1988; Junk et al.
1997).
Germination and growth of seedlings are also ¯ood-pulse regulated and
¯ood duration and timing determines the success of species in reaching an adult
stage (De Andrade 1991; Saul Flores pers. comm.). Preliminary data from
seedlings by David Williams, Leandro Ferreira and Rita Mesquita (Ferreira
pers. comm.) suggest that there is little di€erence within species in their basic
allometry (height and stem diameter) at di€erent levels on the Igapo ¯oodplain
forest in Central Amazonia. However changes, sometimes abrupt, have been
identi®ed in allometric slope for most species and di€erences have been shown
between species in the height of their allometric regression lines.
Strategies of vegetative reproduction seem to be more ecient in those en-
vironments characterised by high unpredictability in the timing of the ¯ood
season. With regard to germination, Kubitzki and Ziburski (1994) found three
dominant mechanisms in the reproductive biology of several woody species of
the Amazon ¯ooded forests: (i) dormancy, with germination only after the
completion of a period of submersion; (ii) prolonged drift, and (iii) lack of
dormancy, which is common in pioneer or colonising species. Worbes et al.
(1992) di€erentiate successional stages of Amazon VaÂrzea-type forests, com-
bining growth strategies of the species with quantitative aspects of vegetation
analysis. They point out that strategies of tolerance to seasonality in general
can lead these species to support the seasonal ¯ooding of the VaÂrzea envi-
ronments because many of the species typical of VaÂrzea forests are also found
in the terra ®rme forests.
 565

b. Mechanisms of nutrient conservation. Mechanisms of nutrient conservation

have been suggested by Herrera (1985) in Igapo communities of the Amazon
River. The Igapo must compensate for the unfavourable nutrient status of the
soil and strategies for nutrient conservation are driven by a high eciency of
nutrient uptake (Furch and Klinge 1989 in Furch 1997). For the Mapire River
in the Orinoco basin, Barrios and Herrera (1994) report that in synchrony with
the hydrological cycle, the seasonally ¯ooded forests show a nitrogen cycle
where inputs and accumulation are maximised when the system is under
minimum stress during the dry season. During the ¯ood season, the system
enters a period of dormancy with minimal use of nutrients and energy to avoid
or tolerate anaerobiosis.
Biogeochemical di€erences between Igapo and VaÂrzea ¯ooded forests are
reported by Klinge et al. (1983), Klinge and Furch (1991) and Furch (1997) in
the Amazon; the Igapo being related to low nutrient concentrations. Compa-
rable results were found by Rosales (1990) for the Mapire-Igapo ¯ooded for-
ests located in the Orinoco basin (Table 3). The low concentrations of nutrients
in the leaves of Igapo ¯ooded forests, together with their evergreen character,
are evidence of the presence of a mechanism for nutrient conservation under
oligotrophic conditions (Klinge et al. 1983). Parolin (1997) compares the
VaÂrzea and Igapo in terms of the eciency of nutrient uptake associated with
nutrient availability for seedlings of pioneer species. Di€erent survival strate-
gies appear to be operating in these types of forests and she concludes that they
are associated with a higher nutrient availability within the VaÂrzea.

c. Colonisation of exposed riverine sediments. This process has been reported for
the highly dynamic systems that characterise the Upper Amazon basin by
Kalliola et al. (1991) who studied the channel patterns and general relation-
ships between the presence of some colonising species on sediment bars in
sixteen Upper Amazon rivers. Their results show an overall scarcity of co-
lonising species associated with these bars, which are poor in organic matter
and nitrogen. Soil zonation, seasonality in water level ¯uctuation, and dis-
turbance due to siltation, are considered important factors in determining the
species present in the more active environments. Comparable studies are
lacking in the Orinoco basin. However, in an expedition made by researchers of
the Finish Amazon Project to the Middle Orinoco, R. Kalliola (pers. comm.)
observed the presence of similar mechanisms in some herbaceous and shrubs
species that colonise the sandy-channel islands.

d. Tolerance to the anaerobiosis. Profuse lenticel hypertrophy and adventitious

roots are expressions of anatomical and morphological adaptations to avoid
anaerobiosis (Gill 1970). The presence of these morphological adaptations in
species of the Amazon ¯ooded forests has been noted by many authors.
566

Table 3. Chemical composition ± mean (variance) ± of various bioelements in leaves from ¯ooded
forests in the Orinoco and the Amazon.

Site N P K Mg Ca
(%) (%) (%) (%) (%)

Orinoco
(Rosales 1990)
Igapo at the con¯uence
of the Mapire and Orinoco rivers
(see Figure 3)
Community 1, Upstream 2.03 0.10 0.85 0.7 0.18
sedimentation area, 57 ind. 25 sp. (0.62) (0.04) (0.67) (0.07) (0.11)
Community 2, stabilised point-bars, 2.13 0.12 0.65 0.21 0.29
¯ooding 5±8 m 27 ind, 9 sp. (0.37) (0.03) (0.49) (0.15) (0.11)
Community 3, middle zone (see Figure 3) 1.72 0.08 0.50 0.15 0.13
overbank swamp, ¯ooding 3±5 m, 72 ind, 21 sp. (0.56) (0.03) (0.23) (0.06) (0.12)
Community 4, middle zone, overbank swamp, 1.47 0.06 0.35 0.11 0.11
ecotone forest-savanna, ¯ooding 1±3 m, (0.54) (0.02) (0.24) (0.05) (0.05)
23 ind. 10 sp.
Amazon
(Klinge et al. 1983)
Varzea forest ± Marchantaria Amazon, 2.59 0.18 1.32 0.40 1.91
31 ind. 27 sp. (0.53) (0.04) (0.68) (0.19) (0.83)
Varzea forest Iranduba Amazon, 2.49 0.09 0.86 0.40 1.63
13 ind. 13 sp. (0.02) (0.07) (0.66) (0.24) (0.77)
Igapo Praia Grande Amazon, 1.73 0.06 0.63 0.12 0.25
21 ind. 21 sp. (0.40) (0.03) (0.26) (0.06) (0.20)

N = nitrogen, P = phosphorous, K = potassium, Mg = magnesium, Ca = calcium.

However, few studies quantify its importance. In the Mapire Igapo ¯ooded
forests of the Orinoco basin, Rosales (1990) found 87% of the species with
lenticel hypertrophy, and 77% with adventitious roots on their stems. Adven-
titious roots are more frequent in the communities supporting higher ¯ooding
depths and duration as well as soils that have better nutritional conditions and
sandy textures. For the Central Amazon, however, Worbes (1997) did not ®nd
any signi®cant relationship. But Parolin (1997) found that in VaÂrzea, the stress
induced by periodical ¯ooding is compensated for by morphological adapta-
tions (such as adventitious roots, lenticels and stem hypertrophy) which require
high nutrient supply and allow the plants to maintain metabolic processes
under waterlogged conditions. Igapo species on the other hand, which grow in
nutrient poor environments, cannot `a€ord' either the costly production of
morphological adaptations or frequent changes of the leaves. Their strategy is
to remain in a stage of rest during the ¯ood period.
Many authors have also studied physiological adaptations to anaerobiosis
tolerance, for example the role of hormones as ethylene, as well as the
importance of symbiosis either with nitrogen ®xing bacteria, or with fungi
 567

(endo- or ecto-mycorrhizae) for the survival of the species to the ¯ooding con-
ditions (De Andrade 1991; De La Rosa 1988; Izquierdo 1988; Schluter and
Furch 1992; Fernandes Correa and Furch 1992: Moreira et al. 1992; Losos 1995).

Species biodiversity, an integrator factor

Based on the previous discussion we have shown that the common ¯ora found
in the ¯ooded forests of the Orinoco and the Amazon resulted from the ability
of the species to disperse, germinate, and survive under environmental con-
ditions characterised by high ¯ooding intensity and disturbance (erosion,
sedimentation or high water velocities), as well as to conserve nutrients in the
oligotrophic environments. Therefore, the ¯ooded forest communities will be
composed of those plants having morphological, physiological, and/or re-
productive adaptations that enable them to e€ectively grow, compete, repro-
duce, and/or persist under the stresses and disturbances typical of riparian
habitats.
Summary results of species richness in several ¯ooded forests of the Orinoco
and Amazon rivers are presented in Table 4. Di€erences between rivers relate
to catchment types, especially the contrast between white-water (VaÂrzea) and
black-water (IgapoÂ) rivers. The data suggest that in ¯ooded communities,
within similar water types, the number of species (alpha or local diversity)
declines along gradients of increasing ¯ooding depth and duration. Also sim-
ilar habitats in Igapo ¯oodplain forests along the Mapire and Caura rivers in
the Orinoco basin (unpublished data) and along the Jau and Taruma-Mirim
rivers in central Amazonia did not have substantially di€erent species richness
(Ferreira pers. comm.). Abundant tree species showed distinctive local com-
munities and variable forest structures were evident. There was species overlap
within topographic levels and river sites. Ferreira proposes that this may re¯ect
di€erences in ¯ood tolerance among abundant species and local resilience and
persistence among the other plant species. While the former pattern was ex-
pected, the latter pattern is evidence that plant species richness in ¯oodplain
forests may be in¯uenced by a more complex suite of factors than has previ-
ously been suggested.
Prance (1979, 1982) states that Igapo forests have lower alpha diversity than
VaÂrzea forests probably re¯ecting nutrient status. However, the data presented
herein suggests that this is not always the case. Worbes (1997) gives further
support to this observation. Because the examples compared are taken from
di€erent regions and ¯ooding depth and duration or the successional stage are
not speci®ed in all the examples, conclusions about the biogeochemical e€ect
can not be reached. It is also crucial to di€erentiate the level of diversity
observed in the data. If higher di€erentiation has occurred in the Igapo ¯ooded
forests, a higher gamma (regional) diversity could be expected, however
not necessarily a higher alpha (local) or beta (inter-habitats) diversity.
568

Table 4. Woody species richness in di€erent ¯ooded forests of the Orinoco and the Amazon basins.

Sampled area
Forest community N. Species (ha)

Orinoco basin
VaÂrzea forests
1. Middle Orinoco, Laguna de Mamo (Colonello 1990a),
ind. >2 m height, di€erences in geomorphic position
levee 1 21 0.05
levee 2 17 0.05
depression 1 11 0.05
depression 2 11 0.05
Total 34 0.2
Igapo forests
1. Mapire con¯uence in the Orinoco (see Figure 3)
Middle and upper reaches (Rosales 1990) >5 cm height,
e€ects of location
Community 1, upstream sedimentation area 37 0.06
Community 2, stabilised point-bars,, ¯ooding 5±8 m 19 0.07
Community 3, middle zone (see Figure 3), overbank swamp, 36 0.10
¯ooding 3±5 m
Community 4, middle zone, overbank swamp, ecotone 26 0.10
forest-savanna, ¯ooding 1±3 m
Total species number 56 0.33
2. Lower Caura R. Between Los Raudales de la Mura and
Salto Para (Rosales et al. 1997) >10 cm DBH,
averages of alpha-diversity by elevation
Levees ParaÂ-Nichare, infrequently ¯ooded, max. 50 cm 32 (2.73) 7 plots (0.1 ha)
Levees Nichare-la Mura, infrequently ¯ooded, max. 50 cm 30 (5.79) 5 plots (0.1 ha)
Sandy levees Para-Nichare, frequently ¯ooded, max. 1 m 32 (8.63) 4 plots (0.1 ha)
Backswamps Para-Nichare, frequently ¯ooded, 3±5 m 14 (5.22) 5 plots (0.1 ha)
Backswamps Nichare-la Mura, frequently ¯ooded, 3±5 m 20 (2.0) 4 plots (0.1 ha)
Total species number 190 2.8
VaÂrzea to IgapoÂ?
1. Lower Caura R. downstream Los Raudales de La Mura
(Rosales unpublished data) >10 cm DBH (¯ooding
depth 3 to 10 m)
Total species number 41 2.2
Amazon basin
VaÂrzea forests
1. Jurua river, Acre (Campbell et al. (1992)) >10 cm DBH,
e€ects of age
50 year old 106 1.0
14±50 year old 73 1.0
14 year old 20 1.0
Igapo forests
1. Lower Rio Negro (Rodriguez 1961) >1 m height 51 0.035
2. Lower Negro river (Keel and Prance 1979) >1.0 m height 54 1.8
3. Jau National Park (Ferreira 1997a), >10 cm DBH,
e€ects of ¯ooding intensity
depth 8±9 m, 260±270 days 44 1.0
depth 4.6±5 m, 150±177 days 103 1.0
depth 1.5±3.2 m, 65±81 days 137 1.0
 569

Table 4. (Continued)

Sampled area
Forest community N. Species (ha)

4. Xingu river, Para (Campbell et al. 1986) >10 cm DBH 40 0.5

Mixed forests
1. Lower Negro River (do Amaral et al. 1997) >10 cm DAP 66 0.5

Furthermore, a recent work by Ferreira (1997a, 1997b) concluded that for trees
with a minimum DBH of 10 cm, no signi®cant di€erence could be found in
either species richness or tree density between the two ¯oodplain forest types.
Age, related to successional processes, is also a factor in¯uencing diversity;
species alpha diversity increases towards the oldest ¯uvial environments (Salo
et al. 1986; Campbell et al. 1992; Worbes et al. 1992). However, age is also
associated with elevation above the channel along the successional sequences.
Rates of ¯oodplain accretion and overbank sedimentation have to be con-
sidered when successional sequences are analysed. Di€erent successional
stages in a given region might a€ect beta diversity (Puhakka et al. 1992;
Puhakka and Kalliola 1993, 1995). For example, high beta diversity of ¯ooded
forests in the Upper Amazon is explained by Salo et al. (1986) as related to the
successional character of the habitats that follow high disturbance patterns
induced by channel migration. However, the lateral migration of meanders
reduces species-richness and favours rejuvenation through the establishment
of younger successional stages, which have fewer species. Rapid overbank
sedimentation creates more homogeneous edaphic conditions along low gra-
dient white-water ¯oodplains resulting in a low alpha diversity. Kalliola
(1992) states that for the Upper Amazon, ¯oodplains of the most unstable
rivers are also rich in di€erent habitat types. Nevertheless, due to high dis-
turbance intensities, the number of species occupying the habitats may be
relatively low (i.e., severe external disturbance reduces the number of species).
Salo et al. (1986) showed that high beta and gamma diversity of relatively
small white-water ¯oodplain rivers like the Tambopata and Manu in Peru, are
linked with high habitat complexity (oxbow lakes, ¯oodplain depressions and
ridges, river terraces).
Three qualitative factors are then important to explain the patterns of the
riparian forests species diversity in the Orinoco and Amazon river basins: (i)
uniqueness, which relates to the proportion of unique taxa (endemic or rare at
the regional level) associated with unique habitats, (ii) succession which relates
to the dominance of community stages, and (iii) `terrestrialization' or upland
similarity which relates to the ¯oristic resemblance to non-riparian forest
ecosystems dominating the surrounding uplands. The driving forces control-
ling these factors are ¯ooding intensity (i.e. ¯ood duration and depth), channel
570

dynamics (i.e. rate of channel change), and fertility (i.e. nutrient status of the
riparian forest soils). Based on the current theoretical framework around the
ecological signi®cance of disturbance, Junk (1997a) explains how di€erent
levels of riparian ¯ow predictability allow the development of strategies to deal
with environmental changes over evolutionary timescales. Over ecological
timescales it means that established species will only be those that have already
developed strategies to cope with speci®c environments. Along an axis of
predictability, there will be di€erences in the general trends of alpha, beta, and
gamma biodiversity. Intermediate to long-term variability in discharge and/or
sediment loads at di€erent recurrence intervals (from short to long-term) will
a€ect ¯ood timing and then predictability in ¯ooding intensity, i.e. ¯ood du-
ration and depth in each di€erent habitat; either e€ect will occur in channel
dynamics (i.e. predictability in the rates of channel migration) or fertility (i.e.
changes form Igapo to VaÂrzea conditions). These gradients of predictability
can be important in explaining the higher biodiversity associated with tributary
junctions and ¯oodplain-upland ecotones.

Conclusions

A considerable scienti®c e€ort has been expended on describing the ¯oristic,

structural and functional complexity of the riparian forests in speci®c loca-
tions of the Orinoco and Amazon River basins. However, there have been
few attempts to explain the observed patterns in relation to physico-chemi-
cal, geomorphological and hydrological changes along and between rivers.
Such explanations must address di€erent spatio-temporal scales. At the re-
gional scale, contrasts between the Orinoco and the Amazon riparian ¯ooded
forests re¯ect di€erences in climate and geology, and river ¯ows and sedi-
ment loads, but the processes underlying the maintenance of the riparian
ecosystems are similar. Di€erences between the VaÂrzea and Igapo riparian
forests re¯ect trophic conditions, but within each (i.e. at the spatial scale of a
reach), ¯oristic di€erences are related to the hydrological and geomorpho-
logic setting.
The riparian forests of the Orinoco and Amazon basins are currently
threatened as a result of increased human pressure. Biodiversity conservation
and sustainable management are important and urgent medium-term goals.
Nevertheless, in both basins few areas have been selected for the protection of
riparian forests of the lowlands. Although the political division of the Orinoco
and Amazon basins includes countries that are signatories of the Ramsar
Wetlands Convention (IUCN 1971), Ramsar sites for the Orinoco basin for
example include mainly coastal or marine environments. However, the im-
portance of inland water ecosystems for biodiversity conservation and a need
 571

for a watershed approach has been strengthened after the resolutions approved
in the Fourth Meeting of the Conference of the Parties COP4th (Secretariat of
the Convention of Biological Diversity 1998). Therefore, conservationists and
managers have to consider appropriate schemes for conserving and managing
riparian biodiversity.
We have shown that riparian corridors in the Orinoco and Amazon river
basins are highly productive in biologically diverse landscape units. An analysis
of the biota-environment relationships and historical patterns synthesised in
this paper can help to identify areas for conservation. If these relationships
are further quanti®ed, geochemical, hydrological and geomorphological sur-
rogates can be used to aid the conservationist or manager in selecting reserves
and in managing development or restoration operations. Such information
is needed, for example, to enable the prediction of biodiversity values for
remote sites or to derive sustainable operational rules for ¯ow regulation
schemes.
The importance of the di€erent forest communities in terms of local rarity of
the forest habitats has to be further evaluated. Less frequent habitats should be
assigned higher values of importance even if their values of species biodiversity
are not particularly high. Analysis of the representativeness of the selected sites
for the inclusion of potential ¯agships, umbrellas or keystone species should
also be considered, as well as their value for economic, endangered or vul-
nerable species, or for those aspects concerning cultural or religious importance
of the areas. Ethically, and even perhaps more importantly, the value of
¯ooded forest resources to indigenous and local communities must be con-
sidered above the mere assignment of diversity numbers to di€erent areas.
Future basic research should be interdisciplinary and focus on interbasin
comparisons along longitudinal gradients to study: (i) the in¯uence of hydro-
geomorphological dynamics upon successional patterns and beta diversity, and
(ii) the role of fertility and disturbance in explaining taxonomic or genetic
diversity of selected taxa or functional groups which can be of proved im-
portance for conservation (i.e. keystone, economic, restoration species).

Acknowledgements

Prof. Ghillean Prance reviewed the ®rst drafts of this manuscript and allowed
access to his own botanical data and relevant literature for the Amazon. Bruce
Holst and Lourdes Rico reviewed the Myrtaceae and Mimosaceae, respec-
tively, occurring in the Amazon and Orinoco, and Paul Berry, Stanley Greg-
ory, Otto Huber, Claudia Knab-Vispo, and Conrad Vispo made helpful
comments on the manuscript. Leandro Ferreira provided useful information
and valuable comments on the ®nal paper.
572

Appendix. List of species common to both the Venezuelan Orinoco and the Central and Upper
Amazon riparian forests. Location of herbarium specimens for selected sectors in the Venezuelan
Orinoco basin: 1 = Caura R., 2 = Mapire R., 3 = CaronõÂ R., 4 = rivers in the High Plains
(Llanos altos), 5 = Cataniapo R., 6 = Lower Orinoco, 7 = rivers in the Low Plains (Llanos
bajos, Apure), 8 = rivers in the Venezuela Amazonas State, 9 = Atabapo R.

Species Location

ANACARDIACEAE
Anacardium giganteum Hancock ex Engler 1
Astronium lecointei Ducke 1
Spondias mombin L. 1, 2, 4, 6, 7
Tapirira guianensis Aubl. 1, 2, 4, 6, 7, 9
ANNONACEAE
Cymbopetalum brasiliense (Vell.) Benth. 1, 3
Duguetia cauli¯ora R.E.Fries 1, 3, 8
Pseudoxandra polyphleba (Diels) R.E. Fr. 1, 8
Xylopia aromatica Lam. 1, 2, 3, 4, 8
Xylopia sericea A. St.-Hil. 3, 9
APOCYNACEAE
Malouetia glandulifera Miers 9
Malouetia tamaquarina (Aubl.) A. DC. 8, 9
Malouetia virescens Spruce ex Muell. Arg. 8, 9
Parahancornia negroensis Monach. 9
Tabernaemontana macrocalyx MuÈll. Arg. 1, 8
Tabernaemontana siphilitica (L. f.) Leeuwenb. 1, 8
Tabernaemontana undulata Vahl 1, 3, 8
ARACEAE
Montrichardia arborescens (l.) Schott 1, 3, 4, 5, 8, 9
ARECACEAE
Astrocaryum chambira Burret 1, 2
Astrocaryum jauari Mart. 1, 7
Attalea maripa (Aubl.) Mart. 1
Bactris maraja Mart. sin. (Bactris brongnartii Mart.) 1, 2
Euterpe precatoria Mart. var. precatoria 1, 7
Leopoldinia pulchra Mart . 9
Leopoldinia piassaba Wallace 9
Mauritia ¯exuosa L. f. 1, 2, 3, 4
Mauritiella aculeata (Kunth) Burret 1, 3, 8, 9
Oenocarpus bataua Mart. sin (jessenia bataua (Mart.) Burret 1
Socratea exorrhiza (Mart.) H. Wend. 1, 8
BIGNONIACEAE
Crescentia amazonica Ducke 1, 2, 4, 6, 7
Jacaranda copaia subsp. spectabilis (Mart.ex DC.) A.H. Gentry 1, 3
Jacaranda obtusifolia Bonpl. 1, 3, 4, 5, 7
Tabebuia barbata (E. Mey.) Sandwith 8, 9
Tabebuia capitata (Bureau & K. Schum.) Sandwith 1, 3, 8, 9
BIXACEAE
Bixa urucurana Willd. 1, 10
BOMBACACEAE
Ceiba pentandra (L.) Gaertn. 1, 2, 3, 4, 6, 7, 8
Pachira aquatica Aubl. 1, 5, 9
 573

Appendix. (Continued)

Species Location

BORAGINACEAE
Cordia nodosa L. 1, 3
Cordia tetrandra Aubl. 1, 2, 6
BURSERACEAE
Protium heptaphyllum (Aublet) Marchand 1, 3, 4, 8
Protium trifoliolatum Engl. 1, 8
Protium unifoliatum Engl. 1
CAESALPINIACEAE 1
Campsiandra comosa Benth. 3, 6, 7
Campsiandra laurifolia Benth. sin. C. comosa var. laurifolia 2, 3
Cassia grandis L. f. 3
Cynometra bauhiniifolia Benth. 1, 3
Dialium guianense (Aubl.) Steud. 1, 3
Macrolobium acaciifolium (Benth.) Benth. 1, 3, 6, 7
Macrolobium bifolium (Aubl.) Pers. 1, 3, 4, 5, 8
Macrolobium angustifolium (Benth.) Cowan 1, 8, 9
Macrolobium multijugum (DC.) Benth. 1, 2, 6, 9
Macrolobium suaveolens Spruce ex Benth. 8
Sclerolobium guianense Benth. 1, 2, 3, 8
Senna reticulata (Willd.) H.S. Irwin & Barneby 1
CARYOCARACEAE
Caryocar microcarpum Ducke 1, 5, 8
CHRYSOBALANACEAE
Couepia paraensis ssp. glaucescens (Spr.ex Hook f.) Prance 1, 2, 8
Hirtella racemosa var. hexandra (Willd. ex Roem. & Schultz) Prance 2, 8
Licania apetala (E.Mey.) Fritsch. var. aperta 4, 6, 7, 8, 9
Licania apetala (E.Mey.) Fritsch. var. apetala 1, 2, 4, 8, 9
Licania heteromorpha (C. Mart. ex Hook. f.) Benth. 8, 9
Licania parvi¯ora Benth. 8
Parinari excelsa Sabine 1, 3, 5
CLUSIACEAE
Callophyllum brasiliense Cambess. 1, 8, 9
Calophyllum piaroanum 5
Caraipa densifolia Mart. subsp. densifolia 1, 3, 8
Clusia grandi¯ora Splitg. 1, 3, 8
Garcinia macrophylla Mart. 1
Mahurea exstipulata Benth. 1, 3
Symphonia globulifera L. f. 3, 4, 8
Vismia cayennensis (Jacq.) Pers. 1, 3
Vismia japurensis Reich. in Mart. 1, 5, 8
Vismia macrophylla Kunth 1, 8
COMBRETACEAE
Buchenavia oxycarpa (Mart.) Eichl. 1, 2
Buchenavia reticulata Eichler 5
Buchenavia suaveolens Eichler 8
Terminalia amazonia (J. F. Gmel.) Exell 1, 3, 8
574

Appendix. (Continued)

Species Location

CONNARACEAE
Connarus ruber (Poepp.) Planch. 1
ERYTHROXYLACEAE
Erythroxylum mucronatum Benth. 3, 5, 8,
EUPHORBIACEAE
Alchornea castaneifolia (Willd.) Ad. Juss. 1, 2
Alchornea schomburgkii Kl. 1, 2, 6, 8
Alchornea triplinervia (Spreng.) M. Arg. 1
Amanoa guianensis Aubl. 1, 2, 5
Conceveiba guianensis Aubl. 1, 8, 9
Conceveiba latifolia Benth. 8
Croton cuneatus Kl. 1
Drypetes variabilis Uitt. vel. sp. a€. 1
Mabea nitida Spruce ex Benth. 1, 2, 3, 7
Mabea taquari Aublet 1, 3
Maprounea guianensis Aubl. 1, 5, 8
Margaritaria nobilis L.f. 1, 3, 7
Micrandra minor Benth. 1
Micrandra siphonoides Benth. 1
Pera decipiens (Muell.Arg.) Muell. Arg. 1, 9
Pera distichophylla (Mart.) Baill. 9
Piranhea trifoliata Baill. 1, 2, 3, 6, 7
FABACEAE
Acosmium nitens (Vog.) Yakov. 1, 2, 3, 4, 5, 7, 9
Aldina latifolia Spruce ex Benth. 9
Andira inermis (Wright.) Kunth ex DC. 2
Andira surinamensis (Bondt.) Splitg. ex Pulle 1, 3, 5, 8
Dalbergia amazonica (Radlk. ex Kop€) Ducke 1
Dalbergia ecatasphyllum (L.) Taub. 3
Dalbergia glauca (Desv.) Amsh. 1, 2, 6, 7
Dalbergia monetaria var. hygrophylla (Mart. ex Benth.) Macbr. 1, 2, 5
Dalbergia inundata Spruce ex Benth. 2, 8
Dalbergia monetaria L. f. 1, 8
Machaerium inundatum (Mart. ex Benth.) Ducke 3, 5, 6, 7
Machaerium aristulatum (Benth.) Ducke 2
Swartzia cardiosperma Benth. 2, 9
Swartzia laevicarpa Amsho€ 3
Swartzia leptopetala Benth. 1, 7
Swartzia polyphylla DC. 3
Swartzia sericea Vogel 8, 9
Vatairea guianensis Aubl. 9
FLACOURTIACEAE
Banara guianensis Aubl. 8
Casearia aculeata Jacq. 1, 9
Homalium guianense (Aubl.) Oken 1, 3
Homalium racemosum Jacq. 1, 2, 4, 5, 6
Lindackeria paludosa (Benth.) Gilg. 1, 8
Ryania speciosa Vahl 1, 5, 7
 575

Appendix. (Continued)

Species Location

ICACINACEAE
Discophora guianensis Miers 1, 8
LACISTEMATACEAE
Lacistema aggregatum (Berg.) Rusby 1
LAURACEAE
Aniba anis (Meisn.) Mez 8
Ocotea bofo Kunth 1, 2, 5, 6, 8
Ocotea cymbarum H.B.K. 1, 2, 6, 7, 8
Ocotea esmeraldana Moldenke 8, 9
Ocotea pauci¯ora (Nees) Mez 9
LECYTHIDACEAE
Couratari guianensis Aubl . 1, 3
Eschweilera coriacea (DC.) S.A. Mori 3, 8
Eschweilera pedicellata (Richard) Mori 1, 8
Eschweilera subglandulosa (Steud. ex Berg) Miers 1, 3, 5
Eschweilera tenuifolia (Berg.) Miers 1, 2, 9
Gustavia augusta L. 1, 2, 3, 6, 7
Gustavia coriacea Mori 1
Gustavia poeppigiana Berg. 1, 6
Gustavia pulchra Miers 5
Lecythis corrugata Poit ssp. rosea (Spuce ex Berg.) Mori 1, 5, 9
LOGANIACEAE
Strychnos panurensis Sprague & Sandw. 1
MALPIGHIACEAE
Burdachia prismatocarpa C. Martius ex Adr. Jussieu 9
Byrsonima japurensis Adr. Juss 1, 2, 7
Byrsonima leucophlebia Griseb . 3
Byrsonima stipulacea Adr. Juss. 1, 3
MELASTOMATACEAE
Bellucia grossularioides (L.) Triana 1
Bellucia pentamera Naudim 1
Mouriri acuti¯ora Naudim 1
Mouriri sagotiana Triana 1
Mouriri guianensis Aubl. 6
MELIACEAE
Guarea guidonia (L.) Sleumer 1
Trichilia inaequilatera Penn. 1, 6, 7
Trichilia mazanensis Macbr. 1, 2, 7, 8
Trichilia micrantha Benth. 9
Trichilia quadrijuga Kunth ssp. quadrijug 1
MIMOSACEAE
Abarema jupunpa (Willd.) Briton & Killip Sin. 1, 3, 8
(Pithecellobium jupunba (Willd.) Urb.)
Albizia corymbosa (Rich.) G.P.Lewis & Owen.
Sin. Pithecellobium corymbosum (Rich.) Benth. 1, 2, 5
Inga alba (Sw.) Willd 1, 3, 8
Inga bourgoni Scop 1
576

Appendix. (Continued)

Species Location

Inga capitata Desv. 1

Inga coruscans Humb. & Bonpl. ex Willd. 1
Inga edulis Mart. 1
Inga leiocalycina Benth. 1
Inga nobilis Willd. 1, 9
Inga pilosula (Rich.) Macbr. 1
Inga splendens Willd. 1
Inga thibaudiana D.C. 1
Inga umbellifera (Vahl) Steud.? 1
Inga vera Willd. 1, 2, 7
Parkia discolor Spruce ex Benth. 9
Parkia pendula (Willd.) Benth. ex Walp. 1, 3
Pentaclethra macroloba (Willd.) O. Kuntze 1
Pithecellobium divaricatum Benth. Sin. Zygia divaricata Benth. 2, 7
Pithecellobium cauli¯orum (Willd.) Mart. or. Zygia 1, 3, 7
latifolia var. communis Barneby & Grimes
Zygia unifoliolata Benth. Pitt. Sin. Pithecellobium unifoliolatum Benth. 1, 2
Zygia cataractae (Kunth.) L. Rico 1, 2, 3, 8
MORACEAE
Brosimum guianense (Aubl.) Huber 1, 3
Cecropia latiloba Miq. 1, 7
Pseudolmedia laevigata Trecul 1, 8
Maquira coriacea (Karst.) C.C.Berg 1, 7, 9
MYRISTICACEAE
Virola calophylla (Spruce) Warb. 9
Virola elongata (Benth.) Warb., s.l. 1, 3, 8, 9
Virola surinamensis (Rol.) Warb. 1, 3, 7, 8
MYRTACEAE
Calyptranthes fasciculata Berg 1, 3, 8
Calyptranthes multi¯ora Berg. 1, 2, 3, 7, 8
Calyptranthes pullei var. immaculata Mc Vaugh 1, 3, 4, 6, 7
Eugenia bi¯ora (L.)DC. 1, 3, 4
Eugenia egensis DC. 1, 8
Eugenia feijoi Berg. 1, 8
Eugenia ¯avescens DC. 1
Eugenia ¯orida DC. 1, 3, 8
Eugenia lambertiana var. lambertiana DC. 1, 5, 8
Eugenia patrisii Vahl 3, 5, 8
Eugenia punicifolia (H.B.K.) DC. 1, 3, 4, 7, 8
Eugenia tapacumensis Berg 3, 7
Myrcia bracteata (Rich.) DC. 1, 3, 8
Myrcia inaequiloba (DC.) Legrand 3, 8
Myrcia de¯exa (Poir.) DC. 5, 8
Myrcia grandis Mc Vaugh 9
Myrcia pyrifolia (Desv. ex Ham.) Nied. 1, 3, 8
Myrcia splendens (Sw.) DC. 1, 8
Myrciaria dubia (H.B.K.) Mc Vaugh 1, 2, 3, 4, 7, 8
Myrciaria ¯oribunda (West ex Willd.) O. Berg. 1, 5, 7, 8
Psidium maribense Mart. ex DC. 2, 4, 6, 7
 577

Appendix. (Continued)

Species Location

Psidium densicomum DC. (=P. ovatifolium Berg.) 2, 7

Psidium persoonii McVaugh 1, 3, 7
OCHNACEAE
Blastemanthus gemmi¯orus (Mart.) Planch. 9
Ouratea spruceana Engl. 9
Wallacea insignis Spruce ex Benth. 9
OLACACEAE
Cathedra acuminata (Benth.) Miers 1, 2, 8, 9
Chaunochiton loranthoides Benth. 9
Dulacia candida (Poepp.) Kuntze. 2, 9
Minquartia guianensis Aubl. 3, 9
POLYGONACEAE
Coccoloba excelsa Benth. 3, 5, 8, 9
Coccoloba marginata Benth. 1, 3
Ruprechtia tenui¯ora Benth. 1, 2, 6, 7
Symmeria paniculata Benth. 1, 2, 6, 7, 8
Triplaris americana L. 1, 7, 8
PROTEACEAE
Panopsis rubescens (Pohl) Rusby 1, 2, 3
Roupala obtusata Klotzsch 9
QUIINACEAE
Quiina rhytidopus Tul. 2
RUBIACEAE
Aliberia latifolia (Benth.) Schumm. vel.sp.a€. 3, 7
Duroia genipoides Hook. f. ex Schumann 5, 8
Duroia micrantha (Ladbr.) Zarucchi & Krkbride 1, 8
Duroia sprucei Rusby 2, 7
Genipa americana L. 1, 3
Pagamea coriacea Spruce ex Benth . 8, 9
Psychotria capitata R & P 1
Psychotria lupulina Benth . 1, 8
Simira rubescens (Benth.) Bremek. ex. Steyerm. Sin. 5, 8
(Sickingia tinctoria (Kunth) Schumann)
Stachyarrhena spicata Hook. f. 8, 9
SAPINDACEAE
Cupania cinerea Poepp. 1
Matayba elegans Radlk.'' 1, 8, 9
SAPOTACEAE
Ecclinusa guianensis Eyma 1
Elaeoluma glabrescens (Mart. & Eichler)AubreÂv. 8
Micropholis egensis (A. DC.) Pierre 1, 2, 9
Pouteria elegans (A. DC.) Baehni. Sin. 9
(Neoxythece elegans, Sideroxylon elegans)
Pouteria reticulata (Eyl.) Eym. ssp. reticulata 1, 2, 8
SIMAROUBACEAE
Simaba guianensis Aubl. 1, 7
Simaba multi¯ora H.B.K. 6
578

Appendix. (Continued)

Species Location

Simaba obovata Spruce 9

Simaba orinocensis Kunth 2
STERCULIACEAE
Guazuma ulmifolia Lam. 1, 2, 3, 4, 6
TILIACEAE
Mollia speciosa Mart. 8, 9
VIOLACEAE
Leonia glycycarpa Ruiz & Pav. 1, 3, 9
VOCHYSIACEA
Qualea retusa Spruce ex Warming 9
Vochysia obscura Warm. 9

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