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Tree changes in a mature rainforest with high diversity and endemism on the
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Article  in  Biodiversity and Conservation · August 2011

DOI: 10.1007/s10531-011-0067-3

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Biodivers Conserv (2011) 20:1921–1949
DOI 10.1007/s10531-011-0067-3

ORIGINAL PAPER

Tree changes in a mature rainforest with high diversity

and endemism on the Brazilian coast

Felipe Zamborlini Saiter • Frederico Augusto Guimarães Guilherme •

Luciana Dias Thomaz • Tânia Wendt

Received: 17 October 2010 / Accepted: 5 May 2011 / Published online: 18 May 2011
Ó Springer Science+Business Media B.V. 2011

Abstract The tree changes of 1.02 ha of montane forest at the Santa Lúcia Biological
Station, southeastern Brazil, were analyzed using two surveys separated by an interval of
11 years with the aim of confirming the patterns of stability of structure and diversity over
time. In the original survey all trees with diameter at breast height C6.4 cm were sampled.
In second survey (this study), dead trees, survivors and recruits in the same forest were
reported. The data suggest a dynamic balance of the forest structure because mortality
(-1.06% year-1 for number of trees and -0.85% year-1 for basal area) was very close to
recruitment (0.89% year-1) and ingrowth (1.05% year-1). The high diversity of the ori-
ginal survey (H0 [ 5.2) was maintained by the turnover species. The main tree populations
also showed stability of number of trees and basal area. This pattern was shared by most of
the 28 local endemic species, ensuring the maintenance of their populations in the plot.

Keywords Tree mortality  Tree recruitment  Turnover  Forest stability 

Atlantic rainforest

F. Z. Saiter (&)
Instituto Federal de Educação, Ciência e Tecnologia do Espı́rito Santo, Campus Santa Teresa,
Rodovia ES 080, km 21, Santa Teresa, ES 29660-000, Brazil
e-mail: fsaiter@ifes.edu.br

F. A. G. Guilherme
Departamento de Ciências Biológicas, Universidade Federal de Goiás,
Campus Jataı́, Rodovia BR 364, km 192, Jataı́, GO 75801-615, Brazil

L. D. Thomaz
Departamento de Ciências Biológicas, Universidade Federal do Espı́rito Santo,
Avenida Fernando Ferrari, 514, Vitória, ES 29075-900, Brazil

T. Wendt
Departamento de Botânica, IB, Universidade Federal do Rio de Janeiro,
Rio de Janeiro, RJ 21941-590, Brazil

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Introduction

Mature forests are ecosystems with a recognized ability to maintain both structure and
floristic diversity stable over time through the dynamic balance of mortality, recruitment and
growth of plants (Stephenson and van Mantgem 2005). This balance is based on the growth
cycle of the forests, which consists of three phases of development according to Whitmore
(1988): mature, gap and building. The mature phase has mature trees in various layers and a
closed canopy. Eventually canopy trees die or are damaged, knocking down the smaller trees
surrounding them and forming gaps. In a short time, these gaps are filled with herbs, climbers
and treelets which can arise from exposed roots and stumps or a bank of seeds and seedlings.
The building phase corresponds to the growth of plants until the formation of a new canopy
after many years, restoring the mature stage and all its layers (Whitmore 1988).
The turnover rate (the average of tree mortality and recruitment rate) is important data
that has been used with success to summarize the dynamism of forests. In the case of
tropical forests (admittedly more productive than temperate forests), the turnover rates are
higher because the dynamic balance is accelerated, making losses that are offset by gains
more quickly (Rees et al. 2001; Stephenson and van Mantgem 2005).
In the tropics, long-term research projects that monitor dynamic processes in permanent
sample plots have become a central theme in forest ecology in recent years because they
represent an appropriate strategy for gaining an understanding of how tropical forests maintain
their high diversity and how they are affected by global climate patterns (Picard et al. 2010).
Based on these projects, ecologists recognize that treefall gap dynamics is the key process
which maintains diversity in forests (Rees et al. 2001; Machado and Oliveira-Filho 2010) and
atmospheric changes, such as the increasing CO2 concentration and periods of drought and
intense heat, have made tropical forests more dynamic and changed their composition (Phillips
1996; Lewis et al. 2004a; Laurance et al. 2009; Lingenfelder and Newbery 2009).
The Atlantic rainforest is one of the Earth’s biologically richest and most endangered
terrestrial ecoregions (Mittermeier et al. 2004). In Brazil, the Atlantic rainforest has been
severely damaged during the last 50 years; the few remaining areas are covered by mature
forest and are mostly confined to protected areas (Ribeiro et al. 2009). Some of these areas
have already received initial phytosociological studies, but permanent sample plots and
monitoring of floristic composition and structure have only been maintained in a few. In
reflection of this, only the studies of Melo et al. (2000) and Rolim et al. (2005) have
reported on mature forests of the Brazilian coastline.
This study aims to analyze the changes of the community and the populations of trees
and tree palms in a mature forest on the Brazilian coast using two surveys separated by an
interval of 11 years to confirm the patterns of structure and diversity maintenance over
time, as observed in mature tropical forests. The factors that motivated the realization of
the present study were the lack of research on forest dynamics in the Atlantic rainforest, the
great ecological diversity found in the original survey (one of the highest diversity indexes
ever recorded in the world) and the description of many new and locally endemic species
from botanical materials collected in there since the installation of fotest permanent plots.

Methods

Study site

The Santa Lúcia Biological Station (SLBS) is a small protected area (467.89 ha)
located in Santa Teresa, the montane region of Espı́rito Santo state, southeastern Brazil

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Biodivers Conserv (2011) 20:1921–1949 1923

Fig. 1 Location of the Santa

Lúcia Biological Station in the
montane region of the Espı́rito
Santo state, southeastern Brazil

(19°570 1200 –19°590 1000 S; 40°310 1300 –40°320 3200 W, see Fig. 1). The SLBS is supported by
the Museu de Biologia Prof. Mello Leitão and the Universidade Federal do Rio de Janeiro,
but it is not a legally recognized conservation unit. Nevertheless, the SLBS plays an
important role in the fragmented landscape of the region, contributing to the connection (at
least functional) between small surrounding forest fragments (maximum of 100 ha) and the
Augusto Ruschi Biological Reserve (3573 ha), the main protected area in the region.
Recently, important taxonomic and ecology research was undertaken in the SLBS which
identified new plant species of the Atlantic rainforest. Besides this, several research projects
have recorded high levels of local biodiversity in this region (Mendes and Padovan 2000).
The climate is Cfa, according to the Köppen classification, with rainy summers and dry
winters. The average annual temperature at the site is 20°C and the average annual rainfall
(influenced by orographic rainfall) is 1868 mm. November is the wettest month and June is
the only month of the year with precipitation below 60 mm (Mendes and Padovan 2000).
The relief is very undulating forming a valley with several streams that contribute to the
river Timbuı́. There are extensive rocky outcrops at altitudes from 600 to 900 masl.
Shallow dystrophic soil with a high acidity, high levels of exchangeable Al and a low base
saturation dominates the region (Thomaz and Monteiro 1997). According to the classifi-
cation of the Instituto Brasileiro de Geografia e Estatı́stica (1992), the predominant veg-
etation is montane rainforest (500–1500 m in altitude).

Study design and forest surveys

The whole sample stand comprises 1.02 ha and was installed in 1992–1993 during the
initial survey of trees and tree palms with a diameter at breast height (dbh) C6.4 cm. The
stand is located on the northeast side of a slope to the right of the bank of the river Timbuı́,
at 650–850 m high, and is subdivided into three transects of 0.34 ha (see Fig. 1) composed
of 34 contiguous permanent plots 100 m2 (10 9 10 m). Transect 1 is at 650–660 m,
transect 2 is at 675–700 m, and transect 3 is at 820–850 m high (Thomaz and Monteiro
1997). In transect 1 the sequence of quadrats was interrupted due to the dense cluster of

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Merostachys sp. new, a woody bamboo species. Because of this, 10 plots were installed
before this cluster and 24 plots were installed after. The stand is not disturbed by flooding
and there are no reports of human disturbance.
In the second survey (this study), carried out between 2003 and 2004, the dead trees,
survivors and recruits were reported. Recruits were considered as trees which had reached
the minimum dbh after 11 years. Moreover, great effort was expended in the last 5 years to
revise and update the floristic list published by Thomaz and Monteiro (1997), which was
composed of 443 species but more than a half of them (54%) were without a complete
identification due to incipient knowledge of the regional flora at that time. For this, tax-
onomists and the herbarium collections of the Museu de Biologia Mello Leitão Herbarium
(MBML), the Universidade Federal do Espı́rito Santo Herbarium (VIES), the Jardim
Botânico do Rio de Janeiro Herbarium (RB), the Universidade Federal do Rio de Janeiro
Herbarium (RFA), the Missouri Botanical Garden Herbarium (MO) and the New York
Botanical Garden Herbarium (NY) were consulted.
The result of this review was the recognition of 385 species, with only 20% without
complete identification yet (see Table 6 in Appendix). It is noteworthy that 28 of these
species were described as new to science using botanical materials collected in the SLBS
and in its surroundings, and their geographical distributions are only assigned to Santa
Teresa and some adjacent municipalities (totaling only about 3000 km2).
Although the stand design is not ideal, in the present study the three transects were
combined into a single plot. This was enabled by the small distance between transects, the
similarity of soil samples (for details see Thomaz and Monteiro 1997), the small difference
between the richness of the transects in the original survey (208, 203 and 211 species in
transects 1, 2 and 3, respectively), and the high similarity index between the transects
(Sørensen index C0.5, F. Z. Saiter, non-published data).

Data analysis

The Shannon diversity index (H0 ) and the evenness index (J) were calculated for the
original and second surveys according to Magurran (1988). The diversity t test was applied
using the free software PAST (developed by Øyvind Hammer of the University of Oslo and
collaborators; Hammer et al. 2001) to identify possible differences of H0 between surveys
(P B 0.05).
In regard to parameters of forest structure (number of trees and basal area), possible
differences between the two surveys was tested using the Wilcoxon’s signed-rank test
(two-tailed; P B 0.05) according to Zar (1999). This test was applied using the free
software BioEstat 5.0 (developed by Manuel Ayres of the Universidade Federal do Pará
and collaborators; Ayres et al. 2007).
Assuming that changes of communities and tree populations over time are a constant
proportion of their initial conditions, we calculated average annual rates of change (r)
using the exponential equation described by Korning and Balslev (1994):

r ¼ ðCt = C0 Þ1=t 1
For the analysis of the community, r means the average annual rate of death (when r \ 0)
or recruitment/ingrowth (where r [ 0) and C0 is the number of trees or the basal area in the
original survey; Ct is the number or basal area of trees surviving in the second survey
(Ct = C0 - dead trees), in the case of death/loss, and the initial number of trees added by the
number of recruits (Ct = C0 ? recruited trees), in the case of recruitment or the basal area
increasing from initial recruitment and growth of survivors (Ct = C0 ? recruits ? growth

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Biodivers Conserv (2011) 20:1921–1949 1925

of survivors trees) in the case of ingrowth/gain. For change (r) of each population, t is the
period of time in years between the two surveys, C0 is the size of the population in the original
survey and Ct is the size of the population after t years.
The average rate of annual turnover (rate at which trees that die are replaced by recruits)
was calculated from the average of the rates of mortality and recruitment. Turnover was
applied to both number of trees and basal area (as in the method of Oliveira-Filho et al. 1997).
Furthermore, based on the rates of mortality, recruitment and ingrowth, values in years of
half-life (t‘) and doubling time (t2) were obtained, which are, respectively: the time in which
the community will reduce by half (keeping up the rates of mortality), and the time that the
community will double (keeping up the rates of recruitment and ingrowth). According to
Korning and Balslev (1994) the logarithmic expressions for these parameters are:
t1=2 ¼ ln ð0:5Þ=ln ð1 þ rÞ and t2 ¼ ln ð2Þ=ln ð1 þ rÞ
The turnover time and stability time in years were obtained from the mean of t‘ and t2,
and the numerical difference between them, respectively. The turnover time is defined as
the time in years required for the total renovation of the cover in a sample of forest. The
turnover of species and families were presented by the number of species and families
recruited and absent in the second survey.
After the calculations for populations, species were sorted by importance value (IV) in
the second survey, which was obtained from the sum of relative density, relative domi-
nance and relative frequency of each species (Mueller-Dombois and Ellenberg 1974).
The following intervals were used for the analysis of community changes by dbh
classes: 6.4–10, 10–20, 20–40, 40–80 and [80 cm. The classes had increasing intervals in
order to compensate the small number of trees which had dbh[30 cm. For each dbh class,
the number of trees subjected to the following events was obtained: death, ingrowth (given
by the entry of trees into a class by recruitment and growth) and outgrowth (given by the
output of trees of a class by growth). The variations in the rates of mortality, outgrowth and
ingrowth among the dbh classes were tested using the Kolmogorov–Smirnov test (one
sample; P B 0.05) according to Zar (1999).
For analysis of changes of populations the annual rates of change (r), which were
obtained as described above, were classified in thirteen classes (100%; [12 to 24%; [6 to
12%;[3 to 6%;[1.5 to 3%;[0 to 1.5%; 0%;\0 to -1.5%;\-1.5 to -3%;\-3 to -6%;
\-6 to -12%;\-12 to -24%; -100%). The classes ‘100%’ and ‘-100%’ correspond to
recruits and absent species in the second survey, respectively. Spearman’s rank correlation
test (rs; P B 0.05) was used for to identify correlation between annual rates of change and
number of trees recorded for the species which showed changes after 11 years.
The Kolmogorov–Smirnov and the Spearman’s rank correlation tests were carried out
by the software BioEstat 5.0 (Ayres et al. 2007).

Results

Community

Table 1 summarizes the community changes between 1992–1993 and 2003–2004. The
number of trees in 2003–2004 is close to the number of trees in 1992–1993. The difference
in number of trees between the two surveys was small and non-significant (T = 1794;
P = 0.39) because mortality was just a bit higher than recruitment, with 242 dead trees and

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1926 Biodivers Conserv (2011) 20:1921–1949

Table 1 Summary of the community changes in 1.02 ha of montane rainforest at the Santa Lúcia Bio-
logical Station, Espı́rito Santo, southeastern Brazil. Data were obtained from forest surveys carried out in
1992–1993 (Thomaz and Monteiro 1997) and 2003–2004 (this study)
1992–1993 2003–2004 Loss/Dead Gain/ingrowth Survivor

Species 385 384 15 14 370

Families 64 63 2 1 62
Trees 2190 2173 242 225 1948
Basal area (m2) 47.94 49.47 4.28 0.96 48.51
H0 5.27 5.22
J 0.89 0.88

Table 2 Parameters of community changes obtained from forest surveys carried out in 1992–1993
(Thomaz and Monteiro 1997) and 2003–2004 (this study) in 1.02 ha of montane rainforest at the Santa
Lúcia Biological Station, Espı́rito Santo, southeastern Brazil
Number of trees Basal area
-1
Mortality rate (% year ) -1.06 -0.85
Recruitment rate (% year-1) 0.89 1.05
Turnover rate (% year-1) 0.98 0.95
Half life (year) 65.11 81.44
Double time (year) 77.96 66.61
Turnover time (year) 71.54 74.02
Stability time (year) 12.85 14.83

225 recruits accounted after 11 years. These numbers resulted in average annual rates of
mortality and recruitment being very close and an average annual turnover rate of
0.98% year-1, as shown in Table 2. The basal area increased by 1.53 m2 (0.20% year-1),
but the difference between the two surveys was non-significant (T = 2179; P = 0.14). In
total values, the contribution of the basal area increment by recruits (0.96 m2) was about
five times lower than the growth of survivors (4.85 m2), which would be expected since the
recruitment group was composed of trees with a smaller dbh and the surviving group had
the largest number of trees. The community also showed a very close turnover time and
stability time for both periods for the number of trees and the basal area.
The tree richness changed little after 11 years. Of the 385 species in 1992–1993 (number
obtained after reviewing the list of 443 species originally recorded by Thomaz and Monteiro
1997, see Table 6 in Appendix), 15 had disappeared in 2003–2004. However, 14 new species
were recorded as recruits in 2003–2004 (Table 1). The absent species and the recruited
species each had a low abundance (never more than four trees) and are listed in Table 3. For
the families Solanaceae and Winteraceae, which were represented by only one species each
(Solanum sooretamum and Drimys brasiliensis, respectively) in 1992–1993, were absent in
2003–2004, while the family Linaceae was recorded for the first time with the occurrence of
Roucheria cf. dryadica. Melastomataceae was the family with the largest number of
recruited species while Myrtaceae had the largest number of absent species.
The high ecological diversity of the original survey (H0 [ 5.2) was maintained
(t = 1.189; P = 0.23; non-significant), confirming that the SLBS forest has one of the
highest diversity indices recorded in the world. The evenness index, greater than 0.8 in

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Biodivers Conserv (2011) 20:1921–1949 1927

Table 3 Absent and recruited tree species in 1.02 ha of montane rainforest at the Santa Lúcia Biological
Station, Espı́rito Santo, southeastern Brazil, 11 years after the forest survey of Thomaz and Monteiro (1997)

Families Species absent 2003–2004 ND

Annonaceae Rollinia laurifolia 1

Aquifoliaceae Ilex amara 1
Urticaceae Pourouma bicolor 1
Celastraceae Salacia nemorosa 1
Erythroxylaceae Erythroxylum cuspidifolium 1
Euphorbiaceae Croton floribundus 1
Lauraceae Licaria guianensis 2
Melastomataceae Miconia doriana 1
Myrtaceae Eugenia neoglomerata 1
Myrtaceae Myrcia pubipetala 2
Myrtaceae Myrcia racemosa 1
Myrtaceae Siphoneugena dussii 2
Salicaceae Banara serrata 4
Solanaceae Solanum sooretamum 1
Winteraceae Drimys brasiliensis 1

Families Species recruited 2003–2004 NR

Annonaceae Anaxagorea dolichocarpa 1

Annonaceae Annona cacans 1
Lauraceae Aniba firmula 2
Lauraceae Ocotea indecora 2
Melastomataceae Miconia longicuspis 1
Melastomataceae Miconia sellowiana 1
Melastomataceae Miconia sp. new 2
Melastomataceae Miconia tristis 2
Myrtaceae Eugenia sp5 new 1
Myrtaceae Eugenia melanogyna 1
Linaceae Roucheria cf. dryadica 1
Proteaceae Panopsis sp. 1
Sapindaceae Talisia sp. 1
Vochysiaceae Qualea gestasiana 2

Survey held in 2003–2004 (present study)

ND number of dead species, NR number recruited

both surveys, showed a good uniformity of individual distribution in relation to species.

Indeed, in both surveys just over 70% of species were represented by up to five trees.

Diameter classes

The changes of the community by dbh class are shown in Table 4. The recruits were
almost entirely in the 6.4–10 cm class, with a few occurring in the 10–20 cm class (but
these were no larger than 13.7 cm). The 20–40 cm class accumulated the greatest basal
area, but the largest increase of basal area was in the 40–80 cm class (about 1 m2).

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1928 Biodivers Conserv (2011) 20:1921–1949

Table 4 Changes of tree community by diameter class (dbh) in 1.02 ha of montane rainforest at the Santa
Lúcia Biological Station, Espı́rito Santo, southeastern Brazil, surveyed in 1992–1993 (Thomaz and Monteiro
1997) and 2003–2004 (this study)
Dbh classes (cm)

6.4–10 10–20 20–40 40–80 [80 Totals KS

Trees 1992–1993 925 880 348 36 1 2190

2003–2004 900 881 349 42 1 2173
Change -25 1 1 6 0 -17
Deads NT 109 94 38 1 0 242
% year-1 -1.13 -1.02 -1.05 -0.26 0.00 -1.06 0.33 ns
Outgrowth NT 136 52 10 1 0 199
% year-1 -1.44 -0.55 -0.26 -0.26 0.00 -0.86 0.27 ns
Recruits ? Ingrowth NT 216 9 0 0 0 225
4 138 49 8 0 199
% year-1 1.96 1.41 1.20 1.84 0.00 1.62 0.26 ns
BA (m2) 1992–1993 4.68 13.73 20.14 7.96 1.43 47.94
2003–2004 4.55 13.81 20.69 8.99 1.44 49.47

NT number of trees, BA basal area, KS Kolmogorov–Smirnov test (one sample), ns non-significant at

P B 0.05

Fig. 2 Distribution of the number of species per classes of annual rates of change, in number of trees (NT),
after 11 years in 1.02 ha of montane rainforest at the Santa Lúcia Biological Station, Espı́rito Santo,
southeastern Brazil. The classes ‘100%’ and ‘-100%’ correspond to recruits and absent species in the
second survey, respectively

Mortality did not occur in the [80 cm class since this class was represented by only one
live tree of Caryocar edule in both surveys. The variations in the rates of mortality,
outgrowth and ingrowth among classes were not significant and, in general, the decline in
the number of trees in a class was accompanied by a decrease in basal area.

Populations

In general, the size of the populations changed little because 56% of species maintained the
same number of trees and 26% showed annual rates B3% after 11 years (see Fig. 2).
Within the group of 169 species which changed in the number of trees, the annual rates of
change were negatively correlated to the number of individuals (rs = -0.885;

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Biodivers Conserv (2011) 20:1921–1949 1929

Fig. 3 Distribution of the number of species per classes of annual rates of change, in basal area (BA), after
11 years in 1.02 ha of montane rainforest at the Santa Lúcia Biological Station, Espı́rito Santo, southeastern
Brazil. The classes ‘100%’ and ‘-100%’ correspond to recruits and absent species in the second survey,
respectively

P \ 0.0001). With respect to the basal area, the annual rates of change were small and
positive (between 0 and 3%) in most species (see Fig. 3).
The population changes of the 100 species with the highest importance values (IV) are
presented in Table 5. These species were responsible for 66% of the trees and about 72.5%
of the total basal area in both surveys.
The palm tree Euterpe edulis was the most abundant species in the community and
showed a remarkable imbalance in favor of recruitment, which resulted in an increase of 34
individuals in its population. Ocotea aciphylla, Eriotheca macrophylla, Unonopsis sancta-
teresae, Ecclinusa ramiflora and Mezilaurus sp. (new species) were others abundant
species of the community. But they did not show major changes in their populations,
following the pattern of most species (see Table 5).
Unonopsis sancta-teresae and Mezilaurus sp. (new species) are distinguished as local
endemic species. Twenty-six other local endemic species were recorded in plot, but the
size of their populations did not differ much in original and second surveys (see Fig. 4).
Endemic species with special situations were Salacia nemorosa, which was absent in
second survey due to the death of a single tree, and Miconia sp. (new species) and Eugenia
sp. 5 (new species) which were recruited.
Ocotea cryptocarpa, Ocotea odorifera and Maytenus obtusifolia presented negative
growth of basal area, because there was decay of some stems of trees with multiple stem.
Ocotea aciphylla presented the largest growth of survivors and had a positive change of
basal area, even with the decrease in the number of trees. Several others populations also
showed positive changes of basal area and negative changes in number of trees. Caryocar
edule although represented by only two trees had the largest total basal area (1.762 m2).
Vochysia rectiflora showed the greatest loss in basal area (0.349 m2), a fact attributed to
the death of only two trees of great size.

Discussion

Community

The data suggest stability of the forest in SLBS after a period of 11 years as a dynamic
balance was found between the losses and gains of trees and basal areas which corresponds

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 Table 5 Population changes of the 100 species of highest importance value after the second survey in 1.02 ha of montane rainforest at the Santa Lúcia Biological Station,
1930

Espı́rito Santo, southeastern Brazil

Species IV Number of trees Change Basal area Change

123
(% year-1) (% year-1)
1992–1993 D R 2003–2004 1992–1993 Loss Gain 2003–2004

Euterpe edulis 16.62 168 18 52 202 1.69 1.262 0.147 0.333 1.448 1.26
Ocotea aciphylla 8.78 85 13 5 77 -0.89 1.079 0.065 0.225 1.240 1.27
Eriotheca macrophylla 7.57 53 8 5 50 -0.53 1.524 0.171 0.154 1.507 -0.10
Ecclinusa ramiflora 5.20 41 2 2 41 0.00 0.753 0.027 0.086 0.813 0.70
Unonopsis sancta-teresae 4.55 45 4 3 44 -0.20 0.373 0.041 0.037 0.369 -0.10
Mezilaurus sp. new 4.15 41 2 1 40 -0.22 0.588 0.034 0.012 0.565 -0.36
Coussapoa microcarpa 4.02 21 4 2 19 -0.91 0.930 0.048 0.181 1.063 1.22
Vantanea obovata 4.01 19 1 0 18 -0.49 1.148 0.017 0.058 1.189 0.32
Caryocar edule 3.76 2 0 0 2 0.00 1.743 0.000 0.019 1.762 0.10
Guapira opposita 3.48 26 4 6 28 0.68 0.500 0.121 0.126 0.505 0.09
Virola gardneri 3.40 13 0 2 15 1.31 0.874 0.000 0.081 0.955 0.81
Guapira obtusata 3.30 26 5 5 26 0.00 0.424 0.048 0.085 0.461 0.76
Beilschmiedia taubertiana 3.08 17 0 1 18 0.52 0.634 0.000 0.041 0.676 0.58
Inga capitata 2.99 22 0 3 25 1.17 0.251 0.000 0.056 0.306 1.84
Micropholis venulosa 2.91 15 0 0 15 0.00 0.609 0.000 0.104 0.712 1.44
Pseudoxandra spiritus-sancti 2.90 14 2 1 13 -0.67 0.749 0.012 0.070 0.807 0.68
Guapira venosa 2.83 20 3 6 23 1.28 0.315 0.057 0.068 0.326 0.32
Ocotea sp3 2.54 17 0 0 17 0.00 0.431 0.000 0.078 0.509 1.52
Ormosia cf. friburgensis 2.43 16 3 1 14 -1.21 0.556 0.037 0.036 0.555 -0.02
Ocotea cryptocarpa 2.43 16 2 1 15 -0.58 0.593 0.149 0.004 0.448 -2.51
Sloanea guianensis 2.43 17 1 0 16 -0.55 0.479 0.015 0.040 0.505 0.49
Ocotea divaricata 2.42 12 1 0 11 -0.79 0.618 0.003 0.030 0.645 0.39
Siparuna glossostyla 2.37 17 1 5 21 1.94 0.202 0.019 0.071 0.254 2.10
Biodivers Conserv (2011) 20:1921–1949
 Table 5 continued

Species IV Number of trees Change Basal area Change

(% year-1) (% year-1)
1992–1993 D R 2003–2004 1992–1993 Loss Gain 2003–2004

Myrcia laurifolia 2.17 19 1 1 19 0.00 0.191 0.007 0.017 0.201 0.45

Hirtella hebeclada 2.17 14 0 1 15 0.63 0.373 0.000 0.056 0.429 1.27
Trichilia lepidota 2.16 12 2 1 11 -0.79 0.532 0.018 0.028 0.542 0.18
Micropholis guyanensis 2.14 12 0 0 12 0.00 0.465 0.000 0.075 0.539 1.37
Psychotria sessilis 2.11 13 0 2 15 1.31 0.278 0.000 0.096 0.374 2.74
Pouteria sp2 2.10 16 3 0 13 -1.87 0.506 0.098 0.007 0.415 -1.79
Ocotea catharinensis 2.03 18 3 1 16 -1.07 0.330 0.072 0.079 0.338 0.20
Biodivers Conserv (2011) 20:1921–1949

Licania micrantha 1.98 14 2 0 12 -1.39 0.332 0.019 0.065 0.378 1.19

Pouteria aff. reticulata 1.97 16 1 0 15 -0.58 0.241 0.005 0.038 0.274 1.17
Maytenus cestrifolia 1.96 15 3 7 19 2.17 0.070 0.016 0.036 0.094 2.75
Diplotropis incexis 1.95 15 1 0 14 -0.63 0.347 0.084 0.023 0.286 -1.74
Eugenia aff. pruniformis 1.91 9 0 0 9 0.00 0.466 0.000 0.026 0.492 0.50
Swartzia apetala 1.89 14 1 0 13 -0.67 0.281 0.019 0.018 0.280 -0.04
Roupala consimilis 1.82 17 3 0 14 -1.75 0.207 0.023 0.041 0.225 0.78
Myrsine umbellata 1.80 11 2 1 10 -0.86 0.490 0.130 0.026 0.386 -2.13
Trichilia silvatica 1.79 13 0 1 14 0.68 0.193 0.000 0.015 0.208 0.68
Eugenia acutata 1.71 14 1 0 13 -0.67 0.239 0.024 0.031 0.245 0.25
Manilkara cf. longifolia 1.69 9 0 0 9 0.00 0.340 0.000 0.043 0.383 1.08
Geonoma schottiana 1.68 16 3 5 18 1.08 0.055 0.010 0.017 0.063 1.23
Trichilia sp. new 1.66 13 3 0 10 -2.36 0.454 0.106 0.025 0.373 -1.77
Vochysia santaluciae 1.64 13 3 2 12 -0.73 0.248 0.045 0.084 0.291 1.44
Ocotea corymbosa 1.64 11 0 0 11 0.00 0.254 0.000 0.032 0.285 1.08
Lacistema robustum 1.60 12 0 4 16 2.65 0.135 0.000 0.044 0.180 2.61
Amaioua intermedia 1.57 12 2 1 11 -0.79 0.194 0.008 0.035 0.221 1.18
1931

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 Table 5 continued
1932

Species IV Number of trees Change Basal area Change

(% year-1) (% year-1)

123
1992–1993 D R 2003–2004 1992–1993 Loss Gain 2003–2004

Attalea burretiana 1.52 6 0 0 6 0.00 0.443 0.000 0.008 0.451 0.16

Pouteria sp1 1.51 15 0 0 15 0.00 0.097 0.000 0.007 0.105 0.67
Mouriri cf. regeliana 1.50 5 0 1 6 1.67 0.446 0.000 0.022 0.468 0.44
Pouteria bangii 1.47 6 0 2 8 2.65 0.268 0.000 0.055 0.323 1.71
Tovomita leucantha 1.45 12 2 2 12 0.00 0.188 0.071 0.023 0.141 -2.63
Coussapoa pachyphylla 1.38 13 4 1 10 -2.36 0.394 0.208 0.049 0.236 -4.56
Sorocea guilleminiana 1.37 10 0 0 10 0.00 0.176 0.000 0.027 0.203 1.32
Byrsonima variabilis 1.36 11 1 1 11 0.00 0.166 0.013 0.023 0.176 0.53
Tetrastylidium grandifolium 1.33 10 0 0 10 0.00 0.179 0.000 0.032 0.211 1.52
Elvasia capixaba 1.31 13 3 0 10 -2.36 0.249 0.093 0.018 0.174 -3.23
Parinari obtusifolium 1.30 4 0 0 4 0.00 0.422 0.000 0.023 0.444 0.48
Marlierea obscura 1.29 4 0 0 4 0.00 0.393 0.000 0.070 0.463 1.51
Beilschmiedia sp. 1.28 9 0 0 9 0.00 0.167 0.000 0.014 0.181 0.76
Sparattanthelium botocudorum 1.23 3 0 0 3 0.00 0.421 0.000 0.038 0.459 0.80
Schefflera calva 1.21 4 0 0 4 0.00 0.355 0.000 0.042 0.396 1.02
Maytenus obtusifolia 1.21 12 2 0 10 -1.64 0.190 0.044 0.000 0.149 -2.20
Ocotea odorifera 1.19 10 2 0 8 -2.01 0.230 0.044 0.000 0.185 -1.92
Hirtella angustifolia 1.18 10 2 0 8 -2.01 0.192 0.019 0.034 0.207 0.68
Hymenaea aurea 1.14 8 0 0 8 0.00 0.199 0.000 0.016 0.214 0.69
Myrcia plusiantha 1.12 10 2 0 8 -2.01 0.187 0.019 0.013 0.181 -0.28
Chrysophyllum splendens 1.12 11 1 0 10 -0.86 0.134 0.007 0.008 0.136 0.10
Citronella paniculata 1.11 6 1 0 5 -1.64 0.299 0.003 0.004 0.299 0.02
Micropholis sp. 1.07 5 1 0 4 -2.01 0.322 0.000 0.000 0.328 0.18
Barnebya dispar 1.06 9 1 0 8 -1.07 0.167 0.056 0.013 0.124 -2.66
Biodivers Conserv (2011) 20:1921–1949
 Table 5 continued

Species IV Number of trees Change Basal area Change

(% year-1) (% year-1)
1992–1993 D R 2003–2004 1992–1993 Loss Gain 2003–2004

Casearia commersoniana 1.06 7 0 1 8 1.22 0.109 0.000 0.011 0.120 0.90

Eugenia copacabanensis 1.04 4 0 0 4 0.00 0.293 0.000 0.021 0.314 0.64
Pourouma guianensis 1.02 7 1 0 6 -1.39 0.151 0.005 0.056 0.204 2.74
Pouteria gardneri 1.01 7 0 0 7 0.00 0.115 0.000 0.033 0.148 2.31
Myrcia splendens 1.01 8 1 1 8 0.00 0.115 0.007 0.017 0.125 0.78
Eugenia xiriricana 1.00 8 1 1 8 0.00 0.117 0.014 0.015 0.118 0.06
Licania parvifolia 0.98 7 1 0 6 -1.39 0.177 0.005 0.012 0.184 0.34
Biodivers Conserv (2011) 20:1921–1949

Micropholis aff. crassipedicellata 0.97 4 0 0 4 0.00 0.255 0.000 0.026 0.281 0.88
Sloanea obtusifolia 0.96 2 0 0 2 0.00 0.341 0.000 0.036 0.377 0.92
Byrsonima sp. 0.95 6 0 1 7 1.41 0.095 0.000 0.051 0.146 3.98
Pouteria sagotiana 0.94 6 0 0 6 0.00 0.176 0.000 0.017 0.193 0.82
Copaifera langsdorffii 0.94 7 1 1 7 0.00 0.228 0.107 0.022 0.142 -4.19
Pouteria bullata 0.94 8 2 1 7 -1.21 0.141 0.010 0.010 0.141 0.05
Pouteria cuspidata 0.93 8 0 0 8 0.00 0.055 0.000 0.004 0.059 0.62
Gomidesia crocea 0.91 8 0 0 8 0.00 0.061 0.000 0.014 0.075 1.83
Bathysa australis 0.91 6 0 2 8 2.65 0.033 0.000 0.013 0.046 3.05
Calyptranthes sp6 0.90 7 1 0 6 -1.39 0.145 0.018 0.017 0.144 -0.06
Neomitranthes warmingiana 0.89 6 0 1 7 1.41 0.080 0.000 0.010 0.091 1.08
Hydrogaster trinervis 0.89 1 0 0 1 0.00 0.358 0.000 0.031 0.389 0.76
Protium heptaphyllum 0.88 6 1 0 5 -1.64 0.159 0.008 0.032 0.184 1.33
Eugenia rugosissima 0.88 8 1 0 7 -1.21 0.113 0.034 0.002 0.082 -2.91
Symplocos nitens 0.87 4 0 0 4 0.00 0.194 0.000 0.037 0.232 1.60
Ocotea dispersa 0.86 7 1 0 6 -1.39 0.131 0.010 0.004 0.126 -0.37
Ocotea elegans 0.85 4 0 0 4 0.00 0.192 0.000 0.030 0.221 1.31
1933

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 Table 5 continued
1934

Species IV Number of trees Change Basal area Change

(% year-1) (% year-1)

123
1992–1993 D R 2003–2004 1992–1993 Loss Gain 2003–2004

Ocotea velutina 0.85 6 0 0 6 0.00 0.112 0.000 0.007 0.119 0.55

Myrsine venosa 0.85 7 1 0 6 -1.39 0.113 0.008 0.039 0.145 2.25
Mollinedia salicifolia 0.84 7 0 0 7 0.00 0.087 0.000 0.003 0.090 0.30
Cupania furfuracea 0.84 7 1 0 6 -1.39 0.110 0.021 0.022 0.112 0.15
Sclerolobium densiflorum 0.84 4 0 0 4 0.00 0.210 0.000 0.030 0.240 1.23
IV importance value, D dead trees, R recruits
Biodivers Conserv (2011) 20:1921–1949
Biodivers Conserv (2011) 20:1921–1949 1935

Fig. 4 Balance of the number of trees (NT) of 28 endemic species between the original survey in
1992–1993 (Thomaz and Monteiro 1997) and the second survey in 2003–2004 (this study) in 1.02 ha of
montane rainforest at the Santa Lúcia Biological Station, Espı́rito Santo, southeastern Brazil. The diagonal
line represents the complete balance between original and second surveys. 1—Salacia nemorosa; 2—
Eugenia sp. 5 new; 3—Miconia sp. new; 4—Schefflera kollmannii; 5—Williamodendron cinnamomeum,
Myrcia sp. 6 new; 6—Schefflera grandigemma, Euterpe espiritosantensis, Cryptocarya velloziana, Myrcia
sp. 5 new; 7—Schefflera ruschiana; 8—Myrsine sp. new, Calyptranthes sp. 1 new, Myrcia sp. 3 new; 9—
Xylopia decorticans, Cryptocarya sp. new, Ocotea pluridomatiata; 10—Miconia capixaba; 11—Mollinedia
sp. new; 12—Ocotea sp. 7 new; 13—Eugenia rugosissima; 14—Trichilia sp. new, Elvasia capixaba; 15—
Vochysia santaluciae; 16—Pseudoxandra spiritus-sancti; 17—Ocotea cryptocarpa; 18—Mezilaurus sp.
new; 19—Unonopsis sancta-teresae

to the pattern expected for mature tropical forests (Swaine et al. 1987; Ayyappan and
Parthasarathy 2004; Stephenson and van Mantgem 2005).
The rates of mortality, recruitment and ingrowth are close to those found in various
mature tropical forests worldwide (Swaine et al. 1987; Phillips 1996; Ayyappan and
Parthasarathy 2004; Lewis et al. 2004a). The small differences between these rates found
in this study were non-significant and can be attributed to the cyclical rhythms of dynamic
processes, whereby mortality occurs first and opens the way for the establishment and
growth of recruits, creating a small temporary imbalance of these mechanisms (Oliveira-
Filho et al. 1997).
However, it should be noted that the slight increase in basal area could also be explained
by the CO2 fertilization hypothesis, which relates the increase in atmospheric CO2 con-
centrations to a greater productivity of forests and a consequent increase in biomass
(Phillips 1996). However, confirmation of a trend of increasing biomass would only be
possible if a sequence of periodic inventories had been carried out, as has been done in
other forests where this trend was analyzed (Laurance et al. 2009; Lewis et al. 2004a).
Thus, the occurrence of cyclical rhythms of dynamic processes was accepted as the best
explanation at the time.
Although balanced, the dynamics of the forest have been slow (a slow forest according
to Lewis et al. 2004a). The turnover time of about 70 years revealed that the replacement
of trees and the basal area of the forest are long processes. The turnover rates for trees or
basal areas were slightly lower than in other mature tropical forests, where normally the

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turnover rates reach 1–3% year-1 (Phillips 1996). However, comparisons among dynamics
studies with different time intervals are inappropriate because, according to Lewis et al.
(2004b), the values of turnover rates tend to decrease when the time intervals increase.
Thus, small differences when compared to other mature forests may only be mathematical
artifacts.
The high ecological diversity remained after 11 years because there was balance between
the numbers of species and families in both surveys and the turnover of floristic composition
was performed by the less abundant (rare) species of the community. This role of less
abundant species in community dynamics has been confirmed by other studies (Ayyappan
and Parthasarathy 2004; Werneck and Franceschinelli 2004; Guedes-Bruni et al. 2009).
The large participation of the family Myrtaceae in the species turnover was surely a
result of its large number of species (N = 83) in the survey, many of which were repre-
sented by only one or two trees. In the case of Melastomataceae, the large participation,
especially in species recruitment, can be attributed to the ability of some Melastomataceae
species (named gap-dependent species) to respond with rapid growth in height and width of
the canopy under more intense conditions of natural light caused by the opening of natural
gaps (Daws et al. 2007).
These characteristics of the community show that the natural disturbances (especially
the death of trees by various causes) generally expected in any section of a mature tropical
forest did not modify the structure of the forest SLBS over time. It is true that during the
interval of the study, more precisely in the years 1997 and 1998, various tropical regions
experienced a period of drought and intense heat due to the El Niño Southern Oscillation
(ENSO). Although there are no detailed records of the structure of the SLBS forest during
that period, it is likely that mortality rates increased significantly in the community, as
reported by studies in the Amazon rainforest (Nascimento et al. 2007; Laurance et al.
2009), in southeastern Brazil (Rolim et al. 2005) and in Borneo (Lingenfelder and New-
bery 2009). Assuming that these effects did actually occur in the SLBS forest, the results
show that after the ENSO the losses were offset after only 5 years.

Diameter classes

For this study, no relationship was found between mortality rates and diameter class. This
situation also was found in other studies carried out in preserved tropical forests
(Manokaran and Kochummen 1987; Carey et al. 1994; Korning and Balslev 1994). This
contrasts with the higher mortality rates in smaller size classes (usually with dbh between 5
and 20 cm), which has been recorded for tropical forests in regeneration after anthropo-
genic disturbances (Guilherme et al. 2004; Werneck and Franceschinelli 2004; Higuchi
et al. 2008; Taylor et al. 2008).
In regard to recruitment, it was expected that the recruits would be smaller trees, con-
centrated in the 6.4–10 cm class. However, this concentration did not result in an increase of
basal area in this class due to the migration of trees to other classes for ingrowth.

Populations

The main populations also showed stability, following the general pattern of balance
between mortality and recruitment for mature tropical forests. As for the community, small
differences in abundance or in the basal area of each population between surveys may also
be the result of dynamic cycles involving the mechanisms of mortality and recruitment/
growth.

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One of the few species that did not fit into this pattern was the palm Euterpe edulis,
which had expanded its participation due to the large number of recruits. This may be
linked to the reproductive success of this species, as already recorded by Silva-Matos
and Watkinson (1998) and Portela et al. (2010), and the absence of disturbances able to
modify the structure of the community. Euterpe edulis is important in Atlantic rain-
forest because of its large production of fruit, which constitutes a major food resource
for wildlife at certain periods of the year (Silva-Matos and Watkinson 1998; Portela
et al. 2010). However, it is a species which is still exploited illegally for its edible palm
heart.
In the other cases of species which showed large changes (annual rates above 6%,
positive or negative), the small size of these populations may explain why the death/
recruitment of few trees or the loss/gain of very small portions of basal area resulted in
seemingly high rates.
In the case of local endemic species in particular, the balance of trees in the original and
second surveys was important in ensuring the continuity of its populations in the plots.
However, to say that these species are protected from the risk of extinction is still hasty,
since the time scale of this study may have been inappropriate for this interpretation.
Moreover, since the geographical distribution of these species was limited to a few square
kilometers in region, this shows that their future also depends on the conservation of other
populations located in neighboring forest fragments.
The slight increase in basal area experienced by the total community was also confirmed
in most populations because they had small positive changes. Negative changes occurred in
a small number of species and this was not sufficient to fully balance gains and losses of
the community.

Conclusion

The data on the tree changes in this study showed an affinity with the pattern of dynamic
balance for both structure and diversity expected in mature tropical forests. This is
interesting, mainly because the balance in this study was in a forest with high levels of
diversity and endemism.
In addition, this study provides a basis for further work on the dynamics of the forest
studied. Such investigations may be conducted over the coming years in other surveys,
perhaps with shorter intervals and also considering other components such as lianas and
saplings of trees species, as has been done in various forests throughout the world. Topics
that might be explored are the changes in basal area and turnover rates, changes in the
participation of groups of species with different requirements for light and moisture and
monitoring of the dynamics of local endemic species in order to create support for
conservation.
In fact, the 1.02 ha of this study may seem small when compared with the large plots in
other dynamic studies around the world. But the absence of data on the dynamics of mature
forests in the Atlantic rainforest region is mainly caused by the scarcity of these areas, by
the large amount of time needed to carry out these surveys, which also requires a large
amount of human and financial resources, and by the difficulty in identifying botanical
species within a very rich flora that are still little known. The small number of research
projects to date are the only existing records about the dynamics of this magnificent
neotropical ecoregion.

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Acknowledgments This study is part of the Master’s dissertation of Felipe Z. Saiter. The authors thank the
National Council for Scientific and Technological Development (CNPq) for financial support (grant n.
690149/01-8) and for a productivity grant to T Wendt and FAG Guilherme, the Museu de Biologia Prof.
Mello Leitão for research permits and logistical support, and FR Scarano, MT Nascimento and two
anonymous reviewers for their helpful suggestions. They also thank the biologists who helped with the field
work: T Senna, V Demuner, R Vervloet, J Tomasini, FA Obermüller, GM Tobón, TS Cóser, ML Dan and
MAS Godinho.

Appendix

See Table 6.

Table 6 List of 399 species (with 28 local endemic) recorded in two successive surveys (1992–1993 and
2003–2004) of trees with dbh C6.4 cm carried out in a stand of 1.02 ha in Santa Lúcia Biological Station,
Santa Teresa, southeastern Brazil. Presence (1) and absence (0) of species in each survey are listed
separately
No. Family Species Presence/absence

1992–1993 2003–2004

1 Anacardiaceae Tapirira guianensis 1 1

2 Annonaceae Anaxagorea dolichocarpa 0 1
3 Annonaceae Annona cacans 0 1
4 Annonaceae Guatteria australis 1 1
5 Annonaceae Guatteria glabrescens 1 1
6 Annonaceae Guatteria sellowiana 1 1
7 Annonaceae Oxandra nitida 1 1
8 Annonaceae Pseudoxandra spiritus-sanctia 1 1
9 Annonaceae Rollinia dolabripetala 1 0
10 Annonaceae Unonopsis sancta-teresaea 1 1
11 Annonaceae Xylopia decorticansa 1 1
12 Apocynaceae Aspidosperma melanocalyx 1 1
13 Apocynaceae Aspidosperma parvifolium 1 1
14 Apocynaceae Himatanthus bracteatus 1 1
15 Apocynaceae Lacmellea pauciflora 1 1
16 Apocynaceae Rauvolfia grandiflora 1 1
17 Aquifoliaceae Ilex amara 1 0
18 Aquifoliaceae Ilex paraguariensis 1 1
19 Araliaceae Schefflera calva 1 1
20 Araliaceae Schefflera grandigemmaa 1 1
21 Araliaceae Schefflera kollmanniia 1 1
22 Araliaceae Schefflera ruschianaa 1 1
23 Arecaceae Attalea burretiana 1 1
24 Arecaceae Euterpe edulis 1 1
25 Arecaceae Euterpe espiritosantensisa 1 1
26 Arecaceae Geonoma schottiana 1 1
27 Arecaceae Syagrus pseudococus 1 1
28 Asteraceae Vernonia discolor 1 1
29 Bignoniaceae Jacaranda microcalyx 1 1

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Biodivers Conserv (2011) 20:1921–1949 1939

Table 6 continued

No. Family Species Presence/absence

1992–1993 2003–2004

30 Bignoniaceae Tabebuia roseo-alba 1 1

31 Boraginaceae Cordia sp. 1 1
32 Boraginaceae Cordia trachyphylla 1 1
33 Burseraceae Protium brasiliense 1 1
34 Burseraceae Protium heptaphyllum 1 1
35 Cardiopteridaceae Citronella paniculata 1 1
36 Caricaceace Jacaratia heptaphylla 1 1
37 Caryocaraceae Caryocar edule 1 1
38 Celastraceae Cheiloclinium cognatum 1 1
39 Celastraceae Maytenus cestrifolia 1 1
40 Celastraceae Maytenus obtusifolia 1 1
41 Celastraceae Salacia elliptica 1 1
42 Celastraceae Salacia nemorosa 1 0
43 Celastraceae Tontelea martiana 1 1
44 Chrysobalanaceae Couepia grandiflora 1 1
45 Chrysobalanaceae Couepia macrophylla 1 1
46 Chrysobalanaceae Couepia venosa 1 1
47 Chrysobalanaceae Hirtella angustifolia 1 1
48 Chrysobalanaceae Hirtella hebeclada 1 1
49 Chrysobalanaceae Licania kunthiana 1 1
50 Chrysobalanaceae Licania leptostachya 1 1
51 Chrysobalanaceae Licania micrantha 1 1
52 Chrysobalanaceae Licania octandra 1 1
53 Chrysobalanaceae Licania parvifolia 1 1
54 Chrysobalanaceae Licania salzmannii 1 1
55 Chrysobalanaceae Parinari aff. littoralis 1 1
56 Chrysobalanaceae Parinari obtusifolium 1 1
57 Clusiaceae Clusia intermedia 1 1
58 Clusiaceae Garcinia brasiliensis 1 1
59 Clusiaceae Garcinia gardneriana 1 1
60 Clusiaceae Kielmeyera occhioniana 1 1
61 Clusiaceae Tovomita brasiliensis 1 1
62 Clusiaceae Tovomita leucantha 1 1
63 Clusiaceae Tovomitopsis saldanhae 1 1
64 Connaraceae Connarus detersus 1 1
65 Cunoniaceae Lamanonia ternata 1 1
66 Dichapetalaceae Stephanopodium blanchetianum 1 1
67 Elaeocarpaceae Sloanea aff. garckeana 1 1
68 Elaeocarpaceae Sloanea obtusifolia 1 1
69 Elaeocarpaceae Sloanea guianensis 1 1
70 Elaeocarpaceae Sloanea monosperma 1 1
71 Elaeocarpaceae Sloanea nitida 1 1

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1940 Biodivers Conserv (2011) 20:1921–1949

Table 6 continued

No. Family Species Presence/absence

1992–1993 2003–2004

72 Elaeocarpaceae Sloanea sp. 1 1

73 Erythroxylaceae Erythroxylum aff. macrophyllum 1 1
74 Erythroxylaceae Erythroxylum cuspidifolium 1 0
75 Erythroxylaceae Erythroxylum pulchrum 1 1
76 Erythroxylaceae Erythroxylum squamatum 1 1
77 Euphorbiaceae Alchornea triplinervia 1 1
78 Euphorbiaceae Aparisthmium cordatum 1 1
79 Euphorbiaceae Croton floribundus 1 0
80 Euphorbiaceae Maprounea guianensis 1 1
81 Euphorbiaceae Pausandra morisiana 1 1
82 Euphorbiaceae Pera leandri 1 1
83 Fabaceae Abarema cf. obovata 1 1
84 Fabaceae Andira fraxinifolia 1 1
85 Fabaceae Copaifera langsdorffii 1 1
86 Fabaceae Dalbergia foliolosa 1 1
87 Fabaceae Dalbergia miscolobium 1 1
88 Fabaceae Diplotropis incexis 1 1
89 Fabaceae Hymenaea aurea 1 1
90 Fabaceae Hymenaea courbaril 1 1
91 Fabaceae Hymenolobium janeirense 1 1
92 Fabaceae Inga capitata 1 1
93 Fabaceae Inga cylindrica 1 1
94 Fabaceae Inga densiflora 1 1
95 Fabaceae Inga dulcis 1 1
96 Fabaceae Inga flagelliformis 1 1
97 Fabaceae Inga lenticellata 1 1
98 Fabaceae Inga subnuda 1 1
99 Fabaceae Inga tenuis 1 1
100 Fabaceae Inga thibaudiana 1 1
101 Fabaceae Inga vestita 1 1
102 Fabaceae Melanoxylon brauna 1 1
103 Fabaceae Ormosia cf. friburgensis 1 1
104 Fabaceae Peltogyne angustiflora 1 1
105 Fabaceae Pithecellobium cochliocarpum 1 1
106 Fabaceae Pseudopiptadenia contorta 1 1
107 Fabaceae Sclerolobium densiflorum 1 1
108 Fabaceae Sclerolobium striatum 1 1
109 Fabaceae Senna multijuga 1 1
110 Fabaceae Swartzia acutifolia 1 1
111 Fabaceae Swartzia apetala 1 1
112 Fabaceae Swartzia myrtifolia 1 1
113 Fabaceae Zollernia ilicifoia 1 1

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Table 6 continued

No. Family Species Presence/absence

1992–1993 2003–2004

114 Fabaceae Zollernia magnifica 1 1

115 Hernandiaceae Sparattanthelium botocudorum 1 1
116 Humiriaceae Vantanea compacta 1 1
117 Humiriaceae Vantanea obovata 1 1
118 Hypericaceae Vismia brasiliensis 1 1
119 Lacistemataceae Lacistema robustum 1 1
120 Lamiaceae Vitex orinocensis 1 1
121 Lamiaceae Vitex sp. 1 1
122 Lauraceae Aniba firmula 0 1
123 Lauraceae Beilschmiedia sp. 1 1
124 Lauraceae Beilschmiedia taubertiana 1 1
125 Lauraceae Cinnamomum riedelianum 1 1
126 Lauraceae Cinnamomum sp. 1 1
127 Lauraceae Cryptocarya saligna 1 1
128 Lauraceae Cryptocarya sp. newa 1 1
129 Lauraceae Cryptocarya vellozianaa 1 1
130 Lauraceae Endlicheria paniculata 1 1
131 Lauraceae Endlicheria sp. 1 1
132 Lauraceae Licaria armeniaca 1 1
133 Lauraceae Licaria guianensis 1 0
134 Lauraceae Ocotea aciphylla 1 1
135 Lauraceae Ocotea catharinensis 1 1
136 Lauraceae Ocotea corymbosa 1 1
137 Lauraceae Ocotea cryptocarpaa 1 1
138 Lauraceae Ocotea daphnifolia 1 1
139 Lauraceae Ocotea dispersa 1 1
140 Lauraceae Ocotea divaricata 1 1
141 Lauraceae Ocotea domatiata 1 1
142 Lauraceae Ocotea elegans 1 1
143 Lauraceae Ocotea indecora 0 1
144 Lauraceae Ocotea lancifolia 1 1
145 Lauraceae Ocotea longifolia 1 1
146 Lauraceae Ocotea odorifera 1 1
147 Lauraceae Ocotea puberula 1 1
148 Lauraceae Ocotea pulchra 1 1
149 Lauraceae Ocotea silvestris 1 1
150 Lauraceae Mezilaurus sp. newa 1 1
151 Lauraceae Ocotea sp1 1 1
152 Lauraceae Ocotea sp2 1 1
153 Lauraceae Ocotea sp3 1 1
154 Lauraceae Ocotea sp4 1 1
155 Lauraceae Ocotea sp5 1 1

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1942 Biodivers Conserv (2011) 20:1921–1949

Table 6 continued

No. Family Species Presence/absence

1992–1993 2003–2004

156 Lauraceae Ocotea sp6 1 1

157 Lauraceae Ocotea sp7 newa 1 1
158 Lauraceae Ocotea pluridomatiataa 1 1
159 Lauraceae Ocotea spixiana 1 1
160 Lauraceae Ocotea teleiandra 1 1
161 Lauraceae Ocotea tenuiflora 1 1
162 Lauraceae Ocotea velutina 1 1
163 Lauraceae Persea caesia 1 1
164 Lauraceae Persea sp. 1 1
165 Lauraceae Williamodendron cinnamomeuma 1 1
166 Linnaceae Roucheria cf. dryadica 0 1
167 Loganiaceae Strychnos sp. 1 1
168 Malphigiaceae Barnebya dispar 1 1
169 Malphigiaceae Byrsonima sp. 1 1
170 Malphigiaceae Byrsonima variabilis 1 1
171 Malvaceae Bombacopsis calophylla 1 1
172 Malvaceae Eriotheca macrophylla 1 1
173 Malvaceae Hydrogaster trinervis 1 1
174 Melastomataceae Meriania tetramera 1 1
175 Melastomataceae Miconia budlejoides 1 1
176 Melastomataceae Miconia capixabaa 1 1
177 Melastomataceae Miconia cinnamomifolia 1 1
178 Melastomataceae Miconia dodecandra 1 1
179 Melastomataceae Miconia doriana 1 0
180 Melastomataceae Miconia latecrenata 1 1
181 Melastomataceae Miconia lepidota 1 1
182 Melastomataceae Miconia longicuspis 0 1
183 Melastomataceae Miconia octopetala 1 1
184 Melastomataceae Miconia polyandra 1 1
185 Melastomataceae Miconia prasina 1 1
186 Melastomataceae Miconia pusilliflora 1 1
187 Melastomataceae Miconia sellowiana 0 1
188 Melastomataceae Miconia sp. newa 0 1
189 Melastomataceae Miconia tristis 0 1
190 Melastomataceae Mouriri cf. regeliana 1 1
191 Melastomataceae Mouriri doriana 1 1
192 Melastomataceae Mouriri glazioviana 1 1
193 Meliaceae Cabralea canjerana 1 1
194 Meliaceae Guarea macrophylla 1 1
195 Meliaceae Trichilia emarginata 1 1
196 Meliaceae Trichilia lepidota 1 1
197 Meliaceae Trichilia silvatica 1 1

123
Biodivers Conserv (2011) 20:1921–1949 1943

Table 6 continued

No. Family Species Presence/absence

1992–1993 2003–2004

198 Meliaceae Trichilia sp. newa 1 1

199 Monimiaceae Mollinedia stenophylla 1 1
200 Monimiaceae Mollinedia aff. engleriana 1 1
201 Monimiaceae Mollinedia fruticulosa 1 1
202 Monimiaceae Mollinedia heteranthera 1 1
203 Monimiaceae Mollinedia salicifolia 1 1
204 Monimiaceae Mollinedia sp. newa 1 1
205 Moraceae Brosimum cf. glaziovii 1 1
206 Moraceae Ficus citrifolia 1 1
207 Moraceae Sorocea guilleminiana 1 1
208 Myristicaceae Virola gardneri 1 1
209 Myrsinaceae Myrsine lancifolia 1 1
210 Myrsinaceae Myrsine sp. newa 1 1
211 Myrsinaceae Myrsine umbellata 1 1
212 Myrsinaceae Myrsine venosa 1 1
213 Myrtaceae Blepharocalyx eggersii 1 1
214 Myrtaceae Calyptranthes aff. clusiifolia 1 1
215 Myrtaceae Calyptranthes aff. grandifolia 1 1
216 Myrtaceae Calyptranthes pauciflora 1 1
217 Myrtaceae Calyptranthes pulchella 1 1
218 Myrtaceae Calyptranthes sp1 newa 1 1
219 Myrtaceae Calyptranthes sp2 1 1
220 Myrtaceae Calyptranthes sp3 1 1
221 Myrtaceae Calyptranthes sp4 1 1
222 Myrtaceae Calyptranthes sp5 1 1
223 Myrtaceae Calyptranthes sp6 1 1
224 Myrtaceae Calyptranthes sp7 1 1
225 Myrtaceae Calyptranthes sp8 1 1
226 Myrtaceae Calyptranthes sp9 1 1
227 Myrtaceae Calyptranthes widgreniana 1 1
228 Myrtaceae Campomanesia aromatica 1 1
229 Myrtaceae Campomanesia guaviroba 1 1
230 Myrtaceae Campomanesia laurifolia 1 1
231 Myrtaceae Eugenia acutata 1 1
232 Myrtaceae Eugenia aff. platysema 1 1
233 Myrtaceae Eugenia aff. pruniformis 1 1
234 Myrtaceae Eugenia aggregata 1 1
235 Myrtaceae Eugenia candolleana 1 1
236 Myrtaceae Eugenia cerasiflora 1 1
237 Myrtaceae Eugenia cf. martiusiana 1 1
238 Myrtaceae Eugenia cf. neomooniana 1 1
239 Myrtaceae Eugenia copacabanensis 1 1

123
1944 Biodivers Conserv (2011) 20:1921–1949

Table 6 continued

No. Family Species Presence/absence

1992–1993 2003–2004

240 Myrtaceae Eugenia egensis 1 1

241 Myrtaceae Eugenia excelsa 1 1
242 Myrtaceae Eugenia itapemirimensis 1 1
243 Myrtaceae Eugenia melanogyna 0 1
244 Myrtaceae Eugenia neoglomerata 1 0
245 Myrtaceae Eugenia neolanceolata 1 1
246 Myrtaceae Eugenia oblongata 1 1
247 Myrtaceae Eugenia persicifolia 1 1
248 Myrtaceae Eugenia piloensis 1 1
249 Myrtaceae Eugenia platyphylla 1 1
250 Myrtaceae Eugenia rostrata 1 1
251 Myrtaceae Eugenia rugosissimaa 1 1
252 Myrtaceae Eugenia sp1 1 1
253 Myrtaceae Eugenia sp2 1 1
254 Myrtaceae Eugenia sp3 1 1
255 Myrtaceae Eugenia sp4 1 1
256 Myrtaceae Eugenia sp5 newa 0 1
257 Myrtaceae Eugenia tinguyensis 1 1
258 Myrtaceae Eugenia xiriricana 1 1
259 Myrtaceae Gomidesia cf. palustris 1 1
260 Myrtaceae Gomidesia cf. pubescens 1 1
261 Myrtaceae Gomidesia cf. schaueriana 1 1
262 Myrtaceae Gomidesia crocea 1 1
263 Myrtaceae Marlierea cf. ovata 1 1
264 Myrtaceae Marlierea excoriata 1 1
265 Myrtaceae Marlierea obscura 1 1
266 Myrtaceae Marlierea regeliana 1 1
267 Myrtaceae Marlierea silvatica 1 1
268 Myrtaceae Marlierea sp. 1 1
269 Myrtaceae Myrcia crocea 1 1
270 Myrtaceae Myrcia laurifolia 1 1
271 Myrtaceae Myrcia montana 1 1
272 Myrtaceae Myrcia plusiantha 1 1
273 Myrtaceae Myrcia pubipetala 1 0
274 Myrtaceae Myrcia racemosa 1 0
275 Myrtaceae Myrcia sp1 1 1
276 Myrtaceae Myrcia sp2 1 1
277 Myrtaceae Myrcia sp3 newa 1 1
278 Myrtaceae Myrcia sp4 1 1
279 Myrtaceae Myrcia sp5 newa 1 1
280 Myrtaceae Myrcia sp6 newa 1 1
281 Myrtaceae Myrcia sp7 1 1

123
Biodivers Conserv (2011) 20:1921–1949 1945

Table 6 continued

No. Family Species Presence/absence

1992–1993 2003–2004

282 Myrtaceae Myrcia splendens 1 1

283 Myrtaceae Myrcia subrugosa 1 1
284 Myrtaceae Myrciaria disticha 1 1
285 Myrtaceae Myrciaria floribunda 1 1
286 Myrtaceae Neomitranthes glomerata 1 1
287 Myrtaceae Neomitranthes warmingiana 1 1
288 Myrtaceae Pimenta pseudocaryophyllus 1 1
289 Myrtaceae Plinia cf. involucrata 1 1
290 Myrtaceae Plinia renatiana 1 1
291 Myrtaceae Plinia rivularis 1 1
292 Myrtaceae Psidium sp1 1 1
293 Myrtaceae Psidium sp2 1 1
294 Myrtaceae Siphoneugena cf. kiaerskoviana 1 1
295 Myrtaceae Siphoneugena dussii 1 0
296 Nyctaginaceae Guapira laxa 1 1
297 Nyctaginaceae Guapira obtusata 1 1
298 Nyctaginaceae Guapira opposita 1 1
299 Nyctaginaceae Guapira venosa 1 1
300 Ochnaceae Elvasia capixabaa 1 1
301 Ochnaceae Ouratea cuspidata 1 1
302 Olacaceae Heisteria cf. silvianii 1 1
303 Olacaceae Heisteria perianthomega 1 1
304 Olacaceae Tetrastylidium grandifolium 1 1
305 Oleaceae Chionanthus micranthus 1 1
306 Pentaphyllacaceae Ternstroemia brasiliensis 1 1
307 Pentaphyllacaceae Ternstroemia sp. 1 1
308 Phyllanthaceae Hyeronima alchorneoides 1 1
309 Phyllanthaceae Hyeronima oblonga 1 1
310 Phyllanthaceae Margaritaria nobilis 1 1
311 Polygonaceae Coccoloba confusa 1 1
312 Polygonaceae Coccoloba declinata 1 1
313 Proteaceae Panopsis sp. 0 1
314 Proteaceae Roupala aff. rhombifolia 1 1
315 Proteaceae Roupala consimilis 1 1
316 Putranjivaceae Drypetes sessiliflora 1 1
317 Quiinaceae Quiina glaziovii 1 1
318 Rosaceae Prunus brasiliensis 1 1
319 Rosaceae Prunus sellowii 1 1
320 Rubiaceae Alibertia sp. 1 1
321 Rubiaceae Amaioua intermedia 1 1
322 Rubiaceae Amaioua pilosa 1 1
323 Rubiaceae Bathysa australis 1 1

123
1946 Biodivers Conserv (2011) 20:1921–1949

Table 6 continued

No. Family Species Presence/absence

1992–1993 2003–2004

324 Rubiaceae Bathysa stipulata 1 1

325 Rubiaceae Faramea oligantha 1 1
326 Rubiaceae Faramea pachyantha 1 1
327 Rubiaceae Ixora sp. 1 1
328 Rubiaceae Posoqueria acutifolia 1 1
329 Rubiaceae Posoqueria latifolia 1 1
330 Rubiaceae Psychotria carthagenensis 1 1
331 Rubiaceae Psychotria sessilis 1 1
332 Rubiaceae Rudgea recurva 1 1
333 Rubiaceae Simira glaziovii 1 1
334 Rubiaceae Simira sampaioana 1 1
335 Rubiaceae Stachyarrhena krukovii 1 1
336 Rutaceae Hortia brasiliana 1 1
337 Sabiaceae Meliosma chartacea 1 1
338 Salicaceae Banara serrata 1 0
339 Salicaceae Casearia arborea 1 1
340 Salicaceae Casearia commersoniana 1 1
341 Salicaceae Casearia decandra 1 1
342 Salicaceae Casearia sp1 1 1
343 Salicaceae Casearia sp2 1 1
344 Sapindaceae Allophylus laevigatus 1 1
345 Sapindaceae Allophylus petiolulatus 1 1
346 Sapindaceae Cupania crassifolia 1 1
347 Sapindaceae Cupania emarginata 1 1
348 Sapindaceae Cupania furfuracea 1 1
349 Sapindaceae Cupania scrobiculata 1 1
350 Sapindaceae Matayba arborescens 1 1
351 Sapindaceae Matayba cf. guianensis 1 1
352 Sapindaceae Talisia cf. cerasina 1 1
353 Sapindaceae Talisia cupularis 1 1
354 Sapindaceae Talisia sp. 0 1
355 Sapotaceae Chrysophyllum flexuosum 1 1
356 Sapotaceae Chrysophyllum gonocarpum 1 1
357 Sapotaceae Chrysophyllum sp1 1 1
358 Sapotaceae Chrysophyllum sp2 1 1
359 Sapotaceae Chrysophyllum splendens 1 1
360 Sapotaceae Diploon cuspidatum 1 1
361 Sapotaceae Ecclinusa ramiflora 1 1
362 Sapotaceae Manilkara cf. longifolia 1 1
363 Sapotaceae Micropholis aff. crassipedicellata 1 1
364 Sapotaceae Micropholis aff. gardneriana 1 1
365 Sapotaceae Micropholis compta 1 1

123
Biodivers Conserv (2011) 20:1921–1949 1947

Table 6 continued

No. Family Species Presence/absence

1992–1993 2003–2004

366 Sapotaceae Micropholis guyanensis 1 1

367 Sapotaceae Micropholis sp. 1 1
368 Sapotaceae Micropholis venulosa 1 1
369 Sapotaceae Pouteria aff. reticulata 1 1
370 Sapotaceae Pouteria bangii 1 1
371 Sapotaceae Pouteria bullata 1 1
372 Sapotaceae Pouteria caimito 1 1
373 Sapotaceae Pouteria cf. coelomatica 1 1
374 Sapotaceae Pouteria cf. guianensis 1 1
375 Sapotaceae Pouteria cuspidata 1 1
376 Sapotaceae Pouteria gardneri 1 1
377 Sapotaceae Pouteria grandiflora 1 1
378 Sapotaceae Pouteria macahensis 1 1
379 Sapotaceae Pouteria sagotiana 1 1
380 Sapotaceae Pouteria sp1 1 1
381 Sapotaceae Pouteria sp2 1 1
382 Sapotaceae Pradosia lactescens 1 1
383 Simaroubaceae Simarouba amara 1 1
384 Siparunaceae Siparuna glossostyla 1 1
385 Solanaceae Solanum sooretamum 1 0
386 Symplocaceae Symplocos celastrinea 1 1
387 Symplocaceae Symplocos frondosa 1 1
388 Symplocaceae Symplocos nitens 1 1
389 Thymaelaceae Daphnopsis martii 1 1
390 Urticaceae Cecropia hololeuca 1 1
391 Urticaceae Coussapoa cf. glaberrima 1 1
392 Urticaceae Coussapoa microcarpa 1 1
393 Urticaceae Coussapoa pachyphylla 1 1
394 Urticaceae Pourouma bicolor 1 0
395 Urticaceae Pourouma guianensis 1 1
396 Vochysiaceae Qualea gestasiana 0 1
397 Vochysiaceae Vochysia rectiflora 1 1
398 Vochysiaceae Vochysia santaluciaea 1 1
399 Winteraceae Drimys brasiliensis 1 0
a
Local endemic species

123
1948 Biodivers Conserv (2011) 20:1921–1949

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