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Alexandre Gabriel Franchin
Laboratory of Ornithology, Biology Institute, Federal University of Uberlândia, Brazil
Rafael de Freitas Juliano
Laboratory of Ornithology, Biology Institute, Federal University of Uberlândia, Brazil
Mieko Ferreira Kanegae
Bioscience Institute, General Ecology Department, São Paulo University, Brazil
Oswaldo Marçal Junior
Laboratory of Ornithology, Biology Institute, Federal University of Uberlândia, Brazil
Keywords: Avifauna, Biodiversity, Conservation, Ecoregions.
1. Introduction
2. Global Ecoregions and the Savanna Biome
3. Tropical Savannas
4. Birds in the Tropical Savannas
5. Cerrado: A Unique Tropical Savanna Hotspot
Related Chapters
Biographical Sketches
The Savanna biome is characterized by landscapes dominated by grasses with scattered trees
and shrublands, covering about 20% of the world’s surface. Tropical Savannas are located
mainly in South America, Africa, Australia and South Asia. In the present chapter, we are
combining information from different sources to offer a more complete description of the bird
diversity in the tropical savannas. We employed data from BirdLife International, articles,
field guides, local journals, newsletters, bird distribution maps, checklists and reports by
ornithologists around the globe. For our analysis, we considered all main terrestrial and
flooded tropical savannas (those savannas occurring in the tropical region). The results show
that 2,951 bird species (25 orders, 152 families and 1,404 genera) are found in the Tropical
Savannas. The Cerrado (with 850 species) and African Savannas (794 species) have the
greatest bird richness. Tyrannidae family has the most diversity, with 162 species which are
exclusively found on the Neotropical savannas. Accipitridae is another prominent bird family
with around 142 species. Three bird species are common to all tropical savannas: the Cattle
Egret Bubulcus ibis, Barn Swallow Hirundo rustica, and the Osprey Pandion haliaetus. The
tropical savannas also share 15 orders, 20 families and 19 genera. Twenty families were
exclusive to only one savanna. Northern Australian savannas had the greater number of
exclusive families (eight families). The similarity analysis shows the following groupings:
African Savannas; Terai-Duar; Northern Australian and Trans-Fly; and Beni, Pantanal,
Cerrado and Llanos. The Neotropical savannas (Llanos, Pantanal, and Cerrado) are those that
can be considered in fact similar, because they presented a minimum 0.70 Sorenson’s index.
There are 2,057 bird species that face some degree of threat (21% of the recognized bird
species), not considering the extinct species or those with insufficient data available. In the
tropical savannas, there are 319 species that face some degree of threat. The Cerrado is a
unique tropical savanna considered a hotspots, with the most threatened bird fauna (58
species), and seven of them are critically endangered.
1. Introduction
There are approximately 9,990 bird species in the Earth, distributed in 29 orders, 195 families,
and 2,113 genera. The Order Passeriformes include 58% of all extant bird species and among
the non-passerines, the Apodiformes is the most diverse order, totaling roughly 443 species,
which is about 4% of the total of world bird species. The most representative passerine family
in species numbers is Tyrannidae, with around 410 species known as the tyrant flycatchers.
Amongst the non-passerine families, Psittacidae, represented by the macaws, parrots, lories,
and lorikeets, has the greater species number, with around 374 species. The gender Nectarinia
(some sunbirds) shows the greater bird species richness in the world with 78 species.
Due to their varied anatomical and ecological adaptations, birds have a worldwide
distribution. The most distinctive zoological region for the avifauna is the Neotropical region,
with around one third of Earth’s species, including a great number of endemics species (e.g.
toucans, manakins, hummingbirds, rheas, among others). Birds are also diversified and
abundant in other tropical areas of Ethiopian, Australian and Oriental Zoological regions.
They occupy different biomes, mainly tropical forests and savannas.
The plants and animals that inhabit grasslands, savannas and shrublands are adapted to survive
the seasonal extremes and are also more dependent on the impact of fire regimes than on the
climate itself, and birds are not an exception. Our current knowledge about tropical savanna
birds is very broad, although the majority of the data is diffuse, resulting from isolated works.
In the present chapter, we are joining different sources of information to offer a more
complete description of the bird diversity in the tropical savannas. We employed data from
BirdLife International, articles, field guides, local journals, newsletters, bird distribution maps,
checklists and reports by ornithologists around the globe.
2. Global Ecoregions and the Savanna Biome
The concept of Global ecoregions was developed by WWF and is defined as: “a large unit of
land or water containing a geographically distinct assemblage of species, natural
communities, and environmental conditions”. In this sense, ecoregions represent ecological
and conservation units in which natural communities shares the greater part of their
biodiversity, ecological processes and similar environmental conditions, producing critical
mechanisms for their long-term persistence. This knowledge provides us with a powerful tool
that aids us in the investigation of the biodiversity from all terrestrial and aquatic ecosystems
on the Earth. A total of 142 terrestrial, 53 freshwater and 43 marine ecoregions are
recognized. This concept is a to the biodiversity conservation at a global scale, since it makes
possible to sustain regional conservation policies and development strategies. Those habitats
include extensive areas and different environmental conditions, ranging from the wettest of
forest types to the driest and hottest deserts, reflecting a broad diversity of species,
communities and ecosystems.
The Savanna biome is characterized by landscapes dominated by grasses with scattered trees
and shrublands. It is a transition between forests and grasses domains. The primary savannas
result from typical climate (climatic savannas), soil (edaphic savannas) or grazing conditions.
A secondary type, called derived savanna, is produced by human activities, including burning
and deforesting. The distribution of savannas includes extensive areas of America, Africa,
Asia and Australia continents (Figure 1).
Grasslands, Savannas and Shrublands are present in a representative number of ecoregions
distributed in three Terrestrial Major Habitat Types, as follows: Tropical and Subtropical
Grasslands, Savannas and Shrublands; Temperate Grasslands, Savannas and Shrublands; and
Flooded Grasslands and Savannas (Table 1).
Table 1. Global ecoregions, according to Terrestrial Major Habitat Types and geographical
2.1. Flooded Grasslands and Savannas
Flooded Grasslands and Savannas are constituted of large mosaics of flooded grasslands and
savannas, gallery forests, and dry forests. In some areas, over 80% of the region can be
flooded during the rainy season. These complexes occur in the following Global ecoregions:
Everglades and Pantanal (Neotropical), Sudd-Sahelian Flooded Grasslands and Savannas and
Zambezian Flooded Savannas (Afrotropical) and Rann of Kutch Flooded Grasslands (Indo-
Malayan). Flooded Grasslands and Savannas sustain great biodiversities, including animal and
plants intimately adapted to the rainfall regime and soil conditions. The Everglades covers
Geographical distribution
Terrestrial Major Habitat types
1. Tropical and Subtropical Moist
Broadleaf Forests 11 8 17 12 - - 2 50
2. Tropical and Subtropical Dry
Broadleaf Forests 1 2 2 4 - - 1
3. Tropical and Subtropical
Coniferous Forests - - - 2 1 - - 3
4. Temperate Broadleaf and Mixed
Forests - 3 2 - 1 2 - 8
5. Temperate Coniferous Forests - - - 1 4 4 - 9
6. Boreal Forests / Taiga - - - - 2 3 - 5
7. Tropical and Subtropical
Grasslands, Savannas and Shrublands 4 1 1 2 - - -
8. Temperate Grasslands, Savannas
and Shrublands - - - 1 1 1 - 3
9. Flooded Grasslands and Savannas 2 - 1 2 - - - 5
10. Montane Grasslands and
Shrublands 4 1 1 2 - 3 -
11. Tundra - - - - 2 3 -
12. Mediterranean Forests,
Woodlands and Scrub 1 2 - 1 1 1 - 6
13. Deserts and Xeric Shrublands 4 2 - 1 2 2 - 11
14. Mangroves 3 1 2 2 - - -
Total 30 20 26 30 14 19 3 142
20,000 sp km of the Southern Florida State (USA) and supports some 11,000 species of seed-
bearing plants, 150 fish species and 300 bird species. The Pantanal spreads across 171,000 sq
km of Bolivia, Brazil, Paraguay (South America) and it has over 260 species of fish, 700
birds, 90 mammals, 160 reptiles, 45 amphibians, 1,000 butterflies, and 1,600 species of plants.
Any change in the precipitation regime, flow and quality of water, integrity of riparian and
gallery habitats or natural fires can produce serious disturbances in the composition and
structure of communities. So, agricultural expansion, water projects, pollution, over fishing,
uncontrolled ecotourism and road construction constitute severe threats for all Flooded
Grasslands and Savannas. These ecoregions are very important for several bird species,
including migratory species, that feed and breed there, like the Jabiru stork (Jabiru mycteria),
Wood stork (Mycteria americana), Hyacinth macaw (Anodorhynchus hyacinthinus), Roseate
spoonbill (Ajaia ajaia), and White ibis (Eudocimus albus).
2.2. Temperate Grasslands, Savannas and Shrublands
Temperate Grasslands, Savannas and Shrublands are defined as areas where trees are lacking,
except for riparian or gallery forests, although some regions do support woody savannas. They
are known as prairies in North America, pampas in South America, veld in Southern Africa,
downlands in Australia and steppe in Asia. In this habitat, the following Global ecoregions are
found: Northern Prairie (Nearctic); Patagonian Steppe (Neotropical) and Daurian Steppe
(Palaearctic). The biodiversity includes numerous arthropods, large grazing mammals and
associated predators, as well as a hundreds of bird species. In North American prairies, about
100 different plant species can be found flourishing and supporting more than 3 million
individual insects per acre. However, whole communities have been destroyed through the
conversion of grasslands to agriculture. Fire and loss of riparian or gallery forest habitats and
water sources are other significant impacts on these animal and plant communities. The
Patagonian Steppe and Grasslands are notable in terms of taxonomic diversity, mainly in
gender and family level. The plant species diversity (alpha, beta, and gamma) are low, except
for some local sites. The majority of species has relatively widespread distributions and
abundance, especially for larger vertebrates.
2.3. Tropical and Subtropical Grasslands, Savannas and Shrublands
Tropical and Subtropical Grasslands, Savannas and Shrublands are found in warm regions
where the Wet-Dry Tropical climate is dominant, been characterized by a seasonal
precipitation accompanied by a dry season. In general, the rainfall varies from 900-1,500 mm
per year, but in the dry season this precipitation can be lower than 50 mm. There is not enough
rainfall to sustain extensive tree cover. The rainfall is concentrated in six or eight months of
the year, when many grasses and trees flourish and then go into a state of dormancy. Plant
species are very well adapted to the long dry season, especially to reduce the loss of water. In
the dry period another important condition becomes present: fire. Fires control some
populations in the savanna communities, affecting or favoring different species. Many groups,
particularly low-speed or short dispersion vertebrate species (e.g. anurans, lizards, rodents,
etc) are killed by fire, while others, like birds, are benefited, eating insects, mice and lizards
killed or driven out by the fire. Dry stems and leaves of grasses are also consumed by fire, but
the grasses' deep roots remain unharmed, sending up new shoots when the soil becomes moist
again. In different savannas, grass composition also varies according to soil conditions. In the
Brazilian savanna or Cerrado, the soil has high aluminum concentrations and many species of
plants are adapted to this condition. Termites are also a major force shaping the plant
composition of many savannas, and their large mounds are a common sight in most open areas
of the tropics. Termites also help reclaim soils damaged by overgrazing and fires. The rock-
hard walls of the mounds are used by a wide range of animals, including many birds, which
use these protective structures to perch, nest, forage and breed.
3. Tropical Savannas
Tropical savannas are distributed on extensive areas being present in the following
geographical regions: Afrotropical, Indo-Malayan, Australasia and South America, totaling
eight Global ecoregions (Table 1). For our analysis, we considered all main terrestrial and
flooded tropical savannas (those savannas occurring in the tropical region). Therefore, we
included the Pantanal and Zambezian Flooded savannas. However, only the Pantanal was
analyzed separately, once the Zambezian savanna has a lack of specific data about the
avifauna. Indeed, we considered four main Global ecoregions in Africa in the analysis, as
follows: Horn of Africa Acacia Savannas; East African Acacia Savannas; Central and Eastern
Miombo Woodlands; and Sudanian Savannas. In the following sections, we will see some of
the most important characteristics of these ecoregions.
3.1. Afrotropical
African savannas occupy about 4.5 million square kilometers distributed in Central and East
Africa. The grasslands are dominant, but other arboreal species, including acacias (Acacia
sp.), baobabs (Adansonia spp.), as well as bush species (usually leguminous plants) are
scattered across the landscape. The seasons are regulated mainly by pluviosity, more than by
the temperature. The savannas maintain a great diversity of species, as much plant as animal
and many species are found in no other places of the earth. The dry and wet seasons are very
well marked, resulting in an occurrence of a great numbers of migratory species, mainly of
mammals and birds. More than 790 bird species are found in that ecoregion.
3.1.1. The Horn of Africa Acacia Savannas
The Horn of Africa Acacia Savannas are located in Eastern Africa, spreading from Eritrea,
Ethiopia, Kenya, Somalia and Sudan (1,053,000 sq. km) and been characterized by large
Acacia woodland areas. This ecoregion occupies the majority of the Horn of Africa to the east
of the Ethiopian highlands, including the Ogaden Desert and northeast Kenyan semi-deserts.
A combination of African, Mediterranean, Asiatic and old Gondwanan influences have shaped
the flora and fauna in this ecoregion. This ecoregion includes different habitats as bush lands,
semi-desert grasslands and shrublands. The typical birds are contains many arid-adapted
species. The riparian habitats support four endemic birds: the Degodi lark (Mirafra
degodiensis), Bulo Burti Boubou (Laniarius liberatus), white-winged collared-dove
(Streptopelia reichenowi), and Salvadori's Weaver (Ploceus dichrocephalus). The sombre chat
(Cercomela dubia) and the scaly babbler (Turdoides squamulatus) are considered near-
endemic. Abyssinian yellow-rumped seedeater (Serinus xanthopygius), short-billed crombec
(Sylvietta philippae), and Sidamo bush lark (Heteromirafra sidamoensis) are also restricted to
this ecoregion.
3.1.2. The East African Acacia Savannas
The East African Acacia Savannas form another important Global ecoregion. Its geographical
area is 572,000 sq. km, and includes the following terrestrial ecoregions: Southern Acacia-
Commiphora bushlands and thickets; Serengeti volcanic grasslands; Northern Acacia-
Commiphora bushlands and thickets. As the previous Global ecoregion, it extends across five
countries (Ethiopia, Kenya, Sudan, Tanzania, and Uganda) and supports an extraordinary
variety of flora and fauna. This ecoregion spreads across grasslands and related woodlands
dominated by Acacia sp. and Commiphora sp. trees. It is strongly affected by seasonal
patterns of rainfall, with most of the rain falling between the second and last quarter of the
year, being highly unpredictable, for both or only one rainy season may fail in a given year.
Elephants also play an important role as architects in this habitat; by routinely knocking down
trees as they feed, they open and transform woodlands into grassland savannas. Historically,
it was the eruption of the Kerimasi volcano some 150,000 years ago in the highlands that
produced the fine volcanic materials that now make up the soils of the Serengeti plains.
Volcanic ash produces a distinct soil type called vertisol that supports characteristic plant
communities. The richness and abundance of bird species is notable, including exclusive
species, such as the Grey-breasted spurfowl (Francolinus rufopictus), Fischer's lovebird
(Agapornis fischeri) and the Rufus-tailed weaver (Histurgops ruficauda).
3.1.3. The Central & Eastern Miombo Woodlands
The Central & Eastern Miombo Woodlands is the major African Global terrestrial ecoregion
with 1,932,500 sq. km, extending from the Central and Southern Africa (Angola, Botswana,
Burundi, DRC, Malawi, Mozambique, Namibia, Tanzania, Zambia and Zimbabwe). This
subregion is made up of the following terrestrial ecoregions: Eastern Miombo woodlands;
Central Zambezian Miombo woodlands; Zambezian Baikiaea woodlands (under domain of the
Central African Plateau). Consisting mainly of broadleaf, deciduous savannas and woodlands,
it is characteristically interspersed with edaphic grassland and semi-aquatic vegetation as well
as areas of evergreen groundwater forest. Some areas are important in terms of species
richness (e.g. Miombo woodlands), including various bird species, such as the Black-faced
waxbill (Estrilda nigriloris), Miombo rock-thrush (Monticola angolensis), and Miombo pied
barbet (Tricholaema frontata). The avifauna in the ecoregion is very rich. Nonetheless,
endemism is low with the only strict endemics being the Ruwet’s masked weaver (Ploceus
ruweti) and the black-faced waxbill (Estrilda nigriloris). Ruwet’s masked weaver is known
only from the swamps bordering Lake Tshangalele/Lufira in southern Democratic Republic of
Congo, while the black-faced waxbill, and is found in the Lualaba River and Lake Upemba
region. Range-restricted species include grey-crested helmetshrike (Prionops poliolophus), an
uncommon species endemic to the woodlands of Kenya and northern Tanzania. Although
some birds tend to avoid Miombo, many isolated populations are confined within moister
Miombo woodland, like Lilian’s lovebird (Agapornis lilianae). The slender-tailed cisticola
(Cisticola melanurus) is restricted to grassy places in well-developed Miombo. Many globally
threatened water-associated birds have also been recorded, including the vulnerable wattled
crane (Grus carunculatus), slaty egret (Egretta vinaceigula), and the corncrake (Crex crex).
Baikiaea woodlands are the typical habitat of the near-endemic Bradfield’s hornbill (Tockus
bradfieldi), to the Southern ground hornbill (Bucorvus leadbeateri) and the rare and
vulnerable black-cheeked lovebird (Agapornis nigrigenis), confined to medium-altitude where
the mopane woodland is also contiguous. The riverine vegetation is home to Pel’s fishing owl
(Scotopelia peli) and many other raptor species abound in this ecoregion, like the long-legged
secretary bird (Sagittarius serpentarius), white-backed vulture (Gyps africanus), lappetfaced
(Torgos tracheliotus), white-headed vulture (Trigonoceps occipitalis), hooded vulture
(Necrosyrtes monachus), lesser kestrel (Falco naumanni), Dickinson’s kestrel (Falco
dickinsoni), African hobby falcon (Falco cuvierii), bateleur (Terathopius ecaudatus), tawny
eagle (Aquila rapax), martial eagle (Polemaetus bellicosus), and African hawk eagle
(Hieraaetus spilogaster).
3.1.4. The Sudanian Savannas
The Sudanian Savannas are stretch in a band across West Africa south of the Sahel, from
Senegal and Gambia to the eastern border of Nigeria, in central and eastern Africa, and is
divided into a western block and an eastern block by the Sudd swamps in the Saharan Flooded
Grasslands ecoregion, extending for 2,556,100 sq. km. As other Afrotropical savannas, is a
hot, dry, wooded savanna composed mainly of large tree species and long elephant grass
(Pennisetum purpureum). It also comprises large areas of acacia woodland expanses. This
ecoregion occupies an area identified as a center of diversity for many plants and animal
groups. The pronounced dry season triggers a significant migration of fauna within the
ecoregion, together with annual passage of millions of migrant birds on the Afrotropical-
Palaearctic flyway, as well as the intra-African migration associated with the seasonal changes
in African weather. Four bird species are considered near-endemic: Dorst's cisticola (Cisticola
dorsti), white-crowned robin-chat (Cossypha albicapilla), Karamoja apalis (Apalis
karamojae), and Mali firefinch (Lagonosticta virata). Five bird species are considered
endemic, including two strict endemics, Reichenow's firefinch (Lagonostica
umbrinodorsalis), and Fox’s weaver (Ploceus spekeoides). The original wooded savanna
habitat has been significantly reduced.
3.1.5. The Saharan Flooded Grasslands
The Saharan flooded grasslands cover 179,700 km2 and are one of the largest wetlands in
Africa. It is located in the borders of the arid Sahelian region, providing watering and feeding
grounds for populations of migratory birds. During the rainy season (from April to September)
the Nile river rises in the headwaters of Lake Victoria in a region of year-round rainfall, runs
through Uganda, and overflows into the vast floodplain surrounding the permanent swamps of
southern Sudan. This flooding creates the extensive Sudd swamps about (about 600 km long
and wide), just 380-450 m above sea level. This cycle brings nutrients and new life to the dry,
cracked landscape. Temperatures vary from 30-18oC during the year. The floodplains provide
important habitat for large populations of many bird species, like the vulnerable shoebill
(Balaeniceps rex), the endangered migratory white pelican (Pelecanus onocrotalus), and the
vulnerable black crowned crane (Balearica pavonina). The main threats to birds include the
reduced conservation efforts, uncontrolled hunting and the building of the Jonglei canal.
3.1.6. The Zambezian Flooded Grasslands
The Zambezian Flooded Grasslands are a discontinuous ecoregion (153,500 km2) embedded
largely within Miombo and mopane woodlands in the Central African Plateau. The ecoregion
is spreads from the Kilombero Valley, Moyowosi/ Malagarasi system, Ugalla River
(Tanzania), Okavango River Delta (Botswana), Lake Chilwa (Malawi), Barotse Floodplain,
Kafue Flats, Busanga/Lukanga Swamps, Lake Mweru, Mweru Marsh and
Bangwuelu/Luapala/Chambezi system (Zambia) and other smaller floodplains and wetlands.
The landscape is characterized by nutrient poor soil and vegetation, but the productivity is
high and food and water are abundant throughout most of the year and this ecoregion supports
a high avifaunal diversity and density. The wetlands and floodplains of this ecoregion provide
good habitats for many endemic and near-endemic bird species, especially during the rainy
season. The only strict endemic is the vulnerable Kilombero weaver (Ploceus burnieri),
known from one locality along the Kilombero River, in the riverside swamps that are fringed
with tall reed beds. Two other birds are nearly endemic to the ecoregion: Chaplin's barbet
(Lybius chaplini) (south central Zambia), mostly found on the woody margins of flooded
grassland areas. The Tanzania masked weaver (Ploceus reichardi) occurs only in a few
swamps (western Tanzania, northern Zambia, and adjacent areas of the Democratic Republic
of Congo). Two further restricted range birds found in this ecoregion are the Katanga masked
weaver (Ploceus katangae) and the grey-crested helmet-shrike (Prionops poliophus). In
addition, this ecoregion falls within the center of distribution of the globally threatened slaty
egret (Egretta vinaceigula). Other vulnerable species include the wattled crane (Bugeranus
carunculatus) corncrake (Crex crex), lesser kestrel (Falco naumanni), great snipe (Gallinago
media), and shoebill stork (Balaeniceps rex). Rare birds found here include the long-toed
flufftail (Sarothrura lugens) and the white-headed plover (Vanellus albiceps). This delicate
ecoregion may face increasing threat due to the lack of specific conservation measures
required for the ongoing survival of these wetland ecosystems.
In the Afrotropical savannas, there are a good number of protected areas and outside formal
protection; habitats remain in reasonable condition in many regions. In spite of that, African
savannas are submitted to several threats, including uncontrolled wild fires, agriculture, illegal
hunting, deforestation, fragmentation, desertification processes, and other environmental
impact factors caused by human activities. Therefore, it is urgent and absolutely necessary the
adoption of conservation measures that prioritize the protection of the biodiversity in these
Global ecoregions.
3.2. Indo-Malayan
3.2.1. Terai-Duar
In the Indo-Malayan Global ecoregion, the Tropical Savanna is well represented by the Terai-
Duar Savanna and Grasslands region, a part of the Terai Arc landscape. This ecoregion (also
known as Bhabar) is a forested belt of rock, gravel, and soil eroded from the Himalayas where
the water table lies from up to 37 meters deep and is situated geographically north of the Terai
desert. The monsoon-flooded rivers of the Himalaya inundate the Terai yearly. Below lies the
great alluvial plain of the Yamuna, Ganges, Brahmaputra, and their tributaries. The annual
monsoon floods deposit silt from the rivers that meander across the grasslands. Alternate
layers of clay and sand are composed and this high water table creates many springs and
wetlands suitable for the local fauna. With only about 25 km wide, and an area of 34,600 km2,
this ecoregion covers a wide range of savanna-like habitats, from southern Nepal's Terai,
Bhabar, and Dun Valleys eastward to Banke and covering the Dang and Deokhuri Valleys
along the Rapti River, into India's states of Uttar Pradesh and Bihar. The tallest grasslands of
the world are found in the region. They are indicators of these humid conditions and nutrient-
rich soils. By the end of the monsoon, low-lying areas inundated for a few days only are
recharged with a yearly stack of nutrients and the areas covered in silt return to tall grasslands.
The Terai's diverse grasslands, riverine woodlands, hill and scrub forests provide a diverse set
of habitats for many birds. Biogeographically, the Terai straddles a transition zone between
the Palaearctic and Indo-Malayan realms, with species from both contributing to its
biodiversity, including rainforest, steppe and desert migrants. More than 550 bird species are
found in this ecoregion. More than ten species are grassland specialists and are found in
Terai’s grasslands or wetlands. These birds include the Bengal florican (Houbaropsis
bengalensis), lesser florican (Sypheotides indica), sarus crane (Grus antigone), and large grass
warbler (Graminicola bengalensis). Many are threatened, like the vulnerable Manipur bush-
quail (Perdicula manipurensis). The population growth pressure in the hills has led to
migration to and settlement (both spontaneously and through government-sponsored
programs). In the southern Terai, more than half of the region is under cultivation, although
northern regions have a lower population density, but poaching and overgrazing are still
problems. Faulty irrigation projects pose another significant threat and much of the savanna
grasslands may be converted by uncontrolled burning and other human intervention.
3.3. Australasia
The Northern Australia & Trans-Fly Savannas are included among tropical savannas
distributed in the Australasia region. They cover about 1,137,000 km2 and are sited in
Australia, Indonesia and Papua New Guinea. While most of Australia is covered with
grasslands and Papua with rainforests, the savannas are limited to moister areas in South
Papua and North Australia. This region is made up of the Australian savannas (Cape York
tropical savanna, Kimberly tropical savanna, Einasleigh upland savanna, Carpentaria tropical
savanna, Arnhem Land tropical savanna, Mitchell grass downs), and the Papuan Trans-Fly
savanna and grasslands. Patches of dry rainforest with high species diversity also occur
throughout the ecoregion.
3.3.1. Northern Australia
The Gulf of Carpentaria is a low, arid division between the wetter, upper Arnhem Land and
Cape York Peninsula. The distributary channels and alluvial fans at the deltas of the Flinders,
Mitchell, Gilbert, Leichardt, McArthur, and Roper Rivers have prolonged the seaward edge of
the plains. Somewhat extensive freshwater and saline wetlands merge along their lower
courses and around the edge of the Gulf. Many species restricted to sandstone substrates and
gorges are found here, including the Sandstone Antechinus (Pseudantechinus bilarni) and the
carpentarian grasswren (Amytornis dorotheae). Many of the wetlands in this region are highly
productive and are recognized as globally significant. They support the world's largest
breeding colonies of the magpie geese (Anseranas semipalmata). The Valley of Lagoons and
Pelican Lakes of Einasleigh, in the upper Burdekin River, are significant water bird habitat on
a national scale. Among the birds, the black grass-wren (Amytornis housei) and white-quilled
rock-pigeon (Petrophassa albipennis) are endemic to Kimberly tropical savanna. One of the
most distinctive features of the Mitchell Grass Downs are the extraordinary irruptions of one
of its most characteristic and key bird, the flock bronze wing (Phaps histrionica). This large
ground-foraging pigeon almost went extinct during the second half of the 19th century. In the
past, it prospered in flocks of hundreds of thousands, widely dispersing in response to rainfall
variation. They can still be seen across the Mitchell Grass Downs thriving in huge flocks.
Wetlands like the freshwater lakes of the Barkly Tableland, Lake Woods, the coolibah
swamps of Lake Tarrabool and Lakes Buchanan and Galillee in the Desert Uplands also
provide important habitat and breeding grounds for tens of thousands of water birds, most
notably the little curlew (Numenius minutus) and the oriental pratincole (Glareola
maldivarum). The Victoria Plains Tropical Savannas consist of extensive plains, dominated by
eucalypt woodland with a grassy understory, with rainforest elements and is punctuated with
several small patches of sandstone outcrops such as the Bungle Ranges, where, small pockets
of mesic vegetation and riparian strips are found in these sheltered gorges. It is delimited by
the Tanami Desert in the south and the Carpentaria and Kimberly tropical savannas to the
north. There is a clinal variation in climate from the north, which receives monsoonal rains to
the south, dry arid landscapes of central Australia. A very rich and abundant assemblage of
granivorous birds thrive here, including the endangered gouldian finch (Erythrura gouldiae)
and the threatened purple-crowned fairy-wren (Malurus coronatus) a riparian vegetation
specialist. There is limited endemism associated with some sandstone outliers, and many
typical savanna species reach their distribution. Despite the remoteness and its wilderness
character, inappropriate fire regimes associated with grazing management and invasive
species in the region are the most serious concerns. Continued overstocking and invasive
species are affecting two of the vital features of the region, the composition of native
grasslands communities, and riparian and wetland habitats. Improper land use and utilization
of remnants by cattle also degrades the associated monsoon rainforests. Exotic weeds like the
Rubber vine (Crytostegia grandiflora) and Prickly Acacia (Acacia nilotica) modify natural
nesting substrates. Introduced feral pigs (Sus scrofa) and other domestic animals consume
bird’s eggs and destroy their nests.
3.3.2. Trans-Fly
The Trans-Fly ecoregion is a center of plant diversity is also an endemic bird area located in
the southern part of New Guinea, spanning the international border of Papua New Guinea and
Indonesia. The area stretches from the Digul River in Papua to the Fly and Aramia Rivers.
With a total area of approximately 26,700,000 km2, this ecoregion encompasses all the
monsoonal vegetation types, including dry tropical forests, savannas, woodlands, grasslands
and southern New Guinea freshwater swamp forests, much of which is, in fact, one of the
driest parts of the savanna covering. The physiography is characterized by the coastal plains,
the dissected plateau of the Fly Digul shelf, the flat to gently undulating plateau and the flood
plains of the major rivers and tributaries. The climate is distinctly monsoonal and roughly
75% of the yearly 1,875 mm of rain falls in the wet season (December to May), the remainder
falls in the dry season (June to November). The region is lightly populated, and most people
are still living a traditional lifestyle of hunting, gathering and small scale agriculture. The
Trans-Fly protected areas are greater than all the remaining protected areas in Papua New
Guinea collectively. These new protected areas cover significant expanses of monsoon forests
and some key wetlands in the south and middle Fly floodplains. Several of the largest and
mostly intact wetlands in the Australasian region are found here, and more than 350 bird
species (about 50% of New Guinea’s total bird diversity) find their homes in the floodplains
of the slow flowing rivers and in the flanking savannas and monsoon forests, which are unique
to the Trans-Fly. These numbers include 80 species endemic to New Guinea and 6 that are
endemic or near-endemic to the Trans-Fly: Spangled kookaburra (Dacelo tyro), Little
paradise-kingfisher (Tanysiptera hydrocharis), Fly River grassbird (Megalurus albolimbatus),
Grey-crowned munia (Lonchura nevermanni), and the Black munia (Lonchura stygia).
3.4. Neotropical
The Neotropical region extends from Central Mexico south to the southernmost tip of South
America. This is the most biologically diverse region on Earth. More species of plants and
birds occur here than in any other region. Many diverse habitats occur in the Neotropics, and
not all of these habitats are tropical. The area of savanna in South America has apparently
expanded and contracted during cool-dry and warm-wet intervals during the Quaternary and
two or more recent episodes of savanna and other dry vegetation type expansion and
concomitant tropical humid forest contraction occurred during glacial periods in the Northern
Hemisphere. Savannas comprise the second most extensive vegetation type in tropical South
America, after tropical humid forests. The largest expanse of savanna is the Cerrado, followed
by the Llanos of eastern Venezuela and Colombia. Smaller areas of savanna include the Beni
Savanna, islands of savanna surrounded by humid forest in the Amazon basin, the Guyanan
Savannas, and disjunct areas of campo rupestre The Pantanal region of Brazil also includes
extensive areas of hyper-seasonal savanna vegetation.
3.4.1. Cerrado
The Cerrado occupies 190,000 sq. km covering the southeast of the Amazon Basin, in Bolivia,
Brazil, and Paraguay. The Brazilian savanna occupies a broad geographical area, extending
from Central Brazil (Mato Grosso, Mato Grosso do Sul, Tocantins, Goias, Distrito Federal) to
North Brazil (Para, Amazonas, Amapa, Rondônia states), Northeast Brazil (Piauí, Bahia,
Ceara Maranhao), Southeast Brazil (Minas Gerais, Sao Paulo) and South Brazil (Parana). The
climate is highly seasonal with a marked dry (May-August) and rainy (September-March)
seasons. This variation shows marked effects on the fauna and flora. It is one of the largest
savanna-forest complexes in the world, forming a gradient from open grasslands to closed
woodlands. The flora and fauna include several endemic species. The Cerrado is considered
one of the most diverse and important tropical savannas in the world. Typical bird species
include the greater Rhea (Rhea americana), Red-legged seriema (Cariama cristata), and the
critically endangered and endemic Spix's macaw (Cyanopsitta spixii). Among the endemic
species are included the White-striped warbler (Basileuterus leucophrys), Cone-billed tanager
(Conothraupis mesoleuca), and the Dwarf tinamou (Taoniscus nanus). The fragmentation of
habitats due to agriculture, cattle breeding, development projects and urbanization are the
major threats to the Cerrado conservation.
Most studies on grassland species richness show very little variation through the year, while
relative abundance was lower in the drier months. During the rainy season, there is a great
increase in insect populations, coinciding with the arrival of migratory species, like the Fork-
tailed Flycatcher (Tyrannus savanna). This species comes in flocks to the Cerrado in August
and September, breeds in the region, and returns to the Amazon between January and
February. During the dry season various species utilize gallery forests as foraging and shelter
sites. The stripe-tailed Yellow-finch (Sicalis citrina) is also an abundant species in the rainy
season, when it is observed in large migratory flocks. It is estimated that more than 850
species are found in the Cerrado, in total 64 families, of which 48 are threatened. More than
90% of these species breed in the biome. Endemic species as the White-rumped Tanager
(Cypsnagra hirundinacea), White-banded Tanager (Neothraupis fasciata) and Black-throated
Saltator (Saltator atricollis - Figure 2) are examples of nuclear species of campo cerrado. In
the past 35 years, more than half of its original area (1 million km2) was transformed mainly
in artificial pastures and annual crops. This represents a deforestation rate which is greater
than that of the Amazon. Recent studies show that the present deforestation rate varies from
22,000 to 30,000 km2/year, and if these rates persist, it is estimated that the remaining Cerrado
natural habitats will be totally destroyed by 2030.
Figure 2. Black-throated Saltator (Saltator atricollis), an endemic bird of the Cerrado and also
a nuclear species in mixed flocks. (Photo by Alexandre Gabriel Franchin)
3.4.2. Llanos
The Llanos Savannas are one of the world's largest wetland complexes (390,000 sq. km)
spreading in Northern South America, from Colombia to Venezuela. Throughout its elongated
range, the Llanos exhibits a high heterogeneity in landscapes and types of vegetation: the
foothills, the alluvial plains and the highlands, that may be divided in well-drained high plains
and floodable high plains. The wetlands compound includes streams, rivers, marshes, and
savannas. Wading and aquatic birds represent a large portion of the total bird fauna in these
flooded savannas. Most of the birds are usually restricted to the gallery forest and habitat
specialization is rare. Many birds thrive in agricultural areas as is the case of almost all
granivorous birds (pigeons, doves, finches, sparrows, etc). Some typical birds found here
include the Black-bellied tree duck (Dendrocygna autumnalis), Fulvous Whistling-duck
(Dendrocygna bicolor - Figure 3), Roseate spoonbill (Ajaia ajaja) and the Dickcissel sparrow
(Spiza americana). There are no endemic birds restricted to the llanos ecoregion, only the
Orinoco piculet (Picumnus pumilus) is a near-endemic species. The main threats affecting this
ecoregion are related to agricultural land misuse, farmland abandonment and rapid
Figure 3. Fulvous Whistling-duck (Dendrocygna bicolor), a typical bird of the Llanos. (Photo
by Alexandre Gabriel Franchin)
3.4.3. Pantanal
The Pantanal is a 171,000 sq. km2 sedimentary Flooded Savanna in western Brazil and the
largest wetland complex in the world. The Pantanal spreads from Central South America
(Bolivia, Brazil and Paraguay). Most of the Pantanal is located in Brazil and the Paraguay
river is the main watercourse of the region. The formation of the Pantanal is a result of the
flooding of the large pre-Andean Depression that forms an enormous internal delta, in which
several rivers flowing from the Plateau merge, depositing their sediments and erosion
residues. During the rainy season, from (December to April) this depression is almost totally
flooded, due to its low declivity and to the large amount of rainfall. During the dry periods
(July to December) the land is dry, though with some small scattered lagoons, several of
which are perennial during this period, constituting shelters for the fauna. When its
floodwaters recede, aquatic life becomes concentrated in scattered pools, and the convergence
of birds and animals to feed on it and breed make a spectacular wildlife phenomenon. The
relief is flat, since the altitudes are very low and soil and annual floods are determinant factors
of the biodiversity. The landscape includes a mosaic of flooded grasslands and savannas,
gallery forests, and dry forests. There are three migration routes bringing birds to the Pantanal
during either the wet or dry season, depending on the guild and the areas from which they
migrate from. At least 86 species including some from the Arctic and Patagonia migrate to the
area. Species such as the Roseate Spoonbill (Platalea ajaja), hyacinth macaw (Anodorhynchus
hyacinthinus) and Wood Stork (Mycteria americana) have their largest breeding populations
in the Pantanal and time their reproduction to coincide with a period of high food abundance.
The flora composition is very much influenced by other biomes, specially the Cerrado (more
than 50%). Aquatic birds are very abundant and some species are among the largest birds
found in wetlands, including the Jabiru stork (Jabiru mycteria - Figure 4). The main threats
for the ecoregion are: agriculture, water projects, mining, non-organized ecotourism, and road
construction. Siltation is pointed out as the worst environmental problem, changing the
hydrology (wet-and-dry to wet), fauna and flora, and the elimination of the riparian forest.
However, severe threats to the flora and fauna of the Pantanal originate outside the
Figure 4. Jabiru stork (Jabiru mycteria) can be found in great abundance in the Pantanal.
(Photo by Alexandre Gabriel Franchin)
3.4.4. Beni Savannas
The Beni savannas (also known as Llanos de Moxos) occupy 126,000 km2 and are situated in
the north-eastern lowlands of the south-western Amazon basin. Extending north-eastward
from the foot of the Andes, most of the Beni savanna spreads in the lowlands of northern
Bolivia (from El Beni, Cochabamba, La Paz, Pando, and Santa Cruz), extending some
portions to Brazil and Peru. The Beni savanna is surrounded by tropical moist forests; the
Southwest Amazon moist forests to the north, west, and south, and the Madeira-Tapajos moist
forests to the east. This ecoregion comprises seasonal savannas and wetlands with inland and
gallery forests on poorly drained alluvial plains with a marked dry season and flooding in the
rainy season. The plains form a complex mosaic of humid seasonal forest in a matrix of
seasonal savanna where yearly floods and fire are the defining factors. There are three major
rivers, the Beni to the west, Guapore (Itenez) to the east, and the Mamore in the central region.
The convergence of these rivers forms the major southern Amazon tributary, the Madeira
River. Typical bird species found here include the white-bellied Nothura (Nothura boraquira),
greater rhea (Rhea americana), southern screamers (Chauna torquata), plumbeous ibis
(Theristicus caerulescens), crowned eagles (Harpyhaliaetus coronatus), long-tailed ground-
doves (Uropelia campestris), golden-collared macaws (Ara auricollis), monk parakeets
(Myiopsitta monachus), burrowing owls (Athene cunicularia - Figure 5) and the critically
endangered and endemic Blue-throated macaw (Ara glaucogularis). The last species feeds
primarily on the fruits of Attalea sp. and Acrocomia sp. palms and it also nests on the palm
forests formed by these species. This macaw has a very low population density (estimates of
500-1,000 individuals). The main cause of their decline is capture for the pet trade and the
replacement of their natural habitat by artificial grasslands. Many other bird species are
suffering from a high hunting pressure and improved practices of forest management should
be applied.
Figure 5. Burrowing owl (Athene cunicularia) is a common owl in the Beni Savannas, as well
as in most regions of the Neotropics. (Photo by Alexandre Gabriel Franchin)
3.4.5. Guyanan Savannas
The Guyanan Savannas ecoregion occupies an area of 104,400 square kilometres, smoothly
rolling high plains, formed by sediments of the Roraima Formation, which overlie
discordantly the rocks of the Guyana Shield. It is distinguished by extensive savannas, many
rivers and scrubby vegetation, spanning northern Brazil, south-eastern Venezuela, and south-
eastern Guyana, southern Suriname and eastern Brazil (north of the Amazon, from near
Macapa to near Calcoene). In this ecoregion, recurrent fires and edaphic conditions are the
most important factors in the advance of savannas over forests. Endemism is low; however, 36
important endemic birds are found, and most of them on the Gran Sabana are primarily
montane species occurring in the humid forest on the piedmont slopes above 600 m. Some
examples are the Tepui Swift (Cypseloides phelpsi) that inhabits montane evergreen forest,
cliffs, rocky canyon, grasslands and savannas. The Tepui Goldenthroat (Polytmus milleri)
lives in the forest edge and low, seasonally wet grasslands and scrubs, and the Tepui Wren
(Troglodites rufulus), occurs in montane evergreen forest edge, elfin forest, scrub and
savanna. The near-threatened Rio Branco Antbird (Cercomacra carbonaria) is found on
riverine islands where it is relatively common in suitable forest. The endangered Hoary-
throated Spinetail (Synallaxis kollari) is very rare, being known from few specimens and a
couple of observations along six different rivers. The near threatened Blue-cheeked Amazon
(Amazona dufresniana) inhabits savannas, moist, gallery and cloud-forests. It is a low density
and very rare species.
3.4.6. Orinoco Wetlands
The Orinoco wetlands ecoregion occupies 6,000 sq. km and consists of flooded grasslands
embedded in a mosaic of mangroves, swamp forest, moist forest, and savannas, formed by the
sediment deposited by the Orinoco River through thousands of years. The ecoregion is located
in the north of the delta of the Orinoco River - the second largest river in the Neotropics - in
eastern Venezuela. It is both a globally important wetland, and a critical habitat to a number of
endangered species. Species compositions are doubtlessly influenced by the surrounding terra
firme moist forests, swamp forests, and mangroves. The monodominant forests of Buriti palm
(Mauritia flexosa) provide important food and also nesting habitat for many bird species.
Typical birds include the Orinoco goose (Neochen jubata), harpy eagle (Harpia harpyja) and
the endemics: Grey-legged Tinamou (Crypturellus duidae), Barred Tinamou (Crypturellus
casiquiare), Orinoco Piculet (Picumnus pumilus), Pelzeln's Tody-tyrant (Hemitriccus
inornatus), Yellow-throated Antwren (Myrmotherula ambigua), Spot-backed Antwren
(Herpsilochmus dorsimaculatus), Yapacana Antbird (Myrmeciza disjuncta), Grey-bellied
Antbird (Myrmeciza pelzelni), Chestnut-crested Antbird (Rhegmatorhina cristata), Orinoco
Softtail (Thripophaga cherriei), Azure-naped Jay (Cyanocorax heilprini), and White-naped
Seedeater (Dolospingus fringilloides). Threats to the area are increasing as water diversion
and damming, oil drilling, and human populations increase.
4. Birds in the Tropical Savannas
We recognize 2,951 bird species in the tropical savannas, representing about 29.54% of all
bird species in the world. A total of 25 bird orders (86% of bird orders), 152 families (77% of
bird families) and 1,404 genera (66.44% of bird genera) were computed. The Cerrado has the
greatest bird diversity, with about 850 species, which is more than 8% of the birds of the
world (Table 2).
Table 2. Number of bird species in the tropical savannas and geographical region, by to
taxonomic category.
Tropical Savannas
Neotropical Afrotropical
Trans –Fly
Species 618 442 850 471 547 610 541 538 392 558 418 359 2,951
Orders 20 19 20 20 19 20 20 20 20 17 18 17 25
Families 66 60 68 63 64 74 75 73 71 75 76 72 152
Genera 383 300 457 323 349 249 240 256 209 271 214 194 1,051
species 128 55 200 22 23 49 128 6 13 451 187 142 1,404
Many typical bird species that thrive in the tropical savannas are birds of open areas and
usually have their ecological counterparts on another savanna, like the south-American Red-
legged Cariama (Cariama cristata) and the African Secretary-bird (Sagittarius serpentarius),
both long-legged ground carnivores; the Neotropical Hummingbirds (Trochilidae) and the Old
World Sunbirds (Nectariniidae), both nectarivorous birds; Australasian emu (Dromaius
novaehollandiae), and Ostrich (Struthio camelus), both cursorial flightless birds.
Among the passerines, the Tyrannidae family has the most diversity, with 162 species
(39.51% of Tyrannidae species) which are exclusively found on the Neotropical savannas
(Figure 6). Another prominent bird family is the Accipitridae, with around 142 species in the
tropical savannas (58.68% of Accipitridae species) (Figure 7). This family has a worldwide
distribution and can be found in all savannas. Two other families which occur in all tropical
savannas were Psittacidae and Columbidae, which show, respectively 100 (26.73% of
Psittacidae species) and 85 (26.41% of Columbidae species) species (Table 3, Figure 8 and 9).
Comparing the representativeness of these four families, Accipitridae can be considered the
most typical bird family in all tropical savannas, because more than half the species
recognized can be found in savannas.
Figure 6. Tropical Kingbird (Tyrannus melancholicus) is a common flycatcher (Tyrannidae
Family) of the Neotropics. (Photo by Alexandre Gabriel Franchin)
Figure 7. White-tailed Hawk (Buteo albicaudatus) is a representative species of the family
Accipitridae. (Photo by Alexandre Gabriel Franchin)
Table 3. Distribution of the number of Tropical Savannas bird species by families.
Tropical Savannas
Bird Families
Acanthizidae 0 0 0 0 0 0 0 0 0 0 15 7
Accipitridae 28 18 29 23 34 38 38 34 36 42 17 21
Aegithinidae 0 0 0 0 0 0 0 0 0 1 0 0
Aegothelidae 0 0 0 0 0 0 0 0 0 0 1 5
Alaudidae 0 0 0 0 0 14 11 15 19 6 1 1
Alcedinidae 5 2 5 5 5 8 8 6 8 10 9 13
Anatidae 11 10 8 12 11 5 5 5 5 27 16 16
Anhimidae 1 1 2 2 2 0 0 0 0 0 0 0
Anhingidae 1 1 1 1 1 0 0 0 0 1 1 2
Anseranatidae 0 0 0 0 0 0 0 0 0 0 1 1
Apodidae 7 7 3 2 9 5 4 3 5 6 5 4
Aramidae 1 1 1 1 1 0 0 0 0 0 0 0
Ardeidae 22 15 10 14 16 9 9 8 9 18 15 15
Artamidae 0 0 0 0 0 0 0 0 0 1 6 2
Balaenicipitidae 0 0 0 0 0 1 1 0 0 0 0 0
Bucconidae 6 2 7 4 13 0 0 0 0 0 0 0
Bucerotidae 0 0 0 0 0 6 4 3 5 4 0 0
Bucorvidae 0 0 0 0 0 2 2 1 1 0 0 0
Burhinidae 1 1 0 0 0 2 2 1 1 2 2 2
Campephagidae 0 0 0 0 0 3 3 3 2 12 7 10
Caprimulgidae 9 11 13 11 17 9 9 7 5 4 3 3
Cardinalidae 6 4 6 7 9 0 0 0 0 0 0 0
Cariamidae 0 0 1 1 1 0 0 0 0 0 0 0
Casuariidae 0 0 0 0 0 0 0 0 0 0 1 1
Cathartidae 4 4 5 4 4 0 0 0 0 0 0 0
Certhiidae 0 0 0 0 0 1 1 1 1 1 0 0
Charadriidae 5 4 4 5 4 8 6 6 6 10 15 7
Chloropseidae 0 0 0 0 0 0 0 0 0 3 0 0
Ciconiidae 3 3 3 3 3 6 5 4 7 8 1 1
Cinclidae 0 0 0 0 0 0 0 0 0 1 0 0
Cinclosomatidae 0 0 0 0 0 0 0 0 0 0 0 1
Cisticolidae 0 0 0 0 0 21 19 13 22 9 2 2
Climacteridae 0 0 0 0 0 0 0 0 0 0 3 0
Coerebidae 1 1 1 1 1 0 0 0 0 0 0 0
Coliidae 0 0 0 0 0 2 2 2 5 0 0 0
Colluricinclidae 0 0 0 0 0 0 0 0 0 0 5 5
Columbidae 12 11 14 12 19 14 12 12 11 15 21 25
Conopophagidae 0 0 0 1 1 0 0 0 0 0 0 0
Coraciidae 0 0 0 0 0 5 4 3 4 2 1 1
Corcoracidae 0 0 0 0 0 0 0 0 0 0 2 0
Corvidae 2 0 3 3 4 4 3 4 2 6 3 2
Cotingidae 15 5 14 7 23 0 0 0 0 0 0 0
Cracidae 7 2 7 6 8 0 0 0 0 0 0 0
Cracticidae 0 0 0 0 0 0 0 0 0 0 6 2
Cuculidae 10 5 9 9 14 13 13 10 13 16 12 16
Dendrocolaptidae 13 8 9 10 21 0 0 0 0 0 0 0
Dicaeidae 0 0 0 0 0 0 0 0 0 5 1 2
Dicruridae 0 0 0 0 0 0 0 1 1 8 1 2
Dromaiidae 0 0 0 0 0 0 0 0 0 0 1 0
Emberizidae 22 24 27 25 45 4 3 3 4 6 0 0
Estrildidae 0 0 0 0 0 31 23 15 26 4 15 11
Eupetidae 0 0 0 0 0 0 0 0 0 0 1 0
Eurylaimidae 0 0 0 0 0 1 1 0 1 2 0 0
Eurypygidae 1 0 1 1 1 0 0 0 0 0 0 0
Falconidae 13 10 12 9 14 6 5 4 8 9 7 6
Falcunculidae 0 0 0 0 0 0 0 0 0 0 1 0
Formicariidae 2 1 1 0 4 0 0 0 0 0 0 0
Fringillidae 2 1 2 1 2 8 6 3 12 1 0 0
Furnariidae 16 9 18 19 40 0 0 0 0 0 0 0
Galbulidae 4 6 1 2 5 0 0 0 0 0 0 0
Glareolidae 0 0 0 0 0 5 5 4 6 2 2 2
Gruidae 0 0 0 0 0 3 3 2 2 1 2 1
Haematopodidae 0 0 0 0 0 0 0 0 0 0 0 1
Heliornithidae 1 1 1 1 1 0 0 0 0 0 0 0
Hemiprocnidae 0 0 0 0 0 0 0 0 0 2 0 1
Hirundinidae 12 12 12 11 14 18 13 8 18 6 5 4
Ibidorhynchidae 0 0 0 0 0 0 0 0 0 1 0 0
Icteridae 21 11 16 17 20 0 0 0 0 0 0 0
Indicatoridae 0 0 0 0 0 6 6 3 6 0 0 0
Irenidae 0 0 0 0 0 0 0 0 0 1 0 0
Jacanidae 1 1 1 1 1 1 1 1 1 2 0 1
Laniidae 0 0 0 0 0 9 10 5 7 5 0 1
Laridae 6 2 3 4 4 1 1 0 1 10 5 5
Machaerirhynchidae 0 0 0 0 0 0 0 0 0 0 1 1
Malaconotidae 0 0 0 0 0 14 15 11 13 0 0 0
Maluridae 0 0 0 0 0 0 0 0 0 0 9 1
Megapodiidae 0 0 0 0 0 0 0 0 0 0 2 3
Melanocharitidae 0 0 0 0 0 0 0 0 0 0 0 2
Meliphagidae 0 0 0 0 0 0 0 0 0 0 42 25
Meropidae 0 0 0 0 0 14 11 5 10 4 1 2
Mimidae 2 2 3 3 3 0 0 0 0 0 0 0
Momotidae 1 0 1 1 4 0 0 0 0 0 0 0
Monarchidae 0 0 0 0 0 4 2 1 4 2 11 7
Motacillidae 1 1 1 1 3 12 13 5 13 13 5 1
Muscicapidae 0 0 0 0 0 30 23 14 26 42 0 1
Musophagidae 0 0 0 0 0 5 3 1 1 0 0 0
Nectariniidae 0 0 0 0 0 21 17 8 18 7 1 2
Neosittidae 0 0 0 0 0 0 0 0 0 0 1 0
Numididae 0 0 0 0 0 1 1 1 1 0 0 0
Nyctibiidae 3 1 2 2 3 0 0 0 0 0 0 0
Odontophoridae 1 1 2 0 3 0 0 0 0 0 0 0
Opisthocomidae 1 0 1 0 1 0 0 0 0 0 0 0
Oriolidae 0 0 0 0 0 2 2 1 2 4 3 4
Orthonychidae 0 0 0 0 0 0 0 0 0 0 1 0
Otididae 0 0 0 0 0 8 8 5 6 2 0 1
Pachycephalidae 0 0 0 0 0 0 0 0 0 0 4 2
Paradisaeidae 0 0 0 0 0 0 0 0 0 0 2 6
Pardalotidae 0 0 0 0 0 0 0 0 0 0 3 0
Paridae 0 0 0 0 0 5 4 2 5 2 0 0
Parulidae 9 6 5 5 10 0 0 0 0 0 0 0
Passeridae 1 0 0 1 1 13 14 9 8 3 1 0
Pelecanidae 0 0 0 0 0 1 1 1 1 3 1 1
Petroicidae 0 0 0 0 0 0 0 0 0 0 9 5
Phalacrocoracidae 1 1 1 1 1 1 1 1 1 3 3 4
Phasianidae 0 0 0 0 0 10 10 5 11 7 3 2
Phoenicopteridae 1 0 0 0 0 0 0 0 0 0 0 0
Phoeniculidae 0 0 0 0 0 5 6 4 3 0 0 0
Picidae 17 11 16 19 25 11 10 7 7 19 0 0
Pipridae 10 9 10 3 15 0 0 0 0 0 0 0
Pittidae 0 0 0 0 0 0 0 0 0 3 1 2
Platysteiridae 0 0 0 0 0 5 5 3 5 0 0 0
Ploceidae 0 0 0 0 0 41 37 25 29 4 1 0
Podargidae 0 0 0 0 0 0 0 0 0 0 2 1
Podicipedidae 2 2 0 3 3 0 0 0 0 2 3 2
Polioptilidae 2 2 1 1 3 0 0 0 0 0 0 0
Pomatostomidae 0 0 0 0 0 0 0 0 0 0 0 1
Promeropidae 0 0 0 0 0 0 0 0 1 0 0 0
Psittacidae 17 15 26 18 33 9 5 4 7 7 18 20
Psophiidae 1 1 0 0 0 0 0 0 0 0 0 0
Pteroclididae 0 0 0 0 0 3 3 3 2 0 0 0
Ptilonorhynchidae 0 0 0 0 0 0 0 0 0 0 5 3
Pycnonotidae 0 0 0 0 0 5 4 1 6 7 0 0
Rallidae 12 13 10 9 16 11 10 8 11 10 13 7
Ramphastidae 9 4 5 4 11 18 11 9 13 4 0 0
Recurvirostridae 1 1 1 1 1 1 1 1 1 2 2 1
Remizidae 0 0 0 0 0 2 3 1 2 1 0 0
Rheidae 0 0 1 1 1 0 0 0 0 0 0 0
Rhinocryptidae 0 0 1 0 2 0 0 0 0 0 0 0
Rhipiduridae 0 0 0 0 0 0 0 0 0 3 4 5
Rostratulidae 0 0 0 0 1 1 1 1 1 1 1 0
Scolopacidae 18 12 9 15 12 9 9 9 9 21 22 20
Sittidae 0 0 0 0 0 0 0 0 0 3 0 0
Strigidae 10 9 10 8 15 9 9 7 7 13 3 2
Struthionidae 0 0 0 0 0 1 1 1 1 0 1 0
Sturnidae 0 0 0 0 0 15 16 15 16 9 1 5
Sylviidae 0 0 0 0 0 24 21 15 18 40 7 5
Thamnophilidae 40 27 26 16 54 0 0 0 0 0 0 0
Thraupidae 37 19 29 20 52 0 0 0 0 0 0 0
Threskiornithidae 8 6 7 6 6 2 2 3 3 3 5 6
Timaliidae 0 0 0 0 0 7 7 3 4 29 0 0
Tinamidae 8 4 8 5 16 0 0 0 0 0 0 0
Trochilidae 25 22 23 17 37 0 0 0 0 0 0 0
Troglodytidae 10 5 5 5 7 0 0 0 0 0 0 0
Trogonidae 4 2 4 2 7 1 1 1 1 0 0 0
Turdidae 10 3 7 3 9 3 2 2 4 9 2 0
Turnicidae 0 0 0 0 0 3 3 3 2 2 7 1
Tyrannidae 78 63 78 61 115 0 0 0 0 0 0 0
Tytonidae 1 1 1 1 1 1 0 0 1 2 4 4
Upupidae 0 0 0 0 0 1 1 1 1 1 0 0
Viduidae 0 0 0 0 0 11 6 4 4 0 0 0
Vireonidae 6 5 3 4 7 0 0 0 0 0 0 0
Zosteropidae 0 0 0 0 0 1 1 1 2 1 1 2
Total 618 442 547 471 850 610 538 392 541 558 418 359
Figure 8. Picazuro Pigeon (Patagioenas picazuro) is a columbid found at the Cerrado. (Photo
by Alexandre Gabriel Franchin)
Figure 9.Red-shouldered Macaw (Diopsittaca nobilis) is one of the a hundred species of
Psittacidae occurrence in the tropical savannas. (Photo by Alexandre Gabriel Franchin)
4.1. Similarity in the Bird Composition among Tropical Savannas
Three birds are common to all tropical savannas: Cattle Egret (Bubulcus ibis), Barn Swallow
(Hirundo rustica), and Osprey (Pandion haliaetus) (Figure 10). The House sparrow (Passer
domesticus) has only recently been colonizing the Trans-Fly region, since the 1980s. The
tropical savannas also share 15 orders, 20 families and 19 genera. Twenty families were
exclusive to only one savanna, and the Pantanal had no exclusive family. The Australian
savannas had the greater number of exclusive families (eight families) (Table 3). When gender
is considered, we can account for 311 exclusive savannic bird genera and the majority occurs
in the Terai-Duar savanna (107 genera). An analysis of similarity of bird genera among the
savannas shows that the taxa occurring in these areas are, truly, distinct, because the
Sorenson’s similarity indexes (Cs) were below 0.40. The Afrotropical savannas (Sudanian,
East, Horn, and Central) are those that can be considered in fact similar, because they
presented a minimum Cs = 0.86. The Neotropical savannas (Llanos, Guyanan, Beni, Pantanal,
and Cerrado) and the Australasian savannas also are those that can be considered in fact
similar, because they presented a minimum Cs = 0.70 (Table 4). Despite of this, a cluster
analysis shows the strong similarity among two distinct groups: (1) African Savannas Terai-
Duar, Australasian Tropical Savannas and (2) South American Tropical Savannas. (Figure
Table 4. Matrix of Similarity with the values of the Index of Sorensen for the presence and
absence of genera of the birds found in the tropical savannas.
Figure 10. Cattle Egret (Bubulcus ibis) is one of the three species that were registered in all of
the tropical savannas of the world. (Photo by Alexandre Gabriel Franchin)
Tropical Savannas
Llanos 1.00
Guyanan 0.79 1.00
Beni 0.77 0.73 1.00
Pantanal 0.75 0.73 0.82 1.00
Cerrado 0.80 0.71 0.83 0.81 1.00
Sudanian 0.13 0.13 0.12 0.13 0.11 1.00
East 0.13 0.14 0.12 0.13 0.11 0.96 1.00
Horn 0.13 0.14 0.12 0.14 0.11 0.88 0.92 1.00
Central 0.13 0.14 0.12 0.13 0.11 0.90 0.88 0.86 1.00
Terai-Duar 0.17 0.18 0.15 0.17 0.15 0.43 0.43 0.43 0.42 1.00
Northern Australia 0.14 0.14 0.11 0.14 0.11 0.28 0.28 0.29 0.29 0.40 1.00
Trans-Fly 0.14 0.14 0.11 0.14 0.10 0.28 0.29 0.30 0.28 0.40 0.70 1.00
All these patterns point to complex historical and biogeographical processes in the
diversification of tropical savanna birds. It shows evidence of a greater similarity among
savannas that are closer geographically, indicating that a good part of the bird populations in
tropical savannas depends mainly on the historic processes and their connectivity with other
In South America, the cluster analysis indicates two groups: one formed by the Llanos and
Guyanan, and another by the Cerrado, Pantanal and Beni. Most of the fauna in the Guyanan
savannas’ birds are not dependent on forest formations and their avifauna shows more
similarities with the fauna of open areas in Llanos than with the fauna of Cerrado or with the
other Amazonian savannas. The Beni, Pantanal and Cerrado show a remarkable similarity
evidencing their close geographic connections. This geographic pattern can also be identified
for the North Australian and Trans-Fly Savannas, which share a many other vertebrate
species, even though the Trans-Fly avifauna composition is strongly influenced by typical
tropical rainforest birds. In Africa, most old species are widespread across a physiognomic
and climatic domain. In contrast, new species have evolved in ecologically equable places
inside geologically complex ecotonal regions (e.g. Horn of Africa). High species richness and
taxonomic diversity, were maintained over wide areas by steady habitat alteration through
patch dynamics (e.g. Sudanian). In Asia, the Terai-Duar savannah and grasslands are a unique
spot where a strongly patchy environment created special conditions for the rich avifauna
thriving in such a very small area. This reality cannot be obscured by the fact that many of the
birds found here are seasonal migrants from the Himalayas or Asian tropical forests.
When we compare similar large Flooded Tropical Savannas from Africa (Zambezian Flooded
Savannas), South America (Pantanal and the Llanos) and Australasia (Trans-Fly), we must do
so carefully. The Trans-Fly is located in an island of very recent origin (Pleistocene), and thus
ecological theory predicts lower diversity. Regionally, the Trans-Fly is very rich in avifauna
and no other site in the region compares to it. The endemism comes from two peculiar features
of the Trans-Fly: its location in an island and both its geomorphology and origins are unique
in Australasia. The Pantanal is also of Pleistocenic origin, but its bird fauna is influenced by
the connectivity to highly diverse ecosystems, such as the Amazon and Brazilian Atlantic
Forests, the Chaco and the Andes’ foothills. The diversity of birds in the Pantanal is higher
than in the Trans-Fly, however, the comparison is inappropriate because the Pantanal is 12
times bigger than the Trans-Fly. Proportionally, the Pantanal contains fewer endemic species
compared to the Trans-Fly because of its connectivity to other systems (e.g. Amazon forest,
Cerrado, Chaco, Chiquitano and Atlantic forest). Another important tropical savanna wetland
is the Zambezian Flooded Savannas, which house hundreds of migratory species that arrive in
their seasonal migrations across the African continent and into Eurasia. The Okavango Delta
is one of the main areas in this complex. Formed largely of Precambrian volcanics, granite,
serpentine and sandstone rocks, this ecosystem is much older than the Trans-Fly and Pantanal,
but because it is surrounded by large areas of high diversity, much like Pantanal,
proportionally it has fewer endemic species than the Trans-Fly (The only strict endemic is the
Kilombero weaver Ploceus burnieri). The Llanos savannas are similarly seasonally flooded
wetlands and a mosaic of habitats. Their origin is recent and is related to the Orinoco and
Magdalena rivers drifting eastwards as a consequence of the Andean uplifting (five m.y.a).
This region is also an important ecosystem for numerous migratory birds, especially wildfowl,
and is adjacent to the region with the highest bird diversity in the world: the Andean foothills.
The Llanos contains bird species typical of Amazonian forests and are usually not a main
route for migratory species from North America. Most species follow the Andes mountain
ranges to South America, including the Pantanal and the South American Pampas. Endemism
is also very low in the Llanos.
4.2. Patterns of Bird Diversity and Speciation in the Tropical Savannas
Bird assemblages are strongly influenced by habitat structure. Fire is an important determinant
of several aspects of the savanna ecosystem, because it alters the quantity or quality of habitat
resources. The variation of fire an area receives directs the response of bird assemblages in the
post-fire habitat. The highest landscape-level bird diversity might have developed through
complex sequences of fire regimes and random patchy burning. Savanna fires may spread
along lines many kilometres long. The abundances of insectivores and granivores may
increase in frequent dry-season fires. Although burning reduces the plant biomass, fire-
adapted remaining plants often exhibit traits that enhance survival in the post-fire
environment, including flushes of new growth via shoots, vegetative regrowth, resprouting,
and flowering events. Alternatively, frequent wet-season burnt sites may favor carnivores, by
greater accessibility to prey following the removal of understory vegetation. Consecutive
years of burning may reduce bird abundances, especially of nectarivores and granivores. The
historical state of bird diversity in tropical savannas is very much linked to fire occurrence and
adaptations of many plant species, because patchy fires have been an important natural
disturbance in tropical ecosystems for ages.
Differences in fire regimes can substantially alter bird assemblages, especially in riparian
zones. Riparian zones are also a key component of many tropical savanna environments. The
spatial patterning, the supplementary assemblages, the occurrence of rare species and the
potential for riparian habitats to serve as refuges reflect the uniqueness of the riparian zones to
the tropical savannas’ bird diversity. Open vegetation enclaves contribute significantly to local
avian species richness. Habitat selection and adaptation to local climate may be the primary
processes structuring bird diversity among landscapes within ecoregions, and dispersal
limitation has a lesser role in influencing beta-diversity among landscapes. This implies the
possibilities that most open country birds maintain higher levels of gene flow than forest
understory birds, and may have expanded to parts of their present-day distribution fairly
The African Savannas encompass almost fully the African Tropical Rain Forests and are
influenced by the Sahara desert to the north and the Kalahari to the southeast. The ecotones
make up a great part of the African Savannas. The mosaic of trees, shrubs and open grassland
in African savannas is highly dynamic and strongly influenced by mammal herbivory and fire.
In the plains, the large mammal herbivores control both the canopy area of subdominant
woody vegetation and the biomass of ground-dwelling arthropods, and both of these factors
are good predictors of the bird diversity. The canopy area of subdominant trees is positively
correlated with the diversity of granivorous birds and the biomass of ground-dwelling
arthropods is positively correlated with the diversity of insectivorous birds. Most native large
herbivores are compatible with an abundant and diverse bird fauna if they are at a relatively
low density. In African Savannas, floristic species composition seems to be less important for
bird communities’ composition than vegetation structure.
In Australia, the savannas border the sea to the North and the Australian Desert to the south,
but encompass a very small region of Tropical Rain Forest. Here, the regional avifaunal
exchange is very low as is also the bird diversity, although the endemism is the highest among
other Tropical Savannas. Probably the most important of these endemic birds are the
Honeyeaters (Meliphagoidea). Australian savannas are characterized mainly by woodlands
dominated by Eucalyptus sp. species and many grasses like Aristida sp., Triodida sp. and
Plectrachne, which are all resinous and mostly unpalatable. These savannas lack the large
mammal herbivores that historically have shaped the African vegetation mosaics. North
Australian Savannas have also been shaped by fire and the current knowledge indicates that
this region has had a long history of regular fire extending back millions of years.
The contrasting higher richness of the Neotropical Savannas in South America is perplexing.
One possible explanation is the greater rates of turnover, but interestingly, many studies have
shown that Tropical Savanna vegetation types within the Neotropics have twice the number of
species as communities in similar vegetation types within the Afrotropics. This is attributed to
greater vertical stratification, largely of arboreal species, which make up a higher proportion
of the Neotropical fauna than the Afrotropical fauna. Termites and harvester ants also play a
great role in the dynamics and functionality both of the Australian and neotropical savannas,
where the large mammal herbivores are mostly absent. Their underground chambers improve
the porosity of the soil, creating better conditions for plant growth especially in soils where
they are historically unfavorable. Termite mounds are a characteristic feature in most savanna
The Cerrado ecoregion occupies a central position in South America and it is limited in the
north by the Amazon forest, northeast with the Caatinga, and east and southeast with the
Atlantic Forest and Southeast with the Chaco and Pantanal. The biodiversity of the Cerrado is
directly related to the faunal exchanges between these adjacent regions, which follow the
climatic and vegetational oscillations of the Quaternary. As a result, Atlantic Forest and
Amazon species expanded their distribution to the Cerrado during the wet periods subsequent
to the riverine and gallery forest expansions. Therewith, during the Quaternary dry periods,
the Caatinga and Chaco faunal elements colonized the Cerrado subsequently to the
development of the dry forests in the peripheral depressions. These connections between north
and south probably took place through two main corridors. First, the Andes corridor connects
the south portion of the savannas to the Oriental Llanos and Roraima Savannas along the
Andes hillsides. Second, a Coastal corridor runs along the Brazilian Atlantic Forest and
connects the south and north through the savanna mosaics found near the Atlantic Coast, like
Marajo and Amapa. Some gallery forest birds have their core area distribution in the Amazon
(202 taxa) and others in south Atlantic Forest (79 taxa). The Amazonian faunal elements
extended their distribution proportionally more into the Cerrado because of the low altitudes
that separate them. Nevertheless, 86% of the Cerrado bird species have their range restricted
to a 250 km radius inside the biome. In this manner, both the distance from the distribution
center and the altitude seem to determine the distribution of birds in the gallery forest
complexes of Central Brazil.
4.3. Bird Conservation in the Tropical Savannas
There are 2,057 bird species in the world that face some degree of threat (21% of the
recognized bird species), not considering the extinct species or those with insufficient data
available. In the tropical savannas, there are 319 species that face some degree of threat. The
Cerrado is the tropical savanna with the most threatened bird fauna (58 species), and seven of
them are critically endangered. The Terai-Duar and Central African savannas also show
alarming numbers in their territory with 49 and 36 species respectively (Table 5).
Table 5. Number of threatened bird species in the Tropical Savannas of the world.
Neotropical Afrotropical
Malayan Australasia
2008 IUCN
(CR) 0 1 2 1 7 1 1 2 0 6 0 0 21
(DD) 0 0 0 0 0 1 1 0 6 0 0 2 10
(EN) 2 2 2 2 5 1 4 3 2 6 4 0 33
(NT) 7 5 11 9 31 18 15 10 15 19 12 14 166
(VU) 0 0 5 3 15 10 8 4 13 18 6 7 89
Total of
Savannas 9 8 20 15 58 31 29 19 36 49 22 23 319
Different mechanisms are responsible for the origin and maintenance of different aspects of
diversity. It is important to evaluate biodiversity concordance and the use of indicator groups
to provide a basis to evaluate priorities for conservation at a regional scale. Highly endangered
ecoregions with many strait endemic species require focused actions to prevent the loss of
further habitat leading to their extinction. Less threatened ecoregions also require maintenance
of large and well-connected habitats that will support large-scale ecological processes.
5. Cerrado: A Unique Tropical Savanna Hotspot
The Brazilian savanna, known as Cerrado, is one of the largest and the richest savanna in the
world, and the most threatened of them all, considered one of the 25 terrestrial biodiversity
“hotspots”. In Brazil, this biome is the one of the prominent in number of threatened and
endemic bird species, second only to the Atlantic Forest. The families with the greatest
species richness are: Tyrannidae (113 species), Thamnophilidae (54 species), Thraupidae (52
species), Emberizidae (45 species), Furnariidae (40 species), Trochilidae (37 species),
Accipitridae (34 species), and Psittacidae (33 species). Most of these species (72%) are
dependent (partially or totally) of forested habitats (cerradao, dry and riparian forests), which
represent less than 10% of the Cerrado phytochorion. Only 25.5% of the birds (218 species)
are forest independent, that is, they feed or reproduce mainly in the cerrado stricto sensu and
in other open vegetation habitats.
One of the most conspicuous Cerrado species in relation to morphology and behavior is the
Cock-tailed Tyrant (Alectrurus tricolor), a tyrannid which occurs in grassland areas of the
Cerrado. The male’s tail has a peculiar shape: the two median rectrices rotate 90º, in a way
that its vexiles remain in a vertical position as the sail on a ship (Figure 12). During the
breeding season, the male performs several displays before the females while in flight, and
numerous males engage in confronting aerial displays between other conspecifics. This
species is vulnerable mainly due to habitat destruction.
Figure 12. The tail of the Cock-tailed Tyrant’s male (Alectrurus tricolor) is one of the most
peculiar among savanna birds. (Photo by Mieko Ferreira Kanegae)
The Greater Rhea (Rhea Americana) is the largest south American bird, and can stand at
1.70m tall. Belonging to the ancient ratite group, the Rheiformes were well represented in this
region during the Upper Paleocene, about 55 million years ago. The male drives away its
rivals and forms a group of about three to six females. The male is also in charge of nest
building, egg incubation and caring of the chicks. The nest may contain 12 to 30 eggs that
weigh about 605g on average, but there are records of nests containing 60 or more eggs.
Formerly, this bird could be seen in flocks of large numbers in the Cerrado (20 to 30
individuals), but at the present time, the flocks are much reduced, mainly because of the
hunting pressure and habitat loss.
One typical Cerrado bird that has a long-range vocalization is the Red-legged Seriema
(Cariama cristata). The “laughing” song of the seriema is also known as the “voice of the
Cerrado”, and this is related to the persistent, well-defined and riotous screams that speed up
and then slow down. They are found in pairs or in small flocks. They consume insects and
other arthropods, small rodents, lizards and amphibians.
The Curl-crested Jay (Cyanocorax cristatellus) is a corvid endemic to the Cerrado and it is
common to the open grassland formations. It is found in flocks of about 8 to 10 individuals.
The acoustic recognition of this species may be accomplished at great distances when it calls
in a repetitive and stridently pattern the “grawn", “grawn", “grawn". During the breeding
season, it exhibits a more distinctive social behavior, with many individuals participating in
cooperative parental care. Predation is very intense for this species with nestling lost up to
The Sharp-tailed Tyrant (Culicivora caudacuta) is an uncommon small tyrannid (10 cm long)
which is found in Southeast and South Brazil. The habitat of this species is campos limpos and
campos sujos. The eradication of campos limpos by agriculture threatens this species, because
this flycatcher does not adapt well in artificial grassland landscapes. Kin groups are found
during winter (3-10 individuals) and they generally participate in mixed flocks, usually
including the Chalk-browed Mockingbird (Mimus saturninus), Cock-tailed Tyrant (Alectrurus
tricolor), White-banded Tanager (Neothraupis fasciata), White-rumped Tanager (Cypsnagra
hirundinacea), and Bearded Tachuri (Polystictus pectoralis). The bird builds a thin, shallow
cup-shaped nest in short bushes, where it incubates 2 to 5 eggs.
Of the total species number recorded for the Cerrado, 759 (90.7%) reproduce inside the
biome, 26 (3.1%) are northern hemisphere migrants, 12 (1.5%) are migrants South America
migrants, 8 (0.9%) are possibly altitudinal migrants of the Brazilian southeast and 32 (3.8%)
have currently unknown status. One of the most common birds in Central Plateau Cerrado,
especially during the rainy season, when it arrives for reproduction is the Lesser Elaenia
(Elaenia chiriquensis). In spite of being regularly the most captured bird in mist nets during
this season, very little is known about its migration routes, primarily due to the permanence of
some populations in the Cerrado. This bird feeds mostly on fruits, and its abundance is related
to its predominant diet. Our current knowledge on this species indicates that it is a key-species
for seed dispersion in the Cerrado.
Many granivorous species are migratory and follow the seed production of native and exotic
grasses in the open grasslands. Despite of the great granivore richness, in the Cerrado,
particularly in the Sporophila genus (18 species - Figure 13), very little is known about its
migratory routes. Species as the Plumbeous Seedeater (Sporophila plumbea) are much more
sensible to habitat alterations, as a rule, being led to extinction as a consequence of the
introduction of exotic grasses. Unfortunately, one of the greatest threats in the Cerrado is the
expansion of these invasive plant species, like the Kenya sheep grass (Brachiaria decumbens)
and Gordura grass (Melinis minutiflora). These African plant species are very well adapted to
the region’s climate, competing with local species. The management of these invasive grasses,
despite demanding a high cost, must be implemented in conservation units de before their
expansion makes these procedures not manageable.
Figure 13. The Lined Seedeater (Sporophila lineola) is one of many Sporophila genus species
found in the Cerrado. (Photo by Alexandre Gabriel Franchin)
In the Cerrado 172 bird species participate in mixed flocks in many phytochoria and these
flocks are constituted of even 16 species and 40 individuals. In general, these are species of
contrasting coloration and occur in conspecific flocks when absent in mixed flocks. The
nuclear species remain much of the time on the watch-out and make alert calls as a result of
the proximity of falcons and hawks. For some typical forest birds that explore the adjacent
savannas, the strategy of joining mixed flocks lowers predation risks associated to the more
open areas.
Despite the very low endemism [36 species (4.3% of total)], many species (39%) are included
in some degree of threat, due mostly to habitat loss. Since 1995, three endemic species were
described to the Cerrado, two of which are already in the IUCN red list.
The Cipo Canastero (Asthenes luizae) is a furnariid that inhabits rock outcrops that are
associated to dry vegetation, in altitudes above 1,100 m and below 1,500 m. It has a very
restrict distribution and it is considered vulnerable due to landscape modification by cattle
grazing, frequent fires and the expansion in the distribution of a nest parasite bird species, the
Shiny Cowbird (Molothrus bonariensis).
The Collared Crescentchest (Melanopareia torquata) is an endemic species which occurs
mainly in campo cerrado and campo sujo. It is a solitary and probably sedentary species,
being less common in observation of pairs. The individuals move about mainly on the ground,
on the lower parts of bushes, usually above tall grasses. It is possible to hear their song during
the whole year, and their detection is more acoustic than by sight. They perch on the highest
part of low bushes and call usually a constant whistling series “feeoo”, “feeoo” that are
strongly resonant and loud (Figure 14). This species is also found in the List of Threatened
Birds in the State of Sao Paulo (Animals Red List of the Sao Paulo State - Decret n° 42.838 of
February 4th, 1998), in the category endangered.
Figure 14. The Collared Crescentchest (Melanopareia torquata), an endemic species of the
Cerrado. (Photo by Mieko Ferreira Kanegae)
In the Cerrado, unfortunately, the conservation efforts are still modest and only 2.2% of this
biome is under legal protection. This low covering does not even reflect the current Forest
Code legislation, which demands that 20% of Cerrado converted to agriculture or cattle
raising be preserved. Depending on the dominant vegetation 70% of the biomass is immersed
in the subsoil. An analogy commonly made to the Cerrado is that it is a forest upside down.
Unfortunately, the view of the Cerrado as an important carbon dioxide sink is only modestly
discussed in the media. Community mobilization, biodiversity, environmental services, and
top-down conservation are severely limited as environmental policy initiatives, given that the
governmental agencies are incapable of inspecting and assuring the legal protection. Not much
importance is given to a vegetation frequently referred as “dry and tortuous”, resulting in an
empty niche occupied by farmers. It is estimated that an area the size of the state of Sao Paulo
is underused by cattle farmers. It is easier and more lucrative to deforest new Cerrado
landscape than to restore degraded areas. Modern simulations modelled the transformation of
the actual Cerrado in artificial grasslands pastures and have shown that the precipitation could
be reduced at least 10%, droughts could become recurrent and the average superficial air
temperature may increase by 0.5ºC, bringing pessimistic impacts for agriculture. Important
bird areas for preservation and sustainable use in the Cerrado that should be protected are
being substituted by sugar cane plantation for ethanol production. According to ISPN institute
(Sociedade, Populaçao e Natureza), it is estimated that a total of 142 thousand ha of Cerrado -
the size of Sao Paulo City - are considered priority conservation units but were converted to
sugar cane plantations in the 2006/2007 harvest. The list is led by Sao Paulo State (86
thousand hectares deforested), followed by Minas Gerais State (25,000), Goias State (13,000),
Mato Grosso State (12,000) e Mato Grosso do Sul State (6,000) states. This deforesting may
compromise the natural resources of rural populations and the food supply in the region.
According to ISPN, the problems of the preservation in the Cerrado start with the lack of
monitoring of vegetation clearing and lack of legal regulation concerning this activity.
Furthermore, in the growing interest in bio fuels, especially in Brazil, the prevision to the
disappearing of Cerrado vegetation outside conservation units has one of the most pessimists.
In case there is not an adequate certification that guarantees the legality of plantation
associated to the maintenance of the legal reserves and permanent protection areas, according
to the Brazilian Forest Code, the importers of these bio fuels in Brazil are conniving with this
Fire is a determinant factor in the formation and patterns of ecological processes in tropical
savannas. The relation of the Cerrado with fire is an ancient one and it is prior to human
occupation (27,100 to 41,700 years ago). Palinological analysis reveals that the longest
intervals between fires in the Cerrado were, on average, 33 to 41 years and the shortest, about
8 years. With increasing human occupation, the frequency of fires has been increasing, being
widely utilized in agriculture and extensive cattle ranching. An “unnatural” fire regime can
interfere negatively on the biota, modifying the evolutionary processes kept under different
fire conditions. Uncontrolled fires may affect birds in a direct way (mortality rate, physical
injuries) and indirectly (changing the habitat, offering resources, abundance of competitors
and predators). Ecologically, birds respond differently, depending on their life history traits, as
length, intensity and duration of fires interval may vary. Species such as the Coal-crested
Finch (Charitospiza eucosma) and the Campo Miner (Geobates poecilopterus) are considered
specialists of burned sites. Other species as the Sedge Wren (Cistothorus platensis) and the
Cock-tailed Tyrant (Alectrurus tricolor) are more sensible, disappearing after fires. Frequently
burnt sites affect negatively the establishment of trees and bushes, in addition to the liberation
of CO2 and other greenhouse gases. Species like the Horned Sungem (Heliactin bilophus),
Wedge-tailed Grass-finch (Emberizoides herbicola), Streamer-tailed Tyrant (Gubernetes
yetapa - Figure 15), and the Cock-tailed Tyrant (Alectrurus tricolor) disappear in many burnt
areas soon after the passage of fire, returning to the site only months later. Others, like
Southern Lapwing (Vanellus chilensis), Spotted Nothura (Nothura maculosa), and the Huayco
Tinamou (Rhynchotus rufescens) increase in number and frequency in September, a few
weeks after the fires.
Figure 15. The Streamer-tailed Tyrant (Gubernetes yetapa), one of the most affected bird
species by action of fire. (Photo by Alexandre Gabriel Franchin)
We thank Dr. Kleber Del-Claro for the invitation to write this chapter, and the colleagues at
the Ornithology Lab, UFU, specially Laíce J. Silva and Diego S. F. Oliveira for the help with
the database. We also thank CAPES for the doctoral scholarships of AGF and MFK and UEG
for the doctoral scholarship of RFJ.
Related Chapters
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Avifauna: The total set of bird species in a given region.
Beta-diversity: A measure of biodiversity which compares the species diversity between ecosystems or
along environmental gradients
Biodiversity: The total of genes, species and ecosystems of a place.
Biogeography: Science studying the spatial distribution of living organisms and its causes, either
ecological or evolutionary.
Biome: A large set of ecosystems, defined by vegetational and climatic characteristics.
Caatinga: Areas of dry shrub lands in northeast Brazil, extend over 800,000 square kilometres.
They have virtually no grass substrate and more dry season moisture availability.
Campo cerrado: A Cerrado type of parkland composed of scattered trees in dense grass.
Campo limpo :
(“clear field”) A treeless grassland with herbs, a few small shrubs, and sub-shrubs.
Campo rupestre:
(“rocky field”) A shrubby or grassland savanna associated with high altitude rocky outcrops.
Campo sujo:
(“dirty field”) An open shrub savanna with very scattered but definitely visible shrubs and tree-like
shrub elements.
Cerradao (“tall:
cerrado”) A woodland with more than 40% woody plant cover and 3000-4000 trees per hectare.
Cerrado: A landscape of savanna-covered plateaus and tablelands separated by broad river valleys
with gallery forests.
Chaco: Thorn scrub, over 900,000 square kilometres, extends from the northern Argentina to
southern Bolivia and across to Paraguay.
Ecosystem: Organisms and physical factors in an environment, as well as the resulting processes of
the interactions among those components.
Gallery Forests: Vegetation bordering streams and rivers, comparable to riparian habitat in North
Miombo: A woodland savanna characterized by the predominant presence of Miombo
(Brachystegia sp.) species, with a range of climates from humid to semi-arid, and
tropical to subtropical or even temperate.
Mopane: A woodland savanna found almost exclusively within the low-lying, hot and arid river
valleys of the major river systems in tropical southern Africa where the mopane
(Colloquius burnham) is the dominant tree species.
Neotropical: Here, we define Neotropical as the biogeographic region of Central and South America.
migration Migrations undertaken by Eurasian birds to the southern hemisphere during the winter.
Extraordinarily long migrations.
Phytochorion :
A phytogeographic area with a relatively uniform composition of plant species.
Threatened :
species Organisms endangered for human activities and that can be in face of extinction.
Tropical region: A set of terrestrial and aquatic natural ecosystems distributed among the Tropic of
Cancer and the Tropic of Capricorn.
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Biographical Sketches
Alexandre Gabriel Franchin is undertaking his PhD degree in Ecology and Conservation of Natural Resources
from the Federal University of Uberlândia. He obtained his Master Degree in Ecology and Conservation of
Natural Resources from Uberlândia Federal University. His studies have focused on Urban Ecology, Bird
Ecology, Behavioral Ecology and Ornithology.
Rafael de Freitas Juliano is Professor of Zoology at the Department of Biology of the Goias State University,
Morrinhos Campus, Brazil. He obtained his Master Degree in Ecology from Goias Federal University and is
undertaking his PhD degree in Ecology and Conservation of Natural Resources from the Federal University of
Uberlândia. His studies have focused on Vertebrate Ecology with emphasis on Animal Communication. His
current research interests focus on natural selection on avian communication systems.
Mieko Ferreira Kanegae is undertaking her PhD degree in Ecology from Sao Paulo University. During her
post-graduate work, she has studied ectoparasites in bird communities in the Cerrado. Currently, her research
focus is on habitat selection, home ranges, and population sizes of endemic species. Working with CEMAVE
(National Center for Research and Conservation of Wild Birds), she implemented the bird monitoring program in
the National Park of Brasília, and the study of the effects of fire in Cerrado areas on bird communities.
Oswaldo Marçal Junior is Professor of Animal Ecology at the Institute of Biology of the Federal University of
Uberlândia. He has a Master Degree and PhD in Ecology. Actually he is coordinating the Postgraduate Program
in Ecology and Conservation of Natural Resources and the Laboratory of Ornithology. His studies have focused
on several themes in Ecology of Ecosystems and Applied Ecology, with special interest in the following areas:
Bird Ecology, Ecology of Parasites and Human Ecology.
To cite this chapter
Alexandre Gabriel Franchin,Rafael de Freitas Juliano,Mieko Ferreira Kanegae,Oswaldo Marcal Junior
,(2008),BIRDS IN THE TROPICAL SAVANNAS, in International Commission on Tropical Biology and
Natural Resources, [Eds. Kleber Del Claro,Paulo S. Oliveira,Victor Rico-Gray,Ana Angelica Almeida
Barbosa,Arturo Bonet,Fabio Rubio Scarano,Francisco Jose Morales Garzon,Gloria Carrion Villarnovo,Lisias
Coelho,Marcus Vinicius Sampaio,Mauricio Quesada,Molly R.Morris,Nelson Ramirez,Oswaldo Marcal
Junior,Regina Helena Ferraz Macedo,Robert J.Marquis,Rogerio Parentoni Martins,Silvio Carlos
Rodrigues,Ulrich Luttge], in Encyclopedia of Life Support Systems (EOLSS), Developed under the Auspices of
the UNESCO, Eolss Publishers,Oxford,UK, [] [Retrieved October 11, 2008]
©UNESCO-EOLSS Encyclopedia of Life Support Systems
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The Hoatzin (Opisthocomus hoazin) is the only extant member of the order Opistho-comiformes. This unique South American bird lives in the riparian lowland vegetation characteristic of the Amazon and Orinoco basins. Hoatzins nest in communal social units close to water bodies; they are strictly folivores being the only bird with pregastric fermentation in the crop. Because of the complex logistics involved in capturing this bird, there is a knowledge gap on its parasites. This study documents two distant lineages of haemosporidian parasites (Plasmodium spp.) in a juvenile and two adults sampled in the Cojedes state, Venezuela. Although negative by microscopy, the parasite identification was possible by using molecular methods. We estimated the phylogenetic relationships on the parasite cytochrome b (cytb, 480 bp) gene and the mitochondrial DNA. We found one of the parasites lineages in two individuals (nestling and adult), and the corresponding fragment of cytb was identical to a one found in Wood Stork (Mycteria americana) from Brazil. The other lineage, found in an adult, has an identity of 469 out of 478 bp (98%) with Plasmodium sp. GAL-2012 (isolate THAMB08) from Brazil. Although a morphological description of these parasites was not possible, this is the first molecular study focusing on Hoatzin haemosporidian parasites and the first documentation of Plasmodium infections in the Hoatzin from Venezuela. Furthermore, we reported microfilaria in two adults as well as hematological parameters for six individuals. Information on hematological parameters could contribute to establishing the necessary baseline to detect underlying conditions, such as infections, in this bird species.
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Many hunter-gatherer-cultivators, including Indigenous Australians, intentionally manipulated their environments via three broad processes — inclusions (plant and animal translocations), exclusions (quarantining of certain plants and animals) and transitions (in situ change of biotas, landforms and hydrologies, often through fire). These environmental changes resulted in deliberately and strategically `constructed landscapes' that not only supported societies but also established new contexts and conditions for cultural change. The structured and structuring nature of constructed landscapes is explored across the islands of Torres Strait of northeast Australia during the mid to late Holocene. Torres Strait Islanders deliberately imported and established the dog (dingo) while actively ensuring that cuscuses, cassowaries and pigs imported from New Guinea never produced viable populations on islands. Floral resources of islands were augmented by introducing a range of food crop staples and raw material plants through long-established contacts with peoples from New Guinea and more recent contacts with outsiders during the colonial era. Fire was a key tool of in situ vegetation change resulting in forest clearance and lowland sedimentation. These terrestrial activities supported, maintained and in some cases entrenched specialized marine lifeways in the region. In many respects, Torres Strait Islanders engineered the deterministic qualities of their own environment.
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Records of birds in and around Dudwa National Park were maintained from 1991 to 1994. The study involved intensive data collection on the bird community structure. Records from previous studies and surveys have been incorporated in this paper also. A total of 330 species were recorded from the park, of which 112 species breed in the area. The impact of the annual grassland burning and other management practices are discussed vis a vis conservation of some of the endangered bird species of the area.
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The savannas (lavrados) of Roraima are located in the far northern portion of the Brazilian Amazon and are part of the "Savannas of Guyana" ecoregion of the Amazonian Biome. They cover an area of approximately 43,358 km2, or 70% of the total area of savannas that straddle the borders of Brazil, Guyana and Venezuela. This ecoregion contains a diverse array of phytophysionomies forming a mosaic of non- forest (open areas) and forest ecosystems associated with different soil types, altitudinal gradients and climates. This diversification of ecosystems generates a great heterogeneity of habitats and is reflected in the diversity of plants and animals. The few existing studies of biological diversity are almost all concentrated along the main highways and therefore fail to indicate the true magnitude of the biodiversity of Roraima's savannas. In spite of Brazilian authorities having classified these ecosystems as having extremely high priority for conservation, no protected areas exist in the savannas of Roraima. However, 57.3% are protected in the form of Indigenous Lands. Large rural properties and settlements occupy 19.4% of the total area. The number of settlements in these savannas can be expected to increase significantly in the coming years due to legal impediments to creation of settlements in forest areas in Amazonia and due to the expansion of soy production, irrigated rice and commercial tree plantations in the state. It is estimated that 23.3% of the savanna area in Roraima is still available for creation of conservation areas.
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An extraordinary abundance of aquatic birds, a highly diversified avifauna and phenomena as the formation of ninhais (nesting colonies) are outstanding peculiarities that contributed to the recognition of the Pantanal wetland as one of the most attractive places in the world to watch or study birds. Its peculiarities, however, have been attracting researchers not only recently, but since the beginning of the nineteenth century, when Johann Natterer coordinated expressive collections of about 150 bird species in the northern portion of Pantanal (Pelzeln 1870, Paynter and Traylor 1991a, b, Vanzolini 1993). In the end of the same century, expeditions coordinated by Borelli (Salvadori 1895, 1900) and Smith (Allen 1891, 1892, 1893) at the surroundings of Corumbá collected a less extensive number of bird species. These and additional records of one or few species published by several authors were joined in a remarkable publication on bird species collected in the current States of Mato Grosso and Mato Grosso do Sul (Naumburg 1930). In the begining of the twentieth century, the most outstanding collections were those coordinated by Roosevelt-Rondon in the former State of Mato Grosso (Naumburg 1930), by J. A. G. Rehn at Descalvados (Stone and Roberts 1934), and by members of the Museu de Zoologia da Universidade de São Paulo at several localities of Pantanal (Pinto 1932, 1938, 1940, 1944, 1945). The mid twentieth century was characterised by greater participation of national institutions which considerably increased the number and distribution of inventories in the Pantanal through several expeditions involving bird collection (Moojen et al.. According to the literature, collections of birds in the wetland occurred until the early 1980s, when Dubs collected specimens in the Rio Negro region for the Zoologisches Museum der Universität Zürich (Dubs 1983a). On the other Bird species of the Pantanal wetland, Brazil
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We compared mitochondrial DNA sequences of portions of the cytochrome b (cyt b) and the NADH dehydrogenase subunit 2 (ND2) genes from samples of ten bird species that occur in both extremes of the Cerrado region (eastern Bolivia and Amapá, Brazil). The species include a wide sampling of taxa from several avian families: Tinamidae (1), Apodidae (1), Dendrocolaptidae (1), Furnariidae (1), Thamnophilidae (1), Tyrannidae (1), Turdidae (1), and Emberizidae (3). The taxa also exhibit a variety of distribution patterns in Neotropical open lands. Levels of genetic divergence within all taxa were low compared to comparable intraspecific values in many other widespread birds. In particular, these data suggest that there is much less genetic differentiation within these Cerrado birds than exists in birds of neighboring Amazonian forest. We suggest this implies the non-mutually exclusive possibilities that these open country birds have maintained higher levels of gene flow than forest understory birds, and that the Cerrado may have expanded to parts of its present-day distribution fairly rapidly. These data also suggest that hypothesized forest connections between Amazonia and Atlantic forest did not isolate open country bird populations from one another to a great extent.
There have been few studies of the fauna of the distinctive vegetation that grows on sandy soil in Amazonia. Leached and nutrient-poor sandy soil is associated with a vegetation type that varies in structure from open fields (campinas) to low canopy forests (campinaranas). During a bird inventory in sandy soil vegetation at Jaú National Park (JNP), I recorded 128 bird species, with 55 in campina and 94 in campinarana. Estimates suggested only 150 bird species should be expected to occur in these habitats, a reduced species diversity compared with other vegetation types in the Amazon region. This low species diversity is probably linked to the low productivity of this habitat and its relatively simple vegetation structure. Despite the relatively low diversity, at least 14 bird species (3% of the entire avifauna) appear to be restricted to white sand vegetation in JNP. In Amazonia as a whole, some 37 bird species are associated with vegetation in sandy soils. Biological inventories of other taxa are needed to understand the contribution of white sandy vegetation to the faunal distribution in Amazonia.