ࡱ> npijklm@ N bjbj "uuN"jjjj\|"tN N N N N N N N $Rl>N N >>N N VVV>N N V>VV)N B  JtGjO l0yXDUD""D$N ` VH+T 5J N N N ""$F$U*""F4Terrestrial habitat types 4.6. Savanna General description and geographic variation The savanna is a structurally simple but spatially patchy ecosystem in tropical and subtropical regions, characterized by a layer of herbaceous plants-mainly C 4 grasses and sedges, and C3 forbs- with varying degrees of shrubs and/or trees. Based on a broad definition of the term, savanna includes associated grasslands and woodlands. The high root/shoot ratio due to a predominant herbaceous layer is a feature that provides savanna ecosystems resistance to stress and disturbance from drought, fire, and herbivory. It is believed that savannas have evolved under disturbance factors like fire, herbivory and drought. Persistence of savannas may depend on such disturbance to preserve stabilizing components and properties (Baruch et al., 1996; Coupland, 1992; Silva, 1996). Savannas occur in hot climates with rainfall varying between 750 to 2000 mm and a dry period of two to six months, forming a transition zone between moist forest and xerophytic vegetation. Rainfall distribution is a major determinant of the savanna vegetation types. Tropical savanna usually develops on nutrient deficient, acidic soils with aluminum toxicity and pronounced alteration of wet and dry conditions (Sarmiento, 1983). In Mexico savannas are best developed in the southeast, in Campeche, Tabasco, Chiapas, and Veracruz. They also occur, much reduced in size, on the Pacific coast from Chiapas to Sinaloa. In Mesoamerica, patches of pine savanna occur in SE Chiapas, Izabl and central Petn of Guatemala, and along the coasts of Belize, eastern Honduras, and eastern Nicaragua. The vegetation is characterized by open stands of Pinus caribaea, Curatella ameircana, and Byrsonima crassifolia in a matrix of grasses, sedges and forbs (mainly legumes and composites) (Greller, 2000). In the Caribbean, sizable savannas are found in Cuba. Seasonal and hyperseasonal savannas occupy between 10 and 20% of Cuba. They occur in western Cuba and the Isla de Pinos, and the highlands (at 400 to 600 m) of the Sierra de Nipe in eastern Cuba. Some of them are floristically similar to continental savannas, but many have a high degree of endemism due to different soil types, either silicious or derived from serpentine (Sarmiento, 1983). In South America, savannas cover an area of approximately 2.5 million km2 [see sum of numbers given in the table below]. They include The Cerrados mostly in the uplands (800 ~ 1,500 m) of central Brazil, extending towards SE and NE in areas with lower elevations. Outliers are found in Paraguay and Bolivia. The Llanos del Orinoco in the central lowlands of Venezuela and northeastern Colombia. The Gran Sabana of southeastern Venezuelan Guayana, the Roraima-Rupununi savannas of Brazil and Guyana), as well as the coastal savannas of the three Guayanas (also called Guianan savannas). The Pantanal region that includes extensive areas of wetlands and savanna in Brazil, Bolivia, and Paraguay. The Llanos de Moxos of northern Bolivia drained by the Beni, Mamor, and Guapor to form the Ro Madeira (covers an area between 11~160S, and 64~690W). Others: Amazonian campos in northern Brazil, Llanos of the Magdalena valley, Colombia (Solbrig, 1996; Daly and Mitchell, 2000). Salient features of several South America savannas reported in Daly (2000), Solbrig (1996), and Sarmiento (1983) are summarized in the following table. SavannaCerrado sensu latoLlanos of Venezuela and ColombiaLlanos de MoxosGran Sabana / Roraima-Rupununi savannasPantanalArea (km2)1,800,000500,000Savannas ca. 150,000 and forests ca. 120,000 km2 54,000 150,000 170,000Elevation range100 ~ 1,500 m0-300 m130-235 m100~1,300 m~100 mPhysiognomic units or savanna typesCampo limpo, campo sujo, campo cerrado, cerrado sensu stricto , cerrado (sclerophyllous woodland).Hyperseasonal savanna, wetlands (esteros) and seasonal savanna (dunes)Hyperseasonal savanna, wetlands (esteros) and seasonal savannaHerbaceous swamp savannas on the tepuys; seasonal savannas and swamps on white sand, lower tablelands; alluvial fans,hyperseasonal savannas on river flat. Physionomically similar to Llanos, a complex mosaic of wetlands, savannas, and forests. It includes cerrado vegetation with an entire continuum from campo limpo to cerrado. Average annual precipitation1,500 mm (rainfall ranges from 750 mm to 2,000 mm, and increases toward NW and SE).1,000 mm (eastern Llanos) ~ 2,200 mm (SW at the Guaviare River in Colombia). Rainfall is highly seasonal.1,300 mm (E) ~ >2,000 mmLow rainfall (1000-2500 mm; northern end of the roughly NW-SE transverse central eastern Amazonia dry belt).1,000-1,400 mm, mostly during the rainy season (November- April).Dry season3-4 to 7 months (May ~ September). The wet period is likely to be interrupted by a short period of drought that may last for 1 to 3 weeks.1-6 months along a SW-NE gradient.2-3 months dry season (June ~ August) in summer. A short to medium dry season (2-6 months, December/January to April/May)3-4 monthsTemperature Frost occurs at southern limit of Cerrado.No frost260C in summer, but can reach 6 0C in winter with cold fronts.No frostSoil typeMost dystrophic oxisols, very poor in N, P, Ca, high in aluminum, low pH and cation exchange capacity. On richer mesotrophic soils, dry forest or cerrado may occur.Mostly alluvial and aeolian sediments, highly leached oxisols and ultisols, low in exchangeable bases and high in aluminum. On richer clays and shales, dry forest occurs. Alluvial sediments, different origin and age, ranging from dystrophic oxisols in the north to less alterated (verwittert) and more nutritive cambi-luvisols, and gleysol, solonetz and hydromorfic soils in the south.Mostly well-drained, but highly oligotrophic soils; many swampy areas with sandy or organic soils and an impermeable hardpan below.Most alluvial and aeolian sediments.Characteristic taxaDominated by woody plants: Dystrophic cerrado: Hirtella glandulosa, Emmotum nitens, Aspidosperma macrocarpon, Vochysia haenkiana, Xylopia sericea Mesotrophic cerrado: Magonia pubescens, Callistheme fasciculata Cerrado sensu stricto, campo cerrado, campo sujo; many species of Fabaceae, Poaceae, Asteraceae, Orchidaceae, Rubiaceae, Myrtaceae, Melastomataceae, Apocynaceae, and Leguminosae. Widespred trees and shrubs are: Qualea grandiflora, Kielmeyera coriacea, Copaifera langsdorffii, Caryocar brasiliense, Byrsonima coccolobifolia, B. crassifolia, Curatella americana. Palms: Butia leiospatha, Syagrus acaulus, Astrocaryum campestre. Grasses: Andropogon, Aristida, Axonopus, Elionurus, Paspalum, Echinolaena and Trachypogon. Dominated by herbaceous plants, especially grasses: Trachypogon, Andropogon, Aristida, Axonopus, Panicum , Paspalum, Mesosetum, Elionurus, Sporobolus, and sedges: Bulbostylis, Rhynchospora, Scleria. Palms: Copernicia tectorum, Mauritia flexuosa. Fauna: waterfowl, ibises, storks, capybaras.Dominated by herbaceous plants and shrubs in southern wetlands: Cyperus giganteus, Thalia geniculata, Paspalum,Panicum, Axonopus and palmares of Copernicia alba. In the northern Llanos cerrado like vegetation with grasslands of Leptocoryphium lanatum, Trachypogon plumosa, Macairea scabra, Mesosetum penicillatum, Bulbostylis juncoides and trees and shrubs as Qualea, Byrsonima, Palicourea rigida, Kielmeyera. In depressiones Palms: Mauritia flexuosa, Mauritiella aculeata and grasslands with Burmanniaceae, Eriocaulaceae, Xyridaceae Termites savanna are commun.In grass savannas: Trachypogon, Axonopus, Echinolaena, Aristida, Panicum, Paspalum, Mesosetum, and sedges: Bulbostylis, Rhynchospora, Scleria, Lagenocarpus, Hypolytrum. Palm swamps: Mauritia flexuosa, Andropogon, Scleria, Xyris, Eriocaulaceae, Rapatea, Drosera, Cephalostemon, Abolboda.In hyperseasonal savanna: Woody plants: Curatella americana, Bactris glaucescens, Licania parvifolia, Copernicia alba. Herbaceous plants: Poaceae (Axonopus purpusii, Paspalum almum, Paratheria prostrata, Reimarochloa brasiliensis, Elionurus muticus), Cyperaceae, Asteraceae, etc. On murundus (forest islands on decayed termite mounds): Tabebuia aurea. Gallery forest: Guarea macrophylla, Abuta grandifolia, Mouriri guianensis, Tapirira guianensis, Hymenaea stilbocarpa, Protium heptaphyllum, Xylopia emarginata, Mauritia flexuosa. Dry deciduos or semideciduos forests on limestone outcrops in Central Brazil: Cavalinesia arborea, Tabebuia impetiginosa, Myracruodruon urundeuva. Savannas that were discussed in the Experts Workshop: Sabanas Centroamericanas Llanos de Colombia y Venezuela Sabana Escudo Guayano Llanos de Moxos Cerrado Community types/zonation and major gradients within the system (patterns) Sarmiento (1983) recognizes three types of Neotropical savannas: Semiseasonal savanna which occurs in a rather humid climate with one or two short dry seasons and less frequent fires (some of the Amazonian and Guianan savannas), Seasonal savanna characterized by a severe dry season and frequent fires (e.g., the Cerrado and the Llanos), Hyperseasonal savanna which is the result of excessive drought and fires during dry season and severe flooding during the wet season. This savanna type is common in the poorly drained bottomlands of the Pantanal, Llanos de Moxos, the Roraima-Rupununi savannas, and part of Llanos. Palm stands are often found in water-logged areas. The savanna system is usually heterogeneous, consisting of a mosaic of pure grasslands, patches of trees or shrubs, dry deciduous or semideciduous forests, gallery forests, and sometimes wetlands (Daly and Mitchell, 2000). The distribution of various plant communities in a savanna landscape often follows edaphic gradients (e.g. soil types or levels of water table) and fire regime. Community types within the eight savanna systems that the experts identified. Sabanas Centroamericanas: 1) hmeda o seca, 2) variacin altitudinal (0-1,500 m), 3) variacin latitudinal Sabanas Antillanas (en Cuba y Repblica Dominicana; palmares en general se encuentran en lugares hmedos, estacionalmente inundados). Llanos de Colombia y Venezuela: 1) Inundable (e.g., Lllanos de Apure-Arauca: palmares, bancos, bajos y esteros), 2) no-inundable (e.g. Llano Alto: sabanas arbustivas con Curatella americana, Byrsonima crassifolia, Bowdichia virgilioides; y sabanas abiertas o lisas: con Trachypogon spicatus, Axonopus anceps, Bulbostylis paradoxa, etc.). Sabanas del Escudo Guayano (con alta frecuencia de termiteros). Sabanas Amaznicas: 1) suelo mal drenado, 2) suelo bien drenado (sabana no-inundable). Llanos de Moxos: 1) Sabana del nordeste de Beni, agua estancada por lluvia, parecido a la flora de cerrado (sensu lato). Tambin incluir bosques abiertos, variacin de pastizal hasta bosque abierto con diferente regimes hdricos. Sobre substrato antiguo, muchos termiteros. 2)Sabana del central y sur de Beni: sabana inundable por ro y lluvia, comunidad vegetal se cambia con nivel topogrfico: i) bajura, ii) semi-altura, iii) altura de algunos metros, con suelo aluviales, algunos semi-alturas con pH hasta 9. Cerrado: 1) herbceo: i) bien drenado: campo limpio, campo sujo, campo cerrado, ii)mal drenada: campo limpo hmedo, vereda, campo de murudum. 2) leosa: i) cerrado sensu stricto: latosol, sandstone, ii) cerradao: mesotrfico, distrfico. Sabanas Andinas: incluye Valle de Magdalena y Valle Sinu. Hay manchas pequeas de Trachypogon plumosus, entre bosque seco y hmedo, zona de colonizacin incaica. Ecological integrity factors for landscape context Table xxx. Ecological integrity factors for landscape context of savanna Key FactorJustification for SelectionNatural Range of Variation and Ecological ThresholdsJustificationIndicatorsFactor RankFire regimeImportant in shaping savanna properties at a local scale. Changes the mix of habitat types by increasing or decreasing the dominance of one type over the other. Modifies community or floristic structure by affecting mortality of many species, especially young plants. Recurrent frequent fires cause nutrient impoverishment.Fire frequency (Frost and Robertson, 1987; Braithwaite, 1996): In the moist savannas fires generally occur at intervals of 1-5 years, whereas in the more arid savannas, the intervals are about 5-50 years. Fires increase patchiness or habitat diversity within the savanna. By optimizing the fire mosaic, savannas can be managed to increase biodiversity. Long-term fire exclusion experiments in Calabozo, Venezuela show (Bulla, 1996): New tree species invade the savanna and increase the diversity of the tree layer. Tree density increases. In the open savanna, evergreen tree species increase rapidly, whereas deciduous trees showed a long lag of at least 16 years. In the groves, evergreen species change little and deciduous species increase in the first 8 years. Fire intensity (Frost and Robertson, 1987; Braithwaite, 1996): Factors that affect fire intensity are fuel types, fuel loads, the occurrence of periods of above-average rainfall, seasons, landscape units and vegetation types. Long intervals between fires increase the fuel load and therefore the probability of a wild fire event. The fire intensity will cause changes in patchiness of habitats and the vertical structure of upper shrub or tree layer and ground herb or shrub layer. Dominance shifts towards to upper shrub or tree layer if fire intensity is low. Conversely, dominance shifts to ground herb or shrub layer if fire intensity is high. Type of fires (Frost and Robertson, 1987): most savanna fires are surface fires, burning through the herbaceous layer, and burn patchily as a result of varying wind speed, topography and distribution of fuel loads. Time of natural fires (Frost and Robertson, 1987; Braithwaite, 1996; Ramos-Neto and Pivello, 2000): Natural fires occur mostly at the beginning of the wet season when the frequency of lightning is highest. Early dry season fires: create greatest patchiness in the ground herb or shrub layer and impact negligibly the upper shrub or tree layer. High intensity late dry season fires typically scorch high into trees and are likely to cause maximum mortality. Impact on fauna (Braithwaite, 1996): Burned sites show increase in ants species diversity due to structural changes in the habitat caused by fire, especially the level of litter accumulation and insulation on the ground. Many bird species that feed on the ground such as granivores, omnivores and carnivores are attracted to areas that have been recently burned. Species of Lepidopterans, herpetofauna, and mammals, respond differently to different fire regimes.Corticolous lichens have been used as potential bioindicators of fire history in the cerrado of central Brazil (Mistry, 1998).Humidity regime This factor should certainly rank in the first place of importance in this Category: Landscape context, before the Fire.Important in shaping savanna structure at a landscape scale. Modifies plant available moisture (PAM). It is related to the length of dry season, precipitation and woody cover (Solbrig, 1996).Pulses in rainfall or drought spells induce important changes in savanna composition. Observations support the following hypotheses. Early onset of rains favors species blooming very early in the wet season (precocious species). A prolonged rainy season favors species blooming at the end of the wet season (late species). A shortening of the rainy season would be detrimental to late species but favorable to species blooming in early or the middle of the wet season (intermediate species) Drought favors deciduous trees or trees with smaller leaf size. A higher rainfall pulse will increase grass biomass during that season, increase standing dead biomass during the next dry season, and increase the probability of fires. [If no fire, shading will increase during the following wet season and reduce grass growth, but increase tree growth and recruitment, alter the tree/grass ratio in the following years, affect grass growth and reproduction, and influence fire and grazing regimes.]Soil type or fertilityImportant in shaping savanna structure at a landscape scale. Modifies plant available nutrients and moisture. Affects savanna productivity.Savanna soils vary widely in particle size, structure, profile, and depth. The nutrient status of the soil is related to the age of the sediments, parent material, and topography. Recent studies show that estimates of productivity of tropical savanna grasslands approximate the figures for tropical forests (Solbrig, 1996).Interactions between plant available moisture (PAM) and nutrients (PAN).Important in shaping savanna structure at a landscape scale.Woody elements dominate where plant available moisture and nutrients have high values. As moisture and/or nutrient levels increase, the savanna gives way to a moist forest eventually. Xerophytic elements prevail where moisture and/or nutrient have low value. If the values of moisture and nutrient get very low, the savanna is replaced by a semidesert (Solbrig, et al., 1996). Interactions between plants and humidity (rainfall, drought) and herbivory during post fire recovery phase (Frost and Robertson, 1987).Cattle grazing (Baruch, 1996): [Cattle were introduced to the Americas in the 16th century.]Important in shaping savanna properties at a local scale. Modifies community or floristic structure.Woody species / grass species ratio or density of trees affects the sensitivity of grass layer to grazing and overgrazing. Grazing pressure by domestic animals plays a role in the dynamics of savanna communities dominated by native and alien grasses, and may favor alien grasses that have higher defoliation tolerance, e.g. African grasses.Connectivity by gallery forest networksImportant corridors linking patches of similar or different ecosystem types, e.g., gallery forest networks of cerrados linking the Amazonian forests to the Atlantic coastal forests of Brazil (Oliveira-Filho and Ratter, 1995; Daly and Mitchell, 2000). Ecological integrity factors for condition Table xxx. Ecological integrity factors for condition of savanna Key FactorJustification for SelectionNatural Range of Variation and Ecological ThresholdsJustificationIndicatorsFactor RankDiversity of above-ground plant functional groups (species that share morphological, chemical, structural or life history characteristics)(Denslow, 1996) Determines the role of biodiversity in ecosystem response to disturbance such as water stress (drought or flood), fire, shading, competition, and herbivory or grazing. Woody species: density and spatial patterns of trees determine the spread of fire and its erratic performance. Herbaceous species: Rapid and homogeneous fires are produced in communities that have a high biomass of grasses and a low fuel biodiversity. The architecture of dominant grasses defines the vertical distribution of fuel. Taller grasses produce higher flame which increases the probability of damage to tree crowns. C4 and C3 plants in savannas result in patchiness of herbaceous layer in terms of productivity and efficiency of water and nutrient use. In seasonal savannas most dominant grasses have the C4 syndrome, and only in very wet environments do C3 grasses become abundant (Baruch et al., 1996) Disturbance-resistant vs. disturbance-sensitive species: Fire-resistant woody species: usually evergreen, sclerophyllous (hard, thick, leathery leaves), and with thick bark or protected buds, or escaping adaptation by underground woody structure such as xylopodia or lignotubers (e.g., Andira humilis and Anacardium humile). They are usually reproductively active during the dry season; and adapt to water or nutrient stress. Fire-sensitive woody species: usually deciduous and mesophyllous. Drought -escaping ephemeral annuals or decicuous perennials: active only in the wet season. Survive the period of drought stress as seed or by going dormant. Drought-enduring evergreens or perennial graminoids and deep-rooted phreatophytes: continue being physiologically active during the period of drought as long as there is sufficient soil moisture. Phreatophytes, represented by the majority of savanna trees and shrubs, can gain access to the water table by their deep roots. Drought-resistant succulents: special morphological and physiological features to maintain physiological activity, even under conditions of drought stress. Species reproduced by seeds vs. vegetative propagules: populations of plants that recover by seeds after fire will be more susceptible to change than those that recover by resprouting. Abundance of dominant species in herbaceous layer and/or woody layer. In Cerrado sensu stricto: 40-100 woody species and three times more herbaceous. One hectareof cerrado contains from 60 to 100 woody species and two times more herbaceous species. One hactare of gallery forest contains from 80 to 200 woody species and a similar number of herbaceous species. One hactare of dry deciduous forest contains 40 to 80 woody species and a similar number of herbaceous species. One hactare of campo limpo contains from 200 to 300 herbaceous species. Dominance of species from the genera Sclerolobium, Mimosa, Solanum and Ouratea can serve as disturbance indicator.Community composition/ Diversity High beta diversity is usually found in areas with mosaic habitats that are created by difference in topography (e.g., valley or ridge crests), or soil types or disturbance of various intensity e.g. fire or flood. The patchy diversity of the community affects fire impacts on species composition (Bilbao et al., 1996).Plant species composition and structureDetermine the fuel mixture, fuel quality and degree of flammability during the event of fire. Affect the types and intensity of fires. Determine the productivity of the savanna. High biodiversity increases the probability of redundancy within functional groups. This may buffer the negative impacts of perturbations to ecosystem processes and products (Denslow, 1996).The available plot data suggests that the range of species diversity for tropical savanna is 7-100 species of trees and shrubs/ha with lower diversity in the Amazonian savannas, e.g. Amap, and higher diversity in central Brazilian cerrado and Roraima-Rupununi savanna (Daly and Mitchell, 2000). Cerrados are the most species rich savannas of the world. About 6.500 species of higher plants have been identified in the Cerrado Dominium (Vania Pivello, Pers. Com.). Plant species composition in savanna often relates to available moisture and nutrients, herbivory, and fire disturbance (Bilbao et al., 1996). Low productivity, stress-driven savanna communities and high productivity, competition dominated ones will have lower species diversity than intermediate ones (Bulla, 1996). Herbaceous plants that are commonly found in the Neotropical savannas are usually members of grass and sedge families. In the Poaceae, Andropogoneae: Trachypogon, Andropogon, Hyparrhenia and Saccharum are usually dominant in the core savanna area; Panicoid: Digitaria, Panicum, Paspalum, Pennisetum, Setaria, dominate in drier environments, and Oryzeae (Oryza, Luziola, Zizania) in very wet and swamp savannas. Cyperaceae includes genera of Cyperus, Rhynchospora, Bulbostylis, and Scleria (Solbrig, 1996). Woody plants widespread throughout the Neotropical savannas are Curatella americana, Byrsonima crassifolia, B. coccolobifolia, B. verbascifolia, Bowdichia virgilioides, Anacardium occidentale, Hancornia speciosa, Salvertia convallariodora (Daly and Mitchell, 2000).Fauna diversityAffect food webs of the savanna system. Neotropical savanna avifauna (Fry, 1983) includes the two most species abundant insectivorous Tyrannidae (tyrant-flycatchers), granivorous Fringillidae (finches), and aboreal frugivorous Aquilidae (parrots), Trochilidae (hummingbirds), Furnariidae (spinetails), Icteridae (caciques) and Thraupidae (tanagers). Endemic bird families: characteristic large ground birds of the open savanna: herbivorous Rheidae (rheas), Anhimidae (screamers), Cariamidae (seriema), and ground-foraging granivorous Tinamidae (tinamous). Reptiles (Barbault, 1983): snakes of Colubridae, Typhlopidae, Boidae, Elapidae and Viperidae, lizards of Gekkonidae, Scincidae and Iguanidae. Rodents and lagomorphs (Happold, 1983): Cricetidae (Calomys, Zygodontomys, Phyllotis, Akodon, Baiomys, and Eligmodontia), Caviomorph rodent (capybara, Hydrochoerus hydrochaeris), and endemic lagomorph, Sylvilagus brasiliensis.Biotic interactions: presence of pollinators (bees, butterflies, moths, bats, and hummingbirds), and seed dispersal agents such as fruit-eating birds (e.g. parrots).Pollinators affect food webs.Wind is important seed dispersal agent: almost 50% of cerrado tree species are dispersed by wind. Birds seem to be the most important pollinators and dispersal agents besides wind. Seedling recruitment for grass species mostly depends on yearly seed production. There is no apparent permanent seed bank of native grasses in the savanna soil.Biotic interactions: presence of large herbivores (e.g., branch-antlered white tailed deer Odocoileus virginianus, pampas deer Ozotoceros bezoarticus, swamp deer Blastocerus dichotomus, and capybara Hydrochoerus hydrochaeris. (Ojasti, 1983). Insects' contribution to consumer biomass is much greater than that of vertebrates in the savannas (Lewinsohn and Price, 1996).Herbivory affects woody/herb plant ratio, vegetation stratification, total cover and biomass (Lewinsohn and Price, 1996).Few large herbivores in Neotropical savanna. Folivores can accelerate nutrient release from living plants. Folivorous insects are much more important than vertebrate herbivores, and their species diversity may be dependent on plant species richness due to host-specificity. Meristem, stem or root feeders can change plant architecture and their susceptibility to fire, drought, and frost damage. Propogule feeders can increase patchiness: In cerrados, seed crops are often almost entirely destroyed by predispersal predation. Peak of insect herbivory: the sunnier or drier part of the year.Biotic interactions: presence of anteaters (Myrmecophaga tridactyla) or armadilos (Dasypus spp., Priodontes maximus) (Rodrguez and Rojas-Surez, 1999) or top predators e.g. jaguar (Felis onca), Orinoco crocodile (Crocodylus intermedius).Savanna is an important habitat for anteaters and armadilos. Top predators control the populations of small mammals and herbivores..etc.Biotic interactions: Presence of decomposers (mycorrhizae, fungi, microbes) and soil macroinvertebrates (termites, nonsocial arthropods, earthearths, leaf-cutter ants)Can increase nutrient supply rates and improve soil structure, including nutrients, moisture and oxygen availability (Denslow, 1996).Biotic interactions: presence of alien vertebrate species: cattle (see grazing)Introduce diseases to native fauna, disseminate exotic herbaceous species, and change species composition of native flora. Its impact can vary from low biomass consumption of natural savanna plant communities during rainy season, to massive plant biomass harvesting of regrowth after burning during dry season, to total substitution of natural savanna plant communities by introduced, artificial fodder grasses. Biotic interactions: presence of alien plant species: e.g. the African grasses that are the most aggressive invaders of Venezuela (Baruch, 1996) and Brazilian savannas, Melinis minutiflora in secondary savannas above 600 m; Hyparrhenia rufa in lowland savannas with poor soils and marked dry season; Panicum maximum in humid and relatively fertile areas; and Brachiaria mutica in periodically flooded savannas. The indigenous community is generally dominated by native grasses Trachypogon, Axonopus, and Bulbostylis.Grass invaders can change composition and structure of the grassland (e.g., shortgrass to tallgrass), and modify factors that control the functioning of savannas such as water and nutrient availability, and fire regime. Often decrease biotic diversityIn Venezuela savannas, the invasive plants usually have higher growth rates, establish themselves rapidly and compete successfully for resources, and displace native species from moist, fertile sites. In contrast, native grasses have higher root/shoot ratios with carbon reserves in underground organs and slower growth rates. They can endure invasion and persist in poorer sites. Studies of Venezuela savannas show that the persistence of some African grasses [depending on the grass species] is sustained by a fire and invasion cycle. Space created after burning is invaded by alien grasses, which in turn promote more frequent and intense fires due to higher biomass, and reinforce the process of colonization and elimination of native competitors. To arrive at thresholds, determine whether the success of invasive species is due to differences in competitive abilities or due to the primary disturbance that removes native species. Invaders may not persist if the disturbance is eliminated. Ecological integrity factors for size Table xxx. Ecological integrity factors for size of savanna Key FactorsJustification for Factor SelectionEcological Thresholds: Min. Dynamic Area Desired Future Condition (Increase in MDA to Rate Good or Very Good) Justifications or Recommendations for Calculating Minimum Dynamic Area (MDA) and Desired Sizes above MDAIndicators for Field-Based MonitoringFactor PriorityEXAMPLE: Minimum Dynamic AreaConsider the average size of the primary disturbance (fire) and the average return intervals (see above). Buffer this by a multiplier depending on the confidence in your data and the variability of the key factor attributes (size, return interval, etc) around the geometric mean.Minimum Population Size of anteaters or armadillos or jaguarsTop predators in this system are critical to control populations of ants or termites, small mammals and herbivores.   Additional information The experts assessed the biodiversity health of seven savanna systems. The results are presented in the following tables. System: Pine Savannas, Central America, Landscape Context, Condition & Size Key FactorsJustification for Factor SelectionEcological Thresholds: (Minimum Integrity Threshold) Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor PriorityFire RegimeSystems dynamic is associated to the fire cycle. The pine savanna requires fire; they are maintained by fire; the species have adaptations to endure the fire; many require the fire to regenerate. Without the appropriate fire regime habitats and species are lost. Fire frequency: 5-20 years for Pinus oocarpa and P. caribaea of Central America. Each 3-10 years for Pinus caribaea in the Bahamas and Cuba, P. tropicalis in Cuba, and P. elliottii in the Keys of Florida. Intensity and duration of the fire and period of occurrence. Size of the natural regeneration. For P. oocarpa and P. caribaea approximately a meter of total height (age: 5-8 years). <5 years: open pine forests in Central America. Pinus caribaea (in Cuba and Bahamas), P. elliottii and P. tropicalis support fires of greater frequency (with P. tropicalis almost annually). Fires of high intensity and duration during times of drought destroy the natural regeneration , the organic material, and define the growth of the pine trees. Regeneration disappears with fires of high intensity and duration.Monitoring through permanent plots. Fire history (place, extension, intensity, and time of the year). Fire mapping. Sources of ignition; areas of high riskHydro regime seasonLimitation of water during certain part of the year.Pinus caribaea: 3-9 months of dry season.Extreme periods of drought weaken the pine trees and make them susceptible to plague and/or illnesses attacks. Records of the periodicity and intensity of the drought. Drainage, depth and fertility of the soil.Density of pine stands and floristic composition of the undergrowth.Age structure Determinant to guarantee the dynamics of the system. The age structure allows maintaining populations of birds that guarantee biological control against endemic plagues. Existence of a diametric structure for all the pine forests in the form of inverted J (ordered: number of trees by hectare and abscisa: diametric classes).Without total diametric structure the system degenerates or tends to disappear (under natural conditions) Through temporary samples to level of all the mass. Diversity of associated species Determinant to maintain systems stability and health. Unknown in Central America and the Caribbean. In Florida there is> 20 species by square meter, mainly herbaceous. In all the pine forest the vital function of microorganisms associated to its roots is recognized (mycorrhiza). Systems susceptibility to the attack of plagues and/or illnesses. Continuous inventories through permanent plots.  Exotic species Displacement of native species, especially to a level of undergrowth. Favors fire incidence. Deteriorate the habitat.Investigations required.Loss of biodiversity. Increment of fire risks.Continuous inventories through permanent plots. Irrational timber exploitationAlter pine structure . Damage biodiversity. Provoke genetic erosion. Induces appearance of exotic species. Alters hydrologic regimes to a soil level.The causes and possible effects are known but not the quantification. Degraded pine trees in genetic and economic terms. Deterioration of habitats. High rates of surface water run-off and reduction of infiltration into soil. Continuous inventories through permanent plots and run-off plots.Grassing Damage biodiversity. Induces appearance of exotic species. Alters hydrologic regimes to a soil level. Alters fire regimes.The causes and possible effects are known but not the quantification. Deterioration of habitats. High rates of surface water run-off and reduction of infiltration into soil. Continuous inventories through permanent plots and run-off plots.Connectivity Loss of speciesSize (minimum dynamic area)Maintains systems stability and health.Depends of the origin and diversity of habitats. Large enough to maintain fire regime and genetic base. For example, if the frequency of the fire is 5 years, should not burn more than one fifth of the pines each year.Need of buffers to facilitate fire management.  Savanna Colombia and Venezuela, Landscape Context Key FactorsJustification for Factor SelectionEcological Thresholds: (Minimum Integrity Threshold) Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor PriorityPrecipitationClimatic evidence Precipitation: 800-1,600 (2,500) mm / year. Dry season: 6 months max. <800 mm: spiny vegetation >1,600mm: semi deciduous forest to evergreen forestsRainfall measure Specific geohydric regime Soil hydric regime and limit soil nutrients reserves 1. Texture: sandy clayey, the gradient can adapt the type of savanna (rank: consult literature). [Experts, please provide literature citation. Thanks.] 2. Hydro edafico regime: Flooded- non-flooded Apure savanna, classified in river bank, low-lying ground and marsh; 6 months of flood min. Other types: with less or non flood (High plains, oriental plains, Mesa de Guanipa, etc.); variables in their woody component: smooth savanna to wooded savanna.In more fertile soils (clayey) semi-deciduous to deciduous forests tend to be installed . In soils less fertile would be a very open and degraded savanna (e.g., Mesa de Guanipa)Soil composition measure Fire regime Maintenance of mosaic of the present vegetation of savanna and forest. Frequency of fire: 5 years to maintain the present equilibrium (little natural fire). If the fire is eliminated, probably the woody plants increase. Monitoring plots of the relationship of woody / non-woody plants.The great beta and range diversity: distributed upon an area of aprox. 300,000 Km2, extensive variety of vegetable communities; great heterogeneity paisajstica. An important and representative biome of the flora and fauna of north South America. Minimum dynamic area: depends on each type of savanna included in this extensive region. For some types less extended, as for example the plain palm groves of Copernicia tectorum, the minimum area can be of 1-2 km2, while for types of savanna more extensive (for example, oriental savannas of Monagas) they can be required not less than 100-200 km2. In the case of plains Morichales, typically riparian, the minimum area is measured in km of river and should cover all the river extension with this type of vegetation (among 200-300 km). Empirical assumptions. Remote sensors. Level of natural state observed in the field.Floristic composition of Plains (see Aristeguieta, 1966; Ramia 1974) dominated by gramineas (Trachypogon spicatus, Axonopus spp., Panicum spp., Andropogon spp. ) in the herbaceous stratum and by Curatella American, Byrsonima crassifolia and Bowdichia virgilioides in the woody stratum. Few endemic species; the same are mostly concentrated in the gallery forests. Floristic typical group of this biome in Venezuela and Colombia. The herbaceous species conform the 60-70% of the savanna Venezuelan flora . The woody species and the epiphytes and parasitic plants conform the rest. 40-60 species are considered as representatives of most of the Venezuelan savanna types. Indicator species and faithful to a basic floristic nucleus of Neotropical savannas (Huber, 1987).Floristic inventories. Savannas and herbaceous of Escudo Guayanes (azonal system), Condition Key FactorsJustification for Factor SelectionEcological Thresholds: (Minimum Integrity Threshold)Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor PriorityLevels of nutrients in the extremely low soil (oligotrofismo edfico extreme). It can be observed in the following three typical and exclusive of the Shield Guayans types of herbaceous ecosystems : Savanna graminous totally open upon soil of very low fertility derived from sandstone (oxisol and ultisol), Herbaceous oligotrophic, upon peat soil (histosol) (relictual vegetation of subtepuy). Herbaceous oligotrophic upon inundated sandy soil (quartzipsamment) Responsible for the presence of this type of savanna in the low and medium soils of the north and east of Escudo Guayanes (Huber, 2000), including the Great Savanna in Venezuela, the state Roraima of Brazil, Savanna Rupununi in Guyana, Sipaliwini Savanna in Surinam (aprox. 60,000 km2). Exclusive and only type of relict subtepuy vegetation in the world; extension of just 1000 km2 in the Great Savanna of Venezuela. Vegetation type exclusive from low and medium soils of the southwest and northwest sector of the Shield Guayanes (aprox. 8,000 km2 in the center of the Amazonas state in Venezuela and Dept. Guainia in southeast of Colombia). (Huber, 1982; Huber, 1995.) Rank of the level of nutrients in the soil should be extremely low. Adequate hydrologic conditions should be assured in the soil that maintain the organic matter in permanent state of saturation. The marked levels of oligotrophisim edaphic should prevail; these levels of soil nutrientes are not measurable with methods of traditional analyses. Obviously the factor nutrients reserve is the principal limiting factor in these ecosystems 1 to 3. With inferior fertility rates, extremely degraded savannas are observed until desertification fenomenous (for example, in Gran Sabana sudeste); with superior fertility rates, savanna forests similar to the savannas in Venezuelan plains or Brazilian cerrados are observed. In case of a prolonged hydric deficits a dry out of the peat is observed, this become inflammable and is frequently destroyed by fire. Seems to be ecosystems adapted to extremely edaphics and environmental conditions in effect in its distribution area. In general: periodic measure of the levels of soil nutrients. Specifically: floristic and ecological inventories for the observation of less evident changes. Floristic composition Highest continuos savannas in Venezuela (e.g., Uper elevation limit of Mauritia flexuosa 990 msnm). High rate of regional endemism. Too high rate of regional endemismo.20-30 species/ gramineas community Herbal subtepuyano should have a minimum of 10-20 spp. of tepuyana flora. The herbal of low soil should have a good representation of endemic species, adapted in their herbaceous and woody behavior to the extreme habitat conditions. Any change in the floristic composition means biologic degradation of the ecosystem. (specially in 2 and 3).Periodic floristic inventories.Intense fire regime High fire frequency (8,000 fires per year in Gran Sabana). 2 y 3. Occasional fires are causing notable impacts.  Apparently, none of the three ecosystems types 1, 2, and 3 are particularly adapted to survive the ecological effects of fire, showing in some cases evident signs of degradation caused by this factor. Therefore, it is estimated.The reduction of the fire frequency seems to be the only way to reduce negative impacts that are observed in these peculiar herbaceous ecosystems.Monitoring of the number of fires.  Cerrado, Landscape Context Key FactorsJustification for Factor SelectionEcological Thresholds: (Minimum Integrity Threshold)Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor PriorityFire regime Controls plant species distribution and balance between herbaceous and woody layers. Controls community composition. Interferes with nutrient cycling. Stimulates flowering, fruiting, and germination, Important for herbivore food availability and distribution. Fire frequency: Herbaceous campo limpo, campo sujo, and campo cerrado: well drained: 2- 5 yrs. badly drained: no fire. Cerrado (sensu stricto): 4-6 yrs. Cerrado and associated gallery forest: no fire. (Coutinho, 1990; Pivello & Norton 1996.) < 2 yrs. : Decrease species diversity and nutrient availability. >5yrs. : Accumulate dead organic matters, slow down nutrient cycling, change structure, and decrease herb diversity. Fires decrease species diversity and change composition and structure. <5yrs. : Reduce woody density and fire sensitive species; decreases nutrient availability. >10yrs. : reduce herbaceous layer and fire prone species. Favors Melinis minutiflora, and increases risks and intensity of wild fires. Fires decrease species diversity and change composition and structure; eliminate fire sensitive tree species.For monitoring changes at large scale: satellite imagery; field studies include: Burned stems or tussocks. Palms age distribution, dead standing individuals; presence or absence of organic matter layer. Burned stems and tree trunks; litter accumulation; density of dead standing woody plants. A normal stand should not have >5% of dead standing woody individuals >5 cm basal diameter (See Felfili et al., 1994, 1997 & 2000). Presence or absence of dead Xylopia and Copaifera trees, >5% of dead standing trees.3Drainage and watertable depth Affects plant community composition. Important for maintenance of seasonal lagoons for birds, amphibians and reptiles. Important for maintenance of pasture for mammals and herbivores during dry season.Herbaceous campos, well drained: moist only during the peak of the rainy season and >1 m watertable depth for the rest of the year. Herbaceous campo, badly drained: flooded 9-10 months and moist during dry season. Cerrado and Cerrado : > 5m watertable depth. Gallery forest : Seasonally flooded, flooded during the rainy season and moist the rest of the year. Well drained: moist during the rainy season and well drained during the rest of the year.Influence structure, floristic and faunal composition.Floristic structure and composition.2Climate: seasonal rainfall and frost frequency. Determines presence or absence of cerrado.5-7 months of dry season (May-Sept.) with sporadic frost events occurring in > 3 yrs.Water deficit prevents the establishment of moist forest. Frequent frost prevents cerrado establishment.Rainfall and frost records.1Soil structure and fertility Determines plant community types.Hydromorphic soils or shallow soils with hardpan: herbaceous campos badly drained and associated gallery forest. Shallow or very poor soils: herbaceous campos, well drained. Deep soils: woodland. Mesotrophic: cerrado. Dystrophic: cerrado or cerrado. Rocky soils (without hardpan): Shallow: high altitude cerrado. Rock outcrops on slopes: woodland cerrado. (Roots penetrate cracked rocks.)1. Humid 8-10 months of the year. 2. < 1m depth. (see Goodland & Pollard, 1973; Lopes & Cox, 1977; Haridasan and Ranzani ?? year for nutrient levels.) [Experts, please complete the citation of Haridasan & Ranzani in the References section. Thanks.] 3. 5 m depth. (see Haridasan ??year for nutrient levels.) [Experts, please complete the citation in the References section. Thanks.] Presence or absence of bare rocks or gravels.Soil Texture and color.3Connectivity by gallery forest networks (gallery forests serve stepping- stones-like connections for cerrado fragments.)Maintenance of genetic interchange among isolated populations.Index of connectivity : see Metzger and Dcamps, 1997. Loss of top predators and rare species.Presence of top predators (e.g.,Jaguar). Cerrado, Condition Key FactorsJustification for Factor SelectionEcological Thresholds: (Minimum Integrity Threshold) Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor PriorityViable and naturally regulated frugivore bats and birds communities and maned wolf.Help seed dispersal.Identify the minimum set of species that disperse all plant species in the ecosystem and the minimum population size for each species. [Experts, if you know of any published studies, please cite. Thanks.]Absence of some animal species may cause diversity erosion in plant communities.Density and diversity of birds, bats, and density of maned wolf.6Pollinators Help plant reproduction.Same as aboveSame as aboveSame as above5Herbivory (ants, termites, and other insects, Rhea, deer..etc.) Improves nutrient cycling. Prevents woody plant encroachment and fuel build up.Same as aboveSame as aboveSame as above4Cattle grazing Introduces exotic grasses and changes community composition; increase fire frequency. May spread diseases to native fauna.Presence or absence of cattle: the approximate carrying capacity in managed areas : 1 head/ 4 ha (This figure may vary depending on the floristic composition and landscape mosaics of the savanna).Compact soil. Grazing increases soil erosion. Reduce population size of plant species sensitive to grazing. Presencia de carcavas, caminos de pisoteo de ganados.Exotic species Competition and displacement of native species. Reduces biodiveristy.Presence or absence of Melinis, Brachiaria, Andropogon, Hyparrhenia, etc. One or more of these species become the dominant herbaceous species in terms of their high Index of Importance Value among herbaceous species.Partial or total replacement of native herbaceous plants.Calculate Index of Importance Value (See: Pivello et al., 1998.)Minimum dynamic area Maintenance of viability of all biotic elements.> 130.000 ha (Parque Nacional Emas)Maintenance of the landscape structure, function at all trophic levels.Areas needed to maintain viable population size of top predators.1 Plains of Moxos - Savannas of north Beni, Landscape Context Key FactorsJustification for Factor SelectionEcological Thresholds: (Minimum Integrity Threshold)Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor PriorityPrecipitationAffects flooding dynamics because of the rainfall in the region and occurrences of termite nests in flooding area. Minimum rainy season: 5-7 months. Flooding period: 3-5 months.The decrease of precipitation and the flooding period change the extension and composition of communities to more xerophytic.Measurement of rainfall regime. Relief and soil (structure: e.g., hard pan, and fertility) Affects flooding dynamic and distribution of the vegetables communities. Depth of hard pan close to the surface: <1 m.High plains with emergence of lateritic crusts and pisolites with campo cerrado vegetation. In ravine dominates the broadleaf evergreen forest partially flooded above lateritic hard pan and fine material.Mosaic of different types of vegetation from the ravine forest to open savannas with cperaceas dominating.Fire regime Affects the mineralization of organic matter and the distribution of vegetable communities. Fire period: <20 years to maintain the system. Change the systems to dryer communities. Domination of pyrophytic species : in the hillock: Trachypogon plumoso (=T. spicata). in the depressions: Paspalum, Leptocoryphium lanatum Plains of Moxos-Savanna of north Beni, Condition Key FactorsJustification for Factor SelectionEcological Thresholds: (Minimum Integrity Threshold)Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor PriorityTermite and nests in plains areas bad drained or old riverbeds forming bogs.Maintain the flow of nutrients. Presence of earth and termite nests: 10-40%.Are there marshy or aquatic vegetation found? It is a supposition not proved!Rate of the termite or earth nests coverage. Plains of Moxos sensu estricto (Savannas of south Beni), Landscape Context Key FactorsJustification for Factor SelectionEcological Thresholds: (Minimum Integrity Threshold) Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor PriorityPrecipitationDetermine the distribution of species. Minimum rain period: 5-7 months. Flooding periods because of river overflowing and local rains: 3-5 months, up to 8 months, relief dependent. Decrease of the flooding period changes the extension and composition of the communities to more xerophytic.Measurement of the rainfall regime. Fire regime Determine the composition and distribution of vegetable communities - aquatics (lagoons, deep streams), marshy (yomomos, curichis), herbaceal (low-lying ground, herbaceous, and woody (mid elevations to tajibales) (Tabebuia spp.) and forest island. Affects the flooding dynamic. Tusicales (Machaerium hirtum), type of mid elevation community, found only in soils with pH> 8. [I modified the sentence. Experts, please verify. Thanks.] Maintenance of other communities requires 3-9 months of flooding. If pH changes and the flooding regime decrease, the vegetable communities tend to be drier. Measurement of the rainfall regime, flooding and pH. Fire regime Affects the dynamic and composition of vegetation. Cyperus giganteus Marsh (Yomomo o Pirizal): fire period >50 years. The bajos de canuela (Luziola, Panicum, Echinochloa, Hymenachne): should not get burned (is source of foraging, important habitat to a marsh deer). Mid elevation with little organic matter, burned sporadically: <10 years. High elevations towards the Andes: burned each year. Ecosystem degradation, decrease of the Tabebuia insignis island and heron nests and caimans and lizards habitats. dem for marsh deer. In places without tusicales, fire destroys the little organic matter, becomes more xerophytic. If there is no fire for 2-5 years in high elevations, the vegetation changes to woody. Fire frequencyDynamic minimum area Maintains the viability of all biotic elements including gallery forests, lagoons and forests island. Larger flooding areas: >200,000 ha.Large flooding events: every 10 years.Presence of top predators.Salinity Responsible for the presence of tusicales (open and espinoso forest of Machaerium hirtum) that serves as refuge for animals that take salt.Salnity level: see Werner Hanagarth (1993). Size of tusicales: de 0.5 a 5 /10 ha.The community depends of salinity and high pH.Dominion of Machaerium hirtum. Information gaps and caveats Vania Pivello: Food sources and feeding behavior of cerrado mammals and birds are not well known. The fire ecology of cerrados needs more studies, though there have been several research projects on this subject. For a list of information gaps on cerrado ecology and management, see Pivello, V.R. and G. Norton, 1996. Firetool: an expert system for the use of prescribed fires in Brazilian savannas. J. Appl. Ecol. 33: 348-356. Jeanine Felfili: 1- Control of weedy species in conservation units. 2 Non-invasive herbaceous species for recovery of degraded areas. 3 Monitoring of vegetation changes in permanent plots. 4 -- biodiversity assessments. 5 Silviculture of native species. Recommended priorities for conservation-driven research agenda and next steps Vania Pivello: 1. Implement fire management in protected areas that harbor representative cerrado ecosystems. 2. Increase connectivity between cerrado ecosystems and forest patches by adding corridors, buffer zones, etc. Literature Cited Aristeguieta, L., 1966. Flrula de la Estacin Biolgica de Los Llanos. Bol. Soc. Venez. Cienc. Nat.,110: 228-307. Barbault, R. 1983. Reptiles in savanna ecosystems. In Bourlire, F. (ed.) Tropical savannas. Ecosystems of the world 13. Elsevier Scientific Publishing Company, New York.Pp. 325-336. Baruch, Z. 1996. Ecophysiological aspects of the invasion by African grasses and their impact on biodiversity and function of neotropical savannas. In Solbrig, O.T., E. Medina and J.F. Silva (eds.) Biodiversity and savanna ecosystem processes: a global perspective. Springer-Verlag, Berlin, Heidelberg. Pp.79-93. Baruch, Z., J.A. Belsky, L. Bulla, A.C. Franco, I. Garay, M. Haridasan, P. Lavelle, E. Medina, and G. Sarmiento. 1996. In Solbrig, O.T., E. Medina and J.F. Silva (eds.) Biodiversity and savanna ecosystem processes: a global perspective. Springer-Verlag, Berlin, Heidelberg. Pp.175-194. Bilbao, B., R. Braithwaite, C. Dall'Aglio, B. Dias, A. Moreira, P. Oliveira, J. F. Ribeiro, and P. Stott. 1996. Biodiversity, fire, and herbivory in tropical savannas. In Solbrig, O.T., E. Medina and J.F. Silva (eds.) Biodiversity and savanna ecosystem processes: a global perspective. Springer-Verlag, Berlin, Heidelberg. Pp.197-203. Braithwaite, R. 1996. Biodiversity and fire in the savanna landscape. In Solbrig, O.T., E. Medina and J.F. Silva (eds.) Biodiversity and savanna ecosystem processes: a global perspective. Springer-Verlag, Berlin, Heidelberg. Pp.121-139. Bulla, L. 1996. Relationships between biotic diversity. In Solbrig, O.T., E. Medina and J.F. Silva (eds.) Biodiversity and savanna ecosystem processes: a global perspective. Springer-Verlag, Berlin, Heidelberg. Pp.97-117. Coupland, R.T. 1992. Overview of South American grasslands. In Coupland, R.T. (ed.) Natural grasslands. Ecosystems of the world 8A. Elsevier Scientific Publishing Company, New York. Pp. 363-366. Coutinho, L. M. 1990. Fire in the ecology of the Brazilian cerrado. In: Goldamer, J.G. (ed.) Fire in the tropical biota. Springer Verlag. Pp. 63-105. Daly, D.C. and J.D. Mitchell. 2000. Lowland vegetation of Tropical South America. In D. L. Lentz (ed.). Imperfect balance: landscape transformations in the Precolumbian Americas. Columbia University Press, New York. Pp. 391-453. Denslow, J.S. 1996. Functional group diversity and responses to disturbance. In Orians, G.H., R. Dirzo, and J.H. Cushman (eds.) Biodiversity and ecosystem processes in tropical forests. Springer-Verlag, Berlin, Heidelberg, New York. Pp. 127-151. Felfili, J.M., A. V. Rezende, M. C.Silva Junior and M.A. Silva. 2000. Changes in the floristic composition of cerrado (sensu stricto) in Brazil over a nine-year period. Journal of Tropical Ecology 16:579-590. Felfili, J.M., M. C. Silva Jr., A. V. Rezende, P. E. Nogueira, B. W. T. Walter, M. A. Silva, J. I. Encinas. 1997. Comparao Florstica e Fitossociolgica do Cerrado nas Chapadas Pratinha e dos Veadeiros. P. 6-11. In: Leite, L. and C. H. Saito. Contribuio ao conhecimento ecolgico do cerrado. Editora Universidade de Braslia. Felfili, J.M., T. S. Filgueiras, M. Haridasan, M. C. Silva Jr., R. Mendona and A. V. Rezende. 1994. Projeto biogeografia do bioma cerrado: Vegetao e solos. Cadernos de geocincias do IBGE.12: 75-166. Frost, P.G.H. and F. Robertson. 1987. The ecological effects of fire in savannas. In Walker, B. H. (ed.). Determinants of tropical savannas. Pp 93-140. IUBS Monograph Series, No.3. IRL Press Limited, Oxford, UK. Fry, C. H. 1983. Birds in savanna ecosystems. In Bourlire, F. (ed.) Tropical savannas. Ecosystems of the world 13. Elsevier Scientific Publishing Company, New York. Pp. 337-357. Goodland, R. and R. Pollard. 1973. The Brazilian cerrado vegetation: a fertility gradient. J. Ecol. 61: 219-224. Greller, A. M. 2000. Vegetation in the floristic regions of North and Central America. In D. L. Lentz (ed.). Imperfect balance: landscape transformations in the Precolumbian Americas. Columbia University Press, New York. Pp. 39-87. Hanagarth,W. 1993. Acerca de la geoecologa de las sabanas del Beni en el Noreste de Bolivia. La Paz. Happold, D.C.D.1983. Rodents and lagomorphs. In Bourlire, F. (ed.) Tropical savannas. Ecosystems of the world 13. Elsevier Scientific Publishing Company, New York. Pp. 363-400. Haridasan ??year in p. 28 Haridasan and Ranzani for nutrient levels in p. 28 Huber, O. 1982. Significance of savanna vegetation in the Amazon Territory of Venezuela. In Prance, G.T. (ed.) Biological diversification in the Tropics. pp. 221-244. Columbia University Press, New York. Huber, O.1987. Neotropical savannas: their flora and vegetation. Trends in Ecology and Evolution (TREE) 2(3): 67-71. Huber, O. 1995. Vegetation. In Berry, P., B.K. Holst and K. Yatskievich (Eds.). 1995. Flora of the Venezuelan Guayana. Vol.I: Introduction. pp. 97-160. Missouri Botanical Garden. St. Louis & Timber Press, Portland, Oregon. Huber, O. and G. Febres (editores). 2000. Gua ecolgica de la gran Sabana. Troncal 10: Piedra de la Virgen Santa Elena de Uairn. The Nature Conservancy EcoGraph, Caracas. 192 pp. Lewinsohn, T.M. and P.W. Price. 1996. Diversity of herbivorous insects. In Solbrig, O.T., E. Medina and J.F. Silva (eds.) Biodiversity and savanna ecosystem processes: a global perspective. Springer-Verlag, Berlin, Heidelberg. Pp.143-157. Lopes, S. and F. R. Cox. 1977. Cerrado vegetation in Brazil: an edaphic gradient. Agronomy Journal 69: 828-831. Metzger, J.P. and H. Dcamps. 1997. The structural connectivity threshold: an hypothesis in conservation biology at the landscape scale. Acta Oecologica 18:1-12. Mistry, J. 1998. Corticolous lichens as potential bioindicators of fire history: A study in the cerrado of the Distrito Federal, central Brazil. Journal of Biogeography 25 (3): 409-441. Ojasti, J. 1983. Ungulates and large rodents of South America. In Bourlire, F. (ed.) Tropical savannas. Ecosystems of the world 13. Elsevier Scientific Publishing Company, New York. Pp. 427-439. Oliveira-Filho, A.T. and J. A. Ratter. 1995. A study of the origin of central Brazilian forests by the analysis of plant species distribution patterns. Edinburgh Journal of Botany. v. 52, p. 141-194. 1995. Pivello, V.R. & Norton, GA. 199. FIRETOOL: an expert system for the use of prescribed fires in Brazilian savannas. J. Appl. Ecol. 33: 348-356. Pivello, V.R.et al. 1998. Abundance and distribution of native and alien grasses in a cerrado (Brazilian savanna) biological reserve. Biotropica, 31: 71-82 Ramia, M. 1974. Plantas de las Sabanas Llaneras. Monte Avla Editores, Caracas, 287 pp. Ramos-Neto, M.B. & Pivello, V.R. 2000 Lightning fires in a Brazilian savanna National Park: re-thinking management strategies. Environ. Manage. 26: 675-684. Rodrguez, J.P. and F.Rojas-Surez. 1999. Libro rojo de la fauna Venezolana (2da. Edicin). PROVITA, Caracas, Venezuela. Sarmiento, G. 1983. The savannas of tropical America. In Bourlire, F. (ed.) Tropical savannas. Ecosystems of the world 13. Elsevier Scientific Publishing Company, New York. Pp. 245-288. Silva, J.F. 1996. Biodiversity and stability in tropical savannas. In Solbrig, O.T., E. Medina and J.F. Silva (eds.) Biodiversity and savanna ecosystem processes: a global perspective. Springer-Verlag, Berlin, Heidelberg. Pp.161-171. Solbrig, O.T. 1996. The diversity of the savanna ecosystem. In Solbrig, O.T., E. Medina and J.F. Silva (eds.) Biodiversity and savanna ecosystem processes: a global perspective. Springer-Verlag, Berlin, Heidelberg. Pp.1-27. Solbrig, O.T., E. Medina, and J.F. Silva 1996. Determinants of tropical savannas. In Solbrig, O.T., E. Medina and J.F. Silva (eds.) Biodiversity and savanna ecosystem processes: a global perspective. Springer-Verlag, Berlin, Heidelberg. Pp. 31-39. Tania Sanaioti, INPA Recommended resources Beard, J.S. 1953. The savanna vegetation of northern tropical America. Ecological Monographs 23(2): 149215. Beck, S.G., J. Samiento, N. Paniagua Z., C. Miranda and M.O. Ribera. 2000. Humedales de Bolivia, una aproximacin a su conocimiento actual. Pp.119-150. In Homenaje al Acadmico Correspondiente Dr.h.c. C.N. Troels Myndel Pedersen. Academia Nacional de Agronomia y Veterinaria. Tomo LIV. Buenos Aires, Argentina. Berry, P., B.K. Holst and K. Yatskievich (Eds.). 1995. Flora of the Venezuelan Guayana. Vol. I: Introduction. Missouri Botanical Garden. St. Louis & Timber Press, Portland, Oregon. Coutinho, L. M. 1990. Fire in the ecology of the Brazilian cerrado. In: Goldammer, J.G. (ed.) Fire in the tropical biota. Springer Verlag. Pp. 63-105. Felfili, J. M. and M. C. SILVA JR. 1993. A comparative study of cerrado (sensu stricto) vegetation in central Brazil. Journal of Tropical Ecology 9(3): 277-289. 1993. Eiten, G. 1982. Brazilian savannas. In: Huntley, B.J. and B. H. Walker (eds.) Ecology of Tropical Savannas. Springer-Verlag. Pp.? Felfili, J.M. 1995. Diversity, structure and dynamics of a gallery forest in central Brazil. Vegetatio 117:1-15. Felfili, J.M., J. F. Ribeiro, C. W. Fagg, and J. W. B. Machado. 2000. Recuperao de matas de galeria. EMBRAPAS-CERRADOS. Planaltina. Doc. 21: 1-45. Felfili, J.M., A. V. Rezende, M. C.Silva Junior and M.A. Silva. 2000. Changes in the floristic composition of cerrado (sensu stricto) in Brazil over a nine-year period. Journal of Tropical Ecology 16:579-590. Filgueiras, T. S., J. M. Felfili, M. C. Silva Junior and P.E. Nogueira. 1998. Floristic and structural comparison of cerrado (sensu stricto) vegetation in central Brasil. Pp. 633-647. In: Dallmeyer, F. (ed.) Measuring and monitoring forest biological diversity. Ed. Smithsonian Foundation/MAB. The Parthenon publishing. New York. Furley, P.A., J. Proctor, and J. A. Ratter (eds.) 1992. Nature and dynamics of forest-savanna boundaries. Chapmen & Hall, London. 614 pages. Hanagarth, W. and S.G. Beck.1996. Biogeographie der Beni-Savannen (Bolivien). Geographische Rundschau 48 (11): 662-668. Braunschweig. Huber, O. 1974. The neotropical savannas. Select bibliography on their plant ecology and phytogeography. Istituto Italo-Latino Americano, Roma (Italia). xlii + 855 pp. Huber, O. 1999. Manejo integral de sabanas tropicales [Propuesta tcnica]. En Jornadas sobre desarrollo sostenible del medio rural: en busca del mejor camino (C. Pardo, coord. gen.), pp. 119120. Fundacin Polar, Caracas. Huber, O. 1990. Savannas and related vegetation types of the Guayana Shield region in Venezuela. In Sarmiento, G. (ed.) Las Sabanas Americanas: Aspectos de su biogeografa, ecologa y utilizacin. pp. 5797. Universidad de los Andes, Mrida, Venezuela. Huber, O.1987. Neotropical savannas: their flora and vegetation. Trends in Ecology and Evolution (TREE) 2(3): 67-71. Mendona, R., J.M. Felfili, B. M. T. Walter, M. C. Silva Jnior, A.V. Rezende, T. S. Filgueiras, and P.E.N. Nogueira. 1998. Flora vascular do bioma Cerrado. pp. 287-556. In: Sano, S. and S. Almeida. Cerrado, Ambiente e Flora. EMBRAPA CERRADOS. Ed. Planaltina, EMBRAPA-CPAC. Medina, E. and O. Huber 1992 [2ed ed. 1994]. The role of biodiversity in the functioning of savanna ecosystems. In Solbrig, O.T., H.M. van Emden and P.G.W.J. van Oordt (eds.). Biodiversity and global change. IUBS Monograph 8: 139158. International Union of Biological Sciences, Paris. Pinto, M.N. 1990 Cerrado: Caraterizao, ocupao e perspectivas. Braslia: Editora Universidade de Braslia, +657pp. Pivello, V.R. and L.. M. Coutinho. 1996. A qualitative successional model to assist in the management of Brazilian cerrados. Forest Ecology and Management 87: 127-138. Pivello, V.R. and G. A. Norton. 1996. Firetool: an expert system for the use of prescribed fires in Brazilian savannas. Journal of Applied Ecology 33: 348-356. Ratter, J. A ., S. Bridgewater, R. Atinkson and J. F. Ribeiro. 1996. Analysis of the floristic composition of the Brazilian vegetation of 98 areas. Edinburgh Journal of Botany 58(2):153-180. Ratter, J. A., J. F. Ribeiro, and S. Bridgewater. 1997. The Brazilian cerrado vegetation and threats to its biodiversity. Annals of Botany London 80(3): 223-230. Redford, K.H. and Fonseca, G. A. B. 1986. The role of gallery forests in the zoogeography of the cerrados non-volant mammalian fauna. Biotropica 18 (2): 126-135. Sano, S. and S. Almeida. 1998. Cerrado, Ambiente e Flora. EMBRAPA CERRADOS. Ed. Planaltina, EMBRAPA-CPAC. Sarmiento, G. 1984. The ecology of neotropical savannas (transl. By O. Solbrig). 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