Professional Documents
Culture Documents
The work presented in this Thesis was mainly funded by Tropenbos Colombia, the Netherlands Organisation for International Cooperation in
Higher Education (NUFFIC) and COLCIENCIAS. Research was
conducted at the Institute for Biodiversity and Ecosystem Dynamics
(IBED), Universiteit van Amsterdam.
ISBN: 978-90-76894-97-3
Lay-out: Ana Catalina Londoo Vega.
Cover: Canopy of the upland plot (left); young leaves of Pseudomonotes
tropenbosii (centre); canopy of the floodplain plot (right).
No part of this dissertation may be reproduced in any format, by print,
photocopy, microfilm or by any other means without the written
permission from the author.
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Contents
1.
Introduction
2.
17
3.
31
49
85
4.
5.
6.
7.
111
Synthesis
141
Literature
149
179
Appendices
191
Agradecimientos
235
Curriculum vitae
239
1
Introduction
Ana Catalina Londoo Vega
1.1
The adequate conservation of tropical forests and wise use of their natural
resources depend to a large extent on our knowledge of the variation in
these forests, both in space and in time. Long-term studies in permanent
forest plots consisting of repeated surveys in well-demarcated and georeferenced plots, allow the quantification of the temporal variation of
forest resources (Campbell et al. 2002). Non-permanent plots to describe
Amazonian forests along environmental gradients have been used
successfully in exploratory surveys (Gentry 1988b; Duivenvoorden and
10
Lips 1993; ter Steege et al. 2003; Duque 2004; Benavides 2010). Yet, such
plots have the disadvantage that they, similar to a photograph, only
provide a limited view at a certain time. In contrast, permanent plots not
only admit a detailed description of the habitat at a particular site, but also
allow detecting temporal changes in the forest. Such information is
indispensable to predict future changes in the diversity and distribution of
species (Losos and Leigh 2004). Beyond the initial inventory of the
vegetation in permanent plots it is necessary to measure forest changes in
the long run, through continuous monitoring of the floristic composition,
structure, growth, mortality and survival of species, preferably at more
than one site (Comiskey et al. 1999).
Permanent plots have been used extensively to study long-term changes of
vegetation and the natural processes that allow the coexistence of species
in relation to the environment (Bakker et al. 1996; Herben 1996; Rees et
al. 2001; Hubbell 2004). Early studies conducted in tropical forests
primarily aimed at measuring the diameter growth for silvicultural
management and timber harvesting (Bell 1971). Recent studies include
other objectives such as quantifying carbon stocks and their relation to
global flows (Dallmeier 1992; Phillips et al. 1998; Orrego et al. 2003). The
oldest plots in tropical forests stem from the first part of the twentieth
century, and were established in Philippine dipterocarp forests (Richards
1952), on the islands of Trinidad and Tobago (Bell 1971), in Peninsular
Malaysia (Manokaran and Swaine 1994) and in Uganda, Africa (Sheil et
al. 2000). The largest plots (25 to 50 ha) are now part of the CTFS network
(Center for Tropical Forest Science) of the Smithsonian Tropical Research
Institute (STRI) (Losos and Leigh 2004), which manages, for example, the
well-known plot on Barro Colorado Island (Panama). Another network is
that by the Amazon Forest Inventory Network (RAINFOR) (Malhi et al.
2002; Malhi and Phillips 2004), which mostly supports 1-ha plots.
In Colombia, permanent forest plots were initially installed to monitor
growth of timber species like Cupressus lusitanica (Tschinkel 1972; del
Valle 1979), Cordia alliodora, Tectona grandis, Eucalyptus saligna, and
Cariniana pyriformis. In the 1980's permanent forest plots were started in
the Choc region (western Colombia) (Gonzlez et al. 1991; Gonzlez
1995; del Valle 1996a, 1998ab, 1999) and Amazonia (Londoo 1993, this
dissertation). In 2000 about 70 permanent plots were being maintained
11
(Vallejo et al. 2005; lvarez et al. 2008). By mid 2006, these plots covered
at least 100 ha. Overall, the two plots described in this dissertation have
particular significance as they are among the oldest permanent forest plots
in Colombian Amazonia, and in Colombia as a whole.
1.3
Floristic and ecological research in the permanent forest plots at
Pea Roja
Between 1986 and 2000 the Tropenbos-Colombia Programme supported
forest ecological research in the mid catchment of the Caquet River, with
its epicenter in Araracuara (Fig. 1-1). The programme started with a land
unit survey of a large area (10,000 km2), east of Araracuara. Based on the
ecological maps (Duivenvoorden and Lips 1993, 1995) in late 1988
several sites were selected as representative for the principal land units, in
order to develop long-term monitoring studies of forest ecosystem
functioning. The following selection criteria were applied: a)
Representation: the land unit should have a wide cover in the Middle
Caquet area and in the Colombian Amazon as a whole, b) Accessibility:
the site should be well accessible by river from Araracuara to allow
efficient transport of personnel and equipment, c) Control, surveillance
and security: sites should offer some protection for permanently installed
research equipment (Duivenvoorden and Lips 1990). The sites selected
were located in the area of Nonuya community of Pea Roja. One site
belonged to the land unit classified as Tertiary Sedimentary Plain and the
other site was located on a rarely inundated flood plain of the Caquet
River (Duivenvoorden and Lips 1990). With all parties involved it was
agreed to develop a long-term research on the flora, structure and
dynamics of the forest vegetation at these two monitoring sites. At each
site a permanent plot of 1.8 ha was established for the long-term study of
the forest. Uniform criteria were used for the establishment, i.e. the sites
were chosen in such a way that very large (multiple tree gaps by wind
throw) canopy gaps were avoided. Also, the physiography, forest
(Duivenvoorden et al. 1988) and soils were as homogeneous as possible.
From that moment on, in and outside these plots, a series of ecological
studies have been successfully carried out by students from Colombia and
other nationalities, for example those regarding plant-animal relationships
12
(van Dulmen 2001; Parrado 2005), hydrology (Tllez 2003) and nutrient
cycling (Tobn 1999; Tobn et al. 2004ab).
one of the most dominant, species rich, and economically important tree
families in tropical forests of Asia (Whitmore 1975; Ashton 1982), was
only represented by one single species in the Neotropics (Pakaraimaea
dipterocarpacea Maguire & Ashton recorded in Guyana). The record of
Pseudomonotes tropenbosii Londoo, lvarez and Forero in the upland
plot near Pea Roja along the Caquet river emphasizes the
phytogeographical links of Colombian Amazonia with the Guiana Shield
region (Duivenvoorden and Lips 1995) and even with Africa and
Madagascar (via the subfamily Monotoideae; Morton 1995), and
underscores the unique position and value of this permanent plot.
The architecture of a plant is the morphological expression of its genome
in a given time, and a result of the balance between endogenous growth
processes and exogenous constraints exerted by the environment
(Barthlmy et al. 1991; Hall 1995). The objective of architectural
analysis is to identify the endogenous processes that control growth and
shape of the whole plant, by means of observation. Three key concepts
have been developed: the architectural model (Hall and Oldeman 1970),
the architectural unit (delin 1977) and the reiteration (Oldeman 1974).
The concepts of model and reiteration unit have provided a valuable tool
for studying the structure and form of plants. Plant architecture further
determines how resources are allocated in plants, for example, regarding
the capture of light, water transport, mechanical stability, and resistance to
wind (Vester 1997; Poorter and Werger 1999). Vester (1997) analyzed
successive architectural development of tree taxa in successional forests
near Araracuara. He showed that on the basis of detailed observations,
architectural analysis yields understanding of the mechanisms of
succession in secondary forest or in old-growth forests. Architectural
analysis thus has the potential to supply significant information regarding
forest dynamics in permanent plots, to complement the demographic
information of recruitment, growth and mortality.
Arguably, two principal results emerged from the ecological surveys of the
Middle Caquet area (Duivenvoorden and Lips 1993, 1995; Urrego 1997;
Quiones 2002; Duque 2004; Snchez 2005; Benavides 2010). The first
was the clear recognition of the high tree species diversity per unit of area
(Valencia et al. 1994; Duivenvoorden 1996), which ranks among the
highest level for tropical forests worldwide. Several explanations for the
14
high upper Amazonian tree species diversity have been put forward
(Duivenvoorden 1995; ter Steege et al. 2006). In most of these, the unique
properties of the geological setting (speciation triggered by the unique
configuration and the highly dynamic environment in the Tertiary; Hoorn
and Wesselingh 2010) and the climatic history of northwestern Amazonia
(continuous high humidity throughout the Pleistocene favouring species
survival; Lips and Duivenvoorden 1994; Mayle et al. 2004) play a key
role. The second result was the evidence and documentation that forest
composition and forest diversity systematically changed across land units,
which differed regarding flooding, drainage and soil nutrient
concentrations. High levels of forest diversity were consistently recorded
in the well-drained uplands (Duivenvoorden 1996; Duivenvoorden and
Duque 2010). On the other hand, poor drainage, seasonal flooding and
extremely low levels of soil nutrient availability were always associated
with low levels of tree species diversity (Urrego 1997; Duivenvoorden and
Duque 2010). Furthermore, litterfall measurements in combination with
studies of the standing stock of litter on the forest floor indicated that the
aboveground decomposition of organic matter (a proxy of net primary
above-ground forest productivity; Vogt et al. 1998) was substantially
higher in floodplains of the Caquet than in well-drained uplands (Lips
and Duivenvoorden 1996). Summarizing, in view of the physiographic
setting of the two permanent forest plots at Pea Roja, a general picture
emerged from the ecological survey that soil fertility, forest disturbances,
and above-ground forest productivity in floodplains of the Caquet River
were higher than in the well-drained uplands. However, because these
survey results were not based on permanent plots, information about forest
dynamics could not be incorporated. In more recent studies in Amazonia at
continental scales, forest dynamics have been correlated with forest
diversity. Wide-scale correlations between tree mortality (Phillips et al.
2004), wood specific gravity (Baker et al. 2004), above-ground biomass
(Baker et al. 2004; Mahli et al. 2006), above-ground wood productivity
(Malhi et al. 2006) and tree species diversity (ter Steege et al. 2006) have
been reported along geological gradients from eastern Amazonia to upper
Amazonian watersheds. In short, low diversity forests were associated to
low levels of forest dynamics (low mortality, low productivity) and low
levels of soil fertility, whereas high diversity forests showed a relatively
15
high mortality (and high wood productivity) and were found on relatively
rich soils. Is this general scheme, obtained from Amazonia as a whole,
useful to predict forest dynamics and forest diversity in a spatially more
limited area like that of Colombian Amazonia?
1.4
16
2
Site description of the upland and
floodplain plots
Ana Catalina Londoo Vega
2.1
Climate
The information about the current climate in the study area is based on two
meteorological stations: one in Araracuara and one in Pea Roja. At
Araracuara daily records were available from 1979-1990 (station code
4413501 located at 160 m altitude above sea level; Duivenvoorden and
Lips 1995). Climate data from Pea Roja were automatically recorded
every 20 minutes between November 1992 and August 1997 (Tobn 1999)
at a station located at 039' S and 725' W at 150 m above sea level, at 2.5
km distance from the upland and floodplain plots. Disregarding slight
differences between the records from both climate stations, which can be
attributed to the different length of the registration period, the climate of
this part of the Middle Caquet Basin is characterized by an annual rainfall
of ca. 3100 mm, an average yearly temperature of 25C and a relative
humidity of 87% (Fig. 2-2). This climate is classified as Afi, equatorial
superhumid with no dry season (Kppen 1936), i.e. more than 60 mm
rainfall in all months and a temperature difference of less than 5C
between the warmest and the coldest month. According to the life zone
system of Holdridge (1982; Holdridge et al. 1971) the Middle Caquet
area is classified as humid tropical forest (bh-T).
18
Figure 2-1. Location of the two permanent plots (M1 = upland plot; M2 = floodplain plot) at Pea Roja, in the middle
part of the Caquet Basin, Colombian Amazonia. The map is derived from Duivenvoorden and Lips (1995).
19
December and February. The wettest month was September not May, as in
Araracuara.
The distribution of rainfall through the year is determined by the trade
winds (Domnguez 1987; Botero 1999). Although Araracuara belongs to
the Southern Hemisphere its rainfall pattern is typical for that of the
Northern Hemisphere, and is furthermore characterized by the absence of a
pronounced dry season (Salati 1985). In Colombian Amazonia the pattern
of rainfall is controlled by the north-south movement of the belt of
intertropical convergence. Most of the rainfall occurs in the afternoon and
evening (Tobn 1999; Tllez 2003). At Pea Roja, the wettest year was
1994, and the driest year was 1995. On average it rained on 197 days per
year, corresponding to 616 hours of rainfall per year (Tobn 1999; Tllez
2003).
2.3
Hydrology
Pea Roja is located in the catchment of the Caquet River. This river
originates in the Andes and, with a length of 2200 km, it is the largest river
in Colombian Amazonia. It is a so-called white-water river, which implies
that its water tends to have a whitish color due to suspended clay. The
river water has a neutral pH (Duivenvoorden and Lips 1993). The water
level of the Caquet River varies annually according to four seasons: low
water, rising water, high water and falling water (Rodrguez 1999). The
variation in river water level recorded at several stations along the river
ranges from 6.5 m to 8.5 m, with peaks during July-August and low water
levels between December-February (Duivenvoorden and Lips 1993;
Urrego 1997). In addition to water levels associated to the annual flood
cycles, occasionally (once every 3 to 11 years) the river water level rises to
exceptional heights. The origin of this phenomenon is unknown, but it has
been linked to global climate change, periods of large sunspot activity, or
to the influence of El Nio events (van der Hammen and Cleef 1992;
Botero 1999).
The floodplain of the Caquet River is called vrzea (Prance 1980; Junk
1984, 1990; Padoch et al. 1999). It is built up by fine sediments which are
deposited during overflows (Eden et al. 1982; Hoorn 1990;
Duivenvoorden and Lips 1993, 1995). Currently there is no evidence of
21
Land units
22
Figure 2-3. Detailed site map of the upland plot (Alarcn 1990;
Tropenbos-Colombia 1990). A: relief of the basin showing 2-m isohypses;
the values denote the altitude relative to an arbitrary reference point; the
total area shown covers 7.2 ha; B: map showing the terrain units and the
location of the permanent plot; C: cross-sections showing representative
soils along the slopes of the U-shaped valley and the V-shaped valley; D:
cover and name of the terrain units.
23
The position of the plots within each land unit was selected such as to
ensure that the terrain characteristics regarding relief were similar.
Because slopes were hardly developed in the floodplain unit, the upland
plot was established at the summit and upper slope positions to minimize
the slope (Fig. 2-3). Both plots were set up in forests that lacked any sign
of recent human activities. Also in the buffer zones of 100 to 300 m
around the plots (Phillips and Baker 2002; Vallejo et al. 2005), no signs of
human activities were present.
The conspicuous presence of charcoal in the soils of the upland plot
(Alarcn 1990) and the remains of pottery found in nearby fluvial terraces
of the Caquet River (Mora et al. 1991; Mora 2003) demonstrate the
ancient human occupation and associated change of the forest cover at the
monitoring sites. However, at the time of the establishment of the plots the
forests did not show any evidence of recent human interventions. This was
also concluded from the presence in the forests of large trees with highly
valuable timber, as Cedrela odorata (Meliaceae, red cedar) in the
floodplain forest and Mezilaurus itauba (Lauraceae, Itaba) in the upland
forest.
At the start, in 1989 (Table 2-1) one square 1-ha plot was installed at the
upland site (part I in Fig. 2-5). Permanent plots of 1 ha have traditionally
been the standard unit of sample area in tropical moist forest inventories,
facilitating comparison and also yielding an optimal ratio area/perimeter
(Synnott 1979, 1991; Jonkers 1987; Alder and Synnott 1992; Dallmeier
1992). However, on the basis of analyses (not shown) of the first data
about the forest diversity using Hill's family of curves (Pielou 1975;
Magurran 1988; Krebs 1989) the plot size was increased to 1.8 ha
(Londoo and Alvarez 1991). To enlarge the original upland plot of 100 x
25
100 m, a strip of 20 m wide was added first (part II in Fig. 2-5). At a later
stage an extra strip of 50 x 120 m (part III in Fig. 2-5) was established to
obtain the final plot of 120 x 150 m (1.8 ha). One year later, in October
1990 (Table 2.1), an equivalent plot of 120 x 150 m (1.8 ha) was
established in floodplain forest, with identical parts I, II and III (Fig 2.5).
The plot inventories included all vascular plant individuals that were
entirely or partially rooted (i.e. with trunks attached to the soil) inside the
plots. Plants rooting precisely on the plot boundaries were included, even
if the aerial parts were outside the plot boundaries. The sampling intensity
was inversely proportional to the abundance of individuals per size class.
Large individuals (DBH 10 cm; DBH is diameter at 130 cm) were
sampled in the entire plot, while the smaller plants were only sampled in a
part (Fig. 2-5). The area of the subplots differed per size class (Dubois
1980; Matteuci and Colma 1982; Jonkers 1987; Campbell et al. 2002;
Oldeman et al. 2006). Large individuals were sampled in 10 x 10 m
subplots, intermediate sized shrubs, lianas and small trees in 5 x 5 m
subplots, and small plants (mostly herbs and seedlings) in 2 x 2 m subplots
(Table 2-2).
To minimise damage to the plants, the inventory was done in the following
sequence: a) demarcation of plots and subplots by setting up the grid of 10
x 10 m; b) inventory of size class C4 (Table 2-2); c) inventory of C1; d)
inventory of plants of size classes C2 and C3; and e) measurement of
additional variables.
Table 2-1. Installment and census dates of the permanent plots.
Activity
Upland
establishmenta
first measurementa and
establishmentb
second measurement
third measurement
fourth measurementc
Floodplain establishment
first measurement
second measurement
a
Date
September 01 1989
September 01 1990
December 31 1993
December 31 1997
March 31 1999
October 01 1990
January 31 1994
April 30 1999
1200 40
1440 48
450 15
3.3
4.0
1.3
1200 40
1890 63
3.3
5.3
only 1 ha; badditional 0.8 ha; conly trunk mortality and recruitment
26
Figure 2-4. Detailed map of the floodplain plot (M2) (Ordez 1990;
Tropenbos-Colombia 1990; Londoo 1993). A: map showing the terrain
units and the location of the permanent plot; the total area shown covers
7.2 ha; B: cross-section, indicating the representative soils and drainage;
C: cover and name of the terrain units.
27
Figure 2-5. Design of the 1.8-ha permanent plots. The roman values I, II,
and III indicate the different stages of plot establishment (see text). A:
general design; B: design and coding of subplots used to sample the
different size classes; C: sampling area for DBH 10 cm (the largest size
class C1); D: sampling area for plants with DBH < 10 cm and DBH 5
cm (size class C2); E: sampling area for plants with DBH < 5 cm and Ht
3 m (size class C3); F: sampling area for plants with Ht < 3 m (size class
C4). Size classes are defined in Table 2-2.
28
Plant size
C1
C2
DBH 10 cm
DBH 5 cm and DBH
< 10 cm
Ht > 3 m, DBH < 5 cm
Ht 0.5 m, Ht 3 m
C3
C4
Subplot size
(m)
10 x 10
5x5
Subplots Area
(ha)
180
1.8
360
0.9
Sampling
intensity (%)
100
50
5x5
2x2
120
300
16.7
6.7
0.3
0.12
29
30
3
A new genus and species of
Dipterocarpaceae from the
Neotropics. I. Introduction,
taxonomy, ecology, and
distribution
Ana Catalina Londoo Vega, Esteban lvarez, Enrique
Forero & Cynthia M. Morton
Brittonia 47: 225-236 (1995)
3.1
Introduction
32
Anatomy-Morphology
34
Phenology
35
36
37
Geography
Ecology
38
3.6
Climate
With respect to soils and geology, this species occurs in upland ("tierra
firme," i.e., not flooded by rivers) that belongs to a dissected sedimentary
plain built up by non-to slightly consolidated, clayey to sandy, Miocene
fluvial sediments (Duivenvoorden and Lips 1993; Hoorn 1994). The
species grows on summits of hills and along shoulders of slopes. The soils
are deep and well-drained. They are loose and sandy at the surface and
become increasingly clayey with depth. According to Duivenvoorden and
Lips (1995: Profile 125)[Appendix 1; addition this Thesis], they are
classified as Typic Kandiudults (SSS 1992) or Xanthic Ferralsols (FAO
1988). They are strong brown (7.5 YR 5/6) in the B horizon, acid (between
3.7 pH in horizon A and 4.9 pH in horizon B), with a very low cation
exchange capacity (ca. 10 cmol(+)/kg clay). The levels of exchangeable
cations and base saturation are very low (less than 15%), and there is a low
level of available phosphorous (1 ppm P, according to Bray II). The
presence of charcoal within the uppermost 30 cm indicates possible sites
of ancient human occupation. The soil is covered by a thick, well-rooted
(10 cm) humus form which consists of an L, F, and discontinuous granular
H horizon.
39
Economic Use
3.9
Vernacular Names
Taxonomic Treatment
only observed in bud). Calyx glabrous, forming a shallow cup at the base,
calyx lobes 5, deltoid, 1.5 mm long, 1.5 mm wide. Corolla glabrous,
greenish white, petals 5, contorted, 5-8 mm long, 1.5-3 mm wide, oblong,
apex obtuse. Stamens numerous, cyclic, hypogynous, free, filaments terete,
anthers basi-versatile, 2-lobed, 4-celled, introrse, connective broad (very
expanded), continued into a triangular appendage one-fourth to one-half as
long as the body of the anther; pollen grains isopolar, tricolporate, rarely
tetracolporate, and sometimes trisyncolporate, exine minutely reticulate to
foveolate, columellate, tectate-perforate. Ovary glabrous, 3-locular,
placentation axial, sub-basal, ovules one per loculus (as far as known),
apparently anatropous; styles terminal, minute. Fruit a dry nut, 3-4 cm
long, 1.5-2 cm wide, glabrous, ovoid, pericarp woody. Calyx in fruit
persistent, all 5 segments accrescent, thinly papyraceous, aliform, 10-16
cm long, 1.5-2.5 cm wide, oblanceolate, apex rounded, base cuneate, lobes
united at base. Seed 1 per fruit.
Distribution: Known only from the type locality in the vicinity of
Araracuara, Department of Amazonas, Colombia, at 200-300 m elevation.
No open flowers have been seen. The plants have been collected in bud in
November and in fruit in April.
Paratypes: COLOMBIA. Amazonas: Araracuara, Villazul, Ro Caquet,
margen izquierda, frente a isla Sumaeta, 0034' S, 7208' W, 200-300 m,
20 Oct 1990 (sterile), lvarez et al. 1184 (COAH); detrs maloca J.
Moreno, 0034' S, 7208' W, 9 Oct 1989 (sterile), Londoo et al. 899 (A,
COAH, NY); frente a isla Sumaeta, 0034' S, 7208' W, 200-300 m, 31
Oct 1989 (sterile), Londoo et al. 983 (COAH, NY), 3 Nov 1989 (sterile),
Londoo et al. 1097 (COAH, JAUM, NY), 7 Nov 1989 (buds), Londoo et
al. 1239 (A, COAH, COL, NY), 1239A (COAH).
42
Dipterocarpoideae
Multiseriate
Stipules
Leaf venation
Pubescence
Trichomes fasciculate,
commonly single-celled or
multi-cellular; often glandular
Sepals
Petals
Stamens
Anthers
5, in flower imbricate or
valvate and connate forming a
tube at base; often becoming
accrescent and aliform in fruit
5, variously pubescent
5-numerous, 1-several-cyclic;
filaments sometimes united
below the middle
2-lobed, basifixed, (2)4-
Monotoideae
Uniseriate (sometimes
biseriate)
Wood without resin canals;
secretory cavities in pith
Parkaraimoideae
Biseriate (sometimes
uniseriate)
Wood without resin
canals; secretory cavities
in pith
Small, fugacious
Secondary veins reticulate
Pseudomonotes
Uniseriate (sometimes
biseriate)
Wood without resin
canals; secretory
cavities in pith
Small, fugacious
Secondary veins parallel
or obscurely reticulate
Trichomes glandular
5, variously pubescent
Numerous, several-cyclic;
filaments free
Trichomes fsciculate,
commonly single-celled;
nither grandular nor
granular
5, in flower imbricate not
connate; becomng
accrescent and shortly
aliform in fruit
5, glabrous
Numerous, several-cyclic;
filaments free
2-lobed, basifixed, 4-
2-lobed, basifixed, 4-
Small, fugacious
Secondary veins obscurely
parallel (transverse), or
reticulate
Trichomes fsciculate,
commonly single-celled;
often granular
5, in flower imbricate not
connate; becoming aliform
and accrescent in fruit
5, in flower connate
forming a shallow cup
at base; accrescent and
aliform in fruit
5, glabrous
Numerous, cyclic;
filaments free
43
Pollen
Ovary
Fruit
Ecology
Geography
44
thecate, basivesatile,
strongly appendaged
thecate, basiversatile;
apically apendaged
Tricolporate, exine ()
layered
3 celled, each locule
probably 1-ovulate;
ovules pendulous,
anatropous; placentation
axial, sub-basal
1-seeded; tardily
loculicidally dehiscent
Trees of savanna or
savanna woodland
Tropical America, one
genus; unispecific
3.11
Relationships
45
clearly nearly basal (sub-basal). The 1-ovuled ovary cells should therefore
be regarded as tentative until more and better flowers are obtained. The
ovary is 3-locular and the ovules are pendulous and anatropous as in the
rest of the family. The style appears as three minute, terminal styles.
The fruit of Pseudomonotes is, as in all members of the family, dry,
usually with a woody pericarp that splits along three sutures at
germination, and 1-seeded.
Morton (Part II, below) compared the anatomy of the wood, bark, and pith
of Pseudomonotes with those of several families of flowering plants
belonging to the subclasses Dilleniidae and Rosidae (Cronquist 1981).
Wood anatomical features indicate close relationships to the Monotoideae
with which it shares the mostly uniseriate wood rays, the lack of resin
canals in the wood, and the presence of secretory cavities in the pith
(Bancroft 1935). Resin canals are absent but mucilage cavities are present
in Pakaraimoideae (De Zeeuw 1977), while resin canals are present and
secretory cavities are absent in Dipterocarpoideae (Guerin 1906, Gottwald
and Parameswaran 1966).
On the basis of the information gathered in the field, herbarium, and
laboratory, it is concluded that the new taxon belongs to subfamily
Monotoideae of the Dipterocarpaceae. These two groups share the
uniseriate (or rarely biseriate) wood rays, presence of secretory cavities in
the pith, lack of resin canals, single gland on the upper surface of the
lamina at the base of the midrib, and basi-versatile anthers.
The new entity differs from the rest of the Dipterocarpaceae in the absence
of fasciculate trichomes and in having sepals in fruit conspicuously
aliform (reaching 10-16 cm in length), and one ovule per locule with
nearly basal (sub-basal) placentation.
The addition of Pseudomonotes tropenbosii to subfamily Monotoideae
makes this the only subfamily of the Dipterocarpaceae with different
taxonomic elements currently growing in the Old as well as the New
World. The new taxon appears to be confined to a small area in the
southwesternmost limit of the Guayana Highland and the superposed
Roraima Formation sediments in Amazonian Colombia.
47
3.12
Acknowledgements
48
4
Arquitectura de Iryanthera
tricornis, Osteophloeum
platyspermum y Virola pavonis
(Myristicaceae)
Eliana Mara Jimnez Rojas, Ana Catalina Londoo Vega &
Henricus F. M. Vester
Caldasia 24(1): 65-94 (2002)
4.1
Introduccin
50
para analizar las reacciones del rbol hacia los otros rboles y su medio
(Vester 1997). El modelo arquitectnico, determinado por el tipo de
crecimiento, la morfologa de los ejes, la ramificacin y la posicin de la
sexualidad, se define como el programa de crecimiento que determina las
fases sucesivas del desarrollo de la planta (Hall et al. 1978). Debido a las
grandes variaciones que puede presentar un modelo dado entre diferentes
especies, se emplea la unidad arquitectnica, definida como la expresin
especfica del modelo para una especie en particular, que incluye mayor
detalle aquitectnico en la descripcin del patrn de ramificacin mediante
una secuencia de diferenciacin (delin 1977) citado por (Barthlmy et
al. 1991). El desarrollo de ejes fuera de la expresin normal de la unidad
arquitectnica se denomina reiteracin (Hall et al. 1978), proceso
morfogentico que duplica total o parcialmente el modelo arquitectnico
(Oldeman 1974). Las fases sucesivas del crecimiento de un rbol se
analizan mediante el diagrama arquitectnico, o secuencia de dibujos que
representa el desarrollo arquitectnico del rbol (Vester 1997), que
comprende los rboles potenciales (juveniles con potencial para expandir
su copa), del presente (rboles maduros con copas plenamente
desarrolladas) y del pasado (rboles viejos con copa debilitadas y daadas)
(Hall et al. 1978). Por ltimo, el plan de organizacin puede ser
jerrquico o polirquico, y est definido por la dominancia apical del eje
principal (delin 1991).
Las Miristicceas del mundo con estudios arquitectnicos son pocas, pero
todos los trabajos previos concuerdan en que stas crecen conforme al
modelo arquitectnico de Massart (Hall et al. 1978; Drnou 1994; Loubry
1994; Loup 1994). Ya desde mediados del siglo XIX, durante sus viajes a
la Amazonia, el botnico Richard Spruce haba hecho detalladas
descripciones sobre la ramificacin y la fisionoma que incluan conceptos
arquitectnicos incipientes sobre las Miristicceas, las cuales llamaron
mucho su atencin por su forma tan caracterstica (Spruce 1861) citado por
(Madrin 1996). Ms tarde, en 1923 Jean Massart estudi y describi un
ejemplar de V. surinamensis en el Jardn Botnico de Ro de Janeiro (Hall
et al. 1978), sentando las bases de lo que despus fue denominado como
modelo arquitectnico de Massart (Hall y Oldeman 1970). Ms
recientemente, esta misma especie ha sido investigada intensivamente
52
(Barthlmy et al. 1991; Drnou 1994; Loup 1994), al igual que Virola
michelii Heckel (Comte 1993) citado por (Loubry 1994).
Adems de ocurrir en la familia Myristicaceae, el modelo de Massart est
bien representado tanto en rboles tropicales (v.g. Ceiba pentandra (L.)
Gaertn., Ocotea guianensis Aubl.), subtropicales (v.g. Araucaria
heterophylla (Salisb.) Franco), como de zonas templadas (v.g. Abies
balsamea (L.) Mill., Sequoia sempervirens (D. Don) Endl.) (Hall et al.
1978). Sin embargo, son pocas las especies con estudios arquitectnicos,
como: Ocotea oblonga Mez (Vester 1997), Symphonia globulifera L.f.
(Loup 1994) y algunas Araucaria spp. (Veillon 1978).
En Colombia los estudios sobre arquitectura de la vegetacin son bastante
escasos. Entre ellos se encuentran los realizados en la regin de Araracuara
en palmas (Echeverri 1993), en el desarrollo de los bosques secundarios y
sus especies ms importantes (Vester y Saldarriaga 1993; Vester 1997) y
sobre Myristicceas (Jimnez 2000). Tambin se han estudiado los
cativales en la regin de Urab (Ros 1996) y la arquitectura del roble
(Quercus humboldtii Bonpl.) en Antioquia (Echeverri 1997).
En este artculo se presenta el anlisis arquitectnico de tres especies de
Miristicceas del dosel en dos bosques de tierra firme. Con base en los
trabajos previos sobre la arquitectura de otras especies de la familia, se
parti de la hiptesis de que las especies, pertenecientes a tres gneros
diferentes, deban exhibir igualmente el modelo de Massart. Se estudi el
desarrollo del modelo arquitectnico en estas especies, la conformacin de
la unidad arquitectnica, los tipos de reiteracin, el plan de crecimiento y
el plan de organizacin arquitectnica a nivel del rbol, as como la
similitud arquitectnica entre las tres especies. Adems, se investig si
existan semejanzas con otras especies de la familia previamente
estudiadas, y cmo el desarrollo arquitectnico favorece la sobrevivencia
de los rboles para alcanzar el dosel.
4.2
rea de estudio
Materiales y Mtodos
Resultados
del eje A1 entre en estado de reposo; i.e. los ejes A2 son silpticos. Luego,
el meristema terminal del eje A1 reanuda su actividad y se produce la
siguiente unidad de crecimiento, y as sucesivamente. Los ejes A2 se
disponen rtmicamente, formando pisos de ramas muy conspicuos,
localizados en posicin acrotnica dentro de las unidades de crecimiento
del eje A1. Los ejes A2 presentan una orientacin horizontal con filotaxia
alterna dstica, acompaada de una pequea torsin de los pecolos que
acenta su carcter plagitropo; son monopdicos, con crecimiento rtmico
y tienen ramificacin difusa, ya que no hay un patrn visible en la
aparicin de los ejes A3. Los A3 exhiben filotaxia alterna dstica y
tambin son plagitropos, en menor grado que los ejes A2, son
monopdicos, con crecimiento rtmico, pero sin ramificacin. Difieren de
los ejes A2 en su origen: los primeros son silpticos, mientras los A3,
prolpticos. Adicionalmente, los ejes A2 estn conformados por un
nmero mayor de unidades de crecimiento en comparacin con los ejes A3
que slo desarrollan dos o tres unidades de crecimiento; en consecuencia,
los ejes A2 son de mayor longitud que los A3. Las estructuras
reproductivas (flores y frutos) se encuentran en los ejes A2 y A3; debido a
su posicin axilar dentro de los ejes, la localizacin de la sexualidad no es
importante en el modelo de Massart, puesto que no interfiere con el
crecimiento vegetativo del rbol (Hall et al. 1978).
Tipos de reiteracin
Las tres especies estudiadas presentaron gran capacidad de reiteracin;
todas las reiteraciones fueron de tipo prolptico. Por una parte, se hallaron
reiteraciones traumticas en individuos pertenecientes a todas las fases de
desarrollo del rbol; por otra, se detectaron reiteraciones adaptativas en
ciertas fases. Siguiendo el desarrollo de las especies, primero surgen las
reiteraciones adaptativas parciales sobre los ejes A2; despus, las totales.
Estas ltimas tambin se localizan principalmente sobre los ejes A2 ms
lignificados, de mayor dimensin, y eventualmente sobre el eje A1. Una
vez que los individuos han alcanzado el dosel, que en el caso de las tres
especies estudiadas corresponde a los rboles del presente, fue posible
identificar reiteraciones arbreas, arbustivas y herbceas.
59
Sitio
Iryanthera tricornis
Osteophloeum
platyspermum
Virola pavonis
PST
TB
Subtotal
PST
Potencial de desarrollo
PO
PR
PA
SU
16
1
0
4
0
3
0
0
16
4
0
4
2
2
1
0
Total
N
21
3
24
5
TB
Subtotal
PST
TB
Subtotal
PST
TB
N
%
7
9
4
0
4
22
7
29
57
14
19
5
3
8
31
20
51
100
1
3
0
3
3
3
7
10
19
5
6
0
0
0
1
5
6
12
1
1
1
0
1
5
1
6
12
%
41
6
47
10
27
37
10
6
16
61
39
100
Tipo
Direccin
Longevidad
Uc
Ramificacin
Simetra
Lmites
No. de hojas
Tipo
Cronologa
Localizacin
Insercin
Hojas
Filotaxia
Sexualidad
Presencia
Localizacin
Eje A1
monopodio
rtmico
-orttropo
-vertical
la vida del
rbol
radial
-hojas
pequeas
-cambio en
corteza
-hilera de ejes
A2
entre 5-9
rtmico
inmediata
acrotnica
alternaespiralada
no
-
Eje A2
Monopodio
Rtmico
-plagitropo
-horizontal
Larga
Eje A3
monopodio
rtmico
-plagitropo
-oblicua
corta
Bilateral
-hojas
pequeas
-cambio en
corteza
-
bilateral
-hojas
pequeas
-cambio en
corteza
-
entre 5-8
Difuso
Retardada
Acrotnica
ca. 90 con eje
A1
alterna-dstica
entre 5-8
no ramificado
40-70 con eje
A2
alterna-dstica
S
-lateral
-en tres
ltimas Uc
s
-lateral
-en tres
ltimas Uc
61
rdenes de ejes diferentes: A1, A2 y A3 (Figura 4-1, Tabla 4-2). Hasta esta
fase, el desarrollo del rbol sigue un plan de organizacin jerrquico, en el
cual existe una fuerte dominancia del meristema apical del eje A1 y una
subordinacin de los ejes A2 y A3.
Fase 4: juvenil grande (Figura 4-1, Figura 4-2E-H). Esta fase an
comprende rboles potenciales, con alturas entre 5-15 m, con la unidad
arquitectnica completa. En este momento el rbol comienza a sobrepasar
el umbral definido por la expansin de la copa conforme al desarrollo del
modelo y reacciona mediante la produccin de numerosas reiteraciones
adaptativas parciales (Rp, Figura 4-2E-H). stas son muy abundantes en
los primeros pisos de ramas (ejes A2), o sea las ms bajas y viejas, las
cuales son igualmente las ms largas y plagitropas, que gradualmente
disminuyen en longitud hacia la parte superior de la copa. Frecuentemente
se observa que el crecimiento de los ejes A2, inicialmente con un carcter
marcadamente monopdico (con crecimiento indefinido), se va tornando
progresivamente definido (monopodios inestables) en virtud de la muerte
de su meristema terminal. La desaparicin de este meristema terminal
ocasiona el desarrollo de yemas laterales, ubicadas en las axilas de las
hojas ms cercanas al pice, que hasta ese momento se encontraban en
estado de latencia (Figura 4-4F, H).
Estas yemas producen complejos reiterados adaptativos, denominados
tenedores (Barthlmy et al. 1991). stos, vistos sobre un plano, pueden
ser sencillos (dos reiteraciones, Figura 4-4A) o mltiples (tres o ms
reiteraciones, Figura 4-4I), que segn su aparicin, pueden sobreponerse
unos a otros modificando totalmente la arquitectura de las ramas. Por otra
parte, en la parte media y proximal de los ejes A2 ms viejos, se observa el
desarrollo de ejes dediferenciados (ejes A2, Figura 4-4C, F-I). En esta
fase se inicia el plan de organizacin polirquico.
Fase 5: transicin entre rbol potencial y rbol del presente (Figura 4-1,
Figura 4-5A-B, F), cuyos individuos se consideraron como rboles del
presente, generalmente con alturas entre 15-25 m, que excepcionalmente
pueden alcanzar hasta 40 m. Sus copas exhiben una arquitectura ms
compleja con gran parte de los caracteres propios de los rboles del
presente, que se encuentran prximas a alcanzar su mxima capacidad de
desarrollo. Las reiteraciones adaptativas parciales de los ejes A2 son ms
63
64
66
67
Figura 4-6. Reiteracin total adaptativa (Rt) sobre eje A2 en rbol del
presente (fase 5).
68
69
70
Discusin
73
75
al. 1991; Sterck et al. 1991; Loup 1994; Vester 1997), pero hasta que no se
efecten las observaciones no es posible determinarlos con precisin. En
este trabajo para las fases iniciales (1-3) se consider la aparicin de los
distintos rdenes de ejes, mientras que para las dems (4-8) se tuvo en
cuenta la reiteracin adaptativa, modificando criterios previamente
empleados en otros estudios sobre arquitectura (Sterck et al. 1991). El plan
de crecimiento de las especies estudiadas puede considerarse en tres
perodos ontognicos del rbol (Oldeman 1990b; Sterck et al. 1991): el de
plntula (fase 1), el de crecimiento conforme al modelo arquitectnico
(fases 2-3, Figura 4-2B-D), y el de reiteracin. ste ltimo comprende
tanto la adaptacin programada mediante reiteracin adaptativa (fases 4-8,
Figura 4-2E-H, Figura 4-5 y Figura 4-7) como el ajuste oportunstico
mediante la reiteracin traumtica (en cualquier etapa de desarrollo desde
la fase 1 hasta la 8, Figura 4-8).
Luego del establecimiento inicial como plntula, durante el perodo
ontognico de crecimiento conforme al modelo, el arbolito comienza a
desarrollar los distintos rdenes de ejes hasta construir su unidad
arquitectnica completa, mediante la secuencia normal de diferenciacin
de los ejes: el eje A1 produce los ejes A2 y stos a su vez, los ejes A3.
Hasta ese momento existe un fuerte control apical del meristema del
tronco (eje A1), con subordinacin de los dems rdenes de ejes, y el plan
de organizacin del rbol es claramente jerrquico. Posteriormente, cuando
se inicia la reiteracin adaptativa (fase 4), se pierde la secuencia inicial de
diferenciacin de los ejes y el plan de organizacin del rbol cambia:
comienza a tornarse polirquico (fase 4 y siguientes). Entonces, los ejes
A2 empiezan a producir ejes dediferenciados (ejes A2 ,Figura 4-4) i e.
ejes que se desarrollan fuera de la secuencia normal de diferenciacin
en vez de generar los del orden siguiente (A3). La dediferenciacin
tambin puede ser vista como una inversin, o un retroceso, de la
produccin de nuevos ejes, la cual llega incluso a avanzar un orden ms,
hacia atrs, cuando se presentan las reiteraciones totales (ejes A1 Figura
4-1 y Figura 4-6) sobre los A2 anteriores, en lo que aqu se ha definido
como el umbral que marca el comienzo de la fase del rbol del presente
(fases 5 y 6). Desde este momento hasta la muerte del rbol, empiezan a
aparecer subcopas dentro de la copa del rbol, como individuos
independientes que compiten entre s (Rt, Figura 4-5 y Figura 4-7)
77
(subcopas) unidas por un tronco que las levanta del suelo y las ubica en el
dosel, permitiendo el acceso directo a los recursos de iluminacin,
facilitando el transporte de polen y de semillas. Mediante la capacidad de
generar reiteraciones estas especies aumentan sus probabilidades de
sobrevivencia durante su desarrollo temprano en el sotobosque
(reiteraciones traumticas) y construyen sus copas durante las etapas
intermedias y finales (reiteraciones adaptativas). De esta manera, en las
tres Miristicceas estudiadas la reiteracin es un mecanismo fundamental
en el desarrollo de la copa, al igual que lo reportado para numerosas
especies (Barthlmy 1991; Barthlmy et al. 1991; Sterck et al. 1991;
Vester 1997).
Comparacin entre las tres especies
A pesar de la gran similitud arquitectnica entre las especies estudiadas,
existen pequeas diferencias entre ellas, al igual que al compararlas con
respecto a V. michelii y V. surinamensis (Drnou 1994; Loubry 1994;
Loup 1994). Por ejemplo, para las tres especies se encontr que la
plagiotropa de los ejes A2 es ms acentuada en los rboles potenciales que
en los del presente, tanto en la orientacin del eje que cambia de horizontal
a oblicua, como en la disposicin de las hojas, que pierden su arreglo en un
solo plano con simetra dorsiventral para obtener una posicin oblicua.
Esta caracterstica se observ mucho ms acentuada en V. pavonis que en
las otras especies. Igualmente se registr variacin en los ngulos
horizontales y verticales de la unin de los ejes. Tambin se observaron
cambios menores en el nmero de hojas que conforman las unidades de
crecimiento; as I. tricornis y V. pavonis generalmente presentaron de 5 a 7
hojas por unidad de crecimiento en los ejes A2 y A3, mientras que O.
platyspermum present de 7 a 8 hojas en los mismos ejes. Por
consiguiente, el nmero de hojas por unidad de crecimiento y la cantidad
de unidades de crecimiento por eje determinan la longitud de estos,
imprimiendo diferencias entre las especies.
Por otra parte, tambin se detectaron diferencias en la forma y la
disposicin de los complejos reiterados adaptativos parciales y totales. En
V. pavonis se encontraron tenedores muy similares a los reportados para V.
surinamensis (Barthlmy et al. 1991; Loubry 1994; Loup 1994), mientras
que en I. tricornis y O. platyspermum los complejos reiterados parciales
79
80
82
Arquitectura y taxonoma
Al igual que la taxonoma, la arquitectura se apoya en el estudio de la
morfologa vegetal (Hall y Oldeman 1970; Hall et al. 1978). Algunos
botnicos pioneros de siglos pasados ya haban incluido conceptos
incipientes de la arquitectura en sus observaciones y descripciones (Spruce
1861) citado por (Madrin 1996). Hoy en da, la taxonoma y la
arquitectura mantienen enfoques independientes y permanecen
relativamente aisladas, con pocos trabajos que las consideran
simultneamente (e.g. Sanoja 1992). Aunque stas comparten una base
morfolgica, la arquitectura considera la planta desde la germinacin hasta
la muerte, y constituye una aproximacin dinmica con nfasis en los
procesos intrnsecos de crecimiento a travs del tiempo. Por el contrario, la
taxonoma considera la planta slo en un momento dado de su ciclo vital, y
tiene un enfoque esttico (Bell 1993; Madrin 1996). No obstante su
importancia (Hall et al. 1978; Barthlmy et al. 1991; Vester 1997), la
arquitectura comnmente se ha visto como un conjunto fijo de
caractersticas morfolgicas, y se ha confundido con la fisionoma o el
hbito de crecimiento de las especies (Hall et al. 1978; Bell 1993;
Madrin 1996; Vester 1997). Por otra parte, en las descripciones
taxonmicas, cada parte de la planta se observa aisladamente de las dems.
Por ejemplo, las hojas se describen detalladamente pero no se tiene en
cuenta dnde se insertan en la planta o el eje donde crecen, lo cual
precisamente origina y explica gran parte de la variacin en algunas
especies. En las tres especies estudiadas, cada unidad de crecimiento
comienza con la produccin de una lmina foliar de tamao muy reducido
(ca. 3.0-4.0 cm x 1.5-2 cm en V. pavonis), fuera del rango de tamao
registrado para la especie (Jimnez 2000). Entonces, al aislar las distintas
partes constitutivas de la planta se pierde la cohesin de la estructura
general de la misma. Por el contrario, la percepcin desde la arquitectura
es fundamentalmente integral puesto que sta concibe la planta como un
sistema en conjunto (Hall et al. 1978). Precisamente por su naturaleza
dinmica e integral, la arquitectura puede enriquecer y complementar los
estudios morfolgicos de la taxonoma.
83
4.6
Agradecimientos
84
5
Composicin florstica de dos
bosques (tierra firme y vrzea) en
la region de Araracuara,
Amazonia colombiana
Ana Catalina Londoo y Esteban lvarez
Caldasia 19(3) 431-463 (1997)
5.1
Introduccin
87
5.2
Ubicacin y acceso
Materiales y mtodos
Mtodos de muestreo
Tanto en tierra firme como en la vrzea se estableci una parcela de 1.8 ha
(120 m x 150 m) donde se muestre la vegetacin vascular terrestre con
altura mayor que 0.5 m. Se consideraron 17 hbitos de crecimiento
diferentes agrupados en cinco categoras: arbrea, arbustiva, escandente
herbcea, escandente leosa y herbcea terrestre (Tabla 5-1). Los criterios
bsicos para definir los diferentes hbitos de crecimiento fueron: la
presencia o ausencia de tejido leoso, la dependencia o independencia del
soporte de otras plantas y, por ltimo, el tamao; se tom como lmite
arbitrario Ht = 3 m para separar las formas arbreas de las arbustivas
[Captulo 2].
Anlisis de los datos
En los bosques inventariados se identificaron las plantas hasta especie
taxonmica (Grant 1989); cuando esto no fue posible se emplearon
morfoespecies en la denominacin de los taxones. Para las familias de
plantas superiores se sigui el sistema de Cronquist (1981), excepto
Caesalpiniaceae, Fabaceae y Mimosaceae, agrupadas en Leguminosae para
facilitar la comparacin con otros estudios. Cecropiaceae se presenta
separada de Moraceae, Hippocrateaceae aparte de Celastraceae y
Mendonciaceae segregada de Acanthaceae. Para las Pteridophyta se sigui
a Mabberley (1990).
Durante la identificacin se consultaron las colecciones del Herbario
Amaznico (COAH), el Jardn Botnico de Nueva York (NY), el Jardn
Botnico de Missouri (MO), el Jardn Botnico de Kew (K), el Herbario
Nacional Colombiano, (COL), la Universidad de Antioquia (HUA) y el
Jardn Botnico Joaqun Antonio Uribe de Medelln (JAUM). A la fecha,
89
Arbreo4
Arbustivo5
90
Hbito de
crecimiento,
cdigo
Arboles, A
Helechos
arbreos, FA
Palmas arbreas
monoestipitadas,
PAM
Palmas arbreas
cespitosas, PAC
Subtotal formas
arbreas
Arbustos, T
Palmas
arbustivas
acaules, PTU
Palmas
arbustivas
cescpitosas, PTC
Palmas
arbustivas
monoestipitadas,
PTM
Subtotal formas
arbustivas
PST
LAI
Comunes
Datos
combinados
No.2
469
1
%3
67.2
0.1
No.
308
1
%
60.3
0.2
No.
43
0
%
71.7
0.0
No.
734
2
%
63.9
0.2
0.6
0.8
0.0
0.7
0.1
0.6
1.7
0.3
475
68.1
316
61.8
44
73.3
747
65.0
45
1
6.4
0.1
34
1
6.7
0.2
1
0
1.7
0.0
78
2
6.8
0.2
0.3
0.2
0.0
0.3
1.1
0.4
0.0
10
0.9
56
8.0
38
7.4
1.7
93
8.1
Escandente
herbceo6
TOTAL
Escandente
herbceo, SH
Hemiepfitas
herbceas, SEH
Helechos
escandentes,
FSH
Subtotal formas
escandentes
herbceas
Escandentes
leosos, SL
Estranguladoras,
SZL
Hemiepfitas
leosas, SEL
Palmas
escandentes, PSL
Subtotal
escandentes
leosas
Helechos
terrestres, FH
Hierbas
terrestres, H
Subtotal hierbas
terrestres
Todos los hbitos
Escandente
leoso7
Herbceo
terrestre8
19
2.7
24
4.7
1.7
42
3.7
13
1.9
13
2.5
1.7
25
2.2
0.3
1.0
1.7
0.5
34
4.9
42
8.2
5.0
73
6.4
110
15.8
84
16.4
11
18.3
183
15.9
0.0
1.0
0.0
0.4
1.1
0.4
0.0
10
0.9
0.0
0.6
0.0
0.3
118
16.9
94
18.4
11
18.3
201
17.5
0.4
0.4
0.0
0.4
12
1.7
19
3.7
1.7
30
2.6
15
2.1
21
4.1
1.7
35
3.0
698
100
511
100
60
100
1149
100
Ver mtodos; Nmero de especies; Porcentaje del total de cada sitio; Plantas con
Trepadoras y hemiepfitas sin tejido leoso; 8Plantas sin tejido leoso, no trepadoras,
5.4
Resultados
91
92
93
Figura 5-1. Composicin florstica de dos bosques en Araracuara (datos combinados de PST y LAI) para plantas
vasculares terrestres con Ht 50 cm (todos los hbitos). A: familias con mayor nmero de especies. B: familias con
mayor nmero de gneros. C: gneros con mayor nmero de especies. Ntese la diferencia entre las escalas de los
grficos.
94
herbceos, y tres familias con igual porcentaje entre las hierbas terrestres
(Figura 5-2).
Aunque la tercera parte (32 de 98) de las familias registradas en el estudio
se encontr exclusivamente en un sitio, el aporte de ellas en nmero de
especies fue bajo (7% y 4% del total de especies en LAI y PST,
respectivamente). Adems, las familias con mayor nmero de especies
fueron similares entre los dos sitios (Figura 5-3A); de las 20 ms
importantes en cada sitio 16 son compartidas, entre las cuales sobresalen
Leguminosae (72 especies en PST y 59 en LAI) y Rubiaceae (32 y 35
especies respectivamente). Por otra parte, entre las familias con mayor
importancia en PST que en LAI estn: Burseraceae (20 y cuatro especies
respectivamente) y Vochysiaceae (14 y una), mientras que Myrtaceae
(cuatro y 17) y Meliaceae (seis y 11) estuvieron mejor representadas en
LAI. Hubo una correlacin altamente significativa (rs = 0.649, p < 0.0001)
entre los dos sitios con respecto a la representacin de especies por
familia.
La cantidad de gneros hallada en cada sitio fue similar: 236 en PST y 235
en LAI. Las familas con mayor nmero de gneros (Figura 5-3B) fueron
Leguminosae (19 y 26 en PST y LAI respectivamente) y Rubiaceae (17 y
16), y Bignoniaceae, Annonaceae y Arecaceae estuvieron entre las seis
primeras. A pesar de las similitudes en cuanto a las familias con mayor
nmero de gneros, los gneros comunes (124) representron apenas 36%
del total. Los dos sitios presentaron una correlacin muy baja (rs = 0.2241,
p = 0.2722, n = 98 familias) con respecto al nmero de gneros por
familia.
Para los datos combinados, entre los 25 gneros con mayor nmero de
especies hubo 20 que estuvieron presentes en los dos sitios (Figura 5-1C y
Figura 5-3C) y se encontr una fuerte correlacin entre stos (rs = 0.6142,
p < 0.0001, n = 25 gneros). Pero entre los 15 gneros con mayor riqueza
de especies, en PST hubo mayor cantidad cuyos taxones son
exclusivamente arbreos y slo se encontraron tres que tienen especies de
otros hbitos: Miconia, Philodendron y Abuta. Por el contrario, en LAI
hubo una mayor cantidad de gneros con especies no arbreas como,
Anthurium, Machaerium, Faramea, Hirtella, Paullinia y Coussarea. Los
96
97
98
99
5.5
Discusin
Figura 5-4 (al lado). Porcentaje de especies por familia para diferentes
hbitos de crecimiento de dos bosques en Araracuara (PST y LAI), para
plantas vasculares terrestres con Ht 50 cm. TOT: todos los hbitos; AR:
formas arbreas; SL: formas escandentes leososas; SH: formas
escandentes herbceas; T: formas arbustivas; H: formas herbceas
terrestres. Categoras de los hbitos de crecimiento segn Tabla 5-1.
Pteridophyta y Leguminosae agrupadas como una sola familia; Moraceae
sensu stricto.
100
101
102
103
104
TOTAL
rs
0.9802
0.6825
0.7317
0.7880
0.7425
0.2949
0.3929
0.5023
0.4127
0.5732
0.5317
p
0.0001**
0.0029**
0.0014**
0.0006**
0.0012**
0.1986
0.0868
0.0286*
0.0720
0.0125*
0.0205
PST
rs
0.9302
0.4805
0.5612
0.7366
0.7075
0.0886
0.2125
0.3136
0.2250
0.3565
0.5232
p
0.0001**
0.0362*
0.0144*
0.0013**
0.0020*
0.6993
0.3543
0.1716
0.3268
0.1202
0.0226*
LAI
rs
0.6825
0.4805
0.5239
0.6205
0.4082
0.5423
0.3596
0.3806
0.4525
0.4788
0.5655
p
0.0029**
0.0362*
0.0224*
0.0068**
0.0752
0.0181*
0.1170
0.0971
0.0486*
0.0369*
0.0137*
106
108
109
(Nelson et al. 1990; Tuomisto et al. 1995). Pero adems subyace el poco
conocimiento florstico de la regin, en particular de la parte colombiana.
Los resultados presentados en este estudio, incluyendo primeros registros
para el pas a nivel de familia y de especies previamente consideradas
endmicas en otros pases de Amazonia, ilustran claramente dos
situaciones. Primero, que an falta mucho por conocer con respecto a la
flora de esta regin y, segundo, que las colecciones botnicas intensivas,
restringidas a parcelas pequeas, son una herramienta importante en la
bsqueda de dicho conocimiento.
5.6
Agradecimientos
6
Forest dynamics in an upland
and a floodplain plot near Pea
Roja, Colombian Amazonia
Ana Catalina Londoo Vega
6.1
Introduction
Tree mortality is a complex process that controls the density of trees in the
forest (Carey et al. 1994; Manokaran and Swaine 1994). Tree death occurs
at different levels of intensity, space and time, reflecting endogenous
processes (e.g. aging) and exogenous disturbances, which involve the
severity, frequency, duration, timescale and interaction with the ecosystem
(Lugo and Scatena 1996). Normal or background mortality rates are
usually below 5% per year, and may range by several orders of magnitude.
This mortality tends to take place at local scales and often occurs gradually
(e.g. the slow death of one single tree) and tends to increase with tree
density (number of individuals per unit area) (Lewis et al. 2004b).
Contrary, catastrophic mortality (> 5% per year) takes place during
massive and often quite sudden events (e.g. by wind throw) (Nelson et al.
1994). These two kinds of mortality have a different effect on the
succession that follows death events (Swaine et al. 1987; Matelson et al.
1995; Lugo and Scatena 1996).
Usually it is difficult to determine the cause of death of an individual tree,
and often several causes act in combination (Lugo and Scatena 1996;
D'Angelo et al. 2004). To quantify tree mortality, types of death can be
defined based on direct field inspections of the remains of the tree after its
death (Gale and Barfod 1999). The type of death determines how the forest
canopy is opened (Hall et al. 1978; Hartshorn 1990b) and defines how the
forest responds regarding species composition (Denslow 1987), diversity
113
(Brnig and Huang 1990; Hartshorn 1990a) and forest structure (Hubbell
and Foster 1986; Swaine 1990). Different types of death create variation in
gaps (Clark 1990; Carey et al. 1994) and in the regeneration opportunities
for the forest (Hartshorn 1990a; Orians 1982; Negrelle 1995).
Figure 6-1. Types of tree death. Left: standing dead; middle: snapped
trunk; right: uprooted.
Three types of death are often distinguished (Fig. 6-1) to quantify forest
dynamics: uprooted tree, snapped trunk, and standing (or upright) death
(Rankin de Merona et al. 1990; Gale 1997; D'Angelo et al. 2004). For the
uprooted tree, the immediate causes of death are often physical (such as
strong wind, heavy rain, shallow soils, steep slope and the fall of trees
interconnected by lianas; D'Angelo et al. 2004), or sometimes biotic (root
rot). Death in snapped trunks is mainly attributed to attacks by fungi and
insects, the effect of which may depend on wood density (Putz et al.
1983). Standing or upright death is mostly due to senility, disease and
pathogen attack, along with shading by other trees (Oldeman 1990b). Light
and drought are the principal abiotic factors associated with upright death.
Recruitment is a manifestation of species fecundity, seed dispersal (Clark
et al. 1999) as well as growth and survival of juveniles in a population
(Swaine et al. 1987). Although in most reports on tropical forests
recruitment is closely linked to mortality, which leads to more or less
constant densities of trees with a certain size, the relationship between
these two opposing processes may be weak for short periods of time or at
small spatial scales (Swaine et al. 1987).
114
6.3
Methods
115
of 9.58 years (115 months) in the upland plot and 8.58 years (103 months)
in the floodplain plot.
During the second and posterior censuses, all previously marked
individuals were checked and verified if they were or alive or dead. If alive
they were checked for damage, such as broken or leaning crown, or
leaning or broken trunks. If they were dead, the type of death was recorded
based on the observation of the type of debris or remains (Fig. 6-1), using
the following categories (Rankin de Merona et al. 1990; Gale 1997): A)
Standing of upright death. This occurred when the dead tree was still
standing without clear signs of having been broken or uprooted. In such
cases, the tree crown was still largely present in situ but fragments of
fallen branches were often observed lying around the dead trunk; B)
Snapped trunk. This occurred when a substantial stump of the trunk was
still standing whereas another part of trunk was found fragmented on the
forest floor. Usually these trunk fragments were oriented in a particular
direction, except when the tree suffered death at an event in which
participated several trees simultaneously; C) Uprooted. This mode of death
was recorded when the trunk was found lying on the forest floor with the
roots exposed and lifted; D) Decomposed. This mode of death was
recorded when trees had disappeared completely leaving no trace at all.
Such trunks were assumed to have died and completely decomposed; E)
Other. This mode referred to the few events in which fragments were
found in a decomposed state that could not be assigned to any of the
aforementioned types of death.
Following standard procedures (Synnott 1979, 1991; Condit 1998; Vallejo
et al. 2005) recruits were defined as those individuals that reached the
minimal surveying size of DBH = 10 cm after the former census. Recruits
were localized, numbered, tagged, mapped, identified, and measured
exactly as was done with the individuals during the initial inventory of the
plots. More information regarding sampling procedures is in Chapter 2.
116
Figure 6-2. Synthesis of forest dynamics in profile diagrams for the upland
(tierra firme) and the floodplain (vrzea) plots in Pea Roja. Shown are
cross sections and projections on the soil surface. Upper diagram: upland
plot. The dotted lines indicate the trees in the back. Lower diagrams:
floodplain plot.
Data processing
The results are presented for each growth habit separately, and for the
whole forest, considering dicot trees (trees sensu stricto without palms or
lianas), palms and lianas. Tree palms do not have vascular cambium and
117
(6-1)
(6-2)
(6-3)
(6-4)
(6-5)
where ADI = annual diametric increment (cm per year), DBH0 = DBH at
the start of the time interval (t0), and DBH1 = DBH at the end period of the
time interval (t1). The time interval (t1 t0) is expressed in years. Records
of ADI < -0.2 cm per year and ADI > 4 cm per year were excluded as
these might be the result of measurement errors (Sheil et al. 1995; Chao et
al. 2008).
Finally, following standards methods (Oldeman et al. 2006), a drawing
was made of a forest transect of 10 m x 100 m in each plot. In these, all
plants with DBH 10 cm were included (trees, palms, lianas), taking into
account their spatial coordinates, DBH, total height, trunk height, and
crown size measured in two perpendicular directions.
119
Trees
Palms
Lianas
Total
257
40
1455
56.47
635.64c
19.2 11.24
10.0 87.4
20.2 5.3
2
1
30
0.44
2.00d
13.6 1.91
10.1 16.8
16.2 3.7
4
4
4
0.05
0.80e
12.3 0.90
11.1 13.2
27.2 4.7
263
44
1489
56.96
638.43
19.0 11.2
10.0 87.4
20.1 5.3
6.5 37.7
0.13 0.14
-0.13 0.13
182
40
1051
39.55
421.3f
18.94
10.0 90.2
16.0
1.0 40.7
0.22 0.31
3
1
104
1.59
7.59d
13.75
10.2 25.1
13.8
4.0 29.0
15
11
25
0.48
56.9e
14.49
10.0 38.0
27.5
16.0 35.0
200
45
1180
41.62
435.8
18.49
10.0 90.2
16.0
1.0 40.7
-0.18 2.53
Data from final inventories (TF 1997, VA 1999) bBiomass (Y, dry weight in kg),
120
6.4
Results
121
1990
66
43
80
58
62
58
30
11
20
29
1504
NI
1993
69
43
78
58
61
59
31
10
20
29
1505
1997
70
44
79
57
60
58
32
10
20
28
1489
1990
8.44
5.07
4.50
4.07
3.58
3.15
2.27
2.00
1.84
1.81
100
IVI (%)
1993
8.66
5.13
4.40
4.10
3.55
3.21
2.34
1.98
1.85
1.82
100
1997
8.66
5.26
4.46
4.11
3.51
3.21
2.44
2.00
1.87
1.79
100
1990
170
75
28
29
31
19
13
32
23
37
1207
NI
1994
172
75
29
29
34
19
14
34
23
39
1268
1999
170
63
25
28
26
19
13
43
21
37
1180
1990
9.68
4.18
3.77
3.10
2.48
2.44
2.36
2.32
2.21
2.08
100
IVI (%)
1994
9.31
3.97
3.73
3.02
2.64
2.33
2.31
2.32
2.13
2.07
100
1999
9.66
3.75
3.37
3.05
2.32
2.48
1.94
2.93
2.11
2.18
100
Floodplain
Brownea macrophylla
Astrocaryum sciophilum
Hyeronima alchorneoides
Eschweilera andina
Inga EA-397
Campsiandra angustifolia
Sterculia apeibophylla
Zygia latifolia
Pachira insignis
Euterpe precatoria
All species
122
Figure 6-3. Diameter distribution regarding trunk density, basal area and
biomass at the 1990 census (diagrams left) and regarding the mortality
(over a cumulative census period of 8.6 y) in trunks, basal area and
biomass (diagrams right). Light-grey colums are from the upland plot and
dark-grey columns from the floodplain plot. The diameter class width is 10
cm (the centre of the class is indicated).
The ten most important species in the upland and floodplain plots are
listed in Table 6-2 (see also Appendix 3). Overall, the rankings of the
species based on their importance values did not change between the
various censuses (1990-1997 in the upland plot and 1990-1999 in the
floodplain plot) despite slight differences in species composition due to
tree mortality and recruitment. In the upland plot Lecythidaceae, Fabaceae,
123
124
Figure 6-4. Family importance index (%) in the upland (left) and
floodplain (right) plots. NI% = relative number of individuals; DI% =
relative number of species; W% = relative above-ground biomass. Data of
the 1990 census.
In the upland plot, for the total period of 8.6 y, most death individuals
were found in Burseraceae (22), followed by Fabaceae (13), Lecythidaceae
(13), Lauraceae (12) and Sapotaceae (10), together corresponding to 71%
of lost individuals, 77% of the species that had died, and 77% of the
biomass of all dead trunks. Dipterocarpaceae, which was only represented
by one species, contributed to 10% of the mortality in biomass. Species
that died often were Protium paniculatum (10 trees), Diplotropis martiusii
(7 trees), Eschweilera parvifolia (5 trees) and Protium hebetatum (5 trees).
In the same time lapse, in the floodplain plot a high mortality was found
among trunks from Arecaceae (27), Mimosaceae (21), Rubiaceae (16),
Euphorbiaceae (14) and Caesalpiniaceae (12). Together these represented
47% of the dead trunks, 70% of the species that had died, and 36% of the
biomass of all dead trunks. Astrocaryum sciophilum (16 palms), Brownea
125
BA (m2)
W (ton)
Floodplain
NI
(individuals)
126
Periods
Period (years of
census)
Initial NI
Interval-1
1990-1993
(3.3)
1504
Interval-2
1993-1999
(5.3)
1505
Total
1990-1999
(8.6)
1504
Dead NI
Recruited NI
Final NI
Mortality (% y-1)
Recruitment (% y-1)
Periods (years of
census)
Initial BA
Dead BA
Recruited BA
Final BA
Growth in BA
Mortality (% y-1)
Recruitment (% y-1)
Initial W
Dead W
Recruited W
Final W
Growth in W
Mortality (% y-1)
Recruitment (% y-1)
Periods (years of
census)
Initial NI
66
67
1505
1.35
1.31
1990-1993
(3.3)
55.524
1.960
0.568
55.878
1.745
0.12
1.22
615.42
19.97
3.87
621.54
22.21
-0.11
1.25
1990-1993
(3.3)
1207
65
49
1489
0.84
0.61
1993-1997
(4.0)
55.877
2.160
0.429
56.963
2.816
-0.29
1.41
621.54
22.08
2.94
638.43
36.04
-0.56
1.52
1993-1999
(5.3)
1185
131
116
1489
1.06
0.87
1990-1997
(7.3)
55.524
4.191
1.080
56.963
4.485
-0.07
1.30
615.42
42.94
7.51
638.43
57.30
-0.32
1.37
1990-1999
(8.6)
1207
Dead NI
Recruited NI
Final NI
83
61
1185
109
104
1180
191
164
1180
BA (m2)
W (ton)
Mortality (% y-1)
Recruitment (% y-1)
Initial BA
Dead BA
Recruited BA
Final BA
Growth in BA
Mortality (% y-1)
Recruitment (% y-1)
Initial W
Dead W
Recruited W
Final W
Growth in W
Mortality (% y-1)
Recruitment (% y-1)
2.14
1.48
40.598
4.192
0.573
39.653
2.674
1.14
2.31
423.77
49.35
4.88
410.41
31.12
1.32
2.45
1.84
1.60
39.653
3.055
1.077
41.622
3.947
-0.42
2.27
410.41
29.80
8.50
435.83
46.72
-0.77
2.40
2.01
1.48
40.598
7.113
1.974
41.622
6.162
0.28
2.13
423.77
77.68
16.70
435.83
73.05
0.13
2.24
Types of death
In the upland plot more trees died standing than in the floodplain plot
(Table 6-4). For the total census period (8.6 y) 29% of the trunks died
standing in the upland plot, and only 17% died as such in the floodplain
plot. Regarding death by uprooting (tree fall) the results were opposite: in
the upland plot only 10% of the trunks had died by uprooting (over 8.6 y)
whereas in the floodplain this proportion was two times higher. All these
differences between the two plots were more pronounced when mortality
was calculated for basal area and biomass. Death through snapping trunks
was the most common mode of death, in both plots. Remarkably, 17% of
the dead trunks had disappeared entirely in the floodplain plot after 8.6 y,
against only 2% in the upland plot.
Recruitment and growth
Both plots showed different degrees of recruitment during the total interval
(1990-1999) of 8.6 years (Table 6.3). In terms of individuals the
recruitment in the upland plot was 7.7% (with respect to initial density, for
stems 10 cm), representing a of 0.87% per year. In the floodplain plot
this recruitment was almost two times higher (13.6% representing a of
1.48% per year). A similar between-plot difference was found for
recruitment in terms of basal area (1.97 m2 and a AB of 1.30% per year in
127
the floodplain plot versus 1.08 m2 and a AB of 2.13% per year in the
upland plot) and biomass (16.7 ton and a W of 1.37% per year in the
floodplain plot versus 7.51 ton and a W of 2.24% per year in the upland
plot).
The average annual growth in terms of diameter increment was 0.13 cm
per year in the upland plot. In the floodplain plot is was almost twice as
high (0.22 cm per year)(Table 6-5). Thin trees showed a wider range in
diameter increments than thick trees (Fig. 6-5). In both plots, newly
recruited individuals showed higher mean and maximum diameter
increments. In the upland plot, sub-canopy individuals of 10-15 m height
showed the highest annual diameter increments. In the floodplain, this
happened with sub-canopy individuals of 15-20 m height. On average,
however, and in both forests, larger individuals showed a tendency of
faster annual growth in trunk diameter.
Of the most important species in the upland plot, Pseudomonotes
tropenbosii, Monopteryx uaucu, Aspidosperma excelsum and Scleronema
micranthum showed the highest average annual diameter increments.
These four species reached the upper canopy. In the floodplain plot, Inga
EA-397, Hyeronima alchorneoides, Sterculia apeibophylla and
Eschweilera andina were the fastest growing species on a yearly basis.
The first species reached the middle part of canopy, and the other three
were upper canopy species. The dominant species in this forest (Brownea
macrophylla) showed a mean annual diameter increment of 0.05, which
represented only 38% of the average of all trunks in the plot.
128
Table 6-4. Modes of death for different census periods in the two plots, in absolute values and in proportion to the first
census (three columns at the right). NI = number of trunks, BA = basal area, W = biomass.
Upland
1990-1993
NI BA (m2)
Uprooted
5
0.19
Snapped
32
0.96
Standing
27
0.79
Decomposed 2
0.03
Other
0
0
Total
66
2.0
1990-1994
NI BA (m2)
Uprooted
19
1.2
Snapped
39
2.6
Standing
13
0.2
Decomposed 12
0.2
Other
0
0
Total
83
4.2
W (ton)
1.8
9.7
8.2
0.2
0
20.0
1993-1997
NI BA (m2)
8
0.35
43 1.13
13 0.67
1
0.02
0
0
65 2.2
W (ton)
4.1
10.7
7.2
0.1
0
22.1
W (ton)
14.0
32.0
2.1
1.4
0
49.4
1994-1999
NI
BA (m2)
19
0.8
43
1.2
20
0.6
20
0.3
7
0.1
109 3.1
W (ton)
8.4
12.5
5.7
2.5
0.7
29.8
Floodplain
1997-1999
NI
BA (m2)
1
0.04
8
0.16
0
0
0
0
0
0
9
0.2
W (ton)
0.3
1.4
0
0
0
1.7
1990-1999
NI (%) BA (%)
10.0
13.1
59.3
52.3
28.6
33.7
2.1
1.0
0
0
100
100
W (%)
14.2
49.8
35.3
0.7
0
100
1990-1999
NI (%) BA (%)
19.8
27.9
42.7
53.3
17.2
11.2
16.7
6.3
3.7
1.2
100
100
W (%)
28.3
56.2
9.8
4.9
0.9
100
129
Table 6-5. Annual diameter increment (cm per year) for in the upland and
floodplain plots in Pea Roja (all data from individuals with DBH 10
cm, found in 1.8 ha).
Annual diameter increment (cm per year)
Maximum
Minimum
Mean one
SD
Upland (1990-1997)
Pseudomonotes tropenbosii
Monopteryx uaucu
Eschweilera laevicarpa
Eschweilera punctata
Eschweilera ovalifolia
Eschweilera parvifolia
Scleronema micranthum
Aspidosperma excelsum
Erisma splendens
Brosimum rubescens
All tree species
Floodplain (1990-1999)
Brownea macrophylla
Astrocaryum sciophilum
Hyeronima alchorneoides
Eschweilera andina
Inga EA-397
Campsiandra angustifolia
Sterculia apeibophylla
Zygia latifolia
Pachira insignis
Euterpe precatoria
All species
130
0.48
0.78
0.44
0.88
0.20
0.89
0.50
0.73
0.29
0.25
1.35
0.00
-0.01
-0.13
0.00
-0.01
-0.01
0.03
0.03
0.01
0.00
-0.13
0.20 0.13
0.19 0.19
0.06 0.07
0.11 0.13
0.06 0.5
0.09 0.13
0.15 0.11
0.18 0.23
0.12 0.08
0.09 0.08
0.13 0.14
0.82
0.74
1.57
1.17
2.07
0.48
0.83
0.79
0.67
0.53
2.53
-0.17
-0.07
0.00
-0.18
0.02
-0.05
0.01
-0.18
0.00
-0.11
-0.18
0.06 0.09
0.11 0.21
0.45 0.40
0.28 0.30
0.80 0.54
0.12 0.14
0.29 0.28
0.12 0.19
0.22 0.22
0.03 0.10
0.22 0.31
Fig. 6-5. Annual diameter increment (ADI) for different growth forms,
DBH classes and height classes (RA: Rasant (Ht 0, < 5 m); UN:
understorey (Ht 5 m, < 10 m); SC-L: subcanopy, low (Ht 10 m, < 15
m); SC-H: subcanopy, high (Ht 15 m, < 20 m); CA-L: canopy, low (Ht
20 m, < 25 m); CA-H: canopy, high (Ht 25 m, < 30 m); EM-L:
emergent, low (Ht 30 m, < 35 m); EM-H: (Ht 35 m, < 40 m); SE:
super emergent (Ht 40 m) in the upland plot (left) and floodplain plot
(right) in Pea Roja. Measurements periods were 1990-1997 for the upland
plot and 1990-1999 for the floodplain plot. The vertical bars denote the
maximum and minimum increments.
131
132
(% y-1)
(% y-1)
1.35
1.31
1.8
5.3
0.84
0.61
1.8
8.6
1.06
0.87
1.8
3.3
2.14
1.48
1.8
5.3
1.84
1.60
1.8
8.4
2.01
1.48
2.6
2.70
2.40
2.85
2.70
1.98
1.78
4.9
2.33
1.93
1.88
2.5
1.04
3.09
0.9
0.95
10
7.75
1.77
1.84
0.81
2.83
7.75
2.85
2.37
11.67
1.97
1.96
7.75
2.70
2.25
5
16
3.2
5
17.2 - 19.5
6
1.15
3.67 0.7
3.5
0.87
2.5
15
1.26 - 3.55
2.7
10
1.18
4.25
10 - 29
1.67 0.67
4.25
10 - 29
1.81 1.10
1
0.5
0.42 - 9
~10
~10
6.4 0.7
2.34 0.31
1.21 0.53
1.50 0.19 1.59 0.19
0.42 - 9
6.4 0.7
4.4
4
4
50
50
13
13
13
3
5
1.80
2.01
2.24
3.4 0.3
2.6 0.2
1.33
1.67
1.80
This dissertation; 2lvarez et al. 2008; 3Korning and Balslev 1994; 4Gentry and
Terborgh 1990; 5Phillips et al. 1994 6Rankin de Merona et al. 1990; 7D'Angelo et al.
2004; 8Felfili 1995; 9Rolim et al. 1999; 10Uhl et al. 1988; 11Carey et al. 1994; 12Chao
et al. 2008; 13Lewis et al. 2004a; 14Lieberman et al. 1985; 15Condit et al. 1995
6.5
Discussion
For both plots the mortality rate based on number of individuals was
higher than recruitment rate in all time lapses, indicating that mortality
preceded recruitment. However, the average of these mortality and
recruitment rates, which represent a measure of turnover, was lower in the
upland plot than in the floodplain plot (Fig. 6-6). Compared to other
Neotropical forests, the turnover in both plots was relatively low (Phillips
et al. 2004).
Reviewed by Phillips et al. (2004), mortality and recruitment rates in
Amazonia across 97 sites showed an average of ca. 2% per year. Higher
turnover rates were found in the northwestern Amazon compared to
eastern and central Amazonia, on rich soils compared to poor soils, and in
areas without marked seasonality in rainfall compared to areas where
climate showed a pronounced dry season. Phillips et al. (2004) further
reported that Pan-Amazonian averages for mortality (at 52 sites) increased
from 1.58 0.1 to 1.91 0.13% per year (corrected for census interval) for
two successive periods between 1976-2001. Also recruitment rates (57
sites) increased from 1.70 0.11 to 2.34 0.15% per year and turnover
rates (55 sites) from 1.65 0.09 to 2.11 0.12% per year, indicating that
the dynamics of these forests has increased over the years.
The mortality rates recorded in the plots near Pea Roja were below 5%,
and therefore corresponded to so-called background mortality (Lugo and
Scatena 1996). Catastrophic events, such as those reported in Peru (Foster
and Terborgh 1998), are not known from the study area at Pea Roja. For
most tropical forest trees the annual mortality ranges between 1% and 3%.
Higher rates were reported from El Verde in Puerto Rico (Crow 1980),
Barro Colorado Island in Panama (Hubbell and Foster 1990), and Aangu
in Ecuador (Korning and Balslev 1994). On Barro Colorado the high
mortality (3%) was attributed to a prolonged and severe dry season
weather event associated with the "El Nio" (Hubbell and Foster 1990).
Reports of mortality rates from before de 1983 drought in Barro Colorado
(Putz and Milton 1983) were much lower (1.83% and 1.06%). On the other
hand, low levels of mortality have been associated with soil substrates that
are low in nutrients (Rankin de Merona et al. 1990; Manokaran and
Swaine 1994, Phillips et al. 2004).
134
1978). In the floodplain plot, the mortality rates regarding basal area and
biomass were low (1.14% and 1.32%, respectively) during the initial
census period, and became negative in the second census period (-0.42%
and -0.77, respectively). This seemed to indicate a shift from a homeostatic
phase to an accumulation phase. Perhaps this was caused by a high
recruitment after the death of some large trees and the subsequent release
of space suitable for the accelerated growth of individuals of intermediate
sizes, as can be deduced from Fig. 6-3. This conclusion would also stem
with the shape of the upper canopy in the floodplain plot, which had a
more open and a more irregular surface than in the upland plot (Fig. 6-2).
Death modes
In the upland plot death by snapping and standing death was most
common. In the floodplain plot tree uprooting was relatively common.
These results suggest that the process of mortality in the upland plot
included events acting on smaller spatial scales, at more gradual pace and
at a lower frequency than in the floodplain plot. Death rates in the upland
plot seem to reflect more a mortality process caused by pathogens,
herbivores, senescence, competition for resources, or a combination
thereof, whereas in the floodplain plot weak rooting due to the poor soil
drainage seems to be more important.
Death by snapping is commonly found as the most frequent type of death
in tropical forests (Londoo and Jimnez 1999; Alcaraz 2001).
Occasionally, other death modes become more important, as in montane
forests of Venezuela (Carey et al. 1994), where standing death (64%) were
most common, followed by tree fall (17%), snapping (11%) and unknown
(11%). Differences in death mode have been associated to edge effects in
large plots. For instance, in central Amazonia (near Manaus) mortality
rates near forest edges (3.67 0.70% per year) were three times higher
than in the forest interior (1.05 0.22% per year). Here, tree death by
snapping was the most common type in edges and interior (34-36%), but
death by uprooting was more abundant in the edge whereas standing death
was more common towards the interior of the plot (D 'Angelo et al. 2004).
In eastern Ecuador, Gale and Barfod (1999) reported that 34% of tree
death was by uprooting, 35% by snapping, 15% by standing death, and
16% unclassified. In that study, the type of death differed between
136
dicotyledonous tree and palms (Iriartea deltoidea). Slope, altitude and the
presence of buttresses influenced death modes. Uprooting and snapping
occurred in a clumped pattern and was explained by steep terrain and high
rainfall. Standing death was randomly distributed over the plots.
Figure 6-6. Turnover rates calculated as the average of the mortality and
recruitment of the tropical forests, shown in Table 6-6. The dot is the
average and the length of the bar represents the difference between
mortality and recruitment rates.
Growth by diameter increment
Overall, the diameter increments in the two forest plots (0.13 cm per year
in the upland plot and 0.21 cm per year in the floodplain plot) were low
compared to those reported from other tropical forests (Taylor et al. 1996).
For example, 0.45 cm per year was measured in Venezuela (Veillon 1985);
0.5 cm per year in Puerto Rico (Crow and Weaver 1977); 0.44 cm per year
in Costa Rica (Lieberman and Lieberman 1985); 0.12 to 0.27 cm per year
in Malaysia (Nicholson quoted by Whitmore 1975); 0.39 to 0.78 cm per
year in so-called cativales in Colombia (Gonzlez 1994); 0.85-0.96 cm per
137
Acknowledgements
140
7
Synthesis
Ana Catalina Londoo Vega
7.1
Architectural analysis
144
The higher species richness in the upland plot compared to floodplain plot
and the low overlap in species composition between the two plots fitted
well into the general scheme of upper Amazonian forest-landscape
associations (Duivenvoorden and Duque 2010). These forest-landscape
associations have two explanations, which take place simultaneously but
which rely on principally different processes. The first explanation
emphasizes niche-filling mechanisms: species tend to show higher
abundances in those environments where they have a competitive
advantage over other species, for example because of species-specific
adaptations related to limited resource availability or to species-specific
abilities to overcome disturbances. This explanation tends to emphasize
the role of the environment in explaining differences in community
assemblage. During and shortly after the installment of the two plots it was
already known that they differed strongly in soils and decomposition of
organic matter. In the upland plot, soil nutrient availability and
decomposition rates were among the lowest levels for mesic upland sites
in Amazonia (Duivenvoorden and Lips 1995). In sharp contrast, the soils
from the floodplain were less nutrient-poor and decomposition rates were
higher, because the parent material was much younger and enriched
through the occasional sedimentation during flooding by the Caquet
River. The results from the permanent plot study in this dissertation
suggest that the two forests also differ substantially regarding forest
dynamics (mortality, recruitment and modes of death). In the upland plot
the tree turnover was relatively slow (comparatively a low mortality and
recruitment) and trees tended to die in upright position yielding a low
spatial spread of canopy disturbances. In the floodplain plot trees died
quicker, grew faster, and tree uprooting was more common leading to
more widespread disturbances of the forest structure through the entire
plot. Clearly these differences agree with the between-plot differences in
decomposition rates and soil nutrient availability of the plots
(Duivenvoorden and Lips 1995). They also line up with results from
analyses of forest dynamics along gradients of soils and forest productivity
in the tropical forests worldwide (Malhi et al. 2006).
The second mechanism to explain the divergent between-plot patterns in
species richness and composition is by random walk processes modulated
through seed dispersal (Hubbell 2001). This explanation emphasizes that
145
plot diversity patterns the results of the permanent plots survey deviated
from the Pan-Amazonian trends. The tree mortality rates in the upland plot
were among the lowest recorded for Amazonian as a whole, and clearly far
lower than would be expected purely because of the location of this plot in
upper Amazonia. The species richness in the upland plot was higher than
that of the floodplain plot, even though its tree turnover was lower. This
suggests that particular conditions of occasional flooding and the presence
of poorly drained soils in Holocene to Lateglacial floodplains together
with the associated assemblage of tree species give rise to a high level of
forest dynamics relative to tree diversity. In general, the results from the
two permanent plots indicate that it is hazardous to apply insights based on
regional studies to local situations without proper knowledge of the local
physiography and terrain conditions.
7.4
147
148
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177
178
Resumen
En la dcada de los aos ochenta del siglo pasado, se establecieron dos
parcelas permanentes en bosques maduros cerca de Araracuara, en la parte
central de la Amazonia colombiana. Las parcelas se instalaron en dos
unidades de paisaje contrastantes: tierra firme y llanura de inundacin del
ro Caquet. El objetivo general de esta tesis es proporcionar
conocimientos bsicos sobre la estructura, la composicin de especies y la
dinmica del bosque de estas dos parcelas permanentes. Las parcelas se
instalaron en la comunidad Nonuya de Pea Roja, a unos 70 km aguas
abajo de Araracuara. En el rea la precipitacin media anual es de
aproximadamente 3000 mm anuales, sin perodo seco. La parcela de tierra
firme se estableci en una unidad de paisaje perteneciente al Plano
Sedimentario Terciario, donde los suelos son Ultisoles muy pobres. La
otra parcela fue localizada en la Llanura de Inundacin Espordica del ro
Caquet, que se caracteriza por unas depresiones poco profundas
alternadas con barras de cauce convexas, de hasta 2 m de altura. Los
suelos son Inceptisoles moderadamente bien drenados. El rea final de
ambas parcelas fue de 1,8 ha.
El Captulo 3 describe Pseudomonotes tropenbosii Londoo et al. Este
nuevo taxn es la segunda aparicin reportada de un miembro de la familia
Dipterocarpaceae en el Neotrpico. La nueva entidad se diferencia del
resto de este familia por la ausencia de tricomas fasciculados y spalos
visiblemente aliformes (llegando a 10-16 cm de longitud) y un vulo por
lculo con placentacin casi basal (sub-basal). La especie apareci como
uno de los rboles dominantes en la parcela de tierra firme. Estudios
posteriores (anatoma de la madera y la filogenia molecular) han
confirmado su posicin taxonmica en la subfamilia Monotoideae de la
familia Dipterocarpaceae.
El Captulo 4 trata sobre el anlisis arquitectnico llevado a cabo en tres
especies de Myristicaceae (Iryanthera tricornis, Osteophloeum
platyspermum y Virola pavonis). Las tres especies mostraron un
crecimiento de acuerdo con el modelo de Massart, al igual que otras
especies de Myristicaceae (V. michelii y V. surinamensis), en el cual se
presentan tres rdenes de ejes. Las reiteraciones juegan un papel
preponderante en el desarrollo de stas tres especies. El anlisis de la
180
differs from the rest of this family in the absence of fasciculate trichomes
and in having sepals conspicuously aliform (reaching 10-16 cm in length)
and one ovule per locule with nearly basal (sub-basal) placentation. The
species appeared as one of the dominant trees in the upland plot.
Subsequent studies (wood anatomy and molecular phylogeny) confirmed
its taxonomic position in the subfamily Monotoideae of the family
Dipterocarpaceae.
Chapter 4 reports on an architectural analysis, which was carried out for
three species of Myristicaceae (Iryanthera tricornis, Osteophloeum
platyspermum and Virola pavonis). The three species showed a growth
according to Massart's Model, just as other species of Myristicaceae (V.
michelii and V. surinamensis), during which three orders of axes were
reached. Reiterations are important in the growth and development of the
three species. The architectural analysis suggested that the growth plan of
these species would contribute to their ability to maintain healthy
populations in the subcanopy forest. The case-study of the three subcanopy
tree species of Myristicaceae shows the large potential of architectural
analysis to build hypotheses why species might be adapted to particular
conditions of forest development, especially regarding abundant
subcanopy (or canopy) species.
A total of 1149 species, 347 genera and 98 families were found in the two
plots (Chapter 5). Arborescent species comprised 65% of the total at both
sites; climbers (woody and herbaceous) contributed 24%, shrubs 8% and
terrestrial herbs 3%. The number of species was larger in the upland plot
(698) than in the floodplain plot (511), but the number of genera was quite
similar (236 in the upland and 235 in the floodplain plot), whereas the
number of families was slightly higher in the floodplain plot (84,
compared to 80 families in the upland plot). Between the plots 67% of the
families were shared, 36% of the genera, and only 5% of the species. In
total, the floristics in the plots showed a strong affinity to the flora
recorded in central Amazonia, near Manaus.
The mortality in the upland plot was almost twice as low as in the
floodplain plot ( = 1.06 versus = 2.0, respectively, over 8.6 y)(Chapter
6). In terms of basal area and biomass, the discrepancies in mortality rates
between the floodplain and upland plot were even more pronounced. Large
184
187
De sterfte in het hoogland perceel was bijna twee keer zo laag als in het
alluviale perceel (respectievelijk = 1.06 en = 2.0, in 8,6 jaar)
(Hoofdstuk 6). In termen van stamoppervlak en biomassa waren de
verschillen in sterftecijfers tussen de twee proefvlakken nog meer
uitgesproken. Grote bomen droegen aanzienlijk bij aan de sterftecijfers
berekend voor stamoppervlak en biomassa. Voor de totale periode (8,6
jaar) stierf 29% van de stammen in staande positie in het hoogland perceel,
en slechts 17% op een dergelijk wijze in het alluviale perceel. Wat betreft
dood door ontworteling waren de resultaten tegenovergesteld: in het
hoogland perceel stierf slechts 10% van de stammen door ontworteling (in
8,6 jaar), terwijl in het alluviale proefvlak dit percentage twee keer hoger
was. In termen van dichtheid was de aanwinst in het hoogland perceel
bijna twee keer lager dan in het alluviale perceel (respectievelijk =
0,87% en = 1,48% per jaar, in 8,6 jaar). Vergelijkbare verschillen tussen
de percelen werden gevonden voor de aanwinst in termen van
stamoppervlak en biomassa.
De gemiddelde jaarlijkse groei in stamomvang was 0,13 cm per jaar in het
hoogland perceel. In het alluviale perceel was deze groei bijna tweemaal
zo hoog (0,21 cm per jaar). De jaarlijkse sterfte, op basis van de dichtheid,
van het hoogland perceel lag in de buurt van de onderste waarden
gerapporteerd voor het noordoostelijk deel van Amazonegebied. De
jaarlijkse sterfte, op basis van de dichtheid, van het alluviale perceel was
dicht bij de ondergrens gemeld voor de bossen in het noordwesten van het
Amazonegebied. Voor beide percelen gold dat deze sterfte hoger was dan
aanwinst, hetgeen betekent dat de sterfte voorafgaat aan de aanwinst. Het
gemiddelde van de sterfte- en aanwinstsnelheden, die een maat vormen
voor de bosomvorming, was lager in het hoogland perceel dan in het
alluviale perceel. Vergeleken met andere neotropische bossen was deze
bosomvorming in beide percelen relatief laag.
De ontdekking van Pseudomonotes tropenbosii in het hoogland proefvlak
illustreert de grote vooruitgang in floristische inventarisaties in het
Colombiaanse Amazonegebied sinds het midden van de jaren tachtig van
de vorige eeuw (Hoofdstuk 7). Op basis van houtanatomie en
fylogenetische analyses is de taxonomische positie van de soort bevestigd.
De architectuuranalyse van de drie Myristicaceae soorten suggereerde dat
het groeiplan van deze soorten bijdraagt aan hun vermogen om gezonde
188
189
190
Appendices
Appendix 1
This appendix contains the descriptions and laboratory analyses of soil
profiles 125 and 126 taken from Duivenvoorden and Lips (1995). Methods
of analyses and sampling procedures are described in Duivenvoorden and
Lips (1995). Mapping units refer to the legend of the landscape ecological
map of the Middle Caquet Basin (Duivenvoorden and Lips 1993).
Profile number: 125
Mapping unit: Sv
Soil classification: clayey, kaolinitic, isohyperthermic typic Kandiudult (SSS 1992),
xanthic Ferralsol (FAO 1988)
Date of examination: February 19 and October 25 1990
Authors: J.M. Lips & J.F. Duivenvoorden
Location: 875 m from the left bank of the Caquet River in front of the Island
Sumaeta, Amazonas, Colombia (039'S, 7205'W)
Elevation: approx. 200 m a.s.l.
Landform: sedimentary Tertiary plain; physiographic position: shoulder straight
slope; surrounding topography: steeply dissected; microtopography: none
Slope: sloping (6); exposition: 222 (SW)
Vegetation: high forest of high biomass, of the Swarzia schomburgkii-Clathrotropis
macrocarpa community
Climate: Afi (3060 mm/year, 25.7C)
General information on the soil
Parent material: Tertiary fluvial deposits, with a cover of Pleistocene? fluvial deposits
Drainage class: well drained (4)
Moisture conditions in profile: moist throughout
Depth of groundwater: not encountered
Presence of surface stones and rock outcrops: none
Evidence of erosion: none
Human influence: none
Description of the individual soil horizons
L
9.78-8.00 cm. 90% leaf, 10% branch; cover 75%; loose to non compact
matted structure; loose consistency; leafy character; abrupt broken
boundary to:
8.00-3.95 cm. 38% leaf, 2% branch, 50% root, 10% fine material; reddish
to dark brown (5-7.5YR4/4) dry; irregular structure; resilient consistency;
fibrous character; abundant roots 5 mm, common roots 6-10 mm; very
few roots >10 mm, random orientation; gradual wavy to irregular boundary
to:
3.95-0 cm. 5% leaf, 42% fine material, 53% root; reddish to dark brown (57.5YR4/4) dry; granular structure; friable consistency; greasy character;
abundant roots 5 mm, common roots 6-10 mm; few roots >10 mm,
random orientation; gradual wavy to irregular boundary to:
192
0-8 cm. Brown (7.5YR4/4) moist loam; weak medium granular; very
friable moist, slightly sticky and slightly plastic wet; common very fine and
fine, frequent medium and coarse roots; clear wavy boundary to:
AB
Bt11
30-70 cm. Bright brown (7.5YR5/6) moist clay; moderate medium and
coarse subangular blocky; friable moist, slightly sticky and slightly plastic
wet; few very fine, fine and medium, very few coarse roots; few faunal
activity; diffuse smooth boundary to:
Bt12
70-160+ cm. Bright brown (7.5YR5/7) moist clay; moderate medium and
coarse subangular blocky; friable moist, slightly sticky and slightly plastic
wet; few very fine, fine and medium, very few coarse roots
Remarks: many coarse angular quartz sand particles present throughout the profile;
data part of Bt12 derived from augering; H horizon discontinuous
Analyses profile 125
horizon
depth
A
AB
Bt11
Bt12 (80-100)
Bt12 (130-160)
cm
0-8
8-30
30-70
70-160+
70-160+
texture
sand silt clay
%
72 10 18
46 14 40
42 8 50
42 10 48
42 10 48
horizon
A
AB
Bt11
Bt12
Bt12
horizon
L
F
H
necromass
meansd
pH (1:1)
H20 KCl
3.7
4.2
4.6
4.7
4.9
C
%
1.97
0.42
0.13
-
exchange properties
Na acidity bases CEC7 ECEC
cmol(+)/kg soil
Ca
Mg
0.2
0.2
0.2
0.2
0.2
2.0
1.7
1.1
1.0
0.9
0.9
0.5
0.8
0.5
0.5
15.3
8.5
6.0
4.8
4.4
2.9
2.2
1.9
1.5
1.4
bulk
density
g/ml
0.8
1.2
1.3
1.3
-
P
avail.
mg/kg
1
1
1
1
1
CECc ECECc
cmol(+)/kg
clay
10.3 3.4
14.1 4.4
10.2 3.5
9.4
3.1
8.9
3.0
BS
%
6
6
13
11
12
totals
LOI
HNO3-H2O2 digestion
Ca Mg K Na
P
N
mg/kg
%
934 617 833 22 145 15664 98
269
112 127 392 68 166 14977 67
199
39 82 150 42 124 11299 46
HF-H2SO4 digestion
Ca Mg K Na
P
g/m2
2000
21600 134 184 488
11700 79 162 243
87
98
193
4.45-2.20 cm. 90% leaf, 10% branch<2% fruit; cover 60%; single particle
structure; loose consistency; leafy character; abrupt broken boundary to:
2.20-0 cm. 80% leaf, 10% branch, 10% root, <2% fruit; brown (107.5YR4/4) dry; single particle to compact matted structure; loose
consistency; leafy character; common roots 5 mm, no roots 6-10 mm; no
roots >10 mm, random orientation; common faunal activity (worm
droppings); clear irregular boundary to:
Ah
0-2 cm. Brown (10YR4/4) moist silty loam; moderate fine and medium
crumb; very friable moist, slightly sticky and slightly plastic wet; common
very fine, fine, medium and coarse roots; common faunal activity; clear
wavy boundary to:
AB
2-22 cm. Yellowish brown (10YR5/6) moist silty clay loam; moderate
medium subangular blocky; very friable moist, slightly sticky and slightly
plastic wet; common very fine and fine, few medium and coarse roots;
common faunal activity; few charcoal fragments; gradual smooth boundary
to:
Bw1
22-45 cm. Yellowish brown (10YR5/6) moist silty clay; moderate medium
subangular blocky; friable to firm moist, slightly sticky and plastic wet;
very few very fine, fine, medium and coarse roots; gradual smooth
boundary to:
194
Bw2
45-75 cm. Yellowish brown (10YR5/6) moist silty clay loam, with
common medium distinct diffuse bright brown (7.5YR5/6) and few
medium distinct diffuse orange grey mottles; moderate medium and coarse
subangular blocky; friable moist, slightly sticky and slightly plastic wet;
very few small soft Fe/Mn nodules; very few very fine, fine, medium and
coarse roots; gradual smooth boundary to:
Bwg
75-155+ cm. Yellowish brown (10YR6/3) moist silty clay loam, with
common medium distinct clear bright brown (7.5YR5/6) and yellowish
brown (10YR5/6) mottles; moderate medium and coarse subangular
blocky; friable moist, slightly sticky and slightly plastic wet; few small and
large soft Fe/Mn nodules; very few very fine, fine, medium and coarse
roots
Remarks: sand size mica fragments visible throughout the profile; data part of B22
derived from augering
Analyses profile 126
horizon
depth
Ah
AB
Bw1
Bw2
Bwg (80-95)
Bwg (140-155)
cm
0-2
2-22
22-45
45-75
75-155+
75-155+
texture
sand silt clay
%
24 46 30
16 44 40
12 38 50
24 36 40
26 40 34
14 36 50
horizon
Ah
AB
Bw1
Bw2
Bwg
Bwg
horizon
L
F
necromass
meansd
pH (1:1)
H20 KCl
3.9
3.9
4.2
4.6
4.7
5.1
C
%
3.26
0.34
0.20
-
exchange properties
Na acidity bases CEC7 ECEC
cmol(+)/kg soil
Ca
Mg
3.2
0.2
0.2
0.2
0.2
3.2
2.8
4.3
6.5
5.8
6.1
5.5
7.4
0.7
0.7
1.5
0.4
6.8
23.8
13.7
14.1
13.3
16.2
15.7
10.2
5.0
7.2
7.3
6.5
12.3
bulk
density
g/ml
0.5
1.3
1.4
1.4
1.4
-
P
avail.
mg/kg
4
1
1
1
1
2
CECc ECECc
cmol(+)/kg
clay
34.1 18.0
30.7 11.2
26.5 13.8
32.6 18.0
47.2 19.0
31.2 24.5
BS
%
31
5
5
11
3
43
totals
LOI
HNO3-H2O2 digestion
Ca Mg K Na
P
N
mg/kg
%
g/m2
3500
11496 2760 2569 42 424 18059 90
9500 8120 2384 4627 683 724
7184 2097 1864 68 463 18541 69
HF-H2SO4 digestion
Ca Mg K Na
P
195
Appendix 2
Listado de plantas vasculares en parcelas permanentes en bosques del
Plano Sedimentario Terciario (PST, tierra firme) y de la Llanura Aluvial
con Inundacin Espordica del ro Caquet (LAI, vrzea) en Pea Roja,
Amazonia colombiana. Publicado en Caldasia 19(3) 431-463 (1997).
Se incluyen plantas vasculares con altura total mayor o igual que 50 cm.
Para cada especie se indica: su nombre; su hbito de crecimiento (ver
abajo); la localidad donde se encontr (L: Llanura Aluvial con inundacin
espordica del ro Caquet; P: Plano Sedimentario Terciario); y una
muestra botnica de referencia (colectores: EA: Esteban lvarez; CL: Ana
Catalina Londoo).
Formas de crecimiento: rboles (A), arbustos (T), escandentes herbceos
(SH), escandentes leosos (SL), estranguladoras (SZL), helechos arbreos
(FA), helechos escandentes (FSH), helechos terrestres (FH), hemiepfitas
herbceas (SEH), hemiepfitas leosas (SEL), hierbas terrestres (H),
palmas arbreas cespitosas (PAC), palmas arbreas monoestipitadas
(PAM), palmas arbustivas acaules (PTU), palmas arbustivas cescpitosas
(PTC), palmas arbustivas monoestipitadas (PTM), palmas escandentes
(PSL).
En los bosques inventariados se identificaron las plantas hasta el nivel de
especie taxonmica (Grant 1989); cuando esto no fue posible se emplearon
morfoespecies en la denominacin de los taxones. Para las familias de
plantas superiores se sigui el sistema de Cronquist (1981), con excepcin
de Caesalpiniaceae, Fabaceae y Mimosaceae, que fueron agrupadas en
Leguminosae en la presentacin de resultados para facilitar la comparacin
con otros estudios. Cecropiaceae se presenta separada de Moraceae,
Hippocrateaceae aparte de Celastraceae y Mendonciaceae segregada de
Acanthaceae. Para las Pteridophyta se emple la nomenclatura de familias
de Mabberley (1990).
ACANTHACEAE: Aphelandra aurantiaca (Scheidweiler) Lindley.; H;
L; CL 1578. Justicia chlorostachya; H; L; EA 258. J. sp. 1; H; P; EA
1046.
ADIANTACEAE: Adiantum latifolium Lam.; FH; L; EA 561.
196
198
203
209
212
venulosa (C. Martius & & Eichler) Pierre; A; L; EA 629. Pouteria Sect.
Franchetella sp. 1; A; P; CL 766. P. Sect. Rivicoa sp. 1; A; P; CL 1155. P.
aff. laviegata (C. Martius) Radlkofer; A; L; EA 666. P. aff. ucuqui Pires &
Schultes; A; L, P; EA 652. P. baehniana Monachino; A; L; EA 435. P.
campanulata Baehni; A; P; EA 896. P. cf. cladantha Sandwith; A; L; EA
428. P. cf. maguirei (Aubrville) Pennington; A; P; CL-1668. P. cf.
multiflora (A. DC.) Eyma; A; P; CL 726. P. cf. oblanceolata Pires; A; P;
CL 237. P. cf. trilocularis Cronquist; A; P; EA 1118. P. cf. williamii
(Aubrville & Pellegrin) Pennington; A; P; EA 864. P. cuspidata (A. DC.)
Baehni; A; L, P; EA 113. P. guianensis Aublet; A; P; EA 852. P. sp. 01;
A; L; EA 708. P. sp. 02; A; P; EA 122. P. sp. 03; A; P; EA 1007. P. sp.
04; A; P; CL 721. P. sp. 05; A; L; EA 15. P. sp. 06; A; P; EA 1111. P. sp.
07; A; P; EA 149. P. sp. 08; A; P; CL 509. P. sp. 09; A; P; CL 340. P. sp.
10; A; P; CL 173. P. sp. 11; A; P; CL 1480. P. sp. 12; A; P; CL 495. P. sp.
13; A; P; CL 1473. P. sp. 14; A; P; CL 916. P. sp. 15; A; P; EA 62. P. sp.
16; A; P; EA 835. P. sp. 17; A; P; EA 1113. P. sp. 19; A; P; EA 1147. P.
sp. 20; A; P; EA 847. P. sp. 21; A; L; EA 659. P. sp. 22; A; L; EA 401. P.
sp. 23; A; L; CL 1628. P. sp. 24; A; P; EA 1029. P. sp. 25; A; P; CL 1003.
P. sp. 26; A; P; EA 183. P. sp. nov. 1; A; P; CL 1017. P. sp. nov. 2; A; P;
EA 862. P. torta (C. Martius) Radlkofer; A; L, P; CL 981; EA 386. P.
vernicosa Pennington; A; P; CL 1335. Pradosia cochlearia (Lecom.)
Pennington ssp. praealta (Ducke) Pennington; A; P; CL 1272. Sapotaceae
sp. 1; A; L; EA 699. S. sp. 2; A; P; CL 584. S. sp. 3; A; P; EA 1035.
SIMAROUBACEAE: Picramnia aff. sellowii Planchon ssp. spruceana
(Engler) Pirani; A; P; CL 1598. P. sp. 1; T; P; EA 1053. P. sp. 2; A; L; CL
1638. P. sp. 3; A; P; CL 5. P. sp. 4; A; P; CL 561. P. sp. nov. aff. P.
caracasana Engler; A; L; EA 1228. Simaba aff. polyphylla (Cavalcante)
W. Thomas; A; P; CL 590. S. guianensis Aubl. ssp. ecaudata (Cronquist)
Cavalcante; A; P; CL 1293. Simarouba amara Aublet; A; L, P; CL 1071.
SMILACACEAE: Smilax aequatorialis A. DC.; SH; L; EA 650. S. sp. 1;
SH; L; EA 1236. S. sp. 2; SH; L; EA 564. S. sp. 3; SH; P; CL 393. S. sp. 4;
SH; P; EA 1062.
SOLANACEAE: Cestrum? sp. 1; T; P; CL 288. Solanum sp. 1; SH; L;
EA 371. S. sp. 2; SH; L; EA 693. S. sp. 3; SH; L; EA 1226. Solanum?
Lycianthes?; SH; L; EA 250.
215
216
Appendix 3
Indice de valor de importancia (IVI en %) para vegetacin con DBH 10
cm, en 1.8 ha en bosques de tierra firme y de vrzea. Inventario incial
(1990). HCR: hbito de crecimiento: A: rboles; PAM: palmas arbreas;
SL: lianas; SZL: estranguladoras. NI: densidad relativa (%) del nmero de
individuos. FR: frecuencia relativa (%). W: biomasa area seca relativa
(%). Se muestran las especies en secuencia alfabtica.
Tierra firme, inventario inicial (ao 1990).
ESPECIE
Abarema adenophora (Ducke)
Barneby & J.W.Grimes
Abarema EA-785
Abarema leucophylla (Spruce ex
Benth.) Barneby & J.W.Grimes
Abuta CL-1161
Aiouea CL-1336
Alchornea discolor Poepp.
Alchornea floribunda (Benth.)
Muell. Arg.
Amaioua CL-1053
Aniba CL-1080
Aniba CL-1659
Aniba CL-2749
Aniba puchury-minor (Mart.) Mez
Aniba williamsii O. C. Schmidt
Annona excellens R. E. Fries
Anthodiscus obovatus Benth.
Aspidosperma CL-1001
Aspidosperma CL-1277
Aspidosperma desmanthum Benth.
ex Mll.Arg.
Aspidosperma EA-180
Aspidosperma excelsum Benth.
Aspidosperma schultesii Woodson
Beilschmiedia aff. brasiliensis
(Kortz.) Kortz.
Bignoniaceae NI-1493
Bocageopsis canescens (Spruce ex
HCR NI
NI%
FR%
W%
IVI%
A
A
2
1
0.133
0.066
0.143
0.072
0.039
0.079
0.105
0.072
A
SL
A
A
9
1
12
3
0.598
0.066
0.798
0.199
0.645
0.072
0.717
0.215
0.462
0.024
0.527
0.307
0.568
0.054
0.681
0.241
A
A
A
A
A
A
A
A
A
A
A
1
1
1
1
1
6
1
1
1
1
1
0.066
0.066
0.066
0.066
0.066
0.399
0.066
0.066
0.066
0.066
0.066
0.072
0.072
0.072
0.072
0.072
0.430
0.072
0.072
0.072
0.072
0.072
0.074
0.010
0.057
0.026
0.012
0.131
0.012
0.011
0.225
0.012
0.011
0.071
0.050
0.065
0.055
0.050
0.320
0.050
0.050
0.121
0.050
0.050
A
A
A
A
1
2
11
11
0.066
0.133
0.731
0.731
0.072
0.143
0.789
0.789
0.010
0.263
4.465
0.977
0.049
0.180
1.995
0.832
A
SL
A
2
1
2
0.133
0.066
0.133
0.143
0.072
0.143
0.069
0.029
0.043
0.115
0.056
0.106
217
ESPECIE
Benth.) R. E. Fries
Bocageopsis multiflora (Mart.) R.
E. Fries
Brosimum longifolium Ducke
Brosimum rubescens Tauber
Brosimum utile (Kunth) Pittier
Buchenavia parvifolia Ducke
Caesalpiniaceae CL-1288
Cariniana decandra Ducke
Caryocar glabrum (Aubl.) Pers.
Caryocar gracile Wittm.
Caryodaphnopsis CL-2520
Caryodaphnopsis EA-906
Casearia oblongifolia Cambess.
Casearia singularis Eichler
Cecropia membranacea Trcul
Chrysobalanaceae EA-827
Chrysophyllum prieurii A. DC.
Chrysophyllum sanguinolentum
(Pierre) Baehni
Chrysophyllum superbum
T.D.Penn.
Clathrotropis macrocarpa Ducke
Compsoneura capitellata (A. DC.)
Warb.
Conceveiba latifolia Benth.
Couepia bracteosa Benth.
Couepia chrysocalyx (Poepp. &
Endl.) Benth. ex Hook. f.
Couepia CL-1321
Couepia CL-2750
Couepia guianensis Aubl.
Dacryodes chimatensis Steyerm. &
Mag.
Dacryodes CL-1166
Dacryodes nitens Cuatrec.
Dacryodes sclerophylla Cuatrec.
Dendrobangia boliviana Rusby
Dendropanax tessmannii (Harms)
Harms
Dicranostyles sericea Gleason
218
HCR NI
NI%
FR%
W%
IVI%
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
1
1
29
16
1
9
11
3
10
1
1
1
1
1
1
1
0.066
0.066
1.928
1.064
0.066
0.598
0.731
0.199
0.665
0.066
0.066
0.066
0.066
0.066
0.066
0.066
0.072
0.072
1.864
1.147
0.072
0.645
0.789
0.215
0.717
0.072
0.072
0.072
0.072
0.072
0.072
0.072
0.022
0.021
1.634
1.176
0.039
0.488
1.750
0.079
1.318
0.181
0.035
0.016
0.035
0.011
0.009
0.022
0.053
0.053
1.809
1.129
0.059
0.577
1.090
0.164
0.900
0.106
0.058
0.052
0.058
0.050
0.049
0.053
19
1.263
1.362
0.966
1.197
A
A
2
14
0.133
0.931
0.143
1.004
0.021
0.264
0.099
0.733
A
A
A
17
1
1
1.130
0.066
0.066
1.147
0.072
0.072
0.297
0.014
0.115
0.858
0.051
0.084
A
A
A
A
2
7
1
1
0.133
0.465
0.066
0.066
0.143
0.502
0.072
0.072
0.072
0.652
0.133
0.010
0.116
0.540
0.090
0.049
A
A
A
A
A
3
3
4
4
2
0.199
0.199
0.266
0.266
0.133
0.215
0.215
0.287
0.287
0.143
0.096
0.093
0.112
0.068
0.108
0.170
0.169
0.222
0.207
0.128
A
SL
4
1
0.266
0.066
0.287
0.072
0.066
0.030
0.206
0.056
ESPECIE
Diplotropis martiusii Benth.
Dipteryx cordata (Ducke) Cowan
Dipteryx polyphylla Huber
Drypetes CL-2707
Duroia saccifera (Mart.ex Roem. &
Schult.) Hook.f. ex Schum.
Elaeoluma glabrescens (Mart. &
Eichler) Aubrv.
Endlicheria sericea (Nees) Mez
Erisma bicolor Ducke
Erisma laurifolium Spruce ex
Warm.
Erisma splendens Slafleu
Erythroxylon CL-1145
Eschweilera andina (Rusby)
Macbride
Eschweilera bracteosa (Poepp. &
Berg.) Miers
Eschweilera laevicarpa S. A. Mori
Eschweilera ovalifolia (DC.) Nied.
Eschweilera parvifolia Mart. ex
DC.
Eschweilera punctata Mori
Eschweilera tessmannii R. Knuth
Euphorbiaceae NA-1265
Fabaceae CL-1121
Fabaceae EA-789
Ferdinandusa chlorantha (Wedd.)
Standl.
Guarea trunciflora C. DC.
Guatteria foliosa Benth.
Guatteria punticulata R. E. Fries
Guatteriella tomentosa R. E. Fries
Heisteria duckei Sleumer
Helicostylis heterotricha Ducke
Helicostylis scabra (Macbr.) C. C.
Berg
Hevea benthamiana Muell. Arg.
Hevea guianensis Aublet
Hevea pauciflora (Spruce ex
Benth.) Muell. Arg.
HCR
A
A
A
A
NI
20
5
3
1
NI%
1.330
0.332
0.199
0.066
FR%
1.219
0.358
0.215
0.072
W%
0.953
0.163
0.426
0.009
IVI%
1.167
0.285
0.280
0.049
0.199
0.215
0.038
0.151
A
A
A
1
10
2
0.066
0.665
0.133
0.072
0.717
0.143
0.010
0.217
0.051
0.049
0.533
0.109
A
A
A
24
20
1
1.596
1.330
0.066
1.649
1.434
0.072
1.157
2.770
0.029
1.467
1.844
0.056
0.066
0.072
0.012
0.050
A
A
A
6
80
62
0.399
5.319
4.122
0.358
4.588
3.943
0.125
3.588
2.687
0.294
4.498
3.584
A
A
A
A
A
A
58
58
1
1
1
1
3.856
3.856
0.066
0.066
0.066
0.066
3.799
3.369
0.072
0.072
0.072
0.072
1.781
4.997
0.028
0.010
0.011
0.012
3.145
4.074
0.056
0.049
0.050
0.050
A
A
A
A
A
A
A
1
1
2
3
2
2
4
0.066
0.066
0.133
0.199
0.133
0.133
0.266
0.072
0.072
0.143
0.215
0.143
0.143
0.287
0.011
0.020
0.227
0.116
0.023
0.025
0.141
0.050
0.053
0.168
0.177
0.100
0.100
0.231
A
A
A
6
11
4
0.399
0.731
0.266
0.430
0.789
0.287
0.200
0.262
0.102
0.343
0.594
0.218
22
1.463
1.434
0.559
1.152
219
ESPECIE
Hirtella brachystachya Spruce &
Hook f.
Huberodendron swietenioides
(Gleason) Ducke
Indet NA-1392
Indet NA-953
Inga brachyrhachis Harms
Inga EA-194
Inga gracilior Sprague
Inga pezizifera Benth.
Iryanthera elliptica Ducke
Iryanthera juruensis Warb.
Iryanthera polyneura Ducke
Iryanthera tricornis Ducke
Iryanthera ulei Warb.
Jacaranda macrocarpa Bureau &
K.Schum. ex K.Schum.
Kotchubaea CL-966
Lacmellea arborescens (Muell.
Arg.) Markgr.
Lacmellea CL-1049
Lacmellea foxii (Stapf) Markgraf
Lacmellea ramosissima (Muell.
Arg.) Markgr.
Ladenbergia amazonensis Ducke
Lauraceae EA-851
Licania apetala (E. Meyer) Fritchs
Licania canescens Benoist
Licania CL-1149
Licania CL-1198
Licania EA-872
Licania heteromorpha Benth.
Licania hypoleuca Benth.
Licania kunthiana Hook. f.
Licania octandra (Hoffmanns. ex
Roem. & Schult.) Kuntze
Licania reticulata Prance
Licaria CL-1310
Licaria guianensis Aubl.
Lueheopsis shultesii Cuatr.
Machaerium CL-1234
220
HCR NI
NI%
FR%
W%
IVI%
0.199
0.215
0.052
0.155
A
A
A
A
A
A
A
A
A
A
A
A
1
1
1
1
2
1
5
5
1
1
13
20
0.066
0.066
0.066
0.066
0.133
0.066
0.332
0.332
0.066
0.066
0.864
1.330
0.072
0.072
0.072
0.072
0.143
0.072
0.358
0.358
0.072
0.072
0.932
1.362
0.056
0.013
0.426
0.026
0.045
0.027
0.101
0.190
0.009
0.014
0.421
0.286
0.065
0.050
0.188
0.055
0.107
0.055
0.264
0.294
0.049
0.051
0.739
0.993
A
A
9
6
0.598
0.399
0.645
0.358
0.091
0.138
0.445
0.298
A
A
A
2
7
3
0.133
0.465
0.199
0.143
0.502
0.215
0.038
0.091
0.040
0.105
0.353
0.152
A
A
A
A
A
A
A
A
A
A
A
1
1
3
12
1
4
9
1
5
1
1
0.066
0.066
0.199
0.798
0.066
0.266
0.598
0.066
0.332
0.066
0.066
0.072
0.072
0.215
0.860
0.072
0.287
0.645
0.072
0.358
0.072
0.072
0.012
0.196
0.221
0.322
0.012
0.269
0.753
0.010
0.245
0.042
0.009
0.050
0.112
0.212
0.660
0.050
0.274
0.665
0.049
0.312
0.060
0.049
A
A
A
A
A
A
1
1
2
2
1
1
0.066
0.066
0.133
0.133
0.066
0.066
0.072
0.072
0.143
0.143
0.072
0.072
0.010
0.041
0.091
0.074
0.022
0.012
0.050
0.060
0.122
0.117
0.053
0.050
ESPECIE
Macoubea guianensis Aubl.
Malouetia CL-1377
Matayba purgans Poepp. & Endlch.
Mezilaurus itauba (Meissn.) Taub.
& Mez
Miconia chrysophylla (Rich.) Urb.
Miconia CL-2476
Miconia CL-2752
Miconia phaeophylla Triana
Miconia poeppigii Triana
Miconia pubipetala Miq.
Micropholis guyanensis (A. DC.)
Pierre
Micropholis madeirensis (Baehni)
Aubr.
Micropholis melinoniana Pierre
Minquartia guianensis Aublet
Monopteryx uaucu Spruce ex
Benth.
Moronobea coccinea Aubl.
Mouriri nigra (DC.) Morley
Mouriri vernicosa Naudin
Naucleopsis caloneura (Hub.)
Ducke
Nectandra cuspidata Nees & Mart.
Nectandra purpurea (Ruiz & Pav.)
Mez
Neea CL-1196
Neea CL-1289
Neea CL-701
Neea divaricata Poepp. & Endl.
Neea macrophylla Poepp. ex Endl.
Ocotea aciphylla (Nees) Mez
Ocotea amazonica (Meissn.) Mez
Ocotea argyrophylla Ducke
Ocotea CL-1259
Ocotea CL-1356
Ocotea CL-1522
Ocotea CL-902
Ocotea cymbarum Kunth
Ocotea EA-193
HCR
A
A
A
NI
2
2
1
NI%
0.133
0.133
0.066
FR%
0.143
0.143
0.072
W%
0.021
0.065
0.090
IVI%
0.099
0.114
0.076
A
A
A
A
A
A
A
4
1
1
1
2
1
8
0.266
0.066
0.066
0.066
0.133
0.066
0.532
0.287
0.072
0.072
0.072
0.143
0.072
0.573
0.574
0.032
0.031
0.012
0.022
0.023
0.198
0.376
0.057
0.056
0.050
0.100
0.054
0.434
11
0.731
0.789
0.335
0.618
A
A
A
1
4
1
0.066
0.266
0.066
0.072
0.287
0.072
0.017
0.222
0.011
0.052
0.258
0.050
A
A
A
A
43
2
1
3
2.859
0.133
0.066
0.199
2.007
0.143
0.072
0.215
10.333
0.042
0.014
0.191
5.066
0.106
0.051
0.202
A
A
1
1
0.066
0.066
0.072
0.072
0.011
0.028
0.050
0.055
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
13
1
1
1
3
1
16
3
1
2
1
5
1
8
1
0.864
0.066
0.066
0.066
0.199
0.066
1.064
0.199
0.066
0.133
0.066
0.332
0.066
0.532
0.066
0.932
0.072
0.072
0.072
0.215
0.072
1.004
0.215
0.072
0.143
0.072
0.358
0.072
0.573
0.072
0.242
0.023
0.013
0.023
0.038
0.035
0.429
0.279
0.013
0.067
0.032
0.190
0.009
0.395
0.018
0.680
0.054
0.050
0.054
0.151
0.058
0.832
0.231
0.050
0.114
0.057
0.294
0.049
0.500
0.052
221
ESPECIE
Ocotea floribunda (Sw.) Mez
Ocotea myriantha (Meisn.) Mez.
Oenocarpus bacaba Mart.
Oenocarpus bataua Mart.
Osteophloeum platyspermum
(Spruce ex A.DC.) Warb.
Pagamea plicata Spruce ex Benth.
Parahancornia peruviana Monach.
Parkia panurensis Benth. &
H.C.Hopkins
Parkia reticulata Ducke
Picramnia sellowii Planch.
Pourouma CL-1455
Pouteria arcuata T. D. Penn.
Pouteria campanulata Baehni
Pouteria CL-1003
Pouteria CL-1017
Pouteria CL-1070
Pouteria CL-1155
Pouteria CL-742
Pouteria EA-1111
Pouteria EA-183
Pouteria EA-835
Pouteria EA-847
Pouteria EA-862
Pouteria glauca T.D.Penn.
Pouteria guianensis Aubl.
Pouteria maguirei (Aubrv.)
T.D.Penn.
Pouteria trilocularis Cronquist
Pouteria ucuqui Pires & Schultes
Pouteria vernicosa Pennington
Pouteria williamii (Aubrv. &
Pelligr.) Penn.
Pradosia cochlearia (Lecom.) Penn.
ssp. praealta (Ducke) Penn.
Protium altsonii Sandw.
Protium apiculatum Swart
Protium gallosum Daly
Protium hebetatum Daly
Protium paniculatum Engl.
222
HCR
A
A
PAM
PAM
NI
1
6
7
24
NI%
0.066
0.399
0.465
1.596
FR%
0.072
0.358
0.502
1.434
W%
0.009
0.540
0.061
0.271
IVI%
0.049
0.432
0.343
1.100
A
A
A
6
1
1
0.399
0.066
0.066
0.430
0.072
0.072
0.286
0.013
0.016
0.372
0.050
0.051
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
5
1
2
1
1
3
1
1
1
2
1
1
2
2
7
6
1
6
0.332
0.066
0.133
0.066
0.066
0.199
0.066
0.066
0.066
0.133
0.066
0.066
0.133
0.133
0.465
0.399
0.066
0.399
0.358
0.072
0.143
0.072
0.072
0.215
0.072
0.072
0.072
0.143
0.072
0.072
0.143
0.143
0.502
0.430
0.072
0.430
0.647
0.109
0.292
0.020
0.255
0.077
0.017
0.746
0.010
0.232
0.011
0.012
0.034
0.029
0.183
0.377
0.042
0.487
0.446
0.082
0.190
0.053
0.131
0.164
0.052
0.295
0.049
0.169
0.050
0.050
0.104
0.102
0.383
0.402
0.060
0.439
A
A
A
A
11
4
1
4
0.731
0.266
0.066
0.266
0.717
0.287
0.072
0.287
0.218
0.053
0.013
0.314
0.556
0.202
0.050
0.289
26
1.729
1.649
0.467
1.281
A
A
A
A
A
A
1
2
4
8
22
23
0.066
0.133
0.266
0.532
1.463
1.529
0.072
0.143
0.287
0.573
1.362
1.577
0.035
0.046
0.182
0.180
0.497
0.551
0.058
0.107
0.245
0.428
1.107
1.219
ESPECIE
Protium polybotryum (Turcz.)
Engl.
Protium rubrum Cuatrec.
Protium subserratum (Engl.) Engl.
Protium urophyllidium Daly
Pseudolmedia laevigata Trc.
Pseudolmedia laevis (R. & P.)
Macbr.
Pseudomonotes tropenbosii
Londoo et al.
Qualea EA-1172
Qualea paraensis Ducke
Remijia pedunculata (H.Karst.)
Flueck.
Rhodostemonodaphne CL-1300
Rhodostemonodaphne EA-198
Roucheria columbiana Hallier f.
Roucheria humiriifolia (Planch.)
Benth.
Saccoglottis CL-2454
Scleronema micranthum (Ducke)
Ducke
Scleronema praecox Ducke
Senefeldera karsteniana Pax. &
Hoffm.
Simaba guianensis Aubl.
Simarouba amara Aublet
Sloanea brevipes Benth.
Sloanea CL-1075
Sloanea CL-886
Sloanea NA-1256
Sloanea NA-1423
Sloanea NA-427
Sloanea NA-635
Sloanea obtusa (Splitg.) K. Schum.
Sloanea steyermarkii Earle Sm.
Sterigmapetalum colombianum
Monachino
Sterigmapetalum obovatum
Kuhlmann
Strychnos peckii B.L.Rob.
HCR NI
NI%
FR%
W%
IVI%
A
A
A
A
A
2
11
1
1
13
0.133
0.731
0.066
0.066
0.864
0.143
0.645
0.072
0.072
0.932
0.032
0.889
0.146
0.015
0.285
0.103
0.755
0.095
0.051
0.694
0.066
0.072
0.010
0.049
A
A
A
66
2
10
4.388
0.133
0.665
3.943
0.143
0.717
16.992
0.634
1.203
8.441
0.304
0.862
A
A
A
A
5
2
3
5
0.332
0.133
0.199
0.332
0.358
0.143
0.215
0.358
0.099
0.034
0.267
0.079
0.263
0.103
0.227
0.256
A
A
2
1
0.133
0.066
0.143
0.072
0.052
0.102
0.110
0.080
A
A
30
1
1.995
0.066
2.151
0.072
2.665
0.014
2.270
0.051
A
A
A
A
A
A
A
A
A
A
A
A
12
3
1
5
4
2
1
1
1
1
3
4
0.798
0.199
0.066
0.332
0.266
0.133
0.066
0.066
0.066
0.066
0.199
0.266
0.717
0.215
0.072
0.287
0.287
0.143
0.072
0.072
0.072
0.072
0.215
0.287
0.144
0.054
0.190
0.071
0.267
0.042
0.224
0.009
0.018
0.010
0.197
0.158
0.553
0.156
0.109
0.230
0.273
0.106
0.121
0.049
0.052
0.049
0.204
0.237
15
0.997
1.075
0.773
0.949
A
SL
16
1
1.064
0.066
1.075
0.072
0.766
0.029
0.968
0.056
223
ESPECIE
Swartzia brachyrachys Harms
Swartzia lamellata Ducke
Swartzia parvifolia Schery
Swartzia schomburgkii Benth.
Swartzia vaupesiana R. S. Cowan
Tachigali CL-1141
Tachigali CL-2734
Tachigali CL-918
Tachigali paniculata Aubl.
Talisia cerasina (Benth.) Radlk.
Terminalia brasiliensis Eichl.
Terminalia oblonga (R. & P.)
Steudel
Tovomita CL-829
Trattinnickia glaziovii Swart
Trichilia micrantha Bentham
Unonopsis CL-1399
Unonopsis guatterioides (A. DC.)
R. E. Fries
Vantanea peruviana J.F.Macbr.
Vantanea spichigeri Gentry
Virola calophylla (Spruce) Warb.
Virola elongata (Benth.) Warb.
Virola multinervia Ducke
Virola pavonis (A. DC.) A. C.
Smith
Vochysia punctata Spruce &
Warm.
Vochysia venulosa Warm.
Warszewiczia schwackei K.
Schum.
Xylopia calophylla R. E. Fries
Xylopia multiflora R. E. Fries
Xylopia parvifolia Diels
Zygia claviflora (Benth.) Barneby
& J.W.Grimes
Zygia racemosa (Ducke) Barneby
& J.W.Grimes
Total general
224
HCR
A
A
A
A
A
A
A
A
A
A
A
NI
2
2
1
10
1
11
2
6
6
2
1
NI%
0.133
0.133
0.066
0.665
0.066
0.731
0.133
0.399
0.399
0.133
0.066
FR%
0.143
0.143
0.072
0.717
0.072
0.645
0.072
0.430
0.430
0.143
0.072
W%
0.031
0.218
0.063
2.511
0.010
0.883
0.196
0.088
0.688
0.064
0.025
IVI%
0.103
0.165
0.067
1.297
0.049
0.753
0.134
0.306
0.506
0.114
0.054
A
A
A
A
A
1
4
1
1
1
0.066
0.266
0.066
0.066
0.066
0.072
0.287
0.072
0.072
0.072
0.009
0.170
0.040
0.020
0.017
0.049
0.241
0.059
0.053
0.052
A
A
A
A
A
A
1
13
5
3
17
1
0.066
0.864
0.332
0.199
1.130
0.066
0.072
0.789
0.358
0.215
1.219
0.072
0.041
0.890
0.298
0.032
0.258
0.134
0.060
0.848
0.330
0.149
0.869
0.091
0.465
0.430
0.110
0.335
A
A
2
22
0.133
1.463
0.143
1.505
0.028
1.779
0.102
1.582
A
A
A
A
1
1
2
1
0.066
0.066
0.133
0.066
0.072
0.072
0.143
0.072
0.012
0.041
0.189
0.038
0.050
0.060
0.155
0.059
0.399
0.358
0.073
0.277
5
0.332
1504 100
0.358
100
0.174
100
0.288
100
HCR
A
A
A
A
A
A
NI
2
6
2
17
3
1
NI%
0.166
0.497
0.166
1.408
0.249
0.083
FR%
0.095
0.571
0.190
1.429
0.286
0.095
W%
0.076
0.252
0.282
1.717
0.178
0.014
IVI%
0.112
0.440
0.213
1.518
0.237
0.064
A
A
A
A
A
A
A
3
1
11
1
1
1
7
0.249
0.083
0.911
0.083
0.083
0.083
0.580
0.286
0.095
0.952
0.095
0.095
0.095
0.476
0.857
0.015
2.413
0.075
0.050
0.027
0.672
0.464
0.064
1.426
0.084
0.076
0.068
0.576
A
A
A
PAM
PAM
SL
SL
A
A
A
3
1
1
5
75
1
4
2
6
1
0.249
0.083
0.083
0.414
6.214
0.083
0.331
0.166
0.497
0.083
0.286
0.095
0.095
0.476
5.238
0.095
0.381
0.190
0.571
0.095
0.154
0.141
0.108
0.330
1.081
0.168
0.243
0.084
0.398
0.046
0.230
0.106
0.095
0.407
4.178
0.115
0.318
0.147
0.489
0.075
A
A
A
A
SL
170
1
1
1
2
14.085
0.083
0.083
0.083
0.166
9.714
0.095
0.095
0.095
0.190
5.231
0.024
0.015
0.054
0.083
9.677
0.067
0.064
0.077
0.146
A
A
4
19
0.331
1.574
A
A
A
3
1
3
0.249
0.083
0.249
225
ESPECIE
Compsoneura sprucei (A.DC.) Warb.
Conceveiba guianensis Aubl.
Couepia EA-72
Coussapoa villosa Poepp. & Endl.
Coussarea hirticalyx Standl.
Coussarea macrophylla Muell. Arg.
Coussarea paniculata (Vahl) Standl.
Croton bilocularis J. Murillo
Croton cuneatus Klotzch.
Cynometra marginata Benth.
Dacryodes cuspidata (Cuatrec.) Daly
Dendropanax EA-657
Diospyros peruviana Hiern.
Dipteryx nudipes Tul
Doliocarpus dentatus (Aublet) Standley
Drypetes amazonica Ducke
Drypetes ampelocerifolia Grndez &
Vsquez
Duguetia odorata (Diels) Macbride
Duguetia spixiana Mart.
Endlicheria krukovii A. C. Smith
Endlicheria williamsii O. C. Schmidt
Eschweilera andina (Rusby) Macbride
Eschweilera bracteosa (Poepp. &
Berg.) Miers
Eschweilera itayensis R. Knuth
Eugenia EA-433
Euterpe precatoria Mart.
Fabaceae EA-625
Faramea multiflora A. Rich.
Faramea parvibractea Steyerm.
Ficus cf. maxima P. Miller
Ficus cf. yaponensis Dugand
Garcinia CL-915
Garcinia EA-688
Garcinia spruceana (Engler) Hammel
Guapira EA-456
Guarea grandiflora DC.
Guarea purusana C. DC.
Guatteria atra Sandw.
226
HCR
A
A
A
SZL
A
A
A
A
A
A
A
SZL
A
A
SL
A
A
NI
1
7
6
1
1
8
7
1
1
11
1
2
3
1
2
7
2
NI%
0.083
0.580
0.497
0.083
0.083
0.663
0.580
0.083
0.083
0.911
0.083
0.166
0.249
0.083
0.166
0.580
0.166
FR%
0.095
0.667
0.571
0.095
0.095
0.762
0.476
0.095
0.095
0.952
0.095
0.190
0.286
0.095
0.190
0.667
0.190
W%
0.014
0.417
0.865
0.019
0.024
0.154
0.192
0.139
0.154
1.127
0.038
0.267
0.095
0.104
0.190
0.180
0.040
IVI%
0.064
0.554
0.644
0.066
0.067
0.526
0.416
0.106
0.111
0.997
0.072
0.208
0.210
0.094
0.182
0.476
0.132
A
A
A
A
A
A
6
1
7
1
29
2
0.497
0.083
0.580
0.083
2.403
0.166
0.571
0.095
0.667
0.095
2.667
0.190
0.196
0.026
0.237
0.017
4.245
1.627
0.422
0.068
0.494
0.065
3.105
0.661
A
A
PAM
A
A
A
A
SZL
A
A
A
A
A
A
A
12
17
37
1
1
5
1
1
1
8
1
1
6
10
1
0.994
1.408
3.065
0.083
0.083
0.414
0.083
0.083
0.083
0.663
0.083
0.083
0.497
0.829
0.083
1.048
1.524
2.667
0.095
0.095
0.476
0.095
0.095
0.095
0.762
0.095
0.095
0.571
0.952
0.095
3.727
0.746
0.508
0.014
0.024
0.193
0.025
0.026
0.014
0.469
0.033
0.020
0.262
0.848
0.062
1.923
1.226
2.080
0.064
0.068
0.361
0.068
0.068
0.064
0.631
0.070
0.066
0.443
0.876
0.080
ESPECIE
Guatteria megalophylla Diels
Gustavia poeppigiana Berg
Heisteria cf. densifrons Engl.
Hirtella davisii Sandw.
Hirtella hispidula Miq.
Hirtella latifolia Prance
Hirtella pilosissima Mart. & Zucc.
Hyeronima alchorneoides Allemo
Hymenaea oblongifolia Huber
Inga EA-381
Inga EA-397
Inga EA-439
Inga EA-457
Inga EA-640
Inga fastuosa (Jacq.) Willd.
Iriartea deltoidea R. & P.
Iryanthera juruensis Warb.
Iryanthera ulei Warb.
Isertia rosea Spruce & K. Schum.
Klarobelia cauliflora Chatrou
Lacmellea gracilis (Muell. Arg.)
Markgraf
Lauraceae EA-402
Leonia crassa L. B. Sm. & A.
Fernndez
Licania arachnoidea Fanshaw &
Maguire
Licania EA-647
Licania EA-654
Licania oblongifolia Standl.
Licania reticulata Prance
Licania triandra Mart. & Hook. f.
Licania vaupesana Killip. & Cuatr.
Lonchocarpus EA-738
Lorostemon bombaciflorum Ducke
Mabea occidentalis Benth.
Machaerium arboreum Benth.
Machaerium EA-727
Machaerium inundatum (Mart.) Ducke
Machaerium paraense Ducke
Macoubea sprucei (Muell. Arg.)
HCR
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
PAM
A
A
A
A
A
NI
11
1
5
1
1
1
1
28
1
1
31
3
1
8
2
2
1
9
1
1
2
NI%
0.911
0.083
0.414
0.083
0.083
0.083
0.083
2.320
0.083
0.083
2.568
0.249
0.083
0.663
0.166
0.166
0.083
0.746
0.083
0.083
0.166
FR%
1.048
0.095
0.476
0.095
0.095
0.095
0.095
2.571
0.095
0.095
2.286
0.286
0.095
0.762
0.190
0.190
0.095
0.762
0.095
0.095
0.190
W%
1.742
0.018
0.210
0.048
0.025
0.018
0.025
6.431
0.031
0.055
2.599
0.058
0.024
0.905
0.047
0.138
0.073
0.177
0.016
0.033
0.050
A
A
1
3
0.083
0.249
10
0.829
A
A
A
A
A
A
A
A
A
A
SL
SL
SL
A
4
1
3
2
4
1
1
1
1
2
3
1
1
7
0.331
0.083
0.249
0.166
0.331
0.083
0.083
0.083
0.083
0.166
0.249
0.083
0.083
0.580
0.381
0.095
0.286
0.190
0.381
0.095
0.095
0.095
0.095
0.190
0.286
0.095
0.095
0.381
0.228
0.022
0.105
0.159
0.183
0.031
0.069
0.036
0.018
0.106
0.102
0.065
0.069
0.244
IVI%
1.234
0.065
0.367
0.075
0.068
0.065
0.068
3.774
0.070
0.078
2.484
0.197
0.067
0.776
0.134
0.165
0.084
0.562
0.065
0.070
0.136
0.314
0.067
0.213
0.172
0.299
0.070
0.082
0.071
0.065
0.154
0.212
0.081
0.082
0.402
227
ESPECIE
Markgr.
Malouetia tamaquarina (Aubl.) A.DC.
Manilkara bidentata (A. DC.) Chev.
Margaritaria nobilis L. f.
Matayba elegans Radlk.
Matisia cf. bracteolosa Ducke
Miconia eleagnoides Cogn.
Micropholis egensis (A. DC.) Pierre
Micropholis venulosa (Mart. & Eichl.)
Pierre
Minquartia guianensis Aublet
Mollia lepidota Spruce
Mouriri myrtifolia Spruce ex Triana
Myrcia cf. magna Legraud
Nectandra EA-452
Nectandra EA-718
Neea divaricata Poepp. & Endl.
Neea floribunda Poepp. & Endl.
Ocotea amazonica (Meissn.) Mez
Ocotea EA-461
Ocotea EA-481
Omphalea diandra L.
Ormosia EA-748
Ouratea kananariensis Sastre
Oxandra polyantha R. E. Fries
Oxandra xylopioides Diels
Pachira insignis (Sw.) Sw. ex Savigny
Parkia multijuga Benth.
Perebea cf. xanthochyma Karsten
Picramnia sellowii Planch.
Posoqueria EA-384
Pourouma cucura Standley & Cuatr.
Pouteria baehniana Monachino
Pouteria cladantha Sandw.
Pouteria EA-401
Pouteria EA-659
Pouteria EA-708
Pouteria laviegata (Mart.) Radlk.
Pouteria NA-80
Pouteria torta (Mart.) Radlk.
228
HCR NI
NI%
FR%
W%
IVI%
A
A
A
A
A
A
A
A
4
3
13
20
1
5
2
3
0.331
0.249
1.077
1.657
0.083
0.414
0.166
0.249
0.381
0.286
1.238
1.810
0.095
0.476
0.190
0.286
0.177
0.913
0.657
0.960
0.027
0.138
0.239
0.227
0.296
0.482
0.991
1.476
0.068
0.343
0.198
0.254
A
A
A
A
A
A
A
A
A
A
A
SL
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
1
1
1
8
2
1
13
2
5
6
2
1
1
2
1
9
23
9
2
5
13
2
25
3
4
1
1
1
1
29
0.083
0.083
0.083
0.663
0.166
0.083
1.077
0.166
0.414
0.497
0.166
0.083
0.083
0.166
0.083
0.746
1.906
0.746
0.166
0.414
1.077
0.166
2.071
0.249
0.331
0.083
0.083
0.083
0.083
2.403
0.095
0.095
0.095
0.762
0.190
0.095
1.238
0.190
0.381
0.571
0.190
0.095
0.095
0.190
0.095
0.857
2.095
0.857
0.190
0.381
1.238
0.190
2.095
0.286
0.381
0.095
0.095
0.095
0.095
2.571
0.049
0.199
0.034
0.162
0.243
0.019
2.563
0.052
0.123
0.142
0.055
0.065
0.019
0.030
0.027
0.923
2.619
2.646
0.061
0.132
0.286
0.073
0.771
0.133
0.184
0.068
0.206
0.055
0.034
1.199
0.076
0.126
0.071
0.529
0.200
0.066
1.626
0.136
0.306
0.404
0.137
0.081
0.066
0.129
0.069
0.842
2.207
1.416
0.139
0.309
0.867
0.143
1.646
0.222
0.299
0.082
0.128
0.078
0.071
2.058
ESPECIE
Pouteria ucuqui Pires & Schultes
Protium krukovii Swart
Protium nodulosum Swart
Pseudolmedia laevis (R. & P.) Macbr.
Pterocarpus amazonicus Huber
Pterocarpus draco L.
Pterocarpus EA-648
Qualea ingens Warm.
Richeria grandis Vahl
Rinorea lindeniana (Tulasne) O.
Kuntze
Rollinia edulis Tr. & Pl.
Salacia EA-752
Salacia impressifolia A. C. Smith
Sciadotecnia toxifera Kruk & Sm.
Simarouba amara Aublet
Sloanea parviflora Planch. & Benth.
Sorocea hirtella Mildbr.
Spondias EA-756
Sterculia apeibophylla Ducke
Sterculia EA-476
Sterigmapetalum colombianum
Monachino
Swartzia cardiosperma Spruce &
Benth.
Swartzia laevicarpa Amsh.
Swartzia racemosa Benth.
Symphonia globulifera L. f.
Tapirira retusa Ducke
Tapura amazonica Poepp.
Tapura capitulifera Spruce ex Baill.
Taralea opositifolia Aubl.
Terminalia EA-635
Theobroma obovatum Klotzsch &
Bernoulli
Theobroma subincanum Martius
Tovomita pyrifolia A. C. Smith
Trichilia EA-669
Triplaris americana L.
Vatairea guianensis Aubl.
Virola calophylla Warb.
HCR
A
A
A
A
A
A
A
A
A
A
NI
1
3
4
9
3
6
1
6
1
2
NI%
0.083
0.249
0.331
0.746
0.249
0.497
0.083
0.497
0.083
0.166
FR%
0.095
0.286
0.381
0.762
0.286
0.571
0.095
0.571
0.095
0.190
W%
0.077
0.155
0.111
2.645
1.279
0.198
0.067
0.733
0.064
0.091
IVI%
0.085
0.230
0.274
1.384
0.604
0.422
0.082
0.600
0.081
0.149
A
SL
A
SL
A
A
A
A
A
A
A
1
1
3
1
4
8
2
1
13
6
6
0.083
0.083
0.249
0.083
0.331
0.663
0.166
0.083
1.077
0.497
0.497
0.095
0.095
0.286
0.095
0.381
0.762
0.190
0.095
1.238
0.476
0.571
0.056
0.036
0.055
0.041
1.165
0.431
0.033
0.107
4.779
0.365
0.148
0.078
0.071
0.196
0.073
0.626
0.618
0.130
0.095
2.365
0.446
0.406
0.331
A
A
A
A
A
A
A
A
A
2
17
1
3
1
1
1
5
12
0.166
1.408
0.083
0.249
0.083
0.083
0.083
0.414
0.994
0.190
1.429
0.095
0.286
0.095
0.095
0.095
0.476
1.143
0.048
1.991
3.126
0.296
0.024
0.225
0.042
1.674
0.336
0.135
1.610
1.101
0.277
0.067
0.134
0.073
0.855
0.824
A
A
A
A
A
A
4
1
1
2
11
19
0.331
0.083
0.083
0.166
0.911
1.574
0.286
0.095
0.095
0.190
1.048
1.524
0.279
0.015
0.017
0.048
1.795
1.897
0.299
0.065
0.065
0.135
1.251
1.665
229
ESPECIE
Virola duckei A. C. Smith
Virola loretensis A. C. Smith
Virola surinamensis Warb.
Vismia macrophylla Kunth
Vitex klugii Moldenke
Warszewiczia coccinea (Vahl) Kl.
Xylopia excellens R. E. Fries
Zygia inequalis (Willd.) Pitt.
Zygia latifolia (L.) Fawc. & Rendle
Total general
230
HCR
A
A
A
A
A
A
A
A
A
NI
2
15
1
8
1
14
6
6
32
1207
NI%
0.166
1.243
0.083
0.663
0.083
1.160
0.497
0.497
2.651
100
FR%
0.190
1.333
0.095
0.667
0.095
1.238
0.571
0.571
2.762
100
W%
0.135
0.784
0.523
0.541
1.058
0.290
0.165
0.845
1.535
100
IVI%
0.164
1.120
0.234
0.623
0.412
0.896
0.411
0.638
2.316
100
Appendix 4
Indice de valor de importancia de familia (IVF%) (Mori et al. 1983), para
parcelas permanentes de 1.8 ha y vegetacin con DBH 10 cm, en
bosques de tierra firme y de vrzea en Pea Roja. NI: nmero de
individuos. W%: dominancia relativa medida en biomasa area seca. S:
nmero de especies. DI%: diversidad relativa (en %). Se muestra
secuencia alfabtica de familias (Mabberley 1990). Fabaceae sensu stricto,
Moraceae aparte de Cecropiaceae.
Tierra firme, inventario inicial (ao 1990).
FAMILIA
ANNONACEAE
APOCYNACEAE
ARALIACEAE
ARECACEAE
BIGNONIACEAE
BOMBACACEAE
BURSERACEAE
CAESALPINIACEAE
CARYOCARACEAE
CECROPIACEAE
CHRYSOBALANACEAE
CLUSIACEAE
COMBRETACEAE
CONVOLVULACEAE
DIPTEROCARPACEAE
ELAEOCARPACEAE
ERYTHROXYLACEAE
EUPHORBIACEAE
FABACEAE
FLACOURTIACEAE
HUGONIACEAE
HUMIRIACEAE
ICACINACEAE
INDET
LAURACEAE
LECYTHIDACEAE
LOGANIACEAE
MELASTOMATACEAE
NI
17
45
4
31
10
32
89
32
14
2
52
6
3
1
66
22
1
56
106
2
7
19
2
2
111
277
1
18
NI%
1.130
2.992
0.266
2.061
0.665
2.128
5.918
2.128
0.931
0.133
3.457
0.399
0.199
0.066
4.388
1.463
0.066
3.723
7.048
0.133
0.465
1.263
0.133
0.133
7.380
18.418
0.066
1.197
W%
0.767
6.023
0.066
0.332
0.120
2.735
2.947
2.147
1.622
0.031
2.754
0.212
0.074
0.030
16.992
0.997
0.029
1.480
15.203
0.052
0.131
1.291
0.108
0.439
4.779
14.968
0.029
0.522
S
11
13
1
2
2
3
14
4
3
2
17
2
3
1
1
9
1
9
14
2
2
3
1
2
29
8
1
8
DI%
4.264
5.039
0.388
0.775
0.775
1.163
5.426
1.550
1.163
0.775
6.589
0.775
1.163
0.388
0.388
3.488
0.388
3.488
5.426
0.775
0.775
1.163
0.388
0.775
11.240
3.101
0.388
3.101
IVF%
2.054
4.685
0.240
1.056
0.520
2.008
4.763
1.942
1.238
0.313
4.267
0.462
0.479
0.161
7.256
1.983
0.161
2.897
9.226
0.320
0.457
1.239
0.209
0.449
7.800
12.162
0.161
1.607
231
MELIACEAE
MENISPERMACEAE
MIMOSACEAE
MORACEAE
MYRISTICACEAE
NYCTAGINACEAE
OLACACEAE
RHIZOPHORACEAE
RUBIACEAE
SAPINDACEAE
SAPOTACEAE
SIMAROUBACEAE
TILIACEAE
VOCHYSIACEAE
Total general
2
1
38
71
91
7
3
31
19
3
121
6
1
82
1504
0.133
0.066
2.527
4.721
6.051
0.465
0.199
2.061
1.263
0.199
8.045
0.399
0.066
5.452
100
0.040
0.024
1.781
3.478
2.038
0.132
0.036
1.539
0.518
0.154
5.203
0.537
0.022
7.622
100
2
1
11
8
11
5
2
2
8
2
27
3
1
7
258
0.775
0.388
4.264
3.101
4.264
1.938
0.775
0.775
3.101
0.775
10.465
1.163
0.388
2.713
100
0.316
0.159
2.857
3.766
4.118
0.845
0.337
1.459
1.627
0.376
7.904
0.699
0.159
5.263
100
232
NI
4
39
25
2
119
2
24
9
213
6
35
21
6
2
2
3
8
70
48
4
7
25
44
NI%
0.331
3.231
2.071
0.166
9.859
0.166
1.988
0.746
17.647
0.497
2.900
1.740
0.497
0.166
0.166
0.249
0.663
5.800
3.977
0.331
0.580
2.071
3.645
W%
0.403
3.259
1.547
0.267
2.057
0.083
2.646
0.319
13.424
0.262
2.297
4.234
1.698
0.249
0.190
0.095
0.431
8.492
5.219
0.091
0.394
0.873
9.617
S
2
11
7
1
4
1
2
4
6
4
12
7
2
2
1
1
1
12
16
2
4
8
4
DI%
1.031
5.670
3.608
0.515
2.062
0.515
1.031
2.062
3.093
2.062
6.186
3.608
1.031
1.031
0.515
0.515
0.515
6.186
8.247
1.031
2.062
4.124
2.062
IVF%
0.588
4.053
2.409
0.316
4.659
0.255
1.888
1.042
11.388
0.940
3.794
3.194
1.075
0.482
0.290
0.286
0.536
6.826
5.815
0.484
1.012
2.356
5.108
MALPIGHIACEAE
MELASTOMATACEAE
MELIACEAE
MENISPERMACEAE
MIMOSACEAE
MORACEAE
MYRISTICACEAE
MYRTACEAE
NYCTAGINACEAE
OCHNACEAE
OLACACEAE
POLYGONACEAE
RHIZOPHORACEAE
RUBIACEAE
SAPINDACEAE
SAPOTACEAE
SIMAROUBACEAE
STERCULIACEAE
TILIACEAE
VERBENACEAE
VIOLACEAE
VOCHYSIACEAE
Total general
1
6
18
1
93
24
48
29
16
2
6
2
6
72
20
77
9
35
12
1
5
6
1207
0.083
0.497
1.491
0.083
7.705
1.988
3.977
2.403
1.326
0.166
0.497
0.166
0.497
5.965
1.657
6.379
0.746
2.900
0.994
0.083
0.414
0.497
100
0.054
0.172
1.474
0.041
8.713
3.317
3.603
0.987
2.635
0.030
0.258
0.048
0.148
3.358
0.960
4.325
1.297
5.759
2.612
1.058
0.272
0.733
100
1
2
4
1
9
8
7
3
3
1
2
1
1
11
1
13
2
4
2
1
2
1
194
0.515
1.031
2.062
0.515
4.639
4.124
3.608
1.546
1.546
0.515
1.031
0.515
0.515
5.670
0.515
6.701
1.031
2.062
1.031
0.515
1.031
0.515
100
0.217
0.567
1.676
0.213
7.019
3.143
3.729
1.645
1.836
0.237
0.595
0.243
0.387
4.998
1.044
5.802
1.024
3.574
1.546
0.552
0.572
0.582
100
233
234
Agradecimientos
237
238
Curriculum vitae
240
Publicaciones
Londoo, A.C., lvarez, E., Forero, E. & Morton, C. M. (1995) A new
genus and species of Dipterocarpaceae from the Neotropics. I.
Introduction, taxonomy, ecology and distribution. Brittonia, 47(3),
225-236.
lvarez, E. & Londoo, A. C. (1995) La etnobotnica cuantitativa: una
herramienta para la valoracin econmica de la biodiversidad (con
nfasis en la Amazonia). Crnica Forestal y del Medio Ambiente,
10, 163-191.
lvarez, E. & Londoo, A. C. (1996) Importancia ecolgica y
etnobotnica de las lianas en un bosque inundable de la Amazonia
colombiana. Cespedesia, 21(67), 373-390.
Londono, A.C. & lvarez, E. (1997) Composicin florstica de dos
bosques (tierra firme y vrzea) en la regin de Araracuara,
Amazonia colombiana. Caldasia, 19(3), 431-463.
Londoo, A.C. & Jimnez, E. (1999) Efecto del tiempo entre los censos
sobre la estimacin de las tasas anuales de mortalidad y de
reclutamiento de rboles (perodos de 1, 4 y 5 aos). Crnica
Forestal y del Medio Ambiente, 14, 41-57.
Jimnez, E., A.C. Londoo & H.F.M. Vester (2002) Descripcin de la
arquitectura de Iryanthera tricornis, Osteophloeum platyspermum
y Virola pavonis (Myristicaceae). Caldasia, 24(1), 65-94.
Vallejo, M. I., Londoo, A. C., Lpez, R., Galeano, G., lvarez, E. &
Devia, W. (2005) Establecimiento de parcelas permanentes en
bosques de Colombia. Mtodos para estudios ecolgicos a largo
plazo, 1. Instituto de Investigacin de Recursos Biolgicos
Alexander von Humboldt, Bogot. 309 pp.
Sin publicar
Alvarez, E., Cogollo, A. A., Melo, O., Rojas, E., Snchez, D., Velsquez,
O., Sarria, E., Jimnez, E., Benitez, D., Velsquez, C., Serna, M.,
Londoo, A. C., Stevenson, P., Galeano, G., Peuela, M. C.,
Garca, F., Ramos, Y., Palacios, J. & Patio, S. Red de parcelas
permanentes para el monitoreo de los bosques nativos de
colombia. Unpublished Paper.
241
242
243