Tailoring Restoration Interventions to the Grassland-Savanna-Forest Complex in Central Brazil Isabel B
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OPINION ARTICLE Tailoring restoration interventions to the grassland-savanna-forest complex in central Brazil Isabel B. Schmidt1,2 , Maxmiller C. Ferreira3, Alexandre B. Sampaio4, Bruno M. T. Walter5,Daniel L. M. Vieira5 , Karen D. Holl6 Defining the reference system for restoration projects in regions characterized by complex vegetation mosaics is challenging. Here we use the Cerrado region of Brazil as an example of the importance of clearly defining multiple natural and anthro- pogenically altered states in grassland-savanna-forest mosaics. We define three main, natural vegetation types–grassland, savanna, and scleromorphic (cerradão) forest–to (1) distinguish between original and degraded states and (2) set appropriate targets for and guide restoration. We contend that the differences in Cerrado vegetation composition originally were driven by soil conditions and secondarily by fire frequency and precipitation patterns that differ from the core to the edge of the Cerrado region. Grasslands are found on the shallowest, least fertile soils and/or in waterlogged soils; scleromorphic forests are gen- erally located on deeper, more fertile soils; and savannas occupy an intermediate position. In recent decades, this biophysical template has been overlain by a range of human land-use intensities that strongly affect resilience, resulting in alternative anthropogenic states. For example, areas that were originally scleromorphic forest are likely to regenerate naturally follow- ing low- or medium-intensity land use due to extensive resprouting of woody plants, whereas grassland restoration requires reintroduction of grass and forb species that do not tolerate soil disturbance and exotic grass competition. Planting trees into historic grasslands results in inappropriate restoration targets and often restoration failure. Correctly identifying original vegetation types is critical to most effectively allocate scarce restoration funding. Key words: Brazilian savanna, Cerrado, natural regeneration, resilience, resprouting, woody encroachment with thresholds between them better describe the dynamics Implications for Practice • Land managers and scientists should collaborate to iden- of many ecosystem types (Westoby et al. 1989; Suding et al. tify the range of natural and anthropogenic states in 2004; Bestelmeyer et al. 2017). Identifying alternative natural grassland-savanna-forest mosaics to choose appropriate vegetation types and isolating the factors driving the transitions targets for restoration. between them and degraded areas are essential to select and • Identifying natural and anthropogenic factors influencing implement compatible management and restoration efforts. Yet, these vegetation types and their degraded states can help there are still few examples where the environmental drivers guide selection of the most suitable and cost-effective of transitions have been clearly identified and used to guide restoration techniques. restoration (Suding et al. 2004). Briske et al. (2005) highlight • The resprouting ability of woody species allows for high the utility of this approach; they describe drivers that influence resilience under low-intensity disturbance regimes; how- the transitions between grassland and woodland ecosystems in ever, herbaceous native species rarely recover naturally North American rangelands, pinpointing ecological features following extensive soil disturbance and exotic grass inva- sion. Author contributions: MCF, DLMV, ABS conceived the paper; IBS, BMTW, MCF • There is an urgent need to improve evidence-based performed the literature search; all authors wrote and edited the manuscript. restoration techniques in the Cerrado grassland-savanna- forest complex, especially how to control invasive grasses, 1Department of Ecology, University of Brasília, Campus Universitário Darcy Ribeiro, Asa Norte, Brasília 70910-900, Brazil reestablish soil conditions, and manage fire, since tech- 2Address correspondence to Isabel B. Schmidt, email [email protected] niques applicable at a large scale are necessary to achieve 3Biotechnology Institute, Federal University of Goiás – Regional Catalão, Avenida Doutor Lamartine Pinto de Avelar, Catalão 75704020, Brazil restoration commitments and targets. 4Centro Nacional de Avaliação da Biodiversidade e de Pesquisa e Conservação do Cerrado, Instituto Chico Mendes de Conservação da Biodiversidade – ICMBio, Brasilia 70635-800, Brazil 5Embrapa Recursos Genéticos e Biotecnologia, Asa Norte, Caixa Postal 02372, Brasília 70770-900, Brazil Introduction 6Environmental Studies Department, University of California, Santa Cruz, CA, U.S.A. Restoration ecologists have long recognized that few degraded ecosystems follow a linear recovery trajectory toward a © 2019 Society for Ecological Restoration doi: 10.1111/rec.12981 reference ecosystem (Trowbridge 2007; Matthews et al. 2009). Supporting information at: Vegetation models that incorporate multiple endpoints or states http://onlinelibrary.wiley.com/doi/10.1111/rec.12981/suppinfo 942 Restoration Ecology Vol. 27, No. 5, pp. 942–948 September 2019 Guide to restoring the Brazilian Cerrado that should be considered to plan and implement management states; and (3) pinpoint the most effective restoration interven- and restoration actions. tions based on the transition controlling variable and natural This need to distinguish among multiple states or vegetation regeneration mechanisms. types is especially true in savanna regions, where a complex mosaic of ecosystems naturally coexists across the landscape, influenced by soil properties, topography, rainfall patterns, nat- Cerrado Vegetation Types ural disturbances, and human activities (Lehmann et al. 2011). The Brazilian Cerrado is a mosaic of grasslands, savannas, Anthropogenic disturbances interact with natural drivers and and forests that originally occupied 2 million km2,mainlyin ecosystem resilience, shaping ecosystem structure and commu- central Brazil (Fig. S1, Supporting Information; Ribeiro & nity composition. In these complex ecosystem mosaics, it can be Walter 2008; MMA 2015). We followed the Cerrado sensu lato challenging to identify the level of ecosystem degradation and concept of Coutinho (1978), that is a gradient of vegetation determine restoration targets. In such systems, vegetation types types of the campo limpo (a grassland) to cerradão (sclero- are commonly classified according to the current vegetation morphic forest). It is a mesic savanna region, with a seasonal structure, which can lead to misinterpretations that jeopardize rainfall pattern varying from 600 to 2,000 mm/year (Sano et al. conservation and restoration (Veldman 2016). Old-growth 2019), 90% of the rainfall occurs between October and April savannas and grasslands (i.e. natural ancient ecosystems with with 2 to 5 months of the year with no precipitation (Agência a continuous herbaceous layer, and in the case of savannas, Nacional de Águas 2019). where grassy and woody species coexist, sensu Veldman et al. The Cerrado is the most biodiverse savanna worldwide 2015), are threatened when they are seen as degraded forests (>12,730 plant species) with high endemism rates (approxi- and targeted for forest restoration or carbon-sequestration mately 40% plant endemics) and ancient flora (Simon et al. initiatives (Veldman 2016). On the flip side, degraded forests 2009; Brazil 2018). Savannas and grasslands originally covered > may structurally resemble savannas or grasslands and be 75% of the Cerrado region (Eiten 1972; Ab’sáber 1981). misidentified as old-growth savannas or grasslands. These In these old-growth savannas, graminoids, forbs, and woody < misidentifications of reference systems can lead to misguided species coexist with crown cover ranging from 5–20% open conservation policies (Abreu et al. 2017) and the selection savanna-grasslands (regionally called cerrado ralo), to 20–50% in typical savannas (cerrado típico), to 50–70% canopy cover of inappropriate restoration targets and methods, leading to in closed savannas (cerrado denso). Herbaceous and shrub restoration failure (Suding 2011). For example, some state species form a continuous layer and represent 60–80% of laws in Brazil require planting trees regardless of the original the species richness (Ribeiro & Walter 2008; Amaral et al. vegetation and do not consider nontree species in revegetation 2017). Shallow and sandy soils are covered by old-growth requirements (e.g. Brazil DF Laws 14.783/1993; 23.585/2003). grasslands with no woody species (campo limpo) or with few This leads to the planting of fast-growing forest trees in areas shrub and tree species (campo sujo), whereas in deeper, mesic, that were originally grasslands and savannas, resulting in high and/or clay soils, tree canopy can cover 70–90%, forming > tree seedling mortality (commonly 60% in the first few years) scleromorphic forest (hereafter referred to as cerradão) with and/or nonrecruiting tree stands (Sousa & Vieira, in review), as a sparse herbaceous layer (Ribeiro & Walter 2008). Several described in other systems (Suding 2011; Veldman et al. 2015). studies link the occurrence of cerradão to deep, fertile soils Environmental laws and international commitments set a tar- with a high proportion of clay, percent organic matter, C, N, get for Brazil to restore approximately 12 million hectares of and K availability, and pH compared to areas of typical cerrado natural habitat of which 5 million hectares are within the Cer- (Furley et al. 1988; Furley & Ratter 1988; Haridasan 1992).