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Tropical moist forests and savannas are iconic biomes. There is, however, a third principal biome in the lowland tropics that is less well known: tropical dry forest. Discussions on responses of vegetation in the tropics to climate and land-use change often focus on shifts between forests and savannas, but ignore dry forests. Tropical dry forests are distinct from moist forests in their seasonal drought stress and consequent deciduousness and differ from savannas in rarely experiencing fire. These factors lead tropical dry forests to have unique ecosystem function. Here, we discuss the underlying environmental drivers of transitions among tropical dry forests, moist forests and savannas, and demonstrate how incorporating tropical dry forests into our understanding of tropical biome transitions is critical to understanding the future of tropical vegetation under global environmental change.
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published: 24 July 2018
doi: 10.3389/fevo.2018.00104
Frontiers in Ecology and Evolution | 1July 2018 | Volume 6 | Article 104
Edited by:
Colin Osborne,
University of Sheffield,
United Kingdom
Reviewed by:
Imma Oliveras,
University of Oxford, United Kingdom
Liam Jude Langan,
Senckenberg Biodiversität und Klima
Forschungszentrum (SBiK-F),
Kyle G. Dexter
Specialty section:
This article was submitted to
Biogeography and Macroecology,
a section of the journal
Frontiers in Ecology and Evolution
Received: 01 May 2018
Accepted: 29 June 2018
Published: 24 July 2018
Dexter KG, Pennington RT,
Oliveira-Filho AT, Bueno ML, Silva de
Miranda PL and Neves DM (2018)
Inserting Tropical Dry Forests Into the
Discussion on Biome Transitions in the
Tropics. Front. Ecol. Evol. 6:104.
doi: 10.3389/fevo.2018.00104
Inserting Tropical Dry Forests Into
the Discussion on Biome Transitions
in the Tropics
Kyle G. Dexter 1,2
*, R. Toby Pennington 2,3 , Ary T. Oliveira-Filho 4, Marcelo L. Bueno 5,
Pedro L. Silva de Miranda 1and Danilo M. Neves 4,6
1School of GeoSciences, University of Edinburgh, Edinburgh, United Kingdom, 2Royal Botanic Garden Edinburgh,
Edinburgh, United Kingdom, 3Geography, University of Exeter, Exeter, United Kingdom, 4Departamento de Botânica,
Universidade Federal de Minas Gerais, Belo Horizonte, Brazil, 5Laboratório de Ecologia e Evolução de Plantas, Universidade
Federal de Viçosa, Viçosa, Brazil, 6Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, AZ,
United States
Tropical moist forests and savannas are iconic biomes. There is, however, a third
principal biome in the lowland tropics that is less well known: tropical dry forest.
Discussions on responses of vegetation in the tropics to climate and land-use change
often focus on shifts between forests and savannas, but ignore dry forests. Tropical dry
forests are distinct from moist forests in their seasonal drought stress and consequent
deciduousness and differ from savannas in rarely experiencing fire. These factors lead
tropical dry forests to have unique ecosystem function. Here, we discuss the underlying
environmental drivers of transitions among tropical dry forests, moist forests and
savannas, and demonstrate how incorporating tropical dry forests into our understanding
of tropical biome transitions is critical to understanding the future of tropical vegetation
under global environmental change.
Keywords: tropical dry forest, tropical moist forest,savanna, biomes, fire, soil fertility, water stress, deciduousness
Predicting vegetation change in the tropics depends on understanding the drivers of transitions
among major vegetation types, or biomes. Climatic factors, such as mean annual precipitation
(MAP) and its seasonality are of obvious importance, but edaphic factors, and disturbance via
fire, humans and herbivores also play key roles. Recent large-scale studies across the tropics have
focused on transitions between forest and savanna (Hirota et al., 2011; Staver et al., 2011; Oliveras
and Malhi, 2016; Xu et al., 2016; Langan et al., 2017). While there is value to simplifying vegetation
concepts in the tropics, we believe the simplification used by these authors in defining “forest”
goes perhaps one step too far. There are two principal kinds of forest in the lowland tropics, moist
forests and dry forests. With very few exceptions (e.g., Hirota et al., 2010; Lehmann et al., 2011),
studies of biome transitions in the tropics have either failed to distinguish them, or have completely
ignored dry forests, focusing solely on moist forests when using the term “forest.” The aim of
this perspective is to discuss biome transitions in the tropics and their underlying drivers, while
including dry forests in the discussion. We focus on transitions among savanna, moist forest and
dry forest, the three biomes in the lowland tropics with a substantial tree component.
Tropical moist forest and savanna are relatively well understood at a global scale compared to
tropical dry forests (Pennington et al., 2018). Moist forest is tall, multi-stratal, and with a closed
canopy. Tropical moist forests include tropical rain forests, as well as forests with lower rainfall
where soil moisture is maintained throughout the year, via edaphic factors such as proximity to
Dexter et al. Tropical Dry Forests and Biome Transitions
rivers, or water recycling, allowing most trees to be evergreen
(Guan et al., 2015). The understory is often dominated by
saplings of taller-statured tree species, although small tree and
shrub species are present. Terrestrial forbs and grasses are
a minor component of diversity and biomass. Savanna is a
more open environment, where tree species are present, but
individuals do not form a closed canopy. There is a significant
understory grass component, which is flammable, and fires are
common. Tree species that occur in savannas are adapted to these
recurring fires (Simon and Pennington, 2012), and regular fires
are necessary for the maintenance of savanna biodiversity (Parr
et al., 2014; Durigan and Ratter, 2016; Abreu et al., 2017). Some
savannas in the paleotropics (e.g., miombo woodlands in Africa,
deciduous dipterocarp forests in southeast Asia) are generally
referred to as dry forests, but we consider them as savannas given
that they have a grassy understory and experience regular fire
(Ratnam et al., 2011; Dexter et al., 2015; Pennington et al., 2018).
Tropical dry forests vary greatly in structure, from tall, closed
canopy forest to short scrub vegetation, occasionally not forming
a closed canopy, especially in drier areas (Pennington et al.,
2000). They are distinct from savanna in not having a significant
grass component and not experiencing regular fires (Murphy and
Lugo, 1986; Gentry, 1995). In fact, regular fires would be lethal
for many of the characteristic life forms and taxa of tropical
dry forest (e.g., cacti; Mooney et al., 1995). This is not to say
that dry forests never experience fires. Even moist forests can
experience fire under extreme drought conditions (Aragão et al.,
2016). Rather, damaging fire is sufficiently rare in dry forests
such that fire-intolerant species can persist in the landscape
as metapopulations (Hanski, 1998). The exact threshold of fire
return interval or intensity involved in the tropical dry forest -
savanna transition is poorly understood and likely to vary with
the broader environmental context (e.g., soil fertility and annual
precipitation; Hoffmann et al., 2012; Murphy and Bowman,
2012), and we suggest that this should be a priority for further
study. Tropical dry forest is distinct from moist forest in its
seasonal drought stress, which leads many tree species to lose
their leaves in the dry season (Reich and Borchert, 1984; Murphy
and Lugo, 1986). The combination of seasonal drought stress
and lack of fire leads to ecosystem function in dry forests that is
markedly different from savannas or moist forests, which justifies
their distinction as a unique biome.
We focus this review on continental lowland tropical South
America (LTSA), where we have conducted most of our research.
In a recent study (Silva de Miranda et al., in press), we
used an unsupervised classification, or hierarchical clustering,
of sites based on their tree species composition (see inset in
Figure 1), followed by interpretation of the resulting cluster
using site information on vegetation physiognomy (savanna vs.
forest) and leaf flush regime (evergreen vs. semideciduous vs.
deciduous) to delimit and map biomes across LTSA east of the
Andes (Figure 1). Moist forests fell in two major groups in the
cluster and occurred in two large geographic blocks, one in the
Amazon basin and another along the Atlantic coast of Brazil.
Semideciduous forests did not form a distinct group in the
cluster and were instead mixed with evergreen, moist forest sites.
Semideciduous forests are often found in drier regions, where
they occur along rivers, lake margins and submontane areas
with orographic precipitation. Savanna formed a single group in
the cluster and was most prevalent in central Brazil, in an area
commonly termed the Cerrado.
Tropical dry forest also formed a single group in the
cluster, which was comprised almost entirely of forest sites
with deciduous phenology. In LTSA, the largest block of dry
forest occurs in the Caatinga region of northeast Brazil. The
Caatinga has been referred to as a biome (Hirota et al., 2010;
Instituto Brasileiro de Geografia e Estatística, 2012), although
as a region it contains non-dry forest habitat (e.g., patches
of savanna). Further, tropical dry forest is found outside of
the Caatinga, in patches throughout the Cerrado, in an area
spanning the Pantanal and Chiquitania in Bolivia (Figure 1),
and scattered more widely across the Neotropics (DRYFLOR,
2016). While Silva de Miranda et al. (in press) broadly assessed
climatic overlaps amongst biomes, they did not focus on the
environmental drivers of transitions between individual biomes.
That is the goal of the present manuscript.
A common view of dry forests in the tropics is that they
are transitional between savannas and moist forests along
precipitation gradients (e.g., Whittaker, 1970; Malhi et al., 2009).
If, however, we examine how the sites featured in Figure 1 are
distributed over variation in MAP (Figure 2), a more complex
picture emerges. Moist forests do occur under wetter conditions
than savanna, but dry forests are largely found under drier
conditions. Below 1,000 mm MAP, savanna quickly disappears
and dry forest becomes the only tree-dominated vegetation type.
The largest area of these arid dry forests is found in the Caatinga
region of northeast Brazil. As discussed above, a key distinction
between savanna and dry forest is the regularity of fire, and in
these dry conditions, there is not sufficient biomass build-up to
sustain regular fires (Van Der Werf et al., 2008). In particular,
this reflects the relative lack of grasses. The tree species that are
present are able to tolerate severe drought, but they do not invest
in adaptations for fire, such as thick bark or underground stems,
characteristic of savanna species (Simon and Pennington, 2012).
There is also extensive occurrence of the dry forest biome
under the same precipitation conditions as savanna. These
are dry forests found in the Cerrado region and around the
Pantanal and Chiquitania regions of Brazil and Bolivia. Within
the Cerrado, dry forests are known to occur on and around
calcareous outcrops, where soils have higher phosphorus and
base cation concentrations (Ratter et al., 1978; Furley and
Ratter, 1988; Oliveira-Filho and Ratter, 2002; Neves et al.,
2015). On these soils, trees can grow more quickly, have better
chances of escaping the “fire trap” and are more likely to
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Dexter et al. Tropical Dry Forests and Biome Transitions
FIGURE 1 | Map of biomes across lowland tropical South America (after Silva de Miranda et al., in press). Inset shows results of unsupervised clustering of 3,331
georeferenced sites based on tree species composition. The biomes of the major groups in the cluster were determined based on vegetation physiognomy and leaf
flush regime. Semideciduous forests were placed in the moist forest biome based on similarities in tree species composition evident in the cluster by Silva de Miranda
et al. (in press), but are distinguished here. We exclude sites south of 23S latitude or above 1,000 m elevation.
form a closed canopy (Hoffmann et al., 2009). There can be
positive feedbacks between tree growth, grass exclusion and fire
mitigation that leads to a forested vegetation (Hoffmann et al.,
2012; Silva et al., 2013; Pausas and Dantas, 2017). Calcareous
outcrops in the Cerrado also have poorly developed, shallow
soils, and vegetation occurring on them may experience greater
drought stress than surrounding vegetation, thus making them
similar to the arid dry forests, which lack fire because of
insufficient fuel build-up. However, it is likely that soil fertility
is relevant for the presence of dry forests around calcareous
outcrops as a different vegetation, cerrado rupestre, which is
floristically related to savanna vegetation, is found on non-
calcareous outcrops in the Cerrado (Ribeiro and Walter, 1998).
Whichever factor is more important (soils with high fertility
or low water-holding capacity), it is evident that the same
drought-tolerant, fire-intolerant tree species and lineages that
dominate vegetation in the arid Caatinga are also found in dry
forest patches in the moister Cerrado (Prado and Gibbs, 1993;
Neves et al., 2015; DRYFLOR, 2016; Silva de Miranda et al.,
in press).
Dry forest and savanna vegetation also intermingle in the
Chiquitania region of Bolivia, but here dry forest predominates
and savanna occurs in patches, which may be because soils in the
Chiquitania are more fertile on average than in the Cerrado (Silva
de Miranda et al., in press). Some of the savannas that are present
in the Chiquitania region may represent dry forest that has been
degraded by logging or anthropogenic fire (Devisscher et al.,
2016), highlighting that human land-use patterns can readily
drive transitions between dry forest and savanna. However, “old-
growth savannas” that would exist independent of anthropogenic
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Dexter et al. Tropical Dry Forests and Biome Transitions
FIGURE 2 | Frequency distribution of sites in Figure 1 over a gradient in mean annual precipitation. For heuristic purposes, semideciduous and moist forests are
distinguished although they may represent the same biome.
influence are also clearly present in Bolivia (Power et al., 2016;
Veldman, 2016).
If fire is excluded from savanna vegetation, it first converts
to a formation with a higher percentage of tree canopy cover
termed “cerradão” (Durigan and Ratter, 2006, 2016), which
translates from the Portuguese as “big cerrado” and is generally
considered as a forest. Cerradão shares some tree species with
tropical dry forest and comparatively few with semideciduous
and evergreen moist forests (Bueno et al., 2018), even though
the latter are present near savanna vegetation along river
courses and lake margins that have year-round water availability
(Ribeiro and Walter, 1998). Tree species from semideciduous
and evergreen forests may be less likely to immigrate into
cerradão than typical dry forest tree species because they are
not adapted to seasonal drought. While cerradão is initially
comprised of fire-adapted tree species from the cerrado, dry
forest tree species colonize this environment if propagules are
available, fire remains absent and soils are sufficiently fertile.
These dry forest tree species may eventually outcompete cerrado
tree species, since they do not invest in fire defense (Ratajczak
et al., 2017), and in the prolonged absence of fire, cerradão
may transition to a dry forest if there are positive feedback
cycles between forest vegetation, lack of fire and soil fertility
(Silva et al., 2013; Pellegrini et al., 2014, 2018). However, if
the underlying soils remain poor and/or if there are high
aluminum concentrations in the soil that do not attenuate
over time, then cerrado tree species, which are adapted to
infertile, aluminum-rich soils may continue to dominate the
Overall, the savanna-dry forest transition is distinct from
the savanna-moist forest transition in two key ways: (1) the
contrasting role of water availability (lowest in dry forest,
intermediate in savanna and highest in moist forest) and (2) the
potentially critical importance of soil fertility for the savanna-
dry forest transition (savanna and moist forests are similar in
generally having infertile, acidic soils).
Both of these biomes are forest, but they function in distinct
ways. Dry forests are found in areas with marked precipitation
seasonality, which leads most species to lose their leaves
during the dry season and has significant implications for
nutrient cycling (Reich and Borchert, 1984; Murphy and Lugo,
1986). Many moist forests also experience seasonality in water
availability (e.g., in the southern and eastern Amazon), but the
dry season is three months or less and subsurface water remains
available to trees (Guan et al., 2015). The systems also differ in the
rate at which they accrue and cycle carbon, with trees in moist
forests growing more quickly and storing more total carbon
(Murphy and Lugo, 1986; Poorter et al., 2017). There are often
differences in soil fertility, with dry forests occurring on more
fertile soils, which facilitates their ability to shed their leaves as
they can readily afford to grow new ones. However, high rainfall
in moist forests results in nutrient leaching, and this correlation
between soil fertility and biome identity may be due to overriding
climatic factors (Webb, 1968; Hall and Swaine, 1976).
In LTSA, there are multiple areas of contact between the
moist and dry forest biomes (Figure 1). One transition zone is in
northeastern Brazil, where evergreen Atlantic forest on the coast
transitions to dry forest in the arid Caatinga. In between the two
lies a band of semideciduous forests. Another transition zone is
found in the Chiquitania region of eastern Bolivia and adjacent
areas of Brazil, where there is a gradual transition over 200+km
of geographic distance, largely covered by semideciduous forest.
We suggest that transitions between dry forest and moist forest
are primarily mediated by water availability and that intermediate
states are possible in zones of intermediate water availability
(Oliveira-Filho and Fontes, 2000; Oliveira-Filho et al., 2006). This
contrasts with transitions between savanna and moist forest that
can be more abrupt and may represent alternative stable states
(although see Lloyd and Veenendaal, 2016).
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Dexter et al. Tropical Dry Forests and Biome Transitions
Previous studies have variously grouped semideciduous forests
with the dry forest biome (Murphy and Lugo, 1986; Pennington
et al., 2000) or the moist forest biome (DRYFLOR, 2016; Silva de
Miranda et al., in press). In fact, as discussed above, these forests
may be transitional between the two. Semideciduous forests in
LTSA have few endemic tree species and instead contain tree
species associated with the dry or moist forest biomes (Oliveira-
Filho and Fontes, 2000). As moist forests contain many more
tree species than dry forests (Esquivel-Muelbert et al., 2017), we
suggest that they may contribute more species to semideciduous
forests simply via mass effects (Shmida and Wilson, 1985), and
this may be why they group with moist forests in clustering
analyses based on presence versus absence of tree species (as
in Silva de Miranda et al., in press). If abundance information
were to be taken into account (e.g., via inventory plot data), we
hypothesize that semideciduous forests may cluster with moist
or dry forest based on the proportion of individuals belonging
to moist versus dry forest tree species. It is clear that future
comparative studies across dry, moist and semideciduous forests
are needed to understand their origins and how they compare
in terms of ecosystem function. Their geographically variable
species composition and lack of endemic species suggests that
semideciduous forests may have been independently and recently
assembled in different ecotonal areas.
As in South America, moist forest—savanna transitions have
been studied extensively on other continents, but transitions to
dry forest have received less attention. This is partly because it is
unclear where dry forest exists outside of the Neotropics (Lock,
2006; Dexter et al., 2015; Pennington et al., 2018). In a recent
study, Linder (2014) delimited and described the main “floras”
of Africa, which are large-scale units of vegetation that have
a distinct evolutionary and biogeographic history and differ in
their present-day plant taxonomic composition. Linder did not
assign the term “biome” to these vegetation units, although his
“floras” correspond to several previously defined biomes. There
is a “savanna flora,” which readily corresponds to the savanna
biome, and a “lowland forest flora” that largely corresponds to
the moist forest biome. Linder postulated an “arid flora” that
is most evident in the Horn of Africa (Somalia, Ethiopia and
northern Kenya), but is also present in arid regions of Angola
and Namibia. The distribution of this “arid flora” largely overlaps
the distribution of the “succulent biome” in Africa, as proposed
by Schrire et al. (2005). The “arid flora” or “succulent biome” is
similar to dry forest in arid regions of the Neotropics in that there
is not adequate water availability to allow for sufficient biomass
build-up to sustain regular fires. Thus, as in the Neotropics,
water availability may be one environmental factor that underlies
transitions between savanna and dry forest in Africa.
Soil fertility is another significant factor that has been shown
to underlie savanna-dry forest transitions in the Neotropics.
An important question for future research in Africa is whether,
amongst its great expanses of savanna, there is distinct vegetation
that does not regularly burn, is found on more fertile soils and
shows greater floristic similarity with the “arid flora” of Linder
(2014) than it does with surrounding savanna vegetation. It
may be that in Africa, a higher abundance of large herbivores,
including elephants, favors grasses over trees, leading to a more
open savanna vegetation with more frequent fires, even in areas
of higher soil fertility (Charles-Dominique et al., 2016; Pellegrini
et al., 2017). If dry forests are not found on fertile soils in
more mesic areas of Africa, there may not be moist forest-dry
forest transitions on this continent, because the areas mapped as
belonging to the “arid flora” or “succulent biome are completely
separated from moist forest regions by large areas of savanna
(Schrire et al., 2005; Linder, 2014).
In the tropical regions of continental Asia and in Malesia,
moist forest is the predominant vegetation type, although drier
forest formations are present (e.g., deciduous forests in the
Western Ghats and dry dipterocarp forests in Indochina). As
we have discussed elsewhere (Dexter et al., 2015; Pennington
et al., 2018), the majority of these drier forest formations have
a significant grassy component in the understory, burn regularly
and may be better considered as savannas (Ratnam et al., 2011).
The succulent biome of Schrire et al. (2005) is mapped as present
in arid regions of northwest India and extending across the coast
of Pakistan and Iran to the Arabian Peninsula. Thus, an arid
form of the dry forest biome may occur in Asia, as in Africa, and
water availability may underlie transitions between vegetation in
seasonally dry areas that regularly burns (what we term savanna)
and that does not regularly burn (what we term dry forest).
As with Africa, future research in Asia should assess if there
are vegetation formations in seasonally dry, yet not arid, areas
that: (1) are found on fertile soils, (2) do not regularly burn
and (3) show greater floristic similarity with arid areas than
with surrounding vegetation that does regularly burn. This will
help determine if soil fertility is also important in understanding
savanna-dry forest transitions in Asia, as it is in the Neotropics.
The aim of this perspective has been to bring the tropical
dry forest biome into discussions of biome transitions in the
tropics. Previous studies of tropical biome transitions have largely
focused on forest-savanna transitions, with all forests being
considered as a single biome. In fact, there are many kinds of
forests in the tropics, some of which are distinct from each other
in species composition and ecosystem function and represent
different biomes (i.e., dry vs. moist forest) and others which are
more difficult to classify (e.g., semideciduous forests, cerradão).
Water availability is a key factor underlying tropical biome
transitions. While forests are often thought to occur under wetter
conditions than savannas, tropical dry forest is actually more
prevalent in areas of lower water availability (<1,000 mm MAP).
Meanwhile, soil fertility, which has received limited attention in
studies of biome transitions, is also critical in the Neotropics,
and merits future research on other continents. More generally,
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Dexter et al. Tropical Dry Forests and Biome Transitions
recognizing tropical dry forest as a distinct biome within the
tropics should improve the accuracy of modeling studies that
aim to predict the future of tropical vegetation and ecosystem
function under global environmental change.
KD and RP wrote the first draft of the manuscript and
contributed to revising the manuscript. AO-F, MB, PS and DN
contributed to revising and improving the manuscript.
KD, RP, and DN thank the Natural Environment Research
Council (UK) for funding via NE/I028122/1. KD thanks
the Leverhulme Trust for supporting him during the time
this study was completed via an International Academic
Fellowship. PS thanks the Coordenação de Aperfeiçoamento
de Pessoal de Nível Superior–Brazil (CAPES) for support via
the Science without Borders Programme (grant 99999.013197/
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Conflict of Interest Statement: The authors declare that the research was
conducted in the absence of any commercial or financial relationships that could
be construed as a potential conflict of interest.
Copyright © 2018 Dexter, Pennington, Oliveira-Filho, Bueno, Silva de Miranda and
Neves. This is an open-access article distributed under the terms of the Creative
Commons Attribution License (CC BY). The use, distribution or reproduction in
other forums is permitted, provided the original author(s) and the copyright owner(s)
are credited and that the original publication in this journal is cited, in accordance
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Frontiers in Ecology and Evolution | 7July 2018 | Volume 6 | Article 104
... 2.6 Ma to 11,000 years) and present climatic and edaphic conditions constrain the establishment of new species, contributing to distinctive species assemblages within the continents (6,10,11). Climatic seasonality has been suggested to be the most important environmental condition defining the main boundaries of Neotropical bioregions (6), the floristic distinctions between neighboring evergreen, deciduous and semideciduous forests (12,13), and the dominant functional traits of moist forests (14). Biogeographical patterns in floristic variation have been documented for old-growth forests (15)(16)(17)(18), but not for the subset of species that colonize successional forests in human-modified landscapes across the Neotropics. ...
... The early successional communities in these regions have a strongly distinct genera and species composition compared to neighboring less-dry, moist, and wet forests (Fig. 1, B and D, and fig. S6), in agreement with the separate phylogenetic origin of dry forests (13,50) and the high levels of endemism found in the Caatinga (17). This supports the idea that a strong deficit in water availability is an important environmental filter defining the floristic composition of deciduous dry forests (13), thus restricting the similarity with neighboring less-dry forests (Fig. 1D). ...
... S6), in agreement with the separate phylogenetic origin of dry forests (13,50) and the high levels of endemism found in the Caatinga (17). This supports the idea that a strong deficit in water availability is an important environmental filter defining the floristic composition of deciduous dry forests (13), thus restricting the similarity with neighboring less-dry forests (Fig. 1D). Conversely, forests under intermediate climatic conditions may be more likely to exchange species and therefore be more susceptible to biotic homogenization. ...
... 2.6 Ma to 11,000 years) and present climatic and edaphic conditions constrain the establishment of new species, contributing to distinctive species assemblages within the continents (6,10,11). Climatic seasonality has been suggested to be the most important environmental condition defining the main boundaries of Neotropical bioregions (6), the floristic distinctions between neighboring evergreen, deciduous and semideciduous forests (12,13), and the dominant functional traits of moist forests (14). Biogeographical patterns in floristic variation have been documented for old-growth forests (15)(16)(17)(18), but not for the subset of species that colonize successional forests in human-modified landscapes across the Neotropics. ...
... The early successional communities in these regions have a strongly distinct genera and species composition compared to neighboring less-dry, moist, and wet forests (Fig. 1, B and D, and fig. S6), in agreement with the separate phylogenetic origin of dry forests (13,50) and the high levels of endemism found in the Caatinga (17). This supports the idea that a strong deficit in water availability is an important environmental filter defining the floristic composition of deciduous dry forests (13), thus restricting the similarity with neighboring less-dry forests (Fig. 1D). ...
... S6), in agreement with the separate phylogenetic origin of dry forests (13,50) and the high levels of endemism found in the Caatinga (17). This supports the idea that a strong deficit in water availability is an important environmental filter defining the floristic composition of deciduous dry forests (13), thus restricting the similarity with neighboring less-dry forests (Fig. 1D). Conversely, forests under intermediate climatic conditions may be more likely to exchange species and therefore be more susceptible to biotic homogenization. ...
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Forests that regrow naturally on abandoned fields are important for restoring biodiversity and ecosystem services, but can they also preserve the distinct regional tree floras? Using the floristic composition of 1215 early successional forests (≤20 years) in 75 human-modified landscapes across the Neotropic realm, we identified 14 distinct floristic groups, with a between-group dissimilarity of 0.97. Floristic groups were associated with location, bioregions, soil pH, temperature seasonality, and water availability. Hence, there is large continental-scale variation in the species composition of early successional forests, which is mainly associated with biogeographic and environmental factors but not with human disturbance indicators. This floristic distinctiveness is partially driven by regionally restricted species belonging to widespread genera. Early secondary forests contribute therefore to restoring and conserving the distinctiveness of bioregions across the Neotropical realm, and forest restoration initiatives should use local species to assure that these distinct floras are maintained.
... Such approaches might underestimate the importance of local adaptations or rather the importance of forest diversity. Under the common assumption that trees operate at their maximum performance by optimizing their trade-off affected trait values in a competitive community (40), any significant deviation from common local environmental conditions poses a stress to the established local community (37,94). It is not that this community can pick from solutions found in the overall pool of trait values in the whole Amazon. ...
... Determining these relative effects is a promising avenue for future research as field-based estimates of local tree adaptations are starting to become available. Specifically, we see several arguments bringing our approach in line with current literature: 1) Adaptation to climatic and environmental conditions is key to survival of certain tree species and, as such, affects the resilience to climatic changes in the Amazon rainforest (17,32); 2) those adaptive capacities have in particular been related to tree strategies to cope with droughts (35,71), but it has been found that tree mortality risks increase when the trees experience climatic conditions outside their adaptive limits (59,94); and 3) it has been found that even occasional strong droughts have long-lasting impacts on drought-induced tree mortality (57), and vegetation conversions have been observed in multiple biomes worldwide including the Amazon (59,66). ...
Tipping elements are nonlinear subsystems of the Earth system that have the potential to abruptly shift to another state if environmental change occurs close to a critical threshold with large consequences for human societies and ecosystems. Among these tipping elements may be the Amazon rainforest, which has been undergoing intensive anthropogenic activities and increasingly frequent droughts. Here, we assess how extreme deviations from climatological rainfall regimes may cause local forest collapse that cascades through the coupled forest–climate system. We develop a conceptual dynamic network model to isolate and uncover the role of atmospheric moisture recycling in such tipping cascades. We account for heterogeneity in critical thresholds of the forest caused by adaptation to local climatic conditions. Our results reveal that, despite this adaptation, a future climate characterized by permanent drought conditions could trigger a transition to an open canopy state particularly in the southern Amazon. The loss of atmospheric moisture recycling contributes to one-third of the tipping events. Thus, by exceeding local thresholds in forest adaptive capacity, local climate change impacts may propagate to other regions of the Amazon basin, causing a risk of forest shifts even in regions where critical thresholds have not been crossed locally.
... Across these five regions, we identified 80 ecoregions based on three criteria that correspond to polygons generated by the World Wide Fund for Nature: (1) whether the site was identified as TDF by Olson et al. (2001) and revised by Dinerstein et al. (2017); (2) whether a site Table S2). Unavoidably, we included disturbance-driven vegetation types, such as savanna, and geomorphologically determined vegetation types, such as riparian forests, which transition into adjacent TDFs that shape TDL dynamics (Dexter et al., 2018). To refine our understanding of environmental controls on TDFs, we compiled diverse climatic and edaphic information with which to quantify variation in NDVI across the five major biogeographical regions and the 80 ecoregions. ...
Spatial patterns in resource supply drive variability in vegetation structure and function, yet quantification of this variability for tropical dry forests (TDFs) remains rudimentary. Several climate‐driven indices have been developed to classify and delineate TDFs globally, but there has not been a climo‐edaphic synthesis of these indices to assess and delineate the extent of TDFs. A statistical climo‐edaphic synthesis of these indices is therefore required. Pantropical. Modern. Vascular plants. We assembled most known prior descriptions of TDFs into a single data layer and assessed statistically how the TDF biome, which we call tropical dry landscapes (TDLs) composed of forest and non‐forest vegetation, varied with respect to the normalized difference vegetation index (NDVI) sensed by MODIS (250 m pixel resolution). We examined how the NDVI varied with respect to mean annual temperature (MAT) and rainfall (MAR), precipitation regime, evapotranspiration and the physical, chemical and biological properties of TDL soils. Overall, the NDVI varied widely across TDLs, and we were able to identify five principal NDVI categories. A regression tree model captured 90% of NDVI variation across TDLs, with 14 climate and soil metrics as predictors. The model was then pruned to use only the three strongest metrics. These included the Lang aridity index, total evapotranspiration (ET) and MAT, which aligned with identified NDVI thresholds and accounted for 70% of the variation in NDVI. We found that across a global TDL distribution, ET was the strongest positive predictor and MAT the strongest negative predictor of the NDVI. The remote sensing‐based approach described here provides a comprehensive and quantitative biogeographical characterization of global TDL occurrence and the climatic and edaphic drivers of these landscapes.
... The Caatinga biome is part of the large group of Seasonally Dry Tropical Forests (FTSS) and comprises one of its most varied vegetation groups (Costa et al. 2015;Dexter et al. 2018) representing an ecosystem of considerable biological The images were grouped on plates using the Inkscape software and the maps were made using ArcGis Pro Student license (2021), using the Geographic Coordinate System, SAD 69. ...
The present study consists of a synopsis of Fabaceae in the Pico do Jabre State Park, Paraíba state, northeastern Brazil. 36 species were recorded, subordinate to 24 genera and three subfamilies (Caesalpinioideae, Cercidoideae and Papilionoideae). The genera with the largest number of species were: Senna (six spp.); Mimosa and Chamaecrista (four spp. each) and Centrosema (two spp.). The other genera were represented by one species each. The treatment includes an identification key, information on geographic distribution, flowering and/or fruiting data and images for the species in the area. Resumo O presente estudo consiste numa sinopse de Fabaceae no Parque Estadual Pico do Jabre, estado da Paraíba, nordeste do Brasil. Foram registradas 36 espécies subordinadas a 24 gêneros e três subfamílias (Caesalpinioideae, Cercidoideae e Papilionoideae). Os gêneros com maior número de espécies foram: Senna (seis spp.); Mimosa e Chamaecrista (quatrospp. cada) e Centrosema (duas spp.). Os demais gêneros foram representados por uma espécie cada. O tratamento inclui uma chave de identificação, informações sobre distribuição geográfica, dados de floração e/ou frutificação e imagens para as espécies da área.
... The prevalence of biome-specialist species in the tropical deciduous woodlands was pointed out in previous works on mammals (Cantalapiedra et al., 2011;Hernández Fernández et al., 2022;Hernández Fernández & Vrba, 2005;Moreno Bofarull et al., 2008). Even though this biome cannot be considered a climatic extreme, it is a markedly heterogeneous environment whose historical dynamic is closely associated with rainforest fluctuations (Dexter et al., 2018;Haffer, 2008;Hoorn et al., 2010). ...
The resource‐use hypothesis, proposed by E.S. Vrba, states that habitat fragmentation caused by climatic oscillations would affect particularly biome specialists (species inhabiting only one biome), which might show higher speciation and extinction rates than biome generalists. If true, lineages would accumulate biome‐specialist species. This effect would be particularly exacerbated for biomes located at the periphery of the global climatic conditions, namely, biomes that have high/low precipitation and high/low temperature such as rainforest (warm‐humid), desert (warm‐dry), steppe (cold‐dry), and tundra (cold‐humid). Here, we test these hypotheses in swallowtail butterflies, a clade with more than 570 species, covering all the continents but Antarctica, and all climatic conditions. Swallowtail butterflies are among the most studied insects, and they are a model group for evolutionary biology and ecology studies. Continental macroecological rules are normally tested using vertebrates, this means that there are fewer examples exploring terrestrial invertebrate patterns at global scale. Here, we compiled a large GIS database on swallowtail butterflies’ distribution maps and used the most complete time‐calibrated phylogeny to quantify diversification rates. In this paper we aim to answer the following questions: 1) Are there more biome‐specialists swallowtail butterflies than biome‐generalists? 2) Is diversification rate related to biome specialisation? 3) If so, do swallowtail butterflies inhabiting extreme biomes show higher diversification rates? 4) What is the effect of species distribution area? Our results showed that swallowtail family presents a great number of biome specialists which showed substantially higher diversification rates compared to generalists. We also found that biome‐specialists are unevenly distributed across biomes. Overall, our results are consistent with the resource‐use hypothesis., species climatic niche and biome fragmentation as key factors promoting isolation.
... In contrast, and as would be expected, given its protective function [50,78], residual OB showed wider ranges and highest variances in fire-prone environments such as savannas (Table 3). TDFs typically do not burn or experience fire over such long intervals that fire is not a significant selective factor [80]. However, some TDF species can reach OBs as thick as in any savanna (e.g., Aralia mexicana or Moringa concanensis), suggesting an important protective role against boring insects or other herbivores. ...
Full-text available
Tropical dry forests (TDFs) represent one of the most diverse and, at the same time, most threatened ecosystems on earth. Restoration of TDFs is thus crucial but is hindered by a limited understanding of the functional diversity (FD) of original communities. We examine the FD of TDFs based on wood (vessel diameter and wood density) and bark traits (total, inner, and outer bark thicknesses) measured on ~500 species from 24 plant communities and compare this diversity with that of seven other major vegetation types. Along with other seasonally dry sites, TDFs had the highest FD, as indicated by the widest ranges, highest variances, and largest trait hypervolumes. Warm temperatures and seasonal drought seem to drive diverse ecological strategies in these ecosystems, which include a continuum from deciduous species with low-density wood, thick bark, and wide vessels to evergreen species with high-density wood, thin bark, and narrow vessels. The very high FD of TDFs represents a challenge to restoring the likely widest trait ranges of any habitat on earth. Understanding this diversity is essential for monitoring successional changes in minimal intervention restoration and guiding species selection for resilient restoration plantings in the context of climate change.
... Also, the record of Caatinga endemic species in the city suggests that the paths of the Preto River are associated with the presence of its channel, rather than the climate. Some endemic species of Caatinga, according to Fernandes et al (2020) Furthermore, it is well known that there is a region of common occurrence of dry forests and savannas (DEXTER et al, 2018). In this sense, and considering that the vegetation map of IBGE indicates several spots of Contact with Savanna/Steppic Savanna for this location, it is suggested that the area should be better classified as part of the transition zone. ...
Full-text available
This work aimed to discuss relevant aspects of the differentiation of semi-arid landscape potential in Brazil at different levels. The discussion started from a comparative reading of maps in the light of concepts of the structure of landscapes. The comparative reading of the maps evidenced the absence of consensus mainly related to the western limits of the Domain and the consideration of the areas of exception as part of the Caatinga. Some of these differences arose due to the nature of the proposals and the concern with cartographic accuracy.
... This edaphic variation exerts strong control over ecosystem functioning and dynamics [4], including in the tropical dry forest (TDF) biome, which exhibits high soil heterogeneity. To date, reviews portray TDFs as being dominated by medium-to high-fertility status soils e.g., [5,6], likely because TDF soils are generally less weathered compared to those found in more humid tropical climates, including tropical rain, wet, or moist forests or in tropical savannas [7,8]. This generalization may obscure the highly heterogeneous nature of TDF soils as a result of the wide variety of climate, parent material, topography, and vegetation that defines the TDF biome [9]. ...
Full-text available
Pantropical variation in soils of the tropical dry forest (TDF) biome is enormously high but has been poorly characterized. To quantify variation in the global distribution of TDF soil physical and chemical properties in relation to climate and geology, we produced a synthesis using 7500 points of data with gridded fields representing lithologic, edaphic, and climatic characteristics. Our analyses reveal that 75 TDF ecoregions across five biogeographic domains (Afrotropical, Australasian, Indo-Malayan, Neotropical, and Oceanian) varied strongly with respect to parent material: sediment (57%), metamorphic (22%), volcanic (13%), and plutonic (7%). TDF ecoregions support remarkably high variability in soil suborders (32), with the Neotropical and Oceanian realms being especially diverse. As a whole, TDF soils trend strongly toward low fertility with strong variation across biogeographic domains. Similarly, the exhibited soil properties marked heterogeneity across biogeographic domains, with soil depth varying by an order of magnitude and total organic C, N, and P pools varying threefold. Organic C and N pool sizes were negatively correlated with mean annual temperature (MAT) and positively correlated with mean annual precipitation (MAP). By contrast, the distribution of soil P pools was positively influenced by both MAT and MAP and likely by soil geochemistry, due to high variations in soil parent material across the biogeographic domains. The results summarized here raise important questions as to how climate and parent material control soil biogeochemical processes in TDFs
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1) The Cerrado Domain of central Brazil houses the largest extent of savanna in the Neotropics, but despite its simple characterisation as a giant savanna, it contains considerable vegetation heterogeneity that is poorly understood. 2) We aimed to determine how vegetation types in the Cerrado diverge in their tree species composition and what role ecological factors play in driving compositional patterns. 3) We used a dataset of 1,165 tree species inventories spread across the Cerrado Domain, which come from six vegetation types that have a substantial arboreal component: woody savannas, dystrophic cerradão, mesotrophic cerradão, seasonally dry tropical forests, semideciduous forests and evergreen forests. We found three extremes in terms of tree species composition, with clear underlying ecological drivers, which leads us to propose a ternary model, the ‘Cerrado Vegetation Triangle’, to characterize woody vegetation in the Cerrado. At one extreme, we found that semideciduous and evergreen forests are indistinguishable floristically and are found in areas with high water availability. At another extreme lie seasonally dry tropical forests which are found on more fertile soils. At the third extreme, we found that all types of savanna, and dystrophic cerradão, are highly similar in tree species composition and are commonly found in areas of poor soils and high flammability. Mesotrophic cerradão is transitional in tree species composition between savannas and seasonally dry tropical forest. 4) The lack of variation in tree species composition attributed to climatic variables indicates that within homogeneous macroclimatic zones, many types of forest and savanna co-exist due to complex mosaics of local substrate heterogeneity and fire history. 5) Synthesis. Our findings highlight the complexity of forest-savanna transitions in the Cerrado Domain, with relevance for understanding the future of Cerrado vegetation under environmental change. If nitrogen deposition is extensive, some savannas may be more likely to transition to mesotrophic cerradão or even seasonally dry tropical forest whereas if water availability increases these same savannas may transition to semideciduous or evergreen forest. Our ‘Cerrado Vegetation Triangle’ model offers a simple conceptual tool to frame discussions of conservation and management.
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Fire frequency is changing globally and is projected to affect the global carbon cycle and climate. However, uncertainty about how ecosystems respond to decadal changes in fire frequency makes it difficult to predict the effects of altered fire regimes on the carbon cycle; for instance, we do not fully understand the long-term effects of fire on soil carbon and nutrient storage, or whether fire-driven nutrient losses limit plant productivity. Here we analyse data from 48 sites in savanna grasslands, broadleaf forests and needleleaf forests spanning up to 65 years, during which time the frequency of fires was altered at each site. We find that frequently burned plots experienced a decline in surface soil carbon and nitrogen that was non-saturating through time, having 36 per cent (±13 per cent) less carbon and 38 per cent (±16 per cent) less nitrogen after 64 years than plots that were protected from fire. Fire-driven carbon and nitrogen losses were substantial in savanna grasslands and broadleaf forests, but not in temperate and boreal needleleaf forests. We also observe comparable soil carbon and nitrogen losses in an independent field dataset and in dynamic model simulations of global vegetation. The model study predicts that the long-term losses of soil nitrogen that result from more frequent burning may in turn decrease the carbon that is sequestered by net primary productivity by about 20 per cent of the total carbon that is emitted from burning biomass over the same period. Furthermore, we estimate that the effects of changes in fire frequency on ecosystem carbon storage may be 30 per cent too low if they do not include multidecadal changes in soil carbon, especially in drier savanna grasslands. Future changes in fire frequency may shift ecosystem carbon storage by changing soil carbon pools and nitrogen limitations on plant growth, altering the carbon sink capacity of frequently burning savanna grasslands and broadleaf forests.
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Tropical forests account for a quarter of the global carbon storage and a third of the terrestrial productivity. Few studies have teased apart the relative importance of environmental factors and forest attributes for ecosystem functioning, especially for the tropics. This study aims to relate aboveground biomass (AGB) and biomass dynamics (i.e., net biomass productivity and its underlying demographic drivers: biomass recruitment, growth and mortality) to forest attributes (tree diversity, community-mean traits and stand basal area) and environmental conditions (water availability, soil fertility and disturbance).
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Tropical savannas have been increasingly viewed as an opportunity for carbon sequestration through fire suppression and afforestation, but insufficient attention has been given to the consequences for biodiversity. To evaluate the biodiversity costs of increasing carbon sequestration, we quantified changes in ecosystem carbon stocks and the associated changes in communities of plants and ants resulting from fire suppression in savannas of the Brazilian Cerrado, a global biodiversity hotspot. Fire suppression resulted in increased carbon stocks of 1.2 Mg ha⁻¹ year⁻¹ since 1986 but was associated with acute species loss. In sites fully encroached by forest, plant species richness declined by 27%, and ant richness declined by 35%. Richness of savanna specialists, the species most at risk of local extinction due to forest encroachment, declined by 67% for plants and 86% for ants. This loss highlights the important role of fire in maintaining biodiversity in tropical savannas, a role that is not reflected in current policies of fire suppression throughout the Brazilian Cerrado. In tropical grasslands and savannas throughout the tropics, carbon mitigation programs that promote forest cover cannot be assumed to provide net benefits for conservation.
Within the tropics, the species richness of tree communities is strongly and positively associated with precipitation. Previous research has suggested that this macroecological pattern is driven by the negative effect of water‐stress on the physiological processes of most tree species. This implies that the range limits of taxa are defined by their ability to occur under dry conditions, and thus in terms of species distributions predicts a nested pattern of taxa distribution from wet to dry areas. However, this ‘dry‐tolerance’ hypothesis has yet to be adequately tested at large spatial and taxonomic scales. Here, using a dataset of 531 inventory plots of closed canopy forest distributed across the western Neotropics we investigated how precipitation, evaluated both as mean annual precipitation and as the maximum climatological water deficit, influences the distribution of tropical tree species, genera and families. We find that the distributions of tree taxa are indeed nested along precipitation gradients in the western Neotropics. Taxa tolerant to seasonal drought are disproportionally widespread across the precipitation gradient, with most reaching even the wettest climates sampled; however, most taxa analysed are restricted to wet areas. Our results suggest that the ‘dry tolerance' hypothesis has broad applicability in the world's most species‐rich forests. In addition, the large number of species restricted to wetter conditions strongly indicates that an increased frequency of drought could severely threaten biodiversity in this region. Overall, this study establishes a baseline for exploring how tropical forest tree composition may change in response to current and future environmental changes in this region.
In the tropics, research, conservation and public attention focus on rain forests, but this neglects that half of the global tropics have a seasonally dry climate. These regions are home to dry forests and savannas (Figures 1 and 2), and are the focus of this Primer. The attention given to rain forests is understandable. Their high species diversity, sheer stature and luxuriance thrill biologists today as much as they did the first explorers in the Age of Discovery. Although dry forest and savanna may make less of a first impression, they support a fascinating diversity of plant strategies to cope with stress and disturbance including fire, drought and herbivory. Savannas played a fundamental role in human evolution, and across Africa and India they support iconic megafauna.
Aim It remains poorly understood why the position of the forest–savanna biome boundary, in a domain defined by precipitation and temperature, differs in South America, Africa and Australia. Process based Dynamic Global Vegetation Models (DGVMs) are a valuable tool to investigate the determinants of vegetation distributions; however, many DGVMs fail to predict the spatial distribution or indeed presence of the South American savanna biome. Evidence suggests that fire plays a significant role in mediating forest–savanna biome boundaries; however, fire alone appears to be insufficient to predict these boundaries in South America. We hypothesize that interactions between precipitation, constraints on tree rooting depth and fire affect the probability of savanna occurrence and the position of the savanna–forest boundary. Location Tropical forest and savanna sites in Brazil and Venezuela north of 23S. Methods We tested our hypotheses using a novel DGVM, aDGVM2, which allows plant trait spectra, constrained by trade‐offs between traits, to evolve in response to abiotic and biotic conditions. Plant hydraulics is represented by the cohesion–tension theory, this allowed us to explore how soil and plant hydraulics control biome distributions and plant traits. The resulting community trait distributions are emergent properties of model dynamics. Results We showed that across much of South America the biome state is not determined by climate alone. Interactions between plant rooting depth, fire and precipitation affected the probability of observing a given biome state and the emergent traits of plant communities. Simulations where plant rooting depth varied in space provided the best match to satellite derived biomass estimates and generated biome distributions that reproduced contemporary biome maps well. Main conclusions Our findings support the contention that areas where multiple vegetation states are possible are widespread and highlight the importance of considering the influence of fire and constraints on plant rooting depth for predicting biome boundaries.