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Biogeographic Patterns and Conservation in The South American Cerrado: A Tropical Savanna Hotspot
Biogeographic Patterns and Conservation in The South American Cerrado: A Tropical Savanna Hotspot
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a b
4. campo sujo, grassland (2–3 m tall) with scattered shrubs have a highly endemic flora (Giullietti and Pirani 1988, Eiten
1990). Narrow fringes of gallery forest often border small rivers
5. campo limpo, grassland with few or no shrubs or taller and streams of the region. Gallery forests on the plateaus
woody plants (Figure 3b)
grow on narrow belts of soils rich in organic matter (Eiten
Tree species common in the cerrado vegetation are Annona 1990). They are evergreen, have trees that are on average
crassiflora, Astronium fraxinifolium, Byrsonima coccolobifolia, 20–30 m tall, and possess a dense understory of low stature
B. verbascifolia, Hancornia speciosa, Kielmeyera coriacea, (Ribeiro and Walter 1998). Associated with gallery forests
Qualea grandifolia, and Q. parviflora, among others (Ratter are stands of a species of palm, Mauritia flexuosa, locally
and Dargie 1992). Besides cerrado, other types of distinct sa- known as veredas.
vanna vegetation, called campos rupestres (Figure 3a), and The peripheral depressions are generally flat surfaces of lit-
miscellaneous lithosolic campos occur on rocky outcrops. tle relief (100–500 m in elevation) that are ocasionally inter-
These habitats are limited to small patches on the plateaus and rupted by the presence of steep-sided hills. The landscape pat-
tern on these depressions is much more heterogeneous than 2000). These species probably originated in adjacent regions
that of the plateaus in that it includes more habitats (river- and maintained viable populations within the Cerrado region
ine forests, tropical dry forests, all types of cerrado, and because their forest habitats did not vanish entirely during the
marshlands) distributed in a mosaic fashion. Riverine forests paleoclimatic period that followed their range expansion. In
(called matas ciliares; Ribeiro and Walter 1998) differ from a fashion similar to what is seen today, large rivers and streams
gallery forests because they are found only along the large rivers must have provided suitable environmental conditions to
in the Cerrado. They are not wide (up to 100 m from each side maintain wide gallery–riverine forests during the dry periods
of the river) and can be semideciduous to evergreen. Trees are (Ab’Saber 1983), whereas large patches of soils derived from
20–30 m tall. Common species are Anadenanthera spp., limestone were important for maintaining dry forests during
Apeiba tibourbou, Aspidosperma spp., Tabebuia spp., and the wet periods (Prado and Gibbs 1993, Pennington et al.
Trema micrantha, among others (Ribeiro and Walter 1998). 2000).
Tropical dry forests (matas secas) are particularly associated
with peripheral depressions and, in some areas (e.g., in the Endemic species: Origin and evolution. The per-
Paranã River valley), are the dominant vegetation type. They centage of species endemic to the Cerrado varies across dif-
are deciduous or semideciduous and grow on patches of ferent groups of organisms: vascular plants (44%), amphib-
moderately productive soils derived from basic rocks such as ians (30%), reptiles (20%), mammals (11.8%), and birds
limestone (Ratter et al. 1978, Prado and Gibbs 1993). Canopy (1.4%) (Silva 1995a, Myers et al. 2000). However, because large
height averages 20–45 m (Ribeiro and Walter 1998). Tree areas of the Cerrado remain unexplored (Silva 1995c), these
species common in dry forests include Astronium urundeuva, numbers will likely change when additional biological in-
Piptadenia macrocarpa, Chorisia speciosa., Tabebuia spp., Ca- ventories are conducted. At present, we still know little about
vanillesia arborea, and Cedrella fissilis (Ratter et al. 1978). how these endemic species are distributed across the Cerrado
or how they evolved. A key question for conservation purposes
Biogeographic patterns is whether one can identify subareas of endemism within
the large Cerrado region.
Forest organisms in a tropical savanna region. To determine if there are subareas of endemism within the
The most basic question about the biota of a region is how Cerrado, Silva (1997) mapped and created overlays of all en-
it is distributed in major vegetation types. In the Cerrado, one demic bird species’ ranges smaller than 60,000 km2. Only 10
might expect that most of the biota would inhabit the savannas of the 30 endemic species met this requirement (Table 1). The
that dominate the region. However, studies of mammals, ranges of these species delimit three main areas:
birds, and plants do not support this hypothesis. Redford and
Fonseca (1986) found that most (56.3%) of the nonvolant • Espinhaço Plateau: Four species that inhabit campos
rupestres are Augastes scutatus (Trochilidae), Asthenes
mammal species in the Cerrado inhabit forests. Similarly,
luizae (Furnariidae), Polystictus superciliaris (Tyran-
Silva (1995b) classified the 759 birds species known or as- nidae), and Embernagra longicauda (Emberizidae).
sumed to breed in the Cerrado as independent (occurring ex-
clusively in forests), semidependent (occurring in both forests • Araguaia River Valley: Three species that inhabit gallery
and open vegetation), or dependent (occurring exclusively in forests are Synallaxis simoni (Furnariidae), Cercomacra
open vegetation). The majority of species (393, or 51.8%) are ferdinandi (Thamnophilidae), and Paroaria baeri
dependent on the various forest habitats, 20.8% (158) are (Emberizidae).
semidependent, and 27.4% (208) are independent species. • Paranã River Valley: Two species found only in tropical
Thus, approximately 82.6% of Cerrado birds require forests dry forests are Pyrrhura pfrimeri (Psittacidae) and
to some degree. Mendonça and colleagues (1998) listed 6671 Knipolegus franciscanus (Tyrannidae) (Figure 4).
plant taxa (species and varieties) for the Cerrado; of these, 38%
occur only in forests. The subareas of endemism identified for birds are at least
In general, these data support the hypothesis that gallery partially supported by studies of other groups of organisms.
and dry forests, which occupy less than 20% of the Cerrado, The Espinhaço Plateau is well known for its unique biota,
are necessary for a large portion of regional biodiversity. composed of hundreds of endemic species of plants (Giulli-
How did this pattern evolve? Silva (1995b) suggested that eti and Pirani 1988, Harley 1988) and some endemic species
biotic diversity of the Cerrado region increased through in- of other vertebrates, such as lizards and amphibians (Haddad
terchange with adjacent regions during the Quaternary et al. 1988, Rodrigues 1988). The Araguaia and Paranã river
climatic–vegetational fluctuations. Thus, Atlantic and Ama- valleys are still poorly known for nonavian groups, but the
zonian forest species expanded their ranges into the Cerrado riverine forests and dry forests likely harbor endemic species
during the wet periods following the expansion of the of other organisms also. For example, a very distinctive species
riverine–gallery forest network (Silva 1995b), whereas Caatinga of rodent of the genus Kerodon was described recently from
and Chaco elements colonized the Cerrado during the Qua- Paranã dry forests (Moojen et al. 1997). Although not known
ternary dry periods as part of the coalescence of dry forests to possess endemic birds, several other areas possess endemic
in peripheral depressions (Ab’Saber 1983, Pennington et al. taxa from other groups. For instance, Erwin and Pogue (1988)
less than 2% of the total area of the Cerrado is protected in recommendations of this workshop and that new protected
reserves (Mittermeier et al. 2000), but there are important dif- areas and ecological corridors will be established. In fact,
ferences in how countries are approaching Cerrado conser- some important actions were taken during 2001, but two
vation. The Bolivian government created Noel Kempff Na- deserve special comments. The first was the expansion of
tional Park in 1988, thereby protecting the country’s largest the National Park of Chapada dos Veadeiros, an important re-
cerrado (Killeen and Schulenberg 1998). These roughly 42,000 serve that includes pristine cerrados and patches of campos
ha of unaltered cerrado near the western edge of the biome rupestres with several endemic plant species, from 60,000 to
is one of the world’s largest continuous protected parcels of 235,000 ha. The second was the creation of the Ecological Sta-
this habitat (Figure 7). Because of the park’s remoteness and tion of Serra Geral do Tocantins with 716,306 ha close to the
Bolivia’s comparatively low human population density (es- State Park of Jalapão with 158,885 ha. Together, these two ar-
pecially in the country’s eastern lowlands), the most sub- eas form the world’s largest continuous area of protected
stantial human threats to this area come from burning asso- Cerrado (Figure 7).
ciated with ranching operations in neighboring Brazil. Bolivia How can the biogeographic information discussed here be
also possesses several smaller and more isolated plateaus with used to help protect and preserve the Cerrado’s biodiversity?
cerrado habitat; these remain unprotected and largely unex- First, we know now that the Cerrado’s biota is not homoge-
plored (Parker and Rocha 1991). neous, so additional distribution data for different groups of
In Paraguay, at the southern limit of the biome, Cerrado organisms must be collected and organized in a retrievable way
is protected by two national parks: Serranías de San Luís and to help conservationists determine whether there are addi-
Cerro Corá (Clay et al. 1998, Robbins et al. 1999). Avian sur- tional unidentified subareas of endemism. Second, even
vey work has been conducted at both sites; however, the re- though the three subareas of endemism identified here as har-
gions are still not considered to be thoroughly surveyed, and boring restricted-range endemic species of birds are partially
both Clay (1998) and Robbins and their colleagues (1999) em- covered by one or more reserves (for comparison, refer to Fig-
phasize that the rapid expansion of agricultural operations ures 4 and 7; Silva 1997), most reserves need to be more for-
threaten all unprotected savanna regions in the country. mally implemented and connected through biodiversity cor-
In Brazil, which possesses the vast majority of Cerrado, the ridors to insure their preservation over the long term. Analyses
situation is drastically worse. There are few large reserves of other, nonavian taxa are certain to uncover additional ar-
(more than 25,000 hectares [ha]), and they are not distributed eas that are not currently protected by reserves. Third, despite
evenly across the biome (Figure 7). Consequently, an im- occupying a relatively small part of the total landscape,
portant part of the Cerrado’s environmental diversity has
not been incorporated in a network of protected areas. Large a b
areas of Brazilian Cerrado and dry forests have been converted
to soybean and rice plantations. Most of this large-scale habi-
tat modification has not followed the most basic principles
of environmental conservation, and problems of erosion and
deterioration of important streams and rivers are increasing.
International support for this alteration has come from ma-
jor development agencies and some senior scientists who
have encouraged increased land use in the Cerrado to reduce
human pressure on Amazonian forests.
The Brazilian government’s appreciation of the uniqueness
of the Cerrado’s biodiversity has developed slowly. The first
major initiative occurred in 1997, when the Brazilian Ministry c d
of Environment promoted a workshop, which included over
200 scientific experts on the region. This group met to define
priority areas for conservation in the Brazilian Cerrado. It fol-
lowed the same methodology applied earlier for the Atlantic
Forest and Amazonia (Cavalcanti 1999). Several criteria for
assessing biodiversity value were used, such as number of
endemic species, species richness, presence of rare or en-
dangered species or both, and sites of unique communities
or key areas for migratory species. The group identified a to-
tal of 87 priority areas. The urgency for conservation action
in a priority area was determined by cross-referencing bio-
diversity data with the human-pressure and land cover- Figure 6. Examples of ranges of bird species supporting
change data (Cavalcanti 1999). As of August 1999, the Brazil- the hypothesis (a, b) of the Andean savanna corridor and
ian government has indicated that it will follow most of the (c, d) of the coastal savanna corridor.