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ABSTRACT The tree flora of seasonally dry forests (SDTF) of eastern tropical and subtropical South America was investigated according to two main aspects: (a) the variations in floristic composition were analyzed in terms of geographical and climatic variables by performing multivariate analyses on 532 existing floristic checklists; and (b) the links among different seasonally dry forest formations, Amazonian forests and cerrados (woody savannas) were assessed. Analyses were performed at the species, genus and family levels. There was a strong spatial pattern in tree species distribution that only receded and allowed clearer climate-related patterns to arise when either the geographical range was restricted or data were treated at the genus and family levels. Consistent floristic differences occurred between rain and seasonal forests, although these were obscured by Strong regional similarities which made the two foresttypes from the same region closer to each other floristically than they were to their equivalents in different regions. Atlantic rain and seasonal forests were floristically closer to each other than to Amazonian rain forests but north-east rain and seasonal forests were both closer to Amazonian rain forests than each other, though only at the generic and familial levels. Atlantic seasonal forests also share a variable proportion of species with caatingas, cerrados and the chaco, and may represent a transition to these open formations. Increasing periods of water shortage, with increases in soil fertility and temperature are characteristic of a transition from semideciduous to deciduous forests and then to the semi-arid formations, either caatingas (tropical) or chaco forests (subtropical), while increasing fire frequency and decreasing soil fertility lead from seasonal forests to either cerrados (tropical) or southern campos (subtropical). The SDTF vegetation of eastern South America may be classified into three floristic nuclei: caatinga, chaco and Atlantic forest (sensu latissimo). Only the last, however, should be linked consistently to the residual Pleistocenic dry seasonal flora (RPDS). Caatinga and chaco represent the extremes of floristic dissimilarity among the three nuclei, also corresponding to the warm-dry and warm-cool climatic extremes, respectively. In contrast to the caatinga and chaco nuclei, the Atlantic SDTF nucleus is poor in endemic species and is actually a floristic bridge connecting the two drier nuclei to rain forests. Additionally, there are few grounds to recognize the Atlantic nucleus flora as a clearly distinct species assemblage, since there is a striking variation in species composition found throughout its wide geographical range. Nevertheless, there is a group of wide-range species that are found in most regions of the Atlantic nucleus, some of which are also part of the species blend of the Caatinga and Chaco floras, though the latter plays a much smaller part. We propose that it is precisely this small fraction of the Atlantic nucleus flora that should be identified with the RPDS vegetation.
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151
7
Floristic Relationships of
Seasonally Dry Forests of Eastern
South America Based on Tree
Species Distribution Patterns
Ary T. Oliveira-Filho, João André Jarenkow, Maria Jesus
Nogueira Rodal
CONTENTS
7.1 Introduction...........................................................................................................................152
7.2 Methods ................................................................................................................................154
7.2.1 Preparation and Revision of the Databases .............................................................154
7.2.2 Vegetation Classification ..........................................................................................157
7.2.3 Multivariate Analyses...............................................................................................158
7.2.4 Condensed Floristic Data .........................................................................................159
7.3 Results...................................................................................................................................160
7.3.1 Multivariate Analyses...............................................................................................160
7.3.2 Analyses of Condensed Floristic Information .........................................................162
7.4 Discussion.............................................................................................................................170
7.5 Conclusion ............................................................................................................................175
Acknowledgements ........................................................................................................................176
References ......................................................................................................................................176
Appendix. Most Frequent Species (>70% of Checklists) in the Tree
flora of Selected SDTF and SDSF Formations of Eastern South America..................................179
ABSTRACT
The tree flora of seasonally dry forests (SDTF) of eastern tropical and subtropical South America
was investigated according to two main aspects: (a) the variations in floristic composition were
analyzed in terms of geographical and climatic variables by performing multivariate analyses on
532 existing floristic checklists; and (b) the links among different seasonally dry forest formations,
Amazonian forests and cerrados (woody savannas) were assessed. Analyses were performed at the
species, genus and family levels. There was a strong spatial pattern in tree species distribution that
only receded and allowed clearer climate-related patterns to arise when either the geographical
range was restricted or data were treated at the genus and family levels. Consistent floristic
differences occurred between rain and seasonal forests, although these were obscured by strong
regional similarities which made the two foresttypes from the same region closer to each other
floristically than they were to their equivalents in different regions. Atlantic rain and seasonal forests
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Neotropical Savannas and Dry Forests: Diversity, Biogeography, and Conservation
were floristically closer to each other than to Amazonian rain forests but north-east rain and seasonal
forests were both closer to Amazonian rain forests than each other, though only at the generic and
familial levels. Atlantic seasonal forests also share a variable proportion of species with caatingas,
cerrados and the chaco, and may represent a transition to these open formations. Increasing periods
of water shortage, with increases in soil fertility and temperature are characteristic of a transition
from semideciduous to deciduous forests and then to the semi-arid formations, either caatingas
(tropical) or chaco forests (subtropical), while increasing fire frequency and decreasing soil fertility
lead from seasonal forests to either cerrados (tropical) or southern campos (subtropical). The SDTF
vegetation of eastern South America may be classified into three floristic nuclei: caatinga, chaco
and Atlantic forest (sensu latissimo). Only the last, however, should be linked consistently to the
residual Pleistocenic dry seasonal flora (RPDS). Caatinga and chaco represent the extremes of
floristic dissimilarity among the three nuclei, also corresponding to the warm-dry and warm-cool
climatic extremes, respectively. In contrast to the caatinga and chaco nuclei, the Atlantic SDTF
nucleus is poor in endemic species and is actually a floristic bridge connecting the two drier nuclei
to rain forests. Additionally, there are few grounds to recognize the Atlantic nucleus flora as a
clearly distinct species assemblage, since there is a striking variation in species composition found
throughout its wide geographical range. Nevertheless, there is a group of wide-range species that
are found in most regions of the Atlantic nucleus, some of which are also part of the species blend
of the Caatinga and Chaco floras, though the latter plays a much smaller part. We propose that it
is precisely this small fraction of the Atlantic nucleus flora that should be identified with the RPDS
vegetation.
7.1 INTRODUCTION
Neotropical seasonally dry tropical forests, or SDTF, are presently an increasing focus of attention
because of both their very threatened status and poorly studied flora and ecology, and this is striking
when compared to traditional flag ecosystems like rain forests and savannas (Mooney et al., 1995).
They occur where annual rainfall is less than 1600 mm and more than 5–6 months receive less than
100 mm (Graham and Dilcher, 1995) and therefore include a diverse array of vegetation formations,
from tall semideciduous forests to thorny woodland with succulents (Murphy and Lugo, 1995).
Despite all this variation, Pennington et al. (2000) argue that the concept of SDTF should exclude
fire-related formations, such as savannas and cerrados, and the non-tropical chaco forests.
The distribution of SDTF in South America forms an arc with the ends positioned at the caatinga
domain of north-eastern Brazil and the Caribbean coast of Colombia and Venezuela and a long
curved route connecting the ends through the seasonal forests of the Atlantic forest domain, the
patches of seasonal forests of the cerrado domain, and the seasonal forests of the Andean piedmont,
inter-Andean valleys, Pacific coast and Caribbean coast. Prado (1991) and Prado and Gibbs (1993)
suggested the hypothesis that this arc is a relic of a much wider distribution of SDTF in South
America reached during the Pleistocene glacial maxima. They based their model on the present
distribution of what they called residual Pleistocenic dry seasonal (RPDS) flora. Since then a number
of studies have analysed species distribution patterns of this flora in order to assess the validity of
the RPDS arc hypothesis in different geographical contexts (e.g. Pennington et al., 2000, 2004;
Prado, 2000; Bridgewater et al., 2003; Linares-Palomino et al., 2003; Spichiger et al., 2004). The
assessment of floristic links among species assemblages of SDTF areas scattered over the putative
RPDS arc has proved a useful tool to elucidate patterns of historical vegetation change. Linares-
Palomino et al. (2003) performed a detailed phytogeographical analysis of SDTF areas of Pacific
South America, i.e. the western section of the RPDS Arc, and found three main groups with a
considerable dissimilarity among them. In the present contribution, we perform a similar analysis
of seasonally dry forest areas of the eastern section of the RPDS arc. As we dealt with both the
tropical and subtropical regions of eastern South America we incorporated seasonally dry subtrop-
ical forests into the SDTF concept.
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Floristic Relationships of Seasonally Dry Forests of Eastern
153
The geographical range of the SDTF areas analysed in the present study is large enough to
include four vast vegetation domains: Atlantic Forest, caatinga, cerrado and chaco (Figure 7.1).
The Atlantic forest domain stretches for >3300 km along the eastern Brazilian coast between the
latitudes of 6
°
S and 30
°
S and makes up the second largest tropical moist forest area of South
America, exceeded only by the vast Amazonian domain. The two forest domains are separated by
the so-called diagonal of open formations, a corridor of seasonally dry formations that includes
another three domains: the caatingas (mostly tropical thorny woodlands), cerrado (mostly woody
savannas), and the chaco (mostly subtropical thorny woodlands). Each domain contains its particular
SDTF formations. The now widely accepted concept of Atlantic forests (sensu lato) attaches
seasonal forests, the Atlantic SDTF, to the coastal rain forests, formerly considered as the true
(sensu stricto) Atlantic forests (Oliveira-Filho and Fontes, 2000; Galindo-Leal and Câmara, 2003).
Caatingas and carrascos (tropical deciduous scrubs) are both SDTF and make up the predominant
vegetation cover of the caatinga domain. SDTF formations are also an important component of the
cerrado domain where they occur as forest patches on more fertile soils and on the freely drained
slopes of gallery forests (Oliveira-Filho and Ratter, 1995, 2002). In the chaco domain, SDTF occur
on peripheral areas and in some internal forest patches (Prado, 2000).
FIGURE 7.1
Map of eastern South America showing the distribution of the predominant vegetation formations
of the South American Atlantic forest domain. Caatingas, cerrados, chaco and campos are the adjacent domains
that make up the “diagonal of open formations”.
4°
8°
16°
28°
32°
12°
64°60°56°52°48°
Atlantic Ocean
44°40° 36°32°
Tropical rain forest
Tropical seasonal semideciduous forest
Tropical seasonal deciduous forest
Subtropical rain forest
Subtropical araucaria rain forest
Subtropical seasonal semideciduous forest
Subtropical seasonal deciduous forest
Atlantic forest formations
68°
20°
24°
Chaco
Cerrados
Cerrados
Cerrados
Caatingas
Caatingas
Chaco
Campos
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Neotropical Savannas and Dry Forests: Diversity, Biogeography, and Conservation
Atlantic SDTF occur as seasonal (semideciduous and deciduous) forests all along the contact
zone between rain forests and the diagonal of open formations, comprising three different scenarios
(see Figure 7.1 for distribution and Table 7.1 for nomenclature). (a) In north-eastern Brazil, SDTF
form a narrow belt (<50 km) in the sharp transition between coastal rain forests and the semiarid
caatingas, but also occur as hinterland montane forest enclaves, the brejos (Rodal, 2002; Rodal and
Nascimento, 2002). (b) The transition between coastal rain forests and cerrados in south-eastern
Brazil involves a much larger extent of SDTF that becomes increasingly wider towards the south
to reach eastern Paraguay and north-eastern Argentina. They also form complex mosaics with
cerrado vegetation to the west so that if the SDTF component of the cerrado domain is seen as an
extension of the Atlantic SDTF, as proposed by Oliveira-Filho and Ratter (1995), a concept of
Atlantic forests sensu latissimo must be created. (c) In the southern subtropical realm, large extents
of hinterland araucaria rain forests are attached to the coastal subtropical rain forests, and SDTF
appear in the west and south as transitions to both chaco forests
and southern campos, or pampa
prairies (Spichiger et al., 1995; Quadros and Pillar, 2002).
In the present study we sought patterns of floristic differentiation among SDTF areas of eastern
tropical and subtropical South America that could be associated with geographical and climatic
variables, and assessed the floristic links of seasonally dry forest formations of different regions,
Amazonian forests and cerrados. We addressed the following questions: (a) How strongly differ-
entiated are Atlantic seasonal and rain forests in different sections of their geographical range?
(b) To what extent is the tree species composition of Atlantic seasonal forests transitional between
those of rain forests and open formations, such as caatingas and cerrados? (c) Is the Atlantic rain
forest flora closer to that of Amazonian rain forests or to that of the Atlantic seasonal forest?
(d) How strong are the floristic links among caatingas, the seasonal forests of the Atlantic and cerrado
domains, and the chaco forests? (e) Does SDTF flora change its composition in response to climatic
variations? and (f) How are the above questions answered at the species, genus and family levels?
7.2 METHODS
7.2.1 P
REPARATION
AND
R
EVISION
OF
THE
D
ATABASES
We selected from the literature a total of 659 papers containing floristic checklists produced by
surveys of the tree component of 532 areas of eastern tropical and subtropical South America. The
geographical range included the Atlantic forest, caatinga, cerrado and chaco domains (Figure 7.1).
Vegetation formations included seasonally dry tropical forests (SDTF), which are the focus of the
present study, as well as tropical and subtropical rain forests, and subtropical araucaria rain forests
(see Table 7.1). SDTF formations (Figures 7.2 and 7.3) are a broad category that contains tropical
and subtropical seasonal forests (both deciduous and semideciduous) as well as caatingas and
carrascos. Mesotrophic cerradões were treated as SDTF-cerrado transition.
Individual areas were defined arbitrarily within a maximum range of 20 km width and 400 m
elevation, and thus included sections of large continuous forest tracts (e.g. Tiradentes), assemblages
of forest fragments (e.g. Santa Maria) and nearby areas at different altitudes (e.g. Lençóis and
Palmeiras). We obtained the following geographical information for each area: latitude and longi-
tude at the centre of the area, median altitude, and shortest distance from the ocean. We also
obtained the annual and monthly means for the temperature and rainfall of each area or the nearest
meteorological stations. When the source of the checklist did not provide the climatic records, they were
obtained from DNMet (1992) and from governmental websites (http://masrv54.agricultura.gov.br/rna;
http://www.inmet.gov.br/climatologia). Some areas required interpolation and/or standard correc-
tion for altitude (Thornthwaite, 1948).
We entered the information from the 532 areas on to spreadsheets using Microsoft Excel 2002
in order to produce two databases. The first consisted of basic information about each area including
locality, forest classification (see below), geographical and climatic variables, and literature sources.
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Floristic Relationships of Seasonally Dry Forests of Eastern
155
TABLE 7.1
Nomenclature Used in the Present Chapter for Vegetation Classification of Eastern Tropical and Subtropical South America.
Vegetation Formations Altitudinal Belt Regions Main Formations
Tropical rain forests
lowland low altitude North-east,east
and south-east
submontane
lower montane high altitude
upper montane
Subtropical rain forests
lowland low altitude
South
Atlantic rain forests
(Atlantic forests sensu stricto)
Atlantic forests
(Atlantic forests
sensu latissimo
;
sensu lato
excludes
CW and SW)
Rain forests
submontane
lower montane
upper montane high altitude
Subtropical araucaria rain forests lower montane high altitude South and
south-east
upper montane
Tropical seasonal semideciduous forests
lowland low altitude North-east, east,
south-east and
central-west Atlantic seasonal forests
(NE/E/SW) and Central-
western seasonal forests (CW)
SDTF –
Seasonally dry
tropical forests
submontane
lower montane high altitude
upper montane
Tropical seasonal deciduous forests
lowland low altitude North-east, east,
south-east and
central-west
submontane
lower montane high altitude
Subtropical seasonal semideciduous forests
lowland low altitude South Atlantic seasonal forests (S)
submontane
lower montane high altitude
Subtropical seasonal deciduous forests
lowland low altitude South and
south-west
Atlantic seasonal forests (S) and
Peripheral Chaco seasonal
forests (SW)
submontane
lower montane high altitude
upper montane
Caatingas (tropical thorny woodlands) lowland and
submontane
low altitude North-east and
east
Carrascos (tropical deciduous scrubs)
Cerrados (
sensu lato:
open savannas to
forests, or cerradões)
lowland to
lower montane
low-high
altitude
Central-west
Chaco (subtropical thorny woodland) lowland to lower
montane
low altitude South-west
Southern campos or pampa prairies lowland low altitude South
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Neotropical Savannas and Dry Forests: Diversity, Biogeography, and Conservation
The second database was a matrix of tree species presence in the 532 areas plus three additional
checklists that we included to compare them to Amazonian rain forests, cerrados (s.l
.
) and chaco
forests. The first of these combined the flora of Reserva Ducke (Ribeiro et al., 1999) with the
22 checklists of Amazonian rain forests compiled by Oliveira-Filho and Ratter (1994) and contained
2190 tree species. The second contained 528 species present in 98 areas of cerrado (Ratter et al.,
1996) and the third 183 chaco species listed by Prado (1991), Lewis et al. (1994) and Spichiger
et al. (1995).
Before reaching its final form, the information contained in the species database underwent a
detailed revision to check all species names cited in the checklists for growth form, synonymy and
geographical distribution. Only species capable of growing to trees or treelets, i.e. producing a free-
standing woody stem >3 m in stature, were maintained in the database. The task involved consultation
of 387 published revisions of families and genera, 32 specialists of various institutions and four
websites (http://www.cnip.org.br; http://www.ipni.org/index.html, 2003-2005; http://www.mobot.org/
W3T/Search/vast.html; http://sciweb.nybg.org/science2/hcol/sebc/index.asp). When these sources
FIGURE 7.2
Seasonally dry tropical forests (SDTF) of eastern South America: (A) caatinga in São Raimundo
Nonato, Piauí; (B) submontane tropical seasonal deciduous forest in the Serra das Confusões, Piauí; (C–F)
submontane tropical seasonal deciduous forest in Três Marias, Minas Gerais in the dry (C and D) and wet
(E and F) seasons (Image credits: F. Filetto [A and B] and M. A. Fontes [C–F]).
AB
CD
EF
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Floristic Relationships of Seasonally Dry Forests of Eastern
157
referred to herbarium specimens unequivocally collected in any of the 532 areas, the species was
added to the database. The final database contained 6598 species, 976 genera, and 128 families. The
species classification into families followed the Angiosperm Phylogeny Group II (APG, 2003).
7.2.2 V
EGETATION
C
LASSIFICATION
We extend here the vegetation classification proposed by Oliveira-Filho and Fontes (2000) for
south-east Brazil to include a much wider geographical range as well as additional vegetation
formations (Table 7.1). This extended classification was based on exploratory multivariate analyses
of both floristic and climatic data (ongoing studies). We defined the top classification level by
combining main thermoclimate (either tropical or subtropical) and rainfall seasonality (rainy,
seasonally rainy and semi-arid) and established the limit between the two thermoclimates at the
latitudes of 25
°
30’S and 26
°
30’S for rain and seasonal forests, respectively. Areas of the chaco
domain and araucaria rain forests were all subtropical; areas of the cerrado and caatinga domains
were all tropical. We classified areas with tropical climates as rain forests, seasonal forests/cerrados
and caatingas/carrasco in which the dry season lasts for up to 30 days (rainy),
>
30
160 days
(seasonally rainy) and
>
160 days (semi-arid), respectively. In subtropical climates, we classified
the areas as chaco forests/peripheral chaco seasonal forests where the dry season lasts for
>
30 days
(semi-arid) and areas below this limit as either rain forests or seasonal forests/campos in which the
difference in mean monthly temperatures between the coolest and warmest months is up to 10
°
C
or
>
10
15
°
C, respectively (rainfall seasonality secondary).
FIGURE 7.3
Seasonally dry tropical forests (SDTF) of eastern tropical and subtropical South America:
(A) lower montane tropical semideciduous forest in Itambé do Mato Dentro, Minas Gerais; (B) submontane
tropical seasonal semideciduous forest in the Chapada dos Guimarães, Mato Grosso; (C) lowland subtropical
seasonal semideciduous forest in Praia do Tigre, Rio Grande do Sul; (D) lowland subtropical seasonal
deciduous forest in Cachoeira do Sul, Rio Grande do Sul (Image credits: A.T. Oliveira-Filho [A, B], J. A.
Jarenkow [D] and J. C. Budke [E]).
CD
AB
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Neotropical Savannas and Dry Forests: Diversity, Biogeography, and Conservation
Seasonal deciduous and semideciduous forests of both tropical and subtropical climates are
commonly distinguished by the amount of leaffall during the slow-growth season (Veloso et al.,
1991). Except for the degree of deciduousness, it is often difficult to tell them apart, particularly
in central Brazil where they commonly form continua determined by local variations of soil moisture
and fertility (Oliveira-Filho and Ratter, 2002). Therefore, in most cases we opted to trust the authors’
experience to classify seasonal forests as either deciduous or semideciduous. Savannas, i.e
.
cerrado
(sensu lato) and campo are also a very important component of the vegetation in seasonal climates
but we included only the mesotrophic cerradão (plural, cerradões) in the analyses because of its
forest-like physiognomy. Again, the authors’ experience was trusted to distinguish this vegetation
from seasonal forests. Another distinction was made between subtropical araucaria rain forests and
other rain forests, based on their geographical location in the inner highlands and the conspicuous
presence of emergent trees of
Araucaria angustifolia
(Bert.) O.Kuntze. Areas of caatinga and
carrasco were distinguished by elevation and substrate, the former occurring on dissected lowlands
and the latter on sandy plateaus (see Queiroz, Chapter 6).
We also used the exploratory multivariate analyses of floristic data to classify the above
vegetation formations according to altitude and geographical region. We defined elevation ranges
as follows. For latitudes <16ºS: lowland, <400 m; submontane, 400
<
800 m; lower montane,
800
<
1200 m; upper montane, >1200 m. For latitudes between 16
°
and
<
23
°
30’S: lowland,
<
300 m; submontane, 300
− <
700 m; lower montane, 700
<
1100 m; upper montane,
>
1100 m.
For latitudes between 23º30’ and <32ºS, lowland, <200 m; submontane, 200 – <600 m; lower
montane, 600
<
1000 m; upper montane, >1000 m. The geographical regions recognized were
north-east, east, south-east, south, central-west and south-west. The resulting classification
categories are given in Table 7.1 and the geographical distribution of the 532 areas are shown
in Figure 7.4. Limited space does not allow us to list the areas, neither to provide here their
description and source references. We intend to make this information available in a forthcoming
publication.
7.2.3 M
ULTIVARIATE
A
NALYSES
We used detrended correspondence analysis, DCA (Hill and Gauch, 1980), processed by the
program PC-ORD 4.0 (McCune and Mefford, 1999) to seek main species distribution gradients
across 341 SDTF areas. We removed rain forests because the patterns within this vegetation
type were not the focus of the present work. An additional DCA was performed with the 243
areas of tropical seasonal forests (subtropical and semiarid formations excluded) to seek more
detailed patterns within the group. Two other DCA were performed separately for the areas of
caatinga and subtropical seasonal forests. We chose DCA coupled to a posteriori interpretation
of ordination results because we aimed at patterns dictated solely by the species without the
interference of environmental variables, as occurs with joint analyses such as CCA (Kent and
Coker, 1992). We used two interpretation tools: the previous vegetation classification of the
areas and 13 geographical and climatic (hereafter geo-climatic) variables. They were both
plotted (a posteriori) on the DCA diagrams as symbols and arrows, respectively. The geo-
climatic variables were latitude, longitude, median altitude, distance from the ocean, mean
annual temperature, mean monthly temperatures in the warmest and coolest months, mean
temperature range obtained from the difference between the two previous variables, mean
annual rainfall, mean monthly rainfall of the dry (June–August) and rainy (December–February)
seasons, rainfall distribution ratio obtained from the proportion between the two previous
variables and mean duration of the dry season obtained from the number of days of water
shortage given by Walter diagrams (Walter, 1985). We also obtained the Pearson correlation
coefficients between the geo-climatic variables and the ordination scores of the areas in each
DCA axis.
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Floristic Relationships of Seasonally Dry Forests of Eastern
159
7.2.4 C
ONDENSED
F
LORISTIC
D
ATA
Because both previous and present multivariate analyses demonstrated that the vegetation classifi-
cation system adopted was highly consistent, we eventually condensed the floristic information
contained in the database by lumping together the species records within main vegetation formations
(Table 7.1). Atlantic rain forests as well as the cerrado, chaco and Amazonian rain forest checklists
were incorporated here. We also merged lowland and submontane categories as low altitude and
lower and upper montane categories as high altitude. The resulting lumped matrix of binary data
of species presence in the main vegetation formations was used to produce two additional matrices,
for genera and families. The generic and familial matrices were both quantitative, as they consisted
of the number of species per genus or family, respectively, in each main vegetation formation. We
performed cluster analyses of the condensed matrices using the program PC-ORD 4.0. Cluster
analyses used Jaccard’s floristic similarity for species and relative squared Euclidian distances for
genera and families (number of species as abundance data); the linkage method was group average
(Kent and Coker, 1992). We also used the condensed data to perform a direct quantitative assessment
of the floristic links between the vegetation formations by plotting the number of shared and
exclusive species in Venn diagrams. The most frequent species, and the richest genera and families
of main vegetation formations were extracted from the matrices.
FIGURE 7.4
Geographical coordinates diagram showing the location and vegetation classification of the 532
areas used in the analyses and the six geographical regions. Forest areas situated in the north-east, east, south-
east and central-west regions are classified as tropical and those in the south and south-west are subtropical.
Longitude (W)X
Latitude (S)X
2
4
6
8
10
12
14
16
18
20
22
24
26
28
30
32
34
323436384042444648505254565860626466
Northeast
East
Central-West
Southwest Southeast
South
Lowland rain forest
Submontane rain forest
Lower montane rain forest
Upper montane rain forest
Lower montane araucaria rain forest
Upper montane araucaria rain forest
Lowland seasonal semideciduous forest
Submontane seasonal semideciduous forest
Lower montane seasonal semideciduous forest
Upper montane seasonal semideciduous forest
Lowland seasonal deciduous forest
Submontane seasonal deciduous forest
Lower montane seasonal deciduous forest
Upper montane seasonal deciduous forest
Caatinga
Carrasco
Mesotrophic cerradão
Vegetation classication
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Neotropical Savannas and Dry Forests: Diversity, Biogeography, and Conservation
7.3 RESULTS
7.3.1 M
ULTIVARIATE
A
NALYSES
The ordination diagrams yielded by DCA are shown in Figures 7.5 and 7.6 for the two assemblages
of vegetation areas, seasonally dry tropical forests (SDTF) and tropical seasonal forests. Their
eigenvalues are first DCA, 0.688 (axis 1) and 0.394 (axis 2); second DCA, 0.400 (axis 1) and 0.475
(axis 2). According to ter Braak (1995), these eigenvalues are relatively high (>0.3), indicating
considerable species turnover along the gradients summarized in the first two axes. In addition,
most DCA axes produce a number of high values of Pearson correlation coefficients between geo-
climatic variables and ordination scores (Table 7.2) giving consistency to the interpretation of the
emerging patterns.
The first ordination axis in the DCA for SDTF is chiefly correlated with latitude, minimum
monthly temperature, duration of the dry season, annual temperature and annual rainfall (Table 7.2).
This indicates that the data structure summarized by the first axis primarily reflects a geographical
gradient based on latitude which corresponds to a major climatic gradient towards the south charac-
terized by decreasing temperatures and duration of the dry season and increasing total rainfall.
Longitude and distance to the ocean are more strongly correlated with the second DCA axis but no
climatic variable accompanies this gradient. The areas of caatinga and carrasco are found at the right-
side of the diagram associated with latitudes near the Equator, longer dry seasons and higher temper-
atures (Figure 7.5). No distinction is made between north-east and east caatingas but the three carrasco
areas are displaced to the top on the second axis together with three areas of caatinga (Serra da
Capivara, São José do Piauí and Ibiraba) which differ from other caatingas in their sandy substrate,
FIGURE 7.5
Diagram yielded by detrended correspondence analysis (DCA) showing the ordination of 341
areas of seasonally dry tropical forests (SDTF) of eastern South America on the first two DCA axes, based
on the presence of 3018 tree species. The areas are classified according to main vegetation formation and
geographical region. The centred straight-lines show the correlation between axes and geoclimatic variables
(only those with r > 0.3 with at least one axis are shown): T
=
temperature, Mth
=
monthly, Min
=
minimum,
R
=
rainfall, D
=
distance.
Longitude
D Ocean
T annual
T mth min
T range
R annual
Dry season
0
0
40 80
20
40
60
80
Axis 1
Axis 2
Latitude
Tropical semideciduous forests:
NE ESE CW
Tropical deciduous forests:
NE CW
Subtropical seasonal Forests:
SSW
Caatingas:
NE E
Carrascos:
NE
Mesotrophic cerradão
CW
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Floristic Relationships of Seasonally Dry Forests of Eastern
161
as do the carrascos. Towards the left-side of the diagram, caatingas and carrascos are followed by
areas of tropical seasonal forests of the north-east and central-west, discriminated at the bottom
and top halves of the diagram, respectively. For the north-east areas, deciduous forests come first
followed by semideciduous forests. For the central-west, however, deciduous and semideciduous
forests are not distinguished from each other and only mesotrophic cerradões form a consistent
clump. Along the latitudinal sequence of the first DCA axis, north-east tropical seasonal forests
are followed by those of the east and then south-east regions. This sequence ends at the left-side
of the diagram where the areas of subtropical seasonal forests lie that correspond to the extremes
of higher latitudes, lower temperatures and shorter dry seasons are situated. In addition, the second
axis discriminates, at the top, the four south-west areas of peripheral chaco seasonal forests.
The DCA performed for tropical seasonal forests shows additional patterns linked to altitude that
do not arise when other seasonally dry forests are included. The first axis is primarily correlated with
distance to the ocean, longitude, temperatures (annual, minimum and maximum) and altitude, while
the second axis is more strongly correlated with latitude, monthly rainfall in DJF, rainfall distribution
ratio and monthly temperature range (Table 7.2). This suggests that the first axis primarily reflects a
geographical gradient based on penetration into the continental interior together with decreasing
altitude, both of which also correspond to a climatic gradient characterized mainly by increasing
temperatures. To a great extent, the second axis repeats patterns already shown by the first axis of
the previous DCA, so that all north-east seasonal forests are strongly discriminated at the top of the
diagram (Figure 7.6). As opposed to the others, north-east areas show stronger correlations with
decreasing latitude, rainfall in DJF and temperature range and with increasing rainfall distribution
ratio. The first ordination axis, however, discriminates north-east areas of deciduous and semidecid-
uous forests to the left- and right-sides of the diagram, respectively, with the single exception of the
oceanic island of Fernando de Noronha, located at the top right corner. At the bottom half of the
TABLE 7.2
Detrended Correspondence Analysis (DCA)
Vegetation Physiognomies and DCA Axes
Geoclimatic variables
SDTF and SDSF
(N
=
341 areas)
Tropical Seasonal Forests
(N
=
243 areas)
Axis 1 Axis 2 Axis 1 Axis 2
Latitude –0.88 0.19 0.08 –0.91
Longitude –0.52 0.72 –0.65 –0.55
Altitude
0.14 –0.26 0.58 –0.13
Distance to the ocean 0.12 0.66 –0.73 –0.07
Annual temperature 0.77 0.15 –0.63 0.49
Minimum monthly temperature 0.81 0.08 –0.54 0.61
Maximum monthly temperature 0.48 0.26 –0.59 0.23
Monthly temperature range –0.66 0.14 0.15 –0.62
Annual rainfall –0.72 0.10 0.08 –0.37
Monthly rainfall in JJA
0.27 –0.17 0.05 0.39
Monthly rainfall in DJF –0.52 0.21 0.07
0.67
Rainfall distribution ratio 0.08 –0.30 0.08 0.60
Duration of the dry season 0.79 0.06 –0.07 0.41
Pearson correlation coefficients between geo-climatic variables and the ordination
scores of N areas of seasonally dry forests of the South American Atlantic forest
domain. Coefficients are given for the first two axes of DCAs performed for species
presence in three different sets of areas. Correlations >0.5 are in bold.
2987_C007.fm Page 161 Thursday, December 1, 2005 7:03 PM
162
Neotropical Savannas and Dry Forests: Diversity, Biogeography, and Conservation
diagram the first ordination axis discriminates central-west seasonal forests from those of the south-
east and east regions in such a way that two concurrent geographical gradients were distinguished,
one related to longitude and distance from the ocean and the other with altitude, both involving
decreasing temperatures. The areas at the extreme left of the diagram are the westernmost low-altitude
seasonal forests of Mato Grosso and Bolivia, while those at the extreme right are mostly the eastern
high-altitude seasonal forests of Bahia and Minas Gerais. As in the previous DCA, central-west
deciduous and semideciduous forests are not discriminated from each other.
Additional DCAs were performed separately for the areas of caatinga and subtropical seasonal
forests, but no relevant additional patterns arose. Deciduous and semideciduous subtropical forests
of the south were not discriminated amongst themselves and only altitude showed a weak correlation
with the floristic patterns.
7.3.2 A
NALYSES
OF
C
ONDENSED
F
LORISTIC
I
NFORMATION
As they were extracted from floristic checklists for specific forest areas, the condensed information
must be regarded as a means of assessing the floristic links between the main forest formations
quantitatively and not as a register of actual figures for number of species, either total or in common.
The classification dendrograms (Figure 7.7) show different patterns for each of the three
taxonomic levels. A clear general trend arising from the species dendrogram is that regional patterns
FIGURE 7.6
Diagram yielded by detrended correspondence analysis (DCA) showing the ordination of 243
areas of tropical seasonal forests of the South America Atlantic forest domain on the first two DCA axes,
based on the presence of 2680 tree species. The areas are classified according to forest formation, geographical
region and altitudinal range. The centred straight-lines show the correlation between axes and geoclimatic
variables (only those with r > 0.3 with at least one axis are shown): T
=
temperature, Mth
=
monthly, Min
=
minimum, Max
=
maximum, D
=
distance, R
=
rainfall, DJF
=
December-January-February, D-Ratio
=
distribution ratio.
Tropical forest formations:
Semideciduous-NE
Semideciduous-E
Semideciduous-SE
Semideciduous-CW
Deciduous-CW
Deciduous-NE
Altitudinal range:
Lowland
Submontane
Lower montane
Upper montane
Latitude
Longitude
D Ocean
T annual
T mth min
T mth max
T range
RD JF
Altitude
R D-ratio
20
30
40 60 80
50
70
Axis 1
Axis 2
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Floristic Relationships of Seasonally Dry Forests of Eastern
163
FIGURE 7.7
Dendrograms produced by group averaging of Jaccard’s floristic similarity for species and relative squared
Euclidian distances for genera and families of the tree flora 23 areas of eastern Amazonian rain forests (Amz), 39 areas of
caatinga (Caa), 3 areas carrasco (Car), 376 areas of cerrado (Cerr), 11 areas of mesotrophic cerradão (Cdm), 5 areas of
chaco forests (Cha) and 479 areas of Atlantic forests
s.l
. merged into 28 main forest formations abbreviated as follows: the
first set of letters stands for either tropical (T) or subtropical (S) forests and for rain (R), araucaria rain (A), seasonal
semideciduous (S) and seasonal deciduous (D) forests; the middle letters stand for low (L) and high (H) altitude; the last
letters stand for north-east (NE), east (E), south-east (SE), central-west (CW), south (S) and south-west (SW) regions.
Scale in dendrograms expresses the remaining information after clustering.
Information remaining (%)
100 75 50 25 0
TR-L-NE
TS-L-NE
TS-H-NE
TR-H-NE
TD-L-NE
Car-NE
Caa-NE
Caa-E
TR-L-E
TR-L-SE
TR-H-SE
TS-L-E
TS-H-E
TS-L-SE
TS-H-SE
TS-L-CW
TS-H-CW
TD-H-CW
TD-L-CW
SR-L-S
SR-H-S
SA-H-S
SA-H-SE
SS-L-S
SD-L-S
Cdm-CW
Cerr-CW
SS-H-S
SD-H-S
Amz
SD-L-SW
SD-H-SW
Cha-SW
Species
TR-L-NE
TS-L-NE
Amz
TR-H-NE
TR-L-E
TS-L-E
TS-L-SE
TS-H-SE
TS-H-E
TS-L-CW
TS-H-CW
TR-L-SE
TR-H-SE
SR-L-S
SR-H-S
SA-H-S
SA-H-SE
SS-L-S
SD-L-S
SS-H-S
SD-H-S
TS-H-NE
TD-L-NE
TD-L-CW
TD-H-CW
Cdm-CW
Cerr-CW
Car-NE
Caa-NE
Caa-E
SD-L-SW
SD-H-SW
Cha-SW
Genera
TR-L-NE
TS-L-NE
Amz
TS-H-NE
TD-L-NE
TD-L-CW
TD-H-CW
Cdm-CW
SD-L-SW
SD-H-SW
Car-NE
Caa-NE
Caa-E
Cha-SW
TR-L-E
TS-L-E
TS-L-SE
TS-L-CW
TS-H-CW
TS-H-E
TS-H-SE
Cerr-CW
TR-H-NE
TR-L-SE
TR-H-SE
SR-H-S
SR-L-S
SA-H-S
SA-H-SE
SS-L-S
SD-L-S
SD-H-S
SS-H-S
Families
2987_C007.fm Page 163 Thursday, December 1, 2005 7:03 PM
164
Neotropical Savannas and Dry Forests: Diversity, Biogeography, and Conservation
are stronger than vegetation formation patterns. Four main geographical groups are discriminated.
The first contains all vegetation formations of the north-east region, including tropical rain and
seasonal forests, caatingas and carrascos. The second, and largest, group contains tropical rain and
seasonal forests of four regions (east, south-east, central-west and south) plus cerrados and cer-
radões. The third main group is composed of the Amazonian rain forests and the fourth, and most
distinct, by chaco forests and peripheral chaco seasonal forests. The north-east main group is split
into two subgroups, the first containing moister formations (tropical rain forests and seasonal
semideciduous forests) and the second drier formations (tropical seasonal deciduous forests, caat-
ingas and carrascos). The second main group is split into five subgroups, all of clear geographical
nature: (a) tropical rain and seasonal semideciduous forests of the east and south-east; (b) tropical
seasonal semideciduous and deciduous forests of the central-west; (c) subtropical low-altitude
seasonal forests and rain forests of the south, plus subtropical high-altitude araucaria rain
forests of both the south and south-east; (d) cerrados (sensu lato) and mesotrophic cerradões; and
(e) subtropical high-altitude seasonal forests of the south.
The dendrogram for genera shows different main groups and an increased role of vegetation
formation over regional patterns. The first main group contains tropical low-altitude rain forests
and semideciduous forests of the north-east and Amazonian regions. The second main group
contains two subgroups: (a) tropical seasonal semideciduous forests of the east, south-east and
central-west, plus tropical rain forests of the east, and (b) tropical rain forests of the south-east,
subtropical rain forests of the South and subtropical Araucaria rain forests of both the south and
south-east. The third, fourth and fifth main groups contain, respectively, subtropical seasonal forests
of the south, tropical seasonal deciduous forests of the north-east and central-west, and cerrados
and cerradões. The last two main groups contain caatingas, carrascos, chaco forests and peripheral
chaco seasonal forests.
The dendrogram for families goes a step further in generating groups with a strong vegetation
formation character. The semi-arid formations, namely the chaco forests, caatingas and carras-
cos, form a clump that merges, at the subsequent level, with a group that includes tropical and
subtropical seasonal deciduous forests and mesotrophic cerradões. An oddity of this group is
the presence of a side subgroup containing low-altitude rain forests and semideciduous forests
of the north-east and Amazonian regions. Tropical seasonal semideciduous forests predominate
in another main group that also includes the cerrado and tropical rain forests of the east and
north-east. The following main group contains tropical and subtropical rain forests and sub-
tropical araucaria rain forests, and the last main group contains subtropical seasonal forests of
the south.
The tree floras represented in the rain and seasonal forest checklists are similar in species
richness: 3009 and 2903 species, respectively. On the other hand, the number of rain forest
checklists, 191, is considerably smaller than that of seasonal forests, 285, therefore suggesting that
the species richness of the latter may actually be lower. In fact, the speciesarea curves of the two
vegetation formations (Figure 7.8) demonstrate that, at any number of areas, the mean cumulative
number of species is much higher in rain than in seasonal forests. The two formations also share
a high proportion of tree species, 1814 out of 4098, or 44.3%, but both also have a considerable
number of putative endemics, 1195 (29.2%) and 1089 (26.6%) for rain and seasonal forests,
respectively.
The Venn diagrams in Figure 7.9 show the relationship of the tree flora of rain and seasonal
forests in different geographical regions. The number of species of both formations is smaller in
the north-east and south and larger in the east and south-east. The non-Atlantic seasonal forests
have higher numbers of species in the central-west and lower in the southwest. Seasonal deciduous
forests of the north-east, despite sharing many species with regional rain and semideciduous forests,
have their own group of putative endemics. The proportions of species shared by Atlantic rain and
seasonal forests are very similar in all regions: 20.4% in the north-east, 21.0% in the east, 22.0%
in the south-east, and 18.8% in the south. The species proportions in rain and seasonal forests,
2987_C007.fm Page 164 Thursday, December 1, 2005 7:03 PM
Floristic Relationships of Seasonally Dry Forests of Eastern
165
however, show opposing trends from the north-east to the south, and are, respectively, 31.8% and
47.8% in the north-east, 39.2% and 39.8% in the east, 48.3% and 29.7% in the south-east, and
58.5% and 22.7% in the south. Subtropical araucaria rain forests share high proportions of species
with both rain and seasonal forests in both the south and South-east regions. However, they also
contain their group of putative endemics, particularly in the south. The central-west seasonal forests
share 76.2% of their species with Atlantic tropical rain- and seasonal forests (north-east, east and
south-east), though 60.6% are present in both formations, 15.6% in seasonal forests only, and none
in rain forests only.
The seasonal forests of all six geographical regions contain 2903 species, of which only 40 are
registered in all regions, 81 in five regions, 257 in four, 414 in three, 630 in two, and 1481 in one.
These putative endemic species are in higher proportion in the floras of the east (619; 35.0%) and
north-east (241; 32.1%), followed by the south-west (68; 27.1%), central-west (310; 24.4%), south
(66; 15.7%) and south-east (177; 14.9%). The relationships among seasonal forests of adjacent
geographical regions are shown in the left-side Venn diagrams of Figure 7.10. The three regions
of Atlantic tropical seasonal forests share a small number of species, 273 out of 2062 (13.2%), but
adjacent regions share larger proportions: 24.8% between the north-east and east and 29.0% between
the east and south-east. Subtropical seasonal forests share a high number of species with the tropical
seasonal forests of the south-east: 76.3% and 50.2% for the south and south-west, respectively. The
latter also share 59.8% of their species with the tropical seasonal forests of the central-west and
67.7% with both the South-east and central-west.
Caatingas are considerably poorer in tree species than are rain- and seasonal forests (Figure 7.8).
The relationships between caatingas and adjacent vegetation formations are shown in the right-side
Venn diagrams of Figure 7.10. A high proportion of their 466 species is shared with adjacent
seasonal forests, 61.2%, but this also leaves 38.8% of putative endemics. The proportion of shared
species is higher with the north-east seasonal forests (49.4%) than with the central-west (39.3%).
The proportion shared with cerrados is much smaller, 17.6%, and most of this is also shared with
central-west seasonal forests. The number of species shared with chaco forests is very small, only
five. In fact, both semi-arid formations, caatingas and chaco forests, share more species with the
central-west seasonal forests than between themselves. The geographical range of the tree flora of
FIGURE 7.8
Mean cumulative number of species in areas of rain forests and seasonally dry tropical forests
of eastern South America with increasing number of areas.
0
500
1000
1500
2000
2500
3000
3500
050 100 150 200 250 300
Number of areas
Number of species
Rain forests Seasonal
forests
Caatingas
2987_C007.fm Page 165 Thursday, December 1, 2005 7:03 PM
166
Neotropical Savannas and Dry Forests: Diversity, Biogeography, and Conservation
those two formations is illustrated in the two flow diagrams of Figure 7.11. Lowland seasonal
deciduous forests and carrascos of the north-east are excluded because they have a strong floristic
identity with the caatingas and are not in the route between the caatinga and chaco domains.
Seasonal forests of the east, South-east and south are merged into the category Austro-Atlantic
seasonal forests. The proportion of endemic to non-endemic species is lower for caatingas than is
for the chaco, 46.1:53.9% and 66.1:33.9%, respectively. The five species that complete the crossing
FIGURE 7.9
Venn diagrams extracted from the checklists showing the number of tree species shared by rain-
and seasonal forests in different geographical regions of eastern South America.
Rain and seasonal forests-Northeast:
82
12
179 290228
65
Semideciduous (665)
Deciduous
(236) 77
Rain (501)
Rain and seasonal forests-East:
834 849954
Semideciduous (1803)
Rain (1778)
Rain and seasonal forests-South:
Deciduous +
semideciduous (431)
458
73
224
Rain (890)
Araucaria rain (577)
165
57
77
131
Rain and seasonal forests-Southeast:
156
23
971 298676
16
Semideciduous (1146)
Araucaria rain (214)
19
Rain (1826)
Tropical rain and seasonal forests:
911
1120
531
771
198
Semideciduous
NE/E/SE (2411)
Deciduous +
semideciduous CW
(1272)
303
Rain NE/E/SE
(2802)
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Floristic Relationships of Seasonally Dry Forests of Eastern
167
between the two formations are Celtis pubescens (Jacquin) Sargent, Ximenia americana L.,
Sideroxylon obtusifolium (Roem. and Schultz) T.D.Penn., Parkinsonia aculeata L. and Aporosella
chacoensis (Morong) Pax and Hoffmg. All five are also present in the peripheral chaco seasonal
forests (south-west), but the former three also cross both the central-west and Austro-Atlantic
FIGURE 7.10 Venn diagrams extracted from the checklists showing the number of tree species shared by
seasonal forests in different geographical regions of eastern South America (left side), and by seasonal forests,
caatingas, cerrados and chaco forests (right side).
Southeast (1146)
236
4
792
280
273
169
Northeast (754)
308
East (1823)
Seasonal forests NE × E × SE:
Seasonal forests SE × S × SW:
93
33
784
87
236
15
South (431)
Southwest (251)
110
Southeast (1146)
Seasonal forests SE × CW × SW:
Central-West (1272)Southeast (1146)
856
295
229
Seasonal-CW (1272)
Seasonal forests and caatingas:
Seasonal-NE (754)
Caatingas (466)
132
55
98
181
Seasonal forests, caatingas and chaco:
Seasonal forests, caatingas and cerrados:
775
174
70
267
12
310
117
Seasonal-CW (1272) Cerrados (1272)
Caatingas (466)
1058
151
4
278
1
27
183
Chaco (183)
Caatingas (466)
453
106
Southwest (251) 81
20 44
555
Seasonal-CW (1272)
567
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168 Neotropical Savannas and Dry Forests: Diversity, Biogeography, and Conservation
seasonal forests while P. aculeata skips these two formations and A. chacoensis is also present in
central-west seasonal forests. Despite this, a much larger number of caatinga species, 55, reach the
peripheral chaco seasonal forests without entering the chaco itself. Similarly, 42 chaco species
reach the Austro-Atlantic and/or central-west seasonal forests without entering the caatingas.
The most species-rich genera and families of each main vegetation formation are given in
Tables 7.3 to 6, and the most frequent species of the same formations are provided in the Appendix.
Some genera rank high in most main Atlantic seasonal forest formations, e.g. Eugenia, Myrcia,
Ocotea and Miconia (Table 7.3). Some trends can be observed with increasing altitude: the relative
importance decreases for some genera such as Eugenia (except in the north-east), Inga and Ficus,
and increases for others such as Miconia and Tibouchina (east and south-east), Ilex and Solanum.
Subtropical seasonal forests of the south are similar to tropical seasonal forests in their generic
profile, but the south-west subtropical seasonal forests, chaco forests and the caatingas have very
particular sets of species-richest genera (Table 7.4). Among the families, Fabaceae is top in all
vegetation formations except the subtropical seasonal forests where it switches places with
Myrtaceae (Tables 7.5 and 7.6). In all other formations, Myrtaceae ranks second, except in the
south-west subtropical seasonal forests (3rd), chaco forests (16th), and caatingas (5th). Other
families ranking high among tropical and subtropical seasonal forests (except in the south-west)
are Rubiaceae, Melastomataceae and Lauraceae. Families showing increasing importance at higher
FIGURE 7.11 Geographical extension of caatinga species towards seasonal forests and chaco
forests (top), and of chaco forest species towards seasonal forests and the caatingas (bottom).
Encircled figures are the number of species shared by the vegetation formations connected by
arrows.
Central-
Western
Seasonal
forests
Peripheral
Chaco
Seasonal
forests
Austro-
Atlantic
Seasonal
forests
1
54
8
Central-
Western
Seasonal
forests
9
Chaco
forests
N = 183
Endemics:
121
3
14
12
Caatingas
Central-
Western
Seasonal
forests
Austro-
Atlantic
Seasonal
forests
Chaco
forests
Central-
Western
Seasonal
forests
Central-
Western
Seasonal
forests
Austro-
Atlantic
Seasonal
forests
Austro-
Atlantic
Seasonal
forests
1
143
6
Caatingas
N = 466
Peripheral
Chaco
Seasonal
forests
1
1
3
Endemics:
215
46
3
63
44
1
2987_C007.fm Page 168 Thursday, December 1, 2005 7:03 PM
Floristic Relationships of Seasonally Dry Forests of Eastern 169
TABLE 7.3
Genera With the Highest Number of Species (S) in the Tree Flora of Tropical Seasonal Forests of the South American Atlantic Forest
Domain Classified into Four Geographical Regions and Two Altitudinal Ranges (N = Number of Areas)
North-East Region East Region South-East Region Central-West Region
Low Altitude S High Altitude S Low altitude S High altitude S Low Altitude S High Altitude S Low Altitude S High Altitude S
(N = 13) 542 (N = 11) 420 ( N = 29) 1317 ( N = 26) 1193 (N = 47) 848 (N = 35) 911 ( N = 74) 1129 ( N = 23) 624
Eugenia 16 Eugenia 17 Eugenia 49 Miconia 43 Eugenia 37 Miconia 36 Miconia 31 Miconia 28
Myrcia 12 Erythroxylum 15 Ocotea 32 Myrcia 37 Ocotea 27 Ocotea 26 Eugenia 26 Myrcia 18
Cordia 9Myrcia 12 Miconia 32 Ocotea 29 Miconia 24 Eugenia 26 Myrcia 23 Machaerium 13
Erythroxylum 9Cordia 9Myrcia 30 Eugenia 29 Myrcia 18 Myrcia 18 Machaerium 20 Eugenia 12
Senna 9Senna 9Machaerium 23 Tibouchina 18 Ficus 16 Solanum 16 Ficus 18 Aspidosperma 11
Inga 9Ocotea 8Inga 22 Machaerium 17 Solanum 15 Ficus 15 Aspidosperma 16 Ocotea 11
Miconia 9Croton 7Ficus 22 Ilex 16 Machaerium 13 Machaerium 14 Inga 15 Ficus 11
Psidium 9Ouratea 7Tabebuia 15 Erythroxylum 16 Nectandra 12 Piper 14 Bauhinia 14 Nectandra 10
Aspidosperma 8Zanthoxylum 7Pouteria 15 Solanum 16 Erythroxylum 11 Ilex 12 Ocotea 14 Casearia 9
Ficus 8Maytenus 6Guatteria 14 Inga 15 Inga 11 Nectandra 11 Erythroxylum 13 Symplocos 9
Casearia 8Acacia 6Psychotria 14 Maytenus 14 Tabebuia 10 Tibouchina 11 Byrsonima 12 Ilex 8
Pouteria 8Byrsonima 6Solanum 14 Psychotria 13 Piper 10 Erythroxylum 10 Casearia 12 Inga 8
Tabebuia 7Helicteres 6Cordia 13 Casearia 13 Senna 9Inga 10 Nectandra 11 Maytenus 7
Bauhinia 7Casearia 6Maytenus 13 Guatteria 12 Psychotria 9Casearia 10 Trichilia 11 Campomanesia 7
Ocotea 7Solanum 6Casearia 13 Psidium 12 Zanthoxylum 9Leandra 9Psidium 11 Vochysia 7
Mimosa 6Aspidosperma 5Aspidosperma 12 Cinnamomum 11 Maytenus 8Trichilia 9Cordia 10 Byrsonima 6
Guapira 6Cyathea 5Trichilia 12 Nectandra 11 Croton 8Myrsine 9Maytenus 10 Trichilia 6
Coccoloba 6Inga 5Rudgea 12 Ouratea 11 Bauhinia 8Psychotria 9Senna 10 Psidium 6
Zanthoxylum 6Psidium 5Erythroxylum 11 Cordia 10 Myrsine 8Aspidosperma 8Acacia 10 Guapira 6
Tabernaemontana 5Psychotria 5Campomanesia 11 Croton 10 Pouteria 8Tabebuia 8Zanthoxylum 10 Tabebuia 5
Licania 5Capparis 4Psidium 11 Byrsonima 10 Cestrum 8Cordia 8Tabebuia 8Licania 5
Clusia 5Pilosocereus 4Croton 10 Ficus 10 Styrax 8Croton 8Dalbergia 8Dalbergia 5
Croton 5Clusia 4Swartzia 10 Campomanesia 10 Aspidosperma 7Mollinedia 8Cupania 8Alibertia 5
Copaifera 5Bauhinia 4Ilex 9Cupania 10 Mollinedia 7Gomidesia 8Vochysia 8Styrax 5
Swartzia 5Caesalpinia 4Nectandra 9Vochysia 10 Casearia 7Symplocos 8Capparis 7Calyptranthes 5
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170 Neotropical Savannas and Dry Forests: Diversity, Biogeography, and Conservation
altitudes are Asteraceae, Melastomataceae (except in the north-east) and Lauraceae (though
unchanged in the east and south-east). Euphorbiaceae are particularly important in most formations
but rank higher in the north-east and central-west low-latitude seasonal forests, as well as in the
caatingas and chaco forests, which are also distinguished by the high ranking of Cactaceae.
7.4 DISCUSSION
An overall pattern emerging from the floristic analyses of the vegetation formations of eastern South
America was the strong influence of distance on tree species distribution. This influence only receded
and allowed clear climate-related patterns to be discerned when either the geographical range con-
sidered was restricted or data were treated at generic and familial levels. Likewise, geographically
restricted analyses of Atlantic forest sections, such as those performed for south-east Brazil by
Oliveira-Filho and Fontes (2000) and north-east Brazil by Ferraz et al. (2004), could clearly detect
species patterns primarily related to the climate. However, analyses performed for wider geographical
ranges, such as the Amazon, could best detect patterns related to climate and vegetation formations when
dealing with genera and families rather than species (ter Steege et al., 2000; Oliveira and Nelson, 2001).
TABLE 7.4
Genera with the Highest Number of Species (S) in the Tree Flora of Subtropical
Seasonal Forests of the South American Atlantic Forest Domain, Chaco Forests,
and Caatingas (N = Number of Areas)
Subtropical Seasonal Forests
South Region S Southwest Rregion S Chaco Forests S Caatingas S
(N = 37) 542 (N = 5) 420 (N = 39) 1193 (N = 6) 1317
Eugenia 18 Acacia 9 Prosopis 19 Croton 14
Ocotea 11 Schinus 7 Acacia 9 Mimosa 13
Myrsine 8 Aspidosperma 6 Lycium 8 Senna 11
Tabebuia 7 Tabebuia 5 Echinopsis 7 Erythroxylum 10
Erythroxylum 7 Zanthoxylum 5 Opuntia 7 Bauhinia 9
Ficus 7 Chloroleucon 4 Jatropha 7 Manihot 8
Myrcia 7 Ficus 4 Senna 6 Eugenia 8
Solanum 7 Eugenia 4 Capparis 5 Aspidosperma 7
Ilex 5 Schinopsis 3 Bougainvillea 5 Cordia 7
Maytenus 5 Tecoma 3 Bulnesia 5 Pilosocereus 7
Sebastiania 5 Maytenus 3 Schinopsis 4 Acacia 7
Machaerium 5 Bauhinia 3 Cereus 4 Helicteres 7
Trichilia 5 Mimosa 3 Harrisia 4 Tabebuia 6
Myrciaria 5 Ceiba 3 Maytenus 4 Maytenus 6
Psychotria 5 Luehea 3 Caesalpinia 4 Caesalpinia 6
Zanthoxylum 5 Cedrela 3 Mimosa 4 Psidium 6
Cestrum 5 Trichilia 3 Aloysia 4 Zanthoxylum 5
Schinus 4 Myrsine 3 Myrcianthes 3 Capparis 4
Cordia 4 Myracrodruon 2 Ruprechtia 3 Pereskia 4
Lonchocarpus 4 Ilex 2 Condalia 3 Cnidoscolus 4
Inga 4 Ruprechtia 2 Zanthoxylum 3 Hymenaea 4
Nectandra 4 Prosopis 2 Quiabentia 2 Chloroleucon 4
Miconia 4 Ziziphus 2 Ceiba 2 Guapira 4
Gomidesia 4 Capparis 2 Aspidosperma 2 Ruprechtia 3
Symplocos 4 Carica 2 Berberis 2 Facheiroa 3
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Floristic Relationships of Seasonally Dry Forests of Eastern 171
TABLE 7.5
Families with the Highest Number of Species (S) in the Tree Flora of Tropical Seasonal Forests of the South American Atlantic
Forest Domain Classified into Four Geographical Regions and Two Altitudinal Ranges (Suffix ‘-aceae’ Omitted from All Families;
N = Number of Areas)
North-East Region East region South-east region Central-west region
Low Altitude S High Altitude S Low Altitude S High Altitude S Low Altitude S High Altitude S Low altitude S High altitude S
(N = 13) 542 (N = 11) 420 (N = 29) 1317 (N = 26) 1193 (N = 47) 848 (N = 35) 911 (N = 74) 1129 (N = 23) 624
Fab 113 Fab 81 Fab 192 Fab 151 Fab 111 Fab 108 Fab 210 Fab 93
Myrt 48 Myrt 44 Myrt 140 Myrt 137 Myrt 101 Myrt 96 Myrt 95 Myrt 62
Rubi 28 Rubi 22 Rubi 83 Melastomat 73 Rubi 55 Melastomat 59 Rubi 58 Rubi 37
Euphorbi 19 Euphorbi 17 Laur 68 Laur 65 Laur 49 Laur 52 Euphorbi 46 Melastomat 34
Apocyn 18 Erythroxyl 15 Melastomat 48 Rubi 58 Melastomat 34 Rubi 44 Melastomat 39 Laur 29
Malv 17 Rut 12 Annon 44 Aster 51 Euphorbi 31 Aster 33 Malv 36 Annon 20
Sapot 16 Malv 11 Euphorbi 44 Euphorbi 38 Solan 29 Euphorbi 32 Annon 32 Malv 17
Annon 15 Laur 10 Mor 35 Annon 32 Rut 24 Solan 27 Laur 32 Mor 16
Mor 14 Solan 10 Sapot 31 Solan 27 Mor 22 Mor 21 Mor 27 Vochysi 15
Clusi 13 Boragin 9 Solan 29 Clusi 24 Annon 18 Malv 20 Sapind 27 Salic 14
Bignoni 11 Malpighi 9 Aster 27 Sapind 24 Bignoni 17 Annon 19 Rut 26 Apocyn 13
Chrysobalan 11 Anacardi 8 Bignoni 27 Malv 21 Malv 16 Rut 19 Apocyn 25 Arec 13
Salic 11 Cact 8 Malv 26 Rut 20 Salic 16 Salic 17 Salic 21 Bignoni 12
Boragin 10 Clusi 8 Sapind 25 Apocyn 19 Sapind 16 Bignoni 15 Nyctagin 18 Celastr 12
Melastomat 10 Salic 8 Arec 24 Chrysobalan 19 Aster 15 Vochysi 15 Meli 18 Clusi 12
Sapind 10 Apocyn 7 Clusi 24 Mor 19 Myrsin 14 Meli 14 Chrysobalan 17 Euphorbi 12
Anacardi 9 Aster 7 Rut 24 Bignoni 18 Meli 12 Piper 14 Arec 17 Meli 12
Combret 9 Bignoni 7 Apocyn 23 Salic 18 Anacardi 11 Sapind 14 Bignoni 16 Chrysobalan 11
Erythroxyl 9 Ochn 7 Celastr 19 Vochysi 18 Celastr 11 Myrsin 13 Vochysi 16 Sapind 11
Laur 9 Celastr 6 Chrysobalan 19 Celastr 17 Erythroxyl 11 Apocyn 12 Celastr 16 Aster 10
Rut 9 Chrysobalan 6 Meli 19 Aquifoli 16 Sapot 11 Aquifoli 12 Aster 16 Myrsin 10
Arec 8 Melastomat 6 Salic 19 Erythroxyl 16 Apocyn 10 Celastr 12 Combret 15 Nyctagin 10
Nyctagin 7 Mor 6 Anacardi 16 Sapot 16 Lami 10 Clusi 10 Malpighi 15 Anacardi 9
Polygon 7 Sapind 6 Nyctagin 14 Malpighi 14 Piper 10 Erythroxyl 10 Sapot 14 Symploc 9
Lecythid 6 Cyathe 5 Boragin 13 Meli 13 Vochysi 10 Cyathe 9 Boragin 13 Aquifoli 8
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172 Neotropical Savannas and Dry Forests: Diversity, Biogeography, and Conservation
The geographical proximity among different vegetation formations within the same region and evo-
lution through adaptive radiation into adjacent habitats could explain much of the strong effect of
distance found in species patterns throughout the geographical range analysed here. On the other
hand, the patterns related to climate and vegetation formations found for genera and families strongly
suggest that climatic variables, particularly temperature and water availability, have had a long
influence on the evolution and speciation of tree taxa in eastern South America. This is not a surprise
since water and temperature are the chief factors determining the distribution of most world’s vegetation
formations, and the history of vegetation and climate of that part of the world during the Quaternary
shows dramatic shifts in both temperature and rainfall regime (Salgado-Labouriau et al., 1997;
Behling, 1998; Ledru et al., 1998, Oliveira et al., 1999).
One important result of the above-mentioned geographical pattern is that there was greater
similarity in species composition between Atlantic rain and seasonal forests of the same region
than between either seasonal or rain forests of disjunct regions, although this holds true only when
east and south-east are merged. In the same region the tree flora of seasonal forests is much less
diverse than that of the rain forests, and is probably composed of species able to cope with relatively
longer dry seasons. Tree species diversity in tropical forests is often correlated with water
TABLE 7.6
Families with the Highest Number of Species (S) in the Tree Flora of Subtropical
Seasonal Forests of the South American Atlantic Forest Domain, Chaco Forests
and Caatingas (Suffix ‘-aceae’ Omitted from All Families; N = Number of Areas)
Subtropical seasonal forests
South Region S South-west Region S Chaco Forests S Caatingas S
(N = 37) 542 (N = 5) 420 (N = 39) 1193 (N = 6) 1317
Myrt 60 Fab 55 Fab 63 Fab 106
Fab 49 Anacardi 14 Cact 29 Euphorbi 36
Laur 21 Myrt 14 Solan 13 Cact 24
Rubi 18 Malv 13 Euphorbi 11 Malv 19
Solan 18 Sapind 12 Zygophyll 9 Myrt 16
Euphorbi 17 Bignoni 10 Anacardi 7 Apocyn 11
Bignoni 11 Apocyn 8 Nyctagin 7 Bignoni 10
Meli 11 Euphorbi 8 Rhamn 6 Erythroxyl 10
Mor 11 Rut 7 Brassic 5 Boragin 9
Aster 10 Meli 6 Celastr 5 Rubi 9
Salic 10 Mor 6 Malv 5 Rut 8
Rut 9 Salic 6 Verben 5 Celastr 7
Myrsin 8 Arec 5 Arec 4 Combret 7
Anacardi 7 Celastr 5 Bignoni 4 Annon 6
Erythroxyl 7 Nyctagin 5 Cannab 4 Nyctagin 6
Malv 7 Sapot 5 Myrt 4 Polygon 6
Melastomat 7 Aster 4 Rut 4 Sapind 6
Sapind 7 Cannab 4 Sapind 4 Arec 5
Urtic 7 Laur 4 Sapot 4 Brassic 5
Celastr 6 Polygon 4 Apocyn 3 Anacardi 4
Sapot 6 Rubi 4 Malpighi 3 Rhamn 4
Apocyn 5 Brassic 3 Mor 3 Salic 4
Aquifoli 5 Rhamn 3 Polygon 3 Solan 4
Arec 5 Myrsin 3 Santal 2 Aster 3
Boragin 5 Phytolacc 3 Aster 2 Malpighi 3
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Floristic Relationships of Seasonally Dry Forests of Eastern 173
consumption and energy uptake, resources that are partitioned among species and limit their number
in forest communities (Hugget, 1995). Water shortage probably plays the chief role in reducing
species richness of seasonal forests compared to rain forests, and even more so of semi-arid
formations such as chaco forests, caatingas and carrascos. Moreover, the structure of seasonal
forests is also less complex, therefore favouring a reduced number of understory species compared
to rain forests (Gentry and Emmons, 1987).
The intimate relationship between the two floras within each geographical region supports the
wider definition of Atlantic forests to include both rain and seasonal forests as physiognomic and
floristic expressions of a single great vegetation domain (Oliveira-Filho and Fontes, 2000; Galindo-Leal
and Câmara, 2003). For all regions but the north-east, there was a greater floristic similarity at all
three taxonomic levels between Atlantic rain and seasonal forests than between either of these and
Amazonian rain forests. The exceptions were the north-east rain and seasonal semideciduous forests,
both closer to Amazonian rain forests though only at the generic and familial levels. As the coastal
north-east is climatically and geographically closer to the Amazon, a stronger past link could have
existed through the so-called north-east bridge (Bigarella et al. 1975; Mori et al., 1981; Andrade-
Lima, 1982; Cavalcanti and Tabarelli, 2004). Nevertheless, as shown by the present results, this
alleged link also includes rain and seasonal forests and, for that reason, there is little floristic ground
for viewing Atlantic rain forests as being closer to Amazonian rain forests than to their adjacent
seasonal forests.
In all four Atlantic regions, seasonal forests and their rain forest neighbours share a similar
proportion of the total species count (c.20%) and are both poorer in species in the north-east and
south and much richer in the east and south-east. Seasonal forests of the central-west were also
comparatively rich. An inspection of the distribution map (Figure 7.1) helps us understand this. Of
all regions, the north-east has the smallest forest area and also lacks the highly rugged relief of
other regions, the latter being a feature that may boost species richness through increased environ-
mental heterogeneity. In addition, the region may have lost much of its primitive species richness
because it was the first to go through mass deforestation, beginning in the sixteenth century. It is
the least known, and most threatened and reduced of all Atlantic forests, now covering only 3.76%
of its original area (Silva and Tabarelli, 2000, 2001). Towards the south, Atlantic seasonal forests
expand increasingly more into the continental interior until reaching Mato Grosso do Sul and
eastern Paraguay, so that they cover a wide area with pronounced variation in relief and climate
(Oliveira-Filho and Fontes, 2000). In addition, seasonal forests also spread towards the west into
the whole of the cerrado domain as galleries and forest patches that are found as far as in the
Bolivian chiquitanía (see Kileen et al., Chapter X). The high environmental heterogeneity of this
large area, combined with the complex contact with the cerrado, certainly explains the high species
richness of the east, south-east and central-west tropical seasonal forests. The comparatively lower
speciesrichness of the subtropical seasonal forests of the south may also be explained by their
comparatively smaller area and modest relief, but these forests are also at the southernmost range
of Atlantic forests where extreme low temperatures in winter coupled with frosts may have already
selected the small proportion of tree species able to cope with this climatic harshness (Rambo,
1980; Leite, 2002; Jurinitz and Jarenkow, 2003).
Surprisingly the proportion of seasonal forest species shared with rainforests remains more or
less constant throughout the geographical range, despite the increase in species numbers of rain-
forest from north-east and north to south-east and south. This is brought about by an increase in
the percentage of seasonal forest species occurring in rainforest from 51% and 52.9% in the north-
east and east, respectively, to 74% and 83% in the south-east and south, thus counterbalancing the
increase in rainforest endemics and maintaining the proportion. Unlike the situation in the other
regions, the number of species in the north-east seasonal forests actually surpasses that of rain
forests. The main contrast between the two pairs of northern and southern regions is that the
former (north-east and east) have warmer temperatures and a much more pronounced variation
in rainfall totals and seasonality than the latter (south-east and south) since they are adjacent to
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174 Neotropical Savannas and Dry Forests: Diversity, Biogeography, and Conservation
the caatinga domain. This is probably why north-east seasonal forests are the only ones to show a
clear distinction between semideciduous and deciduous formations. The wider rainfall gradient is
correlated with a relatively rapid transition from rain- to semideciduous and deciduous forests, and
from those to caatingas, and thus because of ecotonal effects increasing the speciesrichness of
seasonal forests relative to their ‘purer’ rain forest partners. This explains, for example, why
Cactaceae and Euphorbiaceae are so important in the flora of montane semideciduous forests, the
so-called brejo forests, which occur as hinterland forest islands on mountains surrounded by lowland
caatingas, and inevitably share a number of species with the latter (Rodal, 2002; Pôrto et al., 2004).
In addition to this, the assemblage of north-east seasonal forests includes those influenced by other
neighbouring formations, such as the coastal sandy restingas (e.g. at Fernando de Noronha and
Natal) and the cerrado (e.g. at Araripe, Campo Maior and Sete Cidades) that may also boost their
species richness (Farias and Castro, 2004). Similar effects may have occurred in the eastern region
which also combines floristic interactions with both caatingas and cerrados, in addition to the effects
of the rugged relief of the Espinhaço mountain range and the Chapada Diamantina (Guedes and
Orge, 1998; Zappi et al., 2003).
The central-west seasonal forests also have strong floristic links with both the cerrados and
Atlantic forests and share a considerable number of species. In fact, one could extract a continuum
in tree species distribution determined by rainfall seasonality starting at the east and south-east
Atlantic rain and seasonal forests, and extending towards the central-west to reach its seasonal
forests and, lastly, the cerradões and cerrados, as already suggested by Leitão-Filho (1987). However,
this is an oversimplified view of the floristic gradient because it is now known that, under seasonal
climates, more important factors are involved in determining the forest-cerrado transition, and fire
frequency and soil fertility and moisture play the chief role here (Oliveira-Filho and Ratter, 2002). As
a result, it is not uncommon to find in the central-west two or more of those formations on a single
slope (Furley and Ratter, 1988; Furley et al., 1988; Ratter et al., 1978). The complex mosaic of
vegetation formations of the region and the species interchange among them probably explain why
the analyses failed to discriminate deciduous from semideciduous forests floristically. Although the
two formations do form a continuum it is usually easy to tell at least their extremes apart in the
field on the basis of physiognomy and floristic composition (Oliveira-Filho and Ratter, 2002). For
that reason, particular attention should be paid to such nuances in the preparation of checklists for
the region.
Towards the south and south-east, the declining temperatures and related vapor pressure curtail
the water deficit gradient and this probably favors a stronger floristic relationship between rain and
seasonal forests expressed by the much higher proportions of shared species. Extremes of low
temperatures may be important determinants of tree species distribution. Occasional frosts have
been mentioned by Oliveira-Filho et al. (1994) as an important factor limiting species distribution
both in relation to higher elevations and latitudes in south and south-east Brazil. Resistance to
frosts was suggested as a key factor determining the special nature of the chaco flora, together with
their saline to alkaline soils (Pennington et al., 2000). The influence of latitude and altitude on
climate, however, is far more complex than simply that of temperature and frosts. Increasing latitude
also means increasing year-round variation of the daily sunlight period. Rising elevation also
decreases atmospheric pressure, increases solar radiation, accelerates windmovement, promotes
greater cloudiness and boosts rainfall (Jones, 1992). For tropical forests, rainfall seasonality is
apparently more important than annual rainfall in determining presence of rain or seasonal forests,
and the occurrence of at least a 30-day dry season produces effects which can clearly be shown
on a vegetation map (IBGE, 1993). For subtropical forests, however, temperature range prevails
over rainfall seasonality in separating rain and seasonal forests, probably because the contrast
of low winter and high summer temperatures plays an additional role in forest deciduousness
(Holdridge et al., 1971). Low temperatures alone, without the strong annual oscillation, are not
associated with the presence of subtropical seasonal forests, and other formations appear, in particular
araucaria rain forests, in the hinterland highlands, and upper montane rain forests (cloud forests) on
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Floristic Relationships of Seasonally Dry Forests of Eastern 175
the mountain ridges near the coast (Roderjan et al., 2002). Semi-arid formations such as chaco
forests are found only where very strong rainfall seasonality occurs in subtropical climates, but
under these conditions forests also give way to open grasslands (campos or pampas) in many areas
of the south, and this is probably linked to the past history of fire and grazing (Behling, 1995,
1997; Quadros and Pillar, 2002).
Tree species composition of seasonal forests is highly influenced by altitude and associated
temperatures, a well-known fact for mountain vegetation worldwide (Hugget, 1995). Because most
mountain ranges and plateaus in our area of study are concentrated in the east and the lowlands of
the Paraguay river basin lie in the west, the seasonal forest gradient related to decreasing altitude
and increasing temperatures is highly coincident with increasing distance from the ocean. For this
reason, one might speculate that this gradient was another primarily related to distance. Neverthe-
less, altitude-related gradients have already been detected for Atlantic rain and seasonal forests at
more regional scales by Oliveira-Filho and Fontes (2000) and Ferraz et al. (2004) in south-east and
north-east Brazil, respectively, and by Salis et al. 1995, Torres et al. 1997 and Scudeller et al. (2001)
in the state of São Paulo. Moreover, some floristic patterns found with increasing altitude also
coincided with those cited by the above authors and by Gentry (1995) for Andean and Central
American forests. Among these, are the increasing contribution of Melastomataceae to the tree
flora, particularly Miconia and Tibouchina, Solanaceae (Solanum), Lauraceae (Ocotea and Nectan-
dra), Aquifoliaceae (Ilex) and Asteraceae, and the decrease of Eugenia and Ficus with increasing
altitude. A detailed treatment of genera and species diagnostic of montane Atlantic forests is given
by Oliveira-Filho and Fontes (2000).
It is now accepted that the caatinga domain represents the largest, most isolated and species-
rich nucleus of the SDTF and that its flora is made up of a blend of endemic and wide-range species
(Giulietti et al., 2002; Prado, 2003). The strongest internal floristic dichotomy of the semi-arid
vegetation of the caatinga is that linked to soils derived from either crystalline base rock or sandy
deposits (Araújo et al., 1998; Rodal and Sampaio, 2002; Rocha et al., 2004). The present analyses
largely support these findings, which are considered by Queiroz in Chapter X. He hypothesizes
that the vast proportion of caatinga non-endemics results from post-Tertiary migration of wide-
range SDTF species into the region as a result of the progressive retreat of sandy deposits and
exposure of the crystalline bedrock. In fact, most non-endemic caatinga species are found through-
out the Austro-Atlantic and central-west seasonal forests and many reach the peripheral chaco
seasonal forests without entering the chaco itself, as already emphasized by Prado (1991) and Prado
and Gibbs (1993) to demonstrate the strong differentiation of the chaco and caatinga floras.
Interestingly, our analyses also suggest that a similar pattern may also occur in non-endemic chaco
species that are also found in Austro-Atlantic and/or central-west seasonal forests but do not enter
the caatingas. Despite this similarity there is also an important difference in that most non-endemic
chaco species show a more limited distribution outside the chaco domain and do not reach as far
as the caatinga periphery. Thus, chaco and caatinga are well-defined floristic nuclei with very weak
relationships between their floras at both specific and generic levels. Only at the family level do
the two floras show a stronger link, thus suggesting that if a common proto-flora did exist it must
have been in the very remote past. Both floras also show floristic connections with the Atlantic and
central-west seasonal forests but this is probably mainly the result of both species interchange in
transitional areas and expansion of wide-range SDTF species.
7.5 CONCLUSION
We propose here that one can best describe Atlantic seasonal forests as a section of a complex
floristic gradient extending from evergreen forests to semideciduous and deciduous forests (the
SDTF section), and then running in a partial blending of floras to open formations, such as cerrados
and campos or, alternatively, to caatingas and chaco forests. This gradient is chiefly related to
decreasing water availability through either increasing rainfall seasonality and/or decreasing soil
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176 Neotropical Savannas and Dry Forests: Diversity, Biogeography, and Conservation
moisture content, but there is also a strong interference of temperature gradients along the latitudinal
and altitudinal ranges, and of variations in soil fertility and fire frequency. The flora of the Atlantic
seasonal forests occurs mostly in the section of the tropical gradient corresponding to annual periods
of water shortage between 30 and 160 days, and in the section of the subtropical range where
periods of water shortage are below 30 days but year-round monthly temperature oscillation is
above 10°C. Increasing periods of water shortage, soil fertility and temperature range normally
lead from semideciduous to deciduous forests and then to the semi-arid formations, either caatingas
(tropical) or chaco forests (subtropical), while increasing fire frequency and decreasing soil fertility
frequently lead from seasonal forests to either cerrados (tropical) or southern campos (subtropical).
For this reason we suggest here that the definition of SDTF should be reshaped to include both
cerrados and the chaco.
In conclusion, we believe that the best view of the SDTF vegetation of eastern South America
is that of three floristic nuclei: caatinga, chaco and Atlantic forest (sensu latissimo). Only the latter,
however, should be linked consistently to the residual Pleistocenic dry seasonal (RPDS) flora.
Caatinga and chaco form the extremes of floristic dissimilarity among the three SDTF nuclei, also
corresponding to the warm-dry and warm-cool climatic extremes, respectively. In contrast, the
Atlantic SDTF nucleus is poor in endemic species and is actually a floristic bridge connecting the
two drier nuclei to rain forests. Additionally, there is little evidence to describe the Atlantic nucleus
flora as a clearly distinct species assemblage, as are those of the caatinga and chaco nuclei, because
of the striking variation in species composition found throughout the vast geographical extent of
Atlantic seasonal forests. Nevertheless, there is a group of wide-range species that is found in most
regions of the Atlantic nucleus, some of which are also part of the species blend of the caatinga
and chaco floras, though involving the latter to a much lesser extent. We believe that, at least in
eastern South America, it is precisely this small fraction of the Atlantic nucleus flora that should
be identified with the RPDS vegetation. To clarify the past history of neotropical SDTF, we propose
that the focus should now be shifted to the investigation of the distribution patterns of those species
and the past history of their populations in different locations of their geographical range.
ACKNOWLEDGEMENTS
The first author thanks the CNPq and the Royal Society of Edinburgh for the financial support to
the present study and the Royal Botanic Garden Edinburgh for warmly hosting him once more.
We were helped during the taxonomic revision of the database by the following: Marcos Sobral
(Myrtaceae) and João Renato Stehmann (Solanaceae), both from the Federal University of
Minas Gerais; Haroldo Lima (Fabaceae), Alexandre Quinet (Lauraceae), José Fernando Baumgratz
(Melastomataceae) and Angela Studart da Fonseca Vaz (Bauhinia) from the Rio de Janeiro Botanic
Garden; Maria Célia Vianna (Vochysia) from the Alberto Castellanos Herbarium; José Rubens Pirani
(Simaroubaceae, Picramniaceae and Rutaceae) and Pedro Fiaschi (Araliaceae) from the University
of São Paulo; Maria Lúcia Kawazaki (Myrtaceae) and Inês Cordeiro (Euphorbiaceae) from the São
Paulo Botanic Institute; Washington Marcondes-Ferreira (Aspidosperma) from the State University
of Campinas; Germano Guarim Neto (Cupania) from the Federal University of Mato Grosso; and
Toby Pennington and Maureen Warwick (Fabaceae) from the Royal Botanic Garden Edinburgh.
We also thank Toby Pennington, James Ratter and an anonymous reviewer for their critical and
constructive reading of the first draft.
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APPENDIX. MOST FREQUENT SPECIES (>70% OF CHECKLISTS) IN
THE TREE FLORA OF SELECTED SDTF AND SDSF FORMATIONS
OF EASTERN SOUTH AMERICA.
Low altitude tropical seasonal forests — North-east region: Abarema cochliacarpos, Acacia
polyphylla, Albizia pedicellaris, A. polycephala, Allophylus edulis, Alseis pickelii, Anacardium
occidentale, Andira fraxinifolia, A. nitida, Apeiba tibourbou, Apuleia leiocarpa, Aspidosperma
pyrifolium, Astronium fraxinifolium, Bowdichia virgilioides, Brosimum gaudichaudii, B. guian-
ense, Buchenavia capitata, Byrsonima sericea, Caesalpinia ferrea, Campomanesia aromatica, C.
dichotoma, Capparis flexuosa, Casearia sylvestris, Cecropia pachystachya, C. palmata, Cereus
jamacaru, Chamaecrista apoucouita, C. ensiformis, Chrysophyllum rufum, Clusia nemorosa, Coc-
coloba alnifolia, C. cordifolia, Cordia trichotoma, Coutarea hexandra, Cupania revoluta, Curatella
americana, Enterolobium contortisiliquum, Erythrina velutina, Erythroxylum citrifolium, Eschweilera
ovata, Eugenia florida, E. punicifolia, E. uniflora, Guapira noxia, G. opposita, G. pernambucensis,
Guarea guidonia, Guazuma ulmifolia, Guettarda platypoda, Himatanthus phagedaenicus, Hirtella
ciliata, H. racemosa, Hymenaea courbaril, H. rubriflora, Inga capitata, I. ingoides, I. laurina,
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180 Neotropical Savannas and Dry Forests: Diversity, Biogeography, and Conservation
I. thibaudiana, Lecythis pisonis, Luehea ochrophylla, L. paniculata, Manihot epruinosa, Manilkara
salzmannii, Maytenus distichophylla, M. erythroxylon, Miconia albicans, Myrcia multiflora, M.
sylvatica, M. tomentosa, Myrsine guianensis, Ocotea notata, Ouratea hexasperma, Palicourea cro-
cea, Pera glabrata, Pogonophora schomburgkiana, Pouteria grandiflora, Pradosia lactescens, Pro-
tium heptaphyllum, Psidium oligospermum, Pterocarpus rohrii, Rauvolfia ligustrina, Sacoglottis
mattogrossensis, Schefflera morototoni, Spondias mombin, Strychnos parvifolia, Stryphnodendron
pulcherrimum, Swartzia pickelii, Tabebuia impetiginosa, T. roseo-alba, T. serratifolia, Talisia escu-
lenta, Tapirira guianensis, Thyrsodium spruceanum, Trema micrantha, Vismia guianensis, Vitex
triflora, Ximenia americana, Ziziphus joazeiro, Zollernia latifolia.
High altitude tropical seasonal forests — North-east region: Acacia polyphylla, A. riparia, A.
tenuifolia, Albizia polycephala, Allophylus edulis, Anadenanthera colubrina, Bowdichia virgilio-
ides, Buchenavia capitata, Byrsonima sericea, Caesalpinia ferrea, Campomanesia aromatica, Cap-
paris flexuosa, Casearia sylvestris, Ceiba glaziovii, Celtis iguanaea, Clusia nemorosa, Copaifera
langsdorffii, Cordia trichotoma, Coutarea hexandra, Croton rhamnifolius, Cupania revoluta, Cyathea
microdonta, Enterolobium contortisiliquum, Erythroxylum citrifolium, Eugenia punicifolia, Gua-
pira laxiflora, G. opposita, Guazuma ulmifolia, Guettarda sericea, Hymenaea courbaril, Machaerium
hirtum, Manilkara rufula, Maprounea guianensis, Maytenus obtusifolia, Miconia albicans, Myrcia
fallax, M. multiflora, M. sylvatica, M. tomentosa, Myroxylon peruiferum, Myrsine guianensis,
Ocotea duckei, Piptadenia stipulacea, Platymiscium floribundum, Prockia crucis, Psidium
guineense, Randia nitida, Roupala cearensis, Ruprechtia laxiflora, Sapium glandulosum, Schoepfia
brasiliensis, Senna macranthera, S. spectabilis, S. splendida, Tabebuia impetiginosa, T. serratifolia,
Talisia esculenta, Vitex rufescens, Zanthoxylum rhoifolium.
Low altitude tropical seasonal forests — East region: Acacia polyphylla, Aegiphila sellowiana,
Albizia polycephala, Alchornea glandulosa, Allophylus edulis, Amaioua guianensis, Anadenanthera
colubrina, Andira fraxinifolia, Aparisthmium cordatum, Apuleia leiocarpa, Aspidosperma parvifo-
lium, Astrocaryum aculeatissimum, Astronium graveolens, Bathysa nicholsonii, Bauhinia fusco-
nervis, Brosimum guianense, B. lactescens, Byrsonima sericea, Cabralea canjerana, Carpotroche
brasiliensis, Casearia sylvestris, C. ulmifolia, Cassia ferruginea, Cecropia glaziovii, C. hololeuca,
Cedrela fissilis, Copaifera langsdorffii, Croton urucurana, Cyathea delgadii, Dalbergia nigra, Endli-
cheria paniculata, Erythrina verna, Erythroxylum pelleterianum, E. pulchrum, Eugenia florida,
Euterpe edulis, Ficus gomelleira, Gallesia integrifolia, Guapira opposita, Guarea macrophylla,
Guatteria australis, G. villosissima, Guettarda uruguensis, Himatanthus lancifolius, Hortia arborea,
Hymenolobium janeirense, Inga capitata, I. vera, Joannesia princeps, Lacistema pubescens, Lecythis
lurida, L. pisonis, Luehea divaricata, L. grandiflora, Mabea fistulifera, Machaerium brasiliense, M.
hirtum, M. stipitatum, Maclura tinctoria, Maprounea guianensis, Melanoxylon brauna, Miconia
cinnamomifolia, Myrcia fallax, M. rufula, Myrciaria floribunda, Nectandra oppositifolia, Ocotea
dispersa, Pera glabrata, Piptadenia gonoacantha, Plathymenia reticulata, Platymiscium floribundum,
Platypodium elegans, Pogonophora schomburgkiana, Pourouma guianensis, Protium warmin-
gianum, Pseudobombax grandiflorum, Pseudopiptadenia contorta, Pterocarpus rohrii, Pterygota
brasiliensis, Rollinia laurifolia, Senna macranthera, S. multijuga, Siparuna guianensis, Sorocea
guilleminiana, Sparattosperma leucanthum, Stryphnodendron pulcherrimum, Swartzia acutifolia,
S. myrtifolia, Syagrus romanzoffiana, Tabebuia serratifolia, Tabernaemontana hystrix, Tapirira
guianensis, Trichilia lepidota, T. pallida, Urbanodendron verrucosum, Virola bicuhyba, Vismia
guianensis, Xylopia brasiliensis, Xylopia sericea, Zanthoxylum rhoifolium.
High altitude tropical seasonal forests — East region: Aegiphila sellowiana, Alchornea triplinervia,
Amaioua guianensis, Anadenanthera colubrina, Andira fraxinifolia, Apuleia leiocarpa, Aspidosperma
discolor, A. olivaceum, Astronium graveolens, Bauhinia longifolia, Blepharocalyx salicifolius,
Bowdichia virgilioides, Byrsonima sericea, Cabralea canjerana, Campomanesia xanthocarpa, Casearia
arborea, C. decandra, C. obliqua, C. sylvestris, Cassia ferruginea, Cecropia glaziovii, C. hololeuca,
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Floristic Relationships of Seasonally Dry Forests of Eastern 181
C. pachystachya, Cedrela fissilis, Celtis iguanaea, Chrysophyllum gonocarpum, Clethra scabra,
Copaifera langsdorffii, Cordia sellowiana, Croton floribundus, C. urucurana, Cupania paniculata,
C. vernalis, Cyathea corcovadensis, C. delgadii, C. phalerata, Dalbergia frutescens, Dictyoloma
vandellianum, Eugenia florida, Gochnatia polymorpha, Guapira opposita, Guarea macrophylla,
Guatteria australis, G. sellowiana, G. villosissima, Guazuma ulmifolia, Hyptidendron asperrimum,
Inga laurina, I. marginata, I. sessilis, Kielmeyera lathrophyton, Lamanonia ternata, Leandra melas-
tomoides, Luehea divaricata, Machaerium brasiliense, M. hirtum, M. nictitans, M. villosum,
Maprounea guianensis, Matayba elaeagnoides, Maytenus salicifolia, Miconia cinnamomifolia, M.
ligustroides, M. pepericarpa, Myrcia detergens, M. fallax, M. guianensis, M. rostrata, M. tomentosa,
Myrsine coriacea, M. umbellata, Nectandra lanceolata, N. oppositifolia, Ocotea corymbosa, O.
odorifera, O. spixiana, Pera glabrata, Platypodium elegans, Protium heptaphyllum, Prunus myrti-
folia, Psychotria vellosiana, Rollinia laurifolia, Rollinia sylvatica, Roupala brasiliensis, Sapium
glandulosum, Sclerolobium rugosum, Senna macranthera, S. multijuga, Siparuna guianensis, Sipho-
neugena densiflora, Sorocea guilleminiana, Tabebuia serratifolia, Tapirira guianensis, T. obtusa,
Terminalia glabrescens, Tibouchina candolleana, Trichilia pallida, Vitex polygama, Vochysia tucan-
orum, Zanthoxylum rhoifolium.
Low altitude tropical seasonal forests — South-east region: Acacia polyphylla, Actinostemon
klotzschii, Aegiphila sellowiana, Albizia niopoides, Alchornea glandulosa, A. triplinervia, Allophy-
lus edulis, Aloysia virgata, Annona cacans, Apuleia leiocarpa, Aralia warmingiana, Aspidosperma
polyneuron, Astronium graveolens, Balfourodendron riedelianum, Bastardiopsis densiflora,
Cabralea canjerana, Campomanesia guazumifolia, C. xanthocarpa, Cariniana estrellensis, Casearia
gossypiosperma, C. sylvestris, Cecropia pachystachya, Cedrela fissilis, Ceiba speciosa, Celtis igua-
naea, Chrysophyllum gonocarpum, C. marginatum, Colubrina glandulosa, Copaifera langsdorffii,
Cordia ecalyculata, C. trichotoma, Croton floribundus, Cupania vernalis, Dalbergia frutescens,
Dendropanax cuneatus, Diatenopteryx sorbifolia, Endlicheria paniculata, Enterolobium contortisil-
iquum, Esenbeckia febrifuga, Eugenia florida, E. involucrata, Euterpe edulis, Gallesia integrifolia,
Guapira opposita, Guarea guidonia, G. kunthiana, G. macrophylla, Guatteria australis, Gymnanthes
concolor, Heliocarpus americanus, Holocalyx balansae, Inga marginata, I. striata, I. vera, Ixora
venulosa, Jacaranda micrantha, Jacaratia spinosa, Lonchocarpus cultratus, L. muehlbergianus, Lue-
hea divaricata, Machaerium hirtum, M. nictitans, M. paraguariense, M. stipitatum, Maclura tinctoria,
Matayba elaeagnoides, Metrodorea nigra, Myrcia multiflora, Myrciaria floribunda, Myrocarpus
frondosus, Myrsine umbellata, Nectandra megapotamica, Ocotea diospyrifolia, O. puberula, O.
pulchella, Parapiptadenia rigida, Patagonula americana, Peltophorum dubium, Piper amalago,
Prunus myrtifolia, Rollinia emarginata, R. sylvatica, Roupala brasiliensis, Schefflera morototoni,
Sebastiania commersoniana, Seguieria langsdorffii, Sorocea bonplandii, Syagrus romanzoffiana,
Tabernaemontana catharinensis, Terminalia triflora, Trema micrantha, Trichilia catigua, T. clausseni,
T. elegans, T. pallida, Vitex megapotamica, Zanthoxylum caribaeum, Z. fagara, Z. rhoifolium,
Z. riedelianum.
High altitude tropical seasonal forests — South-east region: Aegiphila sellowiana, Albizia
polycephala, Alchornea triplinervia, Amaioua guianensis, Andira fraxinifolia, Annona cacans, Aspi-
dosperma olivaceum, Byrsonima laxiflora, Cabralea canjerana, Calyptranthes clusiifolia, Cam-
pomanesia guazumifolia, Cariniana estrellensis, Casearia decandra, C. lasiophylla, C. obliqua, C.
sylvestris, Cecropia glaziovii, C. pachystachya, Cedrela fissilis, Chrysophyllum marginatum, Cin-
namomum glaziovii, Clethra scabra, Copaifera langsdorffii, Cordia sellowiana, Croton floribundus,
C. verrucosus, Cryptocarya aschersoniana, Cupania vernalis, Cyathea delgadii, C. phalerata, Dal-
bergia villosa, Daphnopsis brasiliensis, D. fasciculata, Dendropanax cuneatus, Endlicheria panic-
ulata, Eugenia florida, Gomidesia affinis, Guapira opposita, Guarea macrophylla, Guatteria australis,
Gymnanthes concolor, Heisteria silvianii, Hyeronima ferruginea, Inga striata, Ixora warmingii,
Jacaranda macrantha, Lamanonia ternata, Leucochloron incuriale, Lithraea molleoides, Luehea
divaricata, L. grandiflora, Machaerium hirtum, M. nictitans, M. stipitatum, M. villosum,
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182 Neotropical Savannas and Dry Forests: Diversity, Biogeography, and Conservation
Maclura tinctoria, Matayba elaeagnoides, Miconia cinnamomifolia, Mollinedia widgrenii, Myrcia
fallax, M. rostrata, Myrciaria floribunda, Myrsine coriacea, M. umbellata, Nectandra grandiflora,
N. oppositifolia, Ocotea corymbosa, O. diospyrifolia, O. odorifera, O. pulchella, Pera glabrata,
Persea pyrifolia, Piptocarpha macropoda, Platycyamus regnellii, Platypodium elegans, Protium
widgrenii, Prunus myrtifolia, Psychotria vellosiana, Rollinia dolabripetala, R. laurifolia, R. sylvat-
ica, Roupala brasiliensis, Sapium glandulosum, Schinus terebinthifolius, Sclerolobium rugosum,
Solanum pseudoquina, Sorocea bonplandii, Syagrus romanzoffiana, Tabebuia serratifolia, Tapirira
guianensis, T. obtusa, Ternstroemia brasiliensis, Trichilia emarginata, T. pallida, Vernonanthura
diffusa, Vismia brasiliensis, Vitex polygama, Vochysia tucanorum, Xylopia brasiliensis, Zanthox-
ylum rhoifolium.
Low altitude tropical seasonal forests — Central-west region: Acacia polyphylla, Acrocomia
aculeata, Albizia niopoides, Alibertia concolor, Anadenanthera colubrina, A. peregrina, Apeiba
tibourbou, Apuleia leiocarpa, Aspidosperma cylindrocarpon, A. olivaceum, A. pyrifolium, A. sub-
incanum, Astronium fraxinifolium, Attalea phalerata, Bauhinia longifolia, Bowdichia virgilioides,
Cabralea canjerana, Callisthene fasciculata, C. major, Calophyllum brasiliense, Cariniana estrel-
lensis, Casearia gossypiosperma, C. rupestris, C. sylvestris, Cecropia pachystachya, Cedrela fissilis,
Ceiba speciosa, Celtis iguanaea, Chrysophyllum gonocarpum, Combretum leprosum, Copaifera
langsdorffii, Cordia glabrata, C. trichotoma, Coutarea hexandra, Cupania vernalis, Dilodendron
bipinnatum, Diospyros hispida, D. sericea, Enterolobium contortisiliquum, Eugenia florida, Genipa
americana, Guapira opposita, Guarea guidonia, Guazuma ulmifolia, Guettarda uruguensis, Hyme-
naea courbaril, Inga laurina, I. marginata, I. vera, Jacaranda cuspidifolia, Licania apetala, Luehea
divaricata, L. paniculata, Machaerium hirtum, M. stipitatum, M. villosum, Maclura tinctoria, Mago-
nia pubescens, Matayba guianensis, Micropholis venulosa, Myracrodruon urundeuva, Myrcia
tomentosa, Myrciaria floribunda, Plathymenia reticulata, Platypodium elegans, Pouteria gardneri,
Protium heptaphyllum, P. spruceanum, Pseudobombax tomentosum, Psidium guineense, Pterogyne
nitens, Qualea multiflora, Randia nitida, Rhamnidium elaeocarpum, Rollinia emarginata, Salacia
elliptica, Sapium glandulosum, Sclerolobium paniculatum, Simira sampaioana, Siparuna guianen-
sis, Sorocea guilleminiana, Spondias mombin, Sterculia striata, Sweetia fruticosa, Tabebuia impe-
tiginosa, T. roseo-alba, T. serratifolia, Talisia esculenta, Tapirira guianensis, Terminalia argentea,
T. glabrescens, Tocoyena formosa, Trichilia catigua, T. clausseni, T. elegans, T. pallida, Triplaris
gardneriana, Unonopsis lindmanii, Vitex cymosa, Zanthoxylum rhoifolium.
High altitude tropical seasonal forests — Central-west region: Aegiphila sellowiana, Alibertia
edulis, Amaioua guianensis, Anadenanthera colubrina, Apuleia leiocarpa, Aspidosperma cylindro-
carpon, A. australe, A. subincanum, Astronium fraxinifolium, Bauhinia longifolia, Cabralea can-
jerana, Callisthene major, Calophyllum brasiliense, Cardiopetalum calophyllum, Cariniana estrel-
lensis, Casearia sylvestris, Cecropia pachystachya, Cedrela fissilis, Cheiloclinium cognatum,
Chrysophyllum marginatum, Copaifera langsdorffii, Cordia sellowiana, C. trichotoma, Cryptocarya
aschersoniana, Cupania vernalis, Dendropanax cuneatus, Diospyros hispida, Emmotum nitens,
Endlicheria paniculata, Eugenia florida, Euplassa inaequalis, Faramea cyanea, Ferdinandusa spe-
ciosa, Gomidesia fenzliana, Guarea guidonia, G. macrophylla, Guatteria sellowiana, Guazuma
ulmifolia, Guettarda uruguensis, Hedyosmum brasiliense, Hirtella glandulosa, H. gracilipes,
Hyeronima alchorneoides, Hymenaea courbaril, Inga alba, I. laurina, I. vera, Ixora warmingii,
Lamanonia ternata, Licania apetala, Luehea divaricata, Machaerium acutifolium, Maprounea guian-
ensis, Matayba guianensis, Mauritia flexuosa, Miconia chamissois, Micropholis venulosa, Mouriri
glazioviana, Myrcia rostrata, M. tomentosa, Myrsine guianensis, M. umbellata, Nectandra cissiflora,
Ocotea corymbosa, O. spixiana, Ormosia fastigiata, Ouratea castaneifolia, Pera glabrata, Piptadenia
gonoacantha, Piptocarpha macropoda, Platypodium elegans, Pouteria gardneri, P. ramiflora, Protium
heptaphyllum, P. spruceanum, Prunus myrtifolia, Pseudolmedia laevigata, Qualea dichotoma, Q.
multiflora, Richeria grandis, Roupala brasiliensis, Schefflera morototoni, Sclerolobium panicula-
tum, Siparuna guianensis, Siphoneugena densiflora, Styrax camporum, Symplocos nitens, Tabebuia
2987_C007.fm Page 182 Thursday, December 1, 2005 7:03 PM
Floristic Relationships of Seasonally Dry Forests of Eastern 183
serratifolia, Talauma ovata, Tapirira guianensis, Terminalia argentea, T. glabrescens, Trichilia
catigua, Virola sebifera, Vitex polygama, Vochysia tucanorum, Xylopia aromatica, Xylopia emar-
ginata, X. sericea, Zanthoxylum rhoifolium.
Subtropical seasonal forests — South region: Aiouea saligna, Alchornea triplinervia, Allophylus
edulis, A. guaraniticus, Apuleia leiocarpa, Banara parviflora, B. tomentosa, Blepharocalyx salici-
folius, Cabralea canjerana, Calyptranthes concinna, Campomanesia xanthocarpa, Casearia decan-
dra, C. sylvestris, Cedrela fissilis, Celtis iguanaea, Chrysophyllum gonocarpum, C. marginatum,
Citronella paniculata, Cordia ecalyculata, C. trichotoma, Cupania vernalis, Dalbergia frutescens,
Daphnopsis racemosa, Dasyphyllum spinescens, Diospyros inconstans, Enterolobium contortisil-
iquum, Erythrina crista-galli, Erythroxylum argentinum, Eugenia hyemalis, E. involucrata, E. opaca,
E. ramboi, E. rostrifolia, E. uniflora, Ficus insipida, F. luschnathiana, F. organensis, Gomidesia
palustris, Guapira opposita, Guettarda uruguensis, Gymnanthes concolor, Helietta apiculata, Ilex
brevicuspis, Inga marginata, I. vera, Jacaranda micrantha, Lithraea brasiliensis, Lonchocarpus
nitidus, Luehea divaricata, Machaerium stipitatum, Matayba elaeagnoides, Maytenus ilicifolia,
Myrcianthes pungens, Myrciaria tenella, Myrocarpus frondosus, Myrsine coriacea, M. loefgrenii,
M. lorentziana, M. umbellata, Nectandra lanceolata, N. megapotamica, Ocotea puberula, O. pul-
chella, Parapiptadenia rigida, Patagonula americana, Phytolacca dioica, Pilocarpus pennatifolius,
Pisonia zapallo, Pouteria gardneriana, P. salicifolia, Prunus myrtifolia, P. subcoriacea, Psidium
cattleianum, Quillaja brasiliensis, Randia nitida, Rollinia emarginata, R. sylvatica, Ruprechtia
laxiflora, Sapium glandulosum, Schefflera morototoni, Schinus terebinthifolius, Sebastiania brasil-
iensis, S. commersoniana, Seguieria americana, Solanum granuloso-leprosum, S. pseudoquina, S.
sanctaecatharinae, Sorocea bonplandii, Strychnos brasiliensis, Styrax leprosus, Syagrus romanzof-
fiana, Terminalia australis, Trema micrantha, Trichilia clausseni, T. elegans, Urera baccifera, Vitex
megapotamica, Xylosma pseudosalzmanii, Zanthoxylum fagara, Z. rhoifolium.
Subtropical seasonal forests — Southwest region: Acacia albicorticata, A. caven, A. praecox,
Acanthosyris falcata, Achatocarpus praecox, Allophylus edulis, Amburana cearensis, Anadenan-
thera colubrina, Aralia angelicifolia, Aspidosperma olivaceum, A quebracho-blanco, Caesalpinia
paraguariensis, Calycophyllum multiflorum, Capparis retusa, Carica quercifolia, Casearia sylvestris,
Celtis pubescens, Chloroleucon tenuiflorum, Chrysophyllum gonocarpum, C. marginatum, Cochlo-
spermum tetraporum, Cordia trichotoma, Crataeva tapia, Cupania vernalis, Diplokeleba floribunda,
Enterolobium contortisiliquum, Erythrina falcata, Eugenia uniflora, Geoffroea decorticans, G. stri-
ata, Gleditsia amorphoides, Holocalyx balansae, Maclura tinctoria, Myracrodruon balansae, M.
urundeuva, Myrcianthes cisplatensis, M. pungens, Myrsine laetevirens, Parkinsonia aculeata, Pat-
agonula americana, Peltophorum dubium, Phyllostylon rhamnoides, Phytolacca dioica, Pilocarpus
pennatifolius, Pisonia aculeata, P. zapallo, Pouteria gardneriana, Prosopis nigra, Pterogyne nitens,
Rollinia emarginata, Ruprechtia laxiflora, Sapindus saponaria, Sapium haematospermum, Schinop-
sis brasiliensis, Schinus polygamus, Scutia buxifolia, Sebastiania brasiliensis, Sideroxylon obtusi-
folium, Solanum granuloso-leprosum, Syagrus romanzoffiana, Tabebuia heptaphylla, T. impetigi-
nosa, Tabernaemontana catharinensis, Terminalia triflora, Tipuana tipu, Ximenia americana,
Zanthoxylum fagara, Z. petiolare, Z. rhoifolium, Ziziphus mistol.
Chaco forests: Acacia aroma, A. caven, A. curvifructa, A. furcatispina, A. praecox, A. tucumanensis,
Acanthosyris falcata, Achatocarpus praecox, Allophylus edulis, Anadenanthera colubrina, Apor-
osella chacoensis, Aralia angelicifolia, Aspidosperma quebracho-blanco, A triternatum, Athyana
weinmanniifolia, Bougainvillea campanulata, B. praecox, Bulnesia bonariensis, Caesalpinia para-
guariensis, Capparis atamisquea, C. retusa, C. salicifolia, C. insignis, Cereus stenogonus, Chryso-
phyllum marginatum, Copernicia alba, Cupania vernalis, Enterolobium contortisiliquum, Eugenia
uniflora, Geoffroea decorticans, Jacaratia corumbensis, Maytenus scutioides, M. spinosa, M. vitis-
idaea, Mimosa castanoclada, M. chacoensis, M. detinens, M. glutinosa, Mimozyganthus carinatus,
Myrcianthes cisplatensis, M. pungens, Myrsine laetevirens, Parkinsonia praecox, Patagonula americana,
2987_C007.fm Page 183 Thursday, December 1, 2005 7:03 PM
184 Neotropical Savannas and Dry Forests: Diversity, Biogeography, and Conservation
Pereskia sacharosa, Phyllostylon rhamnoides, Pisonia zapallo, Prosopis affinis, P. alpataco, P. elata,
P. fiebrigii, P. kuntzei, P. nigra, P. nuda, P. rojasiana, P. ruscifolia, P. sericantha, P. torquata,
Quiabentia chacoensis, Ruprechtia apetala, R. laxiflora, R. triflora, Sapium haematospermum,
Schinopsis balansae, S. cornuta, S. heterophylla, S. quebracho-colorado, Schinus polygamus, Scutia
buxifolia, Sesbania virgata, Sideroxylon obtusifolium, Stetsonia coryne, Tabebuia impetiginosa,
T nodosa, Tessaria dodoneifolia, T integrifolia, Thevetia bicornuta, Trema micrantha, Trithrinax
schizophylla, Ximenia americana, Zanthoxylum coco, Z. petiolare, Ziziphus mistol.
Caatingas: Acacia langsdorffii, A. paniculata, A. polyphylla, Allophylus quercifolius, Amburana
cearensis, Anadenanthera colubrina, Annona spinescens, Aspidosperma pyrifolium, Auxemma
glazioviana, A. oncocalyx, Balfourodendron molle, Bauhinia acuruana, B. cheilantha, B. pentandra,
Bocoa mollis, Brasiliopuntia brasiliensis, Byrsonima gardneriana, Caesalpinia bracteosa, C. ferrea,
C. microphylla, C. pyramidalis, Capparis flexuosa, C. jacobinae, C. yco, Ceiba glaziovii, Cereus
albicaulis, C. jamacaru, Chloroleucon acacioides, C. foliolosum, C. mangense, Cnidoscolus
bahianus, C. obtusifolius, C. quercifolius, Cochlospermum vitifolium, Combretum leprosum, Com-
miphora leptophloeos, Cordia leucocephala, Coutarea hexandra, Croton rhamnifolius, C. sonderi-
anus, Dalbergia catingicola, D. cearensis, Erythrina velutina, Erythroxylum revolutum, Eugenia
punicifolia, E. tapacumensis, Fraunhoffera multiflora, Geoffroea spinosa, Guapira laxa, Jatropha
mollissima, J. mutabilis, Manihot dichotoma, M. glaziovii, Maytenus rigida, Mimosa arenosa, M.
caesalpinifolia, M. gemmulata, M. malacocentra, M. tenuiflora, Myracrodruon urundeuva,
Parapiptadenia zehntneri, Pilosocereus gounellei, P. pachycladus, P. tuberculatus, Piptadenia obli-
qua, P. stipulacea, Pithecellobium diversifolium, Pseudobombax simplicifolium, Rollinia lepto-
petala, Sapium argutum, Schinopsis brasiliensis, Senna acuruensis, S. spectabilis, Senna splendida,
Sideroxylon obtusifolium, Spondias tuberosa, Tabebuia impetiginosa, Tacinga inamoena, T. palma-
dora, Thiloa glaucocarpa, Zanthoxylum stelligerum, Ziziphus joazeiro.
SDTF ‘Supertramp’ species (present in >100 checklists): Acacia polyphylla, Acrocomia aculeata,
Aegiphila sellowiana, Alibertia concolor, Allophylus edulis, Aloysia virgata, Anadenanthera colu-
brina, Andira fraxinifolia, Apuleia leiocarpa, Aspidosperma olivaceum, A. pyrifolium, Astronium
fraxinifolium, Bauhinia forficata, Bowdichia virgilioides, Brosimum gaudichaudii, Cabralea can-
jerana, Campomanesia xanthocarpa, Casearia decandra, C. sylvestris, Cecropia pachystachya,
Cedrela fissilis, Ceiba speciosa, Celtis iguanaea, C. pubescens, Chrysophyllum gonocarpum, C.
marginatum, Copaifera langsdorffii, Cordia trichotoma, Coutarea hexandra, Cupania vernalis, Dal-
bergia frutescens, Diospyros inconstans, Endlicheria paniculata, Enterolobium contortisiliquum,
Eugenia florida, E. punicifolia, E. uniflora, Garcinia gardneriana, Guapira opposita, Guarea gui-
donia, Guazuma ulmifolia, Guettarda uruguensis, Gymnanthes concolor, Hymenaea courbaril, Inga
marginata, I. vera, Lithraea molleoides, Lonchocarpus campestris, Luehea divaricata, L. grandiflora,
Machaerium acutifolium, M. hirtum, M. stipitatum, Maclura tinctoria, Maprounea guianensis,
Matayba elaeagnoides, M. guianensis, Maytenus ilicifolia, Miconia albicans, Myracrodruon urun-
deuva, Myrcia guianensis, M. multiflora, M. rostrata, M. tomentosa, Myroxylon peruiferum, Pel-
tophorum dubium, Pera glabrata, Piper amalago, Pisonia zapallo, Platypodium elegans, Prockia
crucis, Protium heptaphyllum, Prunus myrtifolia, Pterogyne nitens, Randia nitida, Rollinia emar-
ginata, R. sylvatica, Roupala brasiliensis, Ruprechtia laxiflora, Sapium glandulosum, Schefflera
morototoni, Sebastiania brasiliensis, Sideroxylon obtusifolium, Siparuna guianensis, Solanum
granuloso-leprosum, Sweetia fruticosa, Syagrus oleracea, S. romanzoffiana, Tabebuia impetiginosa,
T. serratifolia, Tapirira guianensis, Terminalia fagifolia, Trema micrantha, Trichilia catigua, T. clausseni,
T. elegans, Urera baccifera, Zanthoxylum fagara, Z. petiolare, Z. rhoifolium.
2987_C007.fm Page 184 Thursday, December 1, 2005 7:03 PM
... Dans la région du Nordeste, elle occupe l'Agreste (terme populaire désignant la région de transition entre le littoral humide et le Sertão sec). Certains auteurs rapprochent ce type de forêt de la forêt dense humide semi-décidue du Planalto Centro-Sul et du Parana (pereirA et al., 2002 ;oLiveirA FiLho et al., 2006). Ces forêts sont fortement conditionnées par la pluviométrie : précipitations oscillant entre 700 mm et 1600 mm par an avec une période de 5 à 6 mois durant lesquels les pluies sont inférieures à 100 mm/mois. ...
... Certaines espèces sont typiques de cette formation telles que Zizyphus joazeiro (Rhamnaceae)(IBGE, 2012). La diversité spécifique de cette forêt sèche est moins importante que celle de la Forêt Atlantique mais reste très largement supérieure à celle de la Caatinga (oLiveirA-FiLho et al., 2006 ...
... Les affleurements rocheux et les inselbergs sont des hotspots de diversité biologique, abritant de nombreuses endémiques (FrANçA et al., 2006 ;FrANçA-rochA et al., 2007 ;gomeS & ALveS, 2009 ;. Parmi les formations décidues brésiliennes, la Caatinga est l'une des plus riches en espèces végétales endémiques à cause des particularités du climat et de la roche-mère (oLiveirA-FiLho et al., 2006). ...
... The Atlantic Forest extends across Brazil's eastern coast and reaches more continental areas in northeastern Argentina and southeastern Paraguay (5-33° S latitude and 35-57° W longitude; Figure 1). The Atlantic Forest has strong gradients of rainfall (annual means varying between 500 and 3000 mm) and temperature (annual means between 10 and 25 °C), which is attributed to its large variation in terms of forest structure, species richness, and composition (Oliveira-Filho et al., 2006), with high alpha and beta diversities of tree species (Bergamin et al., 2017), as well as high tree endemism levels (Lima et al., 2020). ...
... To assess tree density per hectare for the different species along the climatic gradients within the Atlantic Forest domain, we used data from 623 tree phytosociological surveys, distributed from south to north of the Atlantic Forest biome plus forest intrusions into the neighboring Caatinga, Cerrado, and Pampa biomes (Oliveira-Filho et al., 2006). The surveys were chosen to embrace tree community data across considerable environmental gradients, both in terms of temperature and water deficit. ...
Article
Full-text available
Species under milder climates (e.g., warm and wet) tend to experience lower variability in temperature and rainfall regimes and might occur in narrower climatic ranges than species that tolerate harsher conditions (e.g., cold or dry climates). Thus, tree species that occur under harsh conditions should have a broader climatic range, being a small subset of the flora. Here, we assess the influence of climate on species distribution of 1138 tree species from the Atlantic Forest biodiversity hotspot. We investigate their range (or niche breadth), and the “center of gravity” index (or niche optima), along with gradients of mean annual temperature and climatic water deficit (CWD). We further identified those species associated with conditions on different ends of temperature and moisture gradients. We found a small subset of species occurring under colder temperatures or under drier conditions, and these species had a wider niche breadth. The warm or wet‐affiliated species had narrower ranges along with the temperature and the CWD gradients, respectively. Moreover, species affiliated to warm and those to moister conditions had greater densities near their occurrence limits, thus they may be more susceptible to climate changes. We conclude that global climate changes will affect the incidence and abundance distribution patterns of tree species along this threatened biodiversity hotspot, mainly those with narrow niches and within the limit of its distribution. Abstract in Portuguese is available with online material Here, we assess the influence of climate on species distribution of 1,138 tree species from the Atlantic Forest biodiversity hotspot. We found a small subset of species occurring under colder temperatures or under drier conditions, and these species had wider niche breadth. The warm or wet‐affiliated species had narrower ranges along with the temperature and the CWD gradients, respectively. We conclude that global climate changes will affect the incidence and abundance distribution patterns of tree species along this threatened biodiversity hotspot, mainly those with narrow niches and in the limit of its distribution.
... These areas encompass a wide variety of physiognomies, from tall forests to rocky outcrops with cacti and other succulents and shrubs (Prado, 2003;Moro et al., 2016;Queiroz et al., 2017). However, the relationships of some areas are still controversial, such as seasonal forests in southwestern Piauí, western Bahia and northern Minas Gerais (Santos et al., 2012a;Neves et al., 2015;Oliveira et al., 2019) that are sometimes considered Caatinga (Andrade-Lima, 1981;Queiroz et al., 2017;Silva et al., 2017a;Fernandes et al., 2020), sometimes Atlantic Forest (Olson et al., 2001;Silva and Casteleti, 2003;Oliveira-Filho et al., 2006;Neves et al., 2017). Because both designations have their justifications , I delimited the Caatinga in two ways ( Fig. 1): (1) broadly, and includes these seasonal forests (Caatinga sensu lato, hereafter Caatinga-SL), and (2) restricted, which excludes these areas (Caatinga sensu stricto, hereafter Caatinga-SS). ...
... While it is clear that the rainforest and savanna island-like enclaves should not be included in the Caatinga delimitation (Queiroz et al., 2017;Fernandes et al., 2020), the inclusion of other areas that have transitional or ambiguous identity may be controversial Neves et al., 2015;Miranda et al., 2018) and lacks further comprehensive studies encompassing both their flora and fauna. For example, the deciduous and semideciduous seasonal forests on the western and southern edges of the Caatinga-SS are sometimes considered as part of the Caatinga (Andrade-Lima, 1981;Queiroz et al., 2017;Silva et al., 2017a;Fernandes et al., 2020) and sometimes considered as part of the Atlantic Forest (Olson et al., 2001;Silva and Casteleti, 2003;Oliveira-Filho et al., 2006), and both may be valid . For this reason, I proposed two alternative delimitations for the Caatinga, one including (Caatinga-SL) and another excluding (Caatinga-SS) these areas, with a resulting difference of 54 more bird species in the broadest delimitation (Table S3). ...
Article
The Caatinga of northeastern Brazil is the largest nucleus of the seasonally dry tropical forest biome. Number of bird species in lists of the Caatinga often are quite different, mainly due to the existence of enclaves of other biomes embedded in this region, and whose species may or may not be included on those lists. While some consider such enclaves as disjunctions of other biomes, and therefore do not consider their species to be Caatinga-inhabitants, others consider the enclaves as part of the Caatinga and then include their species. In addition, the Caatinga has been widely used as a unit of analysis in biogeographical and macroecological studies, sometimes without a biologically meaningful delimitation. The need for a comprehensive review of these issues motivated me to propose a biologically meaningful checklist of the birds of the Caatinga and to explain why the enclaves of other biomes embedded within this region should not be considered part of the Caatinga. I argue that the previous checklists of birds, among other animals, of the Caatinga have been assembled in a biologically meaningless way, for which I provide a biologically meaningful alternative. This checklist comprises as many as 442 bird species for the Caatinga and is the first to comply with strict criteria as to its accessible, verifiable documentation.
... Este fato ressalta a necessidade de um tratamento específico para cada tipologia vegetacional. Entretanto, em estudos fitogeográficos, as caatingas têm sido tratadas como uma única unidade vegetacional integrante das florestas sazonalmente secas da região neotropical (PRADO, 2000;PENNINGTON et al., 2000;OLIVEIRA-FILHO et al., 2006). Caatinga (RODAL et al., 2008a). ...
Article
Este estudo foi realizado com o objetivo de analisar a composição florística do componente arbustivo-arbóreo em um fragmento de Caatinga pertencente à fazenda São Pedro, no município de Porto da Folha-SE, e verificar suas relações florísticas com outras áreas compostas por esta formação vegetacional. As coletas foram realizadas, em trilhas pré-existentes, nas bordas e no interior, durante 12 meses, visando conhecer a composição florística de toda a área de estudo. Foram selecionadas 34 áreas de Caatingas, distribuídas em estados com esta composição vegetacional para analisar a similaridade florística. Foram identificadas 69 espécies, distribuídas em 57 gêneros e 26 famílias botânicas. A similaridade florística entre o fragmento estudado e outras formações vegetacionais do Semiárido nordestino e Norte de Minas Gerais variou entre 2% e 34%. A análise de agrupamento resultou na formação de quatro grandes grupos (A, B, C e D) a 12,34% de similaridade, mostrando nítida separação entre o fragmento estudado, que está inserido no embasamento do cristalino, com as áreas instaladas em bacia sedimentar, sugerindo a existência de uma flora particular para cada uma dessas áreas.
... Mez. (Lauraceae), an abundant, common specie of riparian vegetation in the region (Oliveira-Filho et al., 2006) just after abscission and being air-dried (~20 • C, 5 days) (see Biasi et al., 2019). We weighted sets of 2.0 ± 0.1 g leaves and enclosed them in litter bags (15 × 20 cm, 0.5 mm mesh). ...
Article
Hyphomycetes are important aquatic organisms for organic matter decomposition, releasing inorganic nutrients to the environment. Understanding distribution patterns of hyphomycetes among different types of habitats can reveal important environmental characteristics that affect their assemblages and, as a consequence, can change stream functioning. We evaluated the effects of environmental predictors over hyphomycete assemblages in a field experiment in 12 subtropical Atlantic Forest streams. We identified 21 species of aquatic hyphomycetes, where Lunulospora curvula, Flagellospora curvula and Aquanectria submersa are dominant species, occurring in 10 streams, while Anguillospora sp., Campylospora parvula and Mycocentrospora acerina were found in only one stream. The variables dissolved oxygen, electrical conductivity and pH-influenced hyphomycete assemblages. Electrical conductivity acts as filter for spore output, while dissolved oxygen, electrical conductivity and phosphate are environmental filters for species richness. Our findings highlight the importance of environmental predictors over aquatic hyphomycete assemblages. The influence of environmental predictors over spore output and species richness may change stream functioning, since these organisms play an important role in leaf breakdown.
... Both the Amazonian TDF hypothesis and the Pleistocene Arc hypothesis seem plausible to some extent (Collevatti et al. 2013) since TDFs could experience different climatic conditions during the Pleistocene because of their wide distribution range (Oliveira-Filho et al. 2006). Besides, species diversity could have increased gradually over time if there was long-term stability of the vegetation community structure of TDFs (Collevatti et al. 2013). ...
Article
en Tropical dry forests (TDFs) are one of the most threatened ecosystems worldwide. Two hypotheses have been proposed to explain the origin of TDFs in South America: the Amazonian TDF hypothesis and the Pleistocene Arc hypothesis (PAH). There is a need to evaluate the distribution patterns of different organisms across the TDF distribution. We tested the following hypotheses: the species composition is determined by historical-evolutionary events, and therefore, the TDFs have a similar species composition as predicted by the PAH. Alternatively, the species composition is determined by recent ecological processes, and therefore, the TDFs have a sharing of species to their respective adjacent dominant habitat, with no support for the PAH. We expect that climatic factors drive the species richness, abundance and species dissimilarity (β-diversity) between TDFs and adjacent habitats across the latitudinal gradient. We sampled dung beetles across six Brazilian states in TDF fragments and adjacent dominant habitats and obtained the climatic conditions across the gradient. We used the β-diversity partition and generalised linear models to test our hypotheses. We sampled 8,625 dung beetles representing 102 species. Sorensen dissimilarity was higher among TDFs than between TDFs and adjacent habitats and was mostly due to the substitution of species. Annual mean temperature had a positive effect on abundance in TDFs but did not affect species richness. Species substitution (Podani’s approach) between TDFs and adjacent habitats decreased with mean diurnal range of temperature, while nestedness patterns (Baselga’s approach) increased with annual precipitation. Depending on the approach used (Baselga’s vs. Podani’s), we can obtain different results across the latitudinal gradient. The composition and structure of dung beetle assemblages in TDFs are mostly determined by more recent regional-to-local ecological processes since each TDF has a unique evolutionary history and a different dung beetle species composition. Our results do not support the Pleistocene Arc hypothesis. RESUMO pt As Florestas Tropicais Secas (FTSs) são um dos ecossistemas mais ameaçados do mundo. Duas hipóteses foram propostas para explicar a origem das FTSs na América do Sul: a hipótese da FTS Amazônica e a hipótese do Arco do Pleistoceno (PAH). É necessário avaliar os padrões de distribuição de diferentes organismos ao longa da distribuição da FTS. Testamos as seguintes hipóteses: a composição das espécies é determinada por eventos histórico-evolutivos e, portanto, as FTSs têm uma composição de espécies semelhante à prevista pela PAH. Alternativamente, a composição das espécies é determinada por processos ecológicos recentes e, portanto, as FTSs têm um compartilhamento de espécies com seus respectivos habitats dominantes adjacentes, sem suporte para a PAH. Esperamos que os fatores climáticos conduzam a riqueza de espécies, abundância e dissimilaridade de espécies (diversidade β) entre FTSs e habitats adjacentes em todo o gradiente latitudinal. Amostramos besouros escarabeíneos em seis estados brasileiros em fragmentos de FTS e habitats dominantes adjacentes e obtivemos as condições climáticas ao longo do gradiente. Usamos a partição de diversidade β e modelos lineares generalizados para testar nossas hipóteses. Amostramos 8.625 escarabeíneos que representam 102 espécies. A dissimilaridade de Sorensen foi maior entre FTSs do que entre FTSs e habitats adjacentes e foi principalmente devido à substituição de espécies. A temperatura média anual teve um efeito positivo sobre a abundância em FTSs, mas não afetou a riqueza de espécies. A substituição de espécies (abordagem de Podani) entre FTSs e habitats adjacentes diminuiu com a variação diurna média de temperatura, enquanto os padrões de aninhamento (abordagem de Baselga) aumentaram com a precipitação anual. Dependendo da abordagem utilizada (Baselga’s vs. Podani’s), podemos obter resultados diferentes em todo o gradiente latitudinal. A composição e a estrutura das assembleias de escarabeíneos em FTSs são principalmente determinadas por processos ecológicos regionais a locais mais recentes, uma vez que cada FTS tem uma história evolutiva única e uma composição de espécies diferente de escarabeíneos. Nossos resultados não suportam a hipótese do Arco do Pleistoceno.
... This predominance suggests that the current patchy distribution of the Caatinga Forest is the outcome of a long-term anthropogenic disturbance in the Atlantic Deciduous Forest, resulting in a fragmented landscape (Lôbo et al., 2011;Antongiovanni et al., 2018). Potentially, the Caatinga-Atlantic Forest transition would be originally composed of a continuum of dry (deciduous) forests, with high beta-diversity at the landscape scale (de Oliveira-Filho et al., 2006;DRYFLOR, 2016) yet poorly distinguishable on a macro scale. Despite the greater similarity to the Atlantic Forest, this transitional zone is not protected by the Atlantic Forest Law (IBGE, 2006), which regulates the biodiversity protection in this biome. ...
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Bioclimatic envelope models have been extensively used to predict the vegetation dynamics in response to climate changes. However, they are prone to the uncertainties arising from General Circulation Models (GCMs), classification algorithms and predictors, with low-resolution results and little detail at the regional level. Novel research has focused on the improvement of these models through a combination of climate and soil predictors to enhance ecological consistency. In this framework, we aimed to apply a joint edaphoclimatic envelope to predict the current and future vegetation distribution in the semiarid region of Brazil, which encompasses several classes of vegetation in response to the significant environmental heterogeneity. We employed a variety of machine learning algorithms and GCMs under RCP 4.5 and 8.5 scenarios of Coupled Model Intercomparison Project Phase 5 (CMIP5), in 1 km resolution. The combination of climate and soil predictors resulted in higher detail at landscape-scale and better distinction of vegetations with overlapping climatic niches. In forecasts, soil predictors imposed a buffer effect on vegetation dynamics as they reduced shifts driven solely by climatic drift. Our results with the edaphoclimatic approach pointed to an expansion of the dry Caatinga vegetation, ranging from an average of 16% to 24% on RCP 4.5 and RCP8.5 scenarios, respectively. The shift in environmental suitability from forest to open and dry vegetation implies a major loss to biodiversity, as well as compromising the provision of ecosystem services important for maintaining the economy and livelihoods of the world's largest semiarid population. Predicting the most susceptible regions to future climate change is the first step in developing strategies to mitigate impacts in these areas.
... A aplicação do índice de Morisita pode ser de grande utilidade na conservação e no manejo florestal, pois a sua interpretação produz subsídios importantes para a elaboração de modelos de restauração de áreas degradadas, enriquecimentos de remanescentes florestais, conservação de germoplasma e outras atividades. Para Oliveira-Filho et al. (2006) o processo de dispersão por si só, no entanto, não garante o sucesso da regeneração, pois é preciso que haja condições de solo e clima apropriadas para o estabelecimento das plantas jovens. Essas ações podem atuar em consonância com a capacidade de suporte do ambiente, tornandose mais efetivas e condizentes com os mecanismos reguladores da formação florestal e manejo Fontes, 2000). ...
... Este fato ressalta a necessidade de um tratamento específico para cada tipologia vegetacional. Entretanto, em estudos fitogeográficos, as caatingas têm sido tratadas como uma única unidade vegetacional integrante das florestas sazonalmente secas da região neotropical (PRADO, 2000;PENNINGTON et al., 2000;OLIVEIRA-FILHO et al., 2006). Caatinga (RODAL et al., 2008a). ...
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RESUMO. Estudar a composição e estrutura das vegetações além de contribuírem para a caracterização da vegetação local, serve ainda para fomentar discussões relacionadas à implementação de políticas públicas específicas para a preservação dos diferentes biomas brasileiros, especialmente em áreas de transição, onde essa classificação de domínios ainda é incipiente. O presente estudo teve como objetivo identificar a flora e a vegetação das florestas estacionais semidecíduas de transição, que se encontram nos municípios de Eliseu Martins, Pavussu e Canto do Buriti, no estado do Piauí, Brasil, e propor uma classificação fitossociológica para estas comunidades. Para isso, foram definidos 12 pontos de coleta de material botânico para a caracterização florística. A estrutura foi caracterizada seguindo o Protocolo de Avaliação Fitossociológica Mínima (PAFM) em uma área total de 1,02 hectares de área amostrada (17 parcelas). Foram estimados os parâmetros fitossociológicos usuais, e a diversidade foi calculada utilizando os índices: riqueza, similaridade e equabilidade. No total foram registrados um total de 105 táxons registrados, distribuídos em 30 famílias e 68 gêneros, sendo apenas 33 táxons completamente determinadas até o nível de espécie. Caesalpiniaceae foi a família que se destacou em riqueza específica (16 spp.), seguida por Papilionaceae (15 spp.), Malpighiaceae (7 spp.) e Erythroxylaceae (4 spp.). Na análise fitossociológica, foram amostradas 87 espécies, sendo que apenas quatro tiveram Frequência Total (de 100%). As espécies Combretum glaucocarpum Mart. e Campomanesia sp. foram as com maiores índices de Importância e Densidades Absolutas. Quanto ao endemismo, apesar da presença de espécies dos Domínios Amazônico, Cerrado e Mata Atlântica, a vegetação é predominantemente indicadora do Bioma CAATINGA. ABSTRACT. Understing the vegetation composition and structure, besides contributing to the characterization of the regional vegetation, it also provides discussions about the implementation of public policies to environment preservation of different Brazilian biomes, especially in transition regions, where this classification of domains is remaining incipient. This study aims to identify flora and structure of the Transitional Semideciduous seasonal forests, located in Eliseu Martins, Pavussu and Canto do Buriti, in the state of Piauí, Brazil, and proposing a phytosociological classification for these communities. Was selected 12 points to realize the collection of the botanical material to floristic characterization. The structure was characterized following the Minimum Phytosociological Assessment Protocol (MPAP) in a total area of 1.02 hectares (17 plots). The usual phytosociological parameters were estimated, and the diversity was calculated using the indices: richness, similarity, and equitability. In total, 105 taxa were registered, distributed to 30 families and 68 genera, with only 33 taxa completely determined down to the species level. Caesalpiniaceae was the richest family (16 spp.), followed by Papilionaceae (15 spp.), Malpighiaceae (seven spp.), and Erythroxylaceae (four spp.). In the phytosociological analysis, 87 species were sampled, however, only four had Frequency of 100%. The Combretum glaucocarpum Mart. and Campomanesia sp. species were the ones with the highest indices of Importance and Absolute Densities. As for endemism, despite the presence of species from the Amazon, Cerrado, and Atlantic Forest domains, the vegetation is predominantly indicative of the CAATINGA Biome.
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Estimates of species extinction due to human impact on tropical forests have previously been based on the relationship between species number and area. Here we use a different approach to estimate loss of tree species in the Atlantic forest of northeast Brazil. We evaluate the characteristics of plant species, their avian dispersers and the distribution of the forest remnants on the landscape to estimate that about 33.9% of tree species in this region will become extinct on a regional scale. Because northeast Brazil is the most threatened sector of South American Atlantic forest, our results highlight the need to change the current conservation paradigm for this region. Rather than focus on the creation of isolated reserves in any medium-to-large forest remnant, a bioregional planning approach is urgently required to rescue this unique biota from extinction.
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