Ecology Ecology of the Atlantic Forest
by
Elise Damstra-Oddy, Cristina Banks-Leite
  • LAST REVIEWED: 12 January 2021
  • LAST MODIFIED: 12 January 2021
  • DOI: 10.1093/obo/9780199830060-0233

Introduction

Extending along the southern coast of Brazil, into Argentina and Paraguay, the Atlantic Forest is a domain that once covered 150 Mha and includes many distinct forest subtypes and ecosystems. Its large latitudinal (29˚) and altitudinal (0–2,800 m above sea level) range, as well as complex topography in the region, has created microclimates within forest subtypes, which has led to biodiversity specifically adapted to narrow ecological ranges. The region is incredibly species-rich and is home to charismatic or economically important species such as the black and golden lion tamarin, the red-browned Amazon parrot, and the highly prized palm heart from Euterpe edulis. Through widespread human-driven change dating back to the arrival of European settlers in 1500, this realm has been extensively reduced, fragmented, and modified. Nowadays, this region is home to about 130 million people (60 percent of the Brazilian population) and is responsible for producing 70 percent of Brazil’s GDP, putting a strain on natural resources and providing challenges to conservation. Due to its high levels of endemic species coupled with a high threat of habitat loss and fragmentation, the Atlantic Forest has been identified as a “biodiversity hotspot.” Numerous studies have assessed the effects of habitat transformation on biodiversity and the consensus is that the majority of species are negatively affected. It is puzzling however that few species have actually gone extinct in the wild, even if some extinctions might have gone undetected. Extinctions do not immediately follow habitat change, there is often a time lag of many decades between habitat transformation and extinction. This may suggest that many species in the Atlantic Forest are “living deads,” as despite their presence the available habitat no longer supports their requirements. It also suggests that there is a window of opportunity to restoring the domain to avert extinctions before they are realized. Current research and policy actions are geared toward optimizing restoration and increasing the extent of native forest cover, bringing hope to the conservation of this unique domain.

General Overview of Forest Ecology and Forest Extent

Several good sources give detailed overviews of the forest ecology. Galindo-Leal and Câmara 2003 is a good general introduction to the history, biodiversity, and human impacts in the region as well as conservation management strategies; however, certain aspects of this text pertaining to current trends and conservation management are likely outdated. Metzger and Sodhi 2009, a special issue in Biological Conservation, focuses on conservation issues in the Atlantic Forest. Joly, et al. 2014 provides a comprehensive review of the history of disturbance, the ecology, the ongoing effects of fragmentation, and how climate change is impacting and will impact the Atlantic Forest. Rates of land use change in the Atlantic Forest tend to be monitored on a country-by-country basis, with remote sensing efforts in Brazil, Argentina, and Paraguay revealing different levels of deforestation, as seen in Azevedo, et al. 2018; Izquierdo, et al. 2008; and Huang, et al. 2009, respectively. In Brazil, low levels of deforestation are mostly matched by reforestation, which means that the amount of forest cover has either been stable or slowly increased in the past decades. The amount of remaining forest cover in Brazil has been measured by several groups and has been repeatedly updated since the 2000s as high-resolution satellite imagery becomes available. Galindo-Leal and Câmara 2003 reported that about 7–8 percent of Atlantic Forest still remained. Ribeiro, et al. 2009 calculated the existence of 11–16 percent of native vegetation, but when secondary forest fragments are excluded, the estimated remaining forest would stand at around 8 percent. More recently, using RapidEye imagery with 5 m of resolution, the authors of Rezende, et al. 2018 have shown that there is actually 26 percent of native vegetation. Because each group used different methods to assess the extent of forest cover, these estimates cannot be compared, and do not indicate that the amount of forest has increased over time.

  • Azevedo, T., C. M. Souza, J. Shimbo, and A. Alencar. 2018. MapBiomas initiative: Mapping annual land cover and land use changes in Brazil from 1985 to 2017. American Geophysical Union, Fall Meeting 2018, abstract #B22A-04, 10–14 December 2018, Washington D.C.

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    MapBiomas is an excellent interactive tool to visualize high-resolution land use change in Brazil as well as a reliable source of free geographical data to use in research.

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  • Galindo-Leal, C. G., and I. G. Câmara, eds. 2003. The Atlantic Forest of South America: Biodiversity status, threats, and outlook. Vol. 1. Washington, DC: Island Press.

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    This book provides an excellent initial overview of the history and threats to the Atlantic Forest, divided into sections by country.

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  • Huang, C., S. Kim, K. Song, et al. 2009. Assessment of Paraguay’s forest cover change using Landsat observations. Global and Planetary Change 67.1–2: 1–12.

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    This analysis of Paraguay’s forest cover shows that most forests outside of protected areas were lost by the 2000s, demonstrating the importance of protected areas.

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  • Izquierdo, A. E., C. D. De Angelo, and T. M. Aide. 2008. Thirty years of human demography and land-use change in the Atlantic Forest of Misiones, Argentina: An evaluation of the forest transition model. Ecology and Society 13.2: 3.

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    This paper examines changes in forest cover in Misiones, showing that there has been increased planting of pine and eucalyptus monocultures associated with a loss in natural forest.

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  • Joly, C. A., J. P. Metzger, and M. Tabarelli. 2014. Experiences from the Brazilian Atlantic Forest: Ecological findings and conservation initiatives. New Phytologist 204.3: 459–473.

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    This article provides a good discussion, with the use of conceptual models, of how large-scale landscape ecological processes can help maintain biota, as well as providing a research agenda that would conserve biodiversity of tropical forests.

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  • Metzger, J. P., and N. Sodhi, eds. 2009. Special issue: Conservation issues in the Atlantic Forest. Biological Conservation 142.6: 1137–1252.

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    This special issue of Biological Conservation contains some key articles on the Atlantic Forest including the highly impactful paper Ribeiro, et al. 2009 on how much of the Atlantic Forest remains and the distribution of the remnants.

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  • Rezende, C. L., F. R. Scarano, E. D. Assad, et al. 2018. From hotspot to hopespot: An opportunity for the Brazilian Atlantic Forest. Perspectives in Ecology and Conservation 16.4: 208–214.

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    This study presents high-resolution (5-m) remote sensing data of the Brazilian Atlantic Forest to reveal that there are in fact 32 Mha of forest within the domain, which corresponds to 28 percent of the original extent (2 percent of the area is planted forest).

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  • Ribeiro, M. C., J. P. Metzger, A. C. Martensen, F. J. Ponzoni, and M. M. Hirota. 2009. The Brazilian Atlantic Forest: How much is left, and how is the remaining forest distributed? Implications for conservation. Biological Conservation 142.6: 1141–1153.

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    This article provides a highly detailed analysis of how much forest cover remains in the Atlantic Forest by taking into account even quite small patches of forest. It discusses how the majority of the remaining patches are small, isolated, and composed of secondary vegetation, and it proposes four main strategies for protecting and restoring the Atlantic Forest.

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Historical Background

The earliest evidence of human activity within the Atlantic Forest is from approximately 500 CE, and consists of indigenous settlements causing modest disturbance. Since the arrival of Portuguese settlers in the 16th century, deforestation for urbanization and agriculture has been rampant. Two of the world’s most populous cities (São Paulo and Rio de Janeiro) are located within the Atlantic Forest realm, and Brazil’s economy also relies heavily on agriculture, including commodities such as coffee, cocoa, sugar, rice, soybean, and cotton, as explained by Martinelli, et al. 2010. Much of Brazil’s agriculture is exported and/or transported by trucks across Brazil, requiring an extensive road network, which, as shown in Freitas, et al. 2010, further exacerbates land use change and forest fragmentation. As for pre-Anthropocene history, there has been some research into the underlying evolutionary and paleoclimatic drivers responsible for the centers of endemism found within this region, such as the study Carnaval and Moritz 2008. Álvarez-Presas, et al. 2011 used planarians as model organisms to understand patterns of biodiversity, whereas the authors of Carnaval, et al. 2009 used frogs as indicators in their paleoclimatic models. Fjeldså and Rahbek 2006 used climate and phylogenetic data to explain the higher diversity of tanagers in the Atlantic Forest around Rio de Janeiro. Batalha-Filho, et al. 2013 used bird data to demonstrate how the Amazonian and Atlantic Forests were connected, linking the connection to geotectonic events.

  • Álvarez-Presas, M., F. Carbayo, J. Rozas, and M. Riutort. 2011. Land planarians (Platyhelminthes) as a model organism for fine‐scale phylogeographic studies: Understanding patterns of biodiversity in the Brazilian Atlantic Forest hotspot. Journal of Evolutionary Biology 24.4: 887–896.

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    Paleoclimatic models have previously had variable success in predicting the observed habitat stability in southern Atlantic Forest. This research uses two land planarians as model organisms because they have low dispersal capability, whereas previous work focused on species with high dispersal ability. Results suggest that there were no recent colonizations or population expansions, indicating a long-term stability scenario.

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  • Batalha-Filho, H., J. Fjeldså, P. H. Fabre, and C. Y. Miyaki. 2013. Connections between the Atlantic and the Amazonian forest avifaunas represent distinct historical events. Journal of Ornithology 154.1: 41–50.

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    This study uses phylogenetic and distribution data of birds to unravel the spatiotemporal dynamics of how the Atlantic and Amazon forests, which used to be connected, may have diverged.

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  • Carnaval, A. C., M. J. Hickerson, C. F. Haddad, M. T. Rodrigues, and C. Moritz. 2009. Stability predicts genetic diversity in the Brazilian Atlantic forest hotspot. Science 323.5915: 785–789.

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    This article uses frogs as indicators to examine different potential scenarios of community responses to climate change in the late-Quaternary period. The results show that the Brazilian Atlantic Forest had a relatively unstable climate, with the exception of three refugia that provided a suitable habitat for many neotropical species in the late Pleistocene. The authors suggest conservation priorities based on their results.

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  • Carnaval, A. C., and C. Moritz. 2008. Historical climate modelling predicts patterns of current biodiversity in the Brazilian Atlantic forest. Journal of Biogeography 35.7: 1187–1201.

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    This research models the range of the Atlantic Forest under current and past climatic scenarios to investigate whether these can predict current patterns of biodiversity distribution. The past range of the forest was verified using fossil pollen data. The authors find evidence for two refugia (Bahia and Pernambuco) and suggest that southern forests may have been more unstable.

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  • Fjeldså, J., and C. Rahbek. 2006. Diversification of tanagers, a species rich bird group, from lowlands to montane regions of South America. Integrative and Comparative Biology 46.1: 72–81.

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    This paper links rates of speciation within tanagers to geological events, and links the elevated levels of speciation in the Rio de Janeiro area and the Andean forelands to mountains being uplifted in the late Pleistocene.

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  • Freitas, S. R., T. J. Hawbaker, and J. P. Metzger. 2010. Effects of roads, topography, and land use on forest cover dynamics in the Brazilian Atlantic Forest. Forest Ecology and Management 259.3: 410–417.

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    This article examines the effect of road density, land use, and topography on forest fragmentation, deforestation, and regrowth. Roads were found to have the strongest links to deforestation and forest fragmentation, as they facilitate both processes by creating increased accessibility.

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  • Martinelli, L. A., R. Naylor, P. M. Vitousek, and P. Moutinho. 2010. Agriculture in Brazil: Impacts, costs, and opportunities for a sustainable future. Current Opinion in Environmental Sustainability 2.5–6: 431–438.

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    This is a thorough article that sets out the current state of Brazilian agriculture, the potential for sustainable development, and the major hurdles to achieving socioeconomic development without further environmental damage. Though it is about the whole of Brazil, it highlights key areas of conflict between economic and environmental priorities.

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Habitat Types: Present and Future

The Atlantic Forest is commonly divided either into forest types (a particular community of plant species that define a region can be referred to as a physiognomy) or into biogeographical subregions. The types of forest present are: dense and open ombrophilous forests, mixed ombrophilous forests, seasonal forests, and semideciduous as well as mangrove forests. The most studied forest physiognomy is the ombrophilous dense forest, which is believed to have higher levels of biodiversity and endemism than other physiognomies, although this could be due to differences in sampling effort. However, Scarano 2009 argues that these peripheral forest subtypes, including restingas and swamp forests, should be given more priority, making the case that they have a high conservation value due to their oligarchic diversity, with a few dominant species but many rare species at local scales. The western extent of the forest that extends into Argentina (an introduction to this forest can be found in Chebez and Hilgert 2003) and Paraguay (Cartes 2003 presents a background to this forest) forms part of the subtropical semideciduous forest. Silva and Casteleti 2003 defined eight biogeographical subregions: Araucaria, Bahia, Brejos Nordestinos, Diamantina, Interior, Pernambuco, Serra do Mar, and São Francisco. Five of these regions are considered to be centers of endemism, as shown by Tabarelli, et al. 2010 (cited under Biodiversity of the Atlantic Forest). Assessments of the potential impacts of climate change on the Atlantic Forest have revealed the area to be extremely vulnerable: Lemes, et al. 2014 found that as species ranges shift under climate change, protected areas will need to shift with them, particularly for vulnerable taxa like amphibians. Scarano and Ceotto 2015 reviewed the vulnerability of both biodiversity and society of the Atlantic Forest to climate change and discusses important adaptive practices.

  • Cartes, J. L. 2003. Brief history of conservation in the Interior Atlantic Forest. In The Atlantic Forest of South America: Biodiversity status, threats, and outlook. Edited by C. G. Leal and I. G. Câmara, 269–287. Washington, DC: Island Press.

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    This book chapter presents a thorough introduction to a lesser-researched area of the Atlantic Forest, the Paraguayan Atlantic Forest. The author details the key differences to Brazilian and Argentinian forests and discusses its history.

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  • Chebez, J., and N. Hilgert. 2003. Brief history of conservation in the Paraná Forest. In The Atlantic Forest of South America: Biodiversity status, threats, and outlook. Edited by C. G. Leal and I. G. Câmara, 141–159. Washington, DC: Island Press.

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    A descriptive history of the Argentinian Atlantic Forest from its early history to current land use in the region as well as an introduction to the local economic concerns and conservation efforts.

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  • Colombo, A. F., and C. A. Joly. 2010. Brazilian Atlantic Forest lato sensu: The most ancient Brazilian forest, and a biodiversity hotspot, is highly threatened by climate change. Brazilian Journal of Biology 70.3: 697–708.

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    This study examines how the distribution of thirty-eight different species of tree that are typical to the Brazilian Atlantic Forest would be affected by different potential future climate scenarios. They use species distribution models to show that at best, they would lose about 25 percent of their distribution, but under the worst scenarios, 50 percent could be lost, demonstrating the sensitivity of the region to climate change.

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  • Lemes, P., A. S. Melo, and R. D. Loyola. 2014. Climate change threatens protected areas of the Atlantic Forest. Biodiversity and Conservation 23.2: 357–368.

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    This article makes the excellent point that as species ranges change with climate change, protected areas need to expand to contain them, particularly in highland areas. By using amphibians as an example, the paper shows that without changes to protected areas, the number of species within them will decline.

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  • Neves, D. M., K. G. Dexter, R. T. Pennington, et al. 2017. Dissecting a biodiversity hotspot: The importance of environmentally marginal habitats in the Atlantic Forest Domain of South America. Diversity and Distributions 23.8: 898–909.

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    This article shows that marginal habitats with poor environmental protection are very important to maintain high species richness in the Atlantic Forest, as 45 percent of Atlantic Forest endemic species only occur within these habitats.

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  • Scarano, F. R. 2009. Plant communities at the periphery of the Atlantic rain forest: Rare-species bias and its risks for conservation. Biological Conservation 142.6: 1201–1208.

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    Scarano makes the case for a wider range of Atlantic Forest subtypes on the periphery of the core forest to be made a higher conservation priority despite lower diversity and endemism.

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  • Scarano, F. R., and P. Ceotto. 2015. Brazilian Atlantic forest: Impact, vulnerability, and adaptation to climate change. Biodiversity and Conservation 24.9: 2319–2331.

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    This review identifies vulnerabilities of biodiversity and society in the Atlantic Forest, as over 60 percent of Brazil’s population live within it. It discusses the role of ecosystem base adaptation strategies and highlights examples of good adaptive practice.

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  • Silva, J. M. C., and C. H. M. Casteleti. 2003. Status of the biodiversity of the Atlantic Forest of Brazil. In The Atlantic Forest of South America: Biodiversity status, threats, and outlook. Edited by C. G. Leal and I. G. Câmara, 43–59. Washington, DC: Island Press.

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    This chapter summarizes the threat posed by deforestation to the Brazilian Atlantic Forest and how much of the biodiversity is now threatened due to habitat loss. It also outlines potential solutions and conservation strategies.

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Biodiversity of the Atlantic Forest

Myers, et al. 2000 showed that nearly half of all plant species and more than a third of mammals, birds, reptiles, and amphibians are found in only 1.4 percent of the Earth’s land surface area, regions which are known as biodiversity hotspots. Hence, by focusing conservation policies and protecting these areas, a disproportionately high level of biodiversity could be protected. A follow-up book about biodiversity hotspots, Mittermeier, et al. 2005, discusses that 40 percent of the 20,000 plant species, 16 percent of the 688 bird species, 27 percent of the 261 mammal species, 31 percent of 200 reptile species, and 60 percent of 280 amphibian species are endemic to the Atlantic Forest, which means they can only be found within this realm. Together, they represent over 8,650 species, 8,000 of which are tree species. The Atlantic Forest has shown some of the highest levels of biodiversity in the world. Martini, et al. 2007 identified an area containing 144 species of trees (above diameter at breast height > 4.8 cm) within 0.1 ha in southern Bahia, which is the second highest concentration of tree species in the world. Among animal species, one notable example is the golden lion tamarin, an endangered primate species that Lapenta and Procópio-de-Oliveira 2008 found to have a role in the seed dispersal of ninety-seven species of plants. The Atlantic Forest has high levels of endemism and of habitat loss, making it one of the most endangered biodiversity hotspots. The species that are still present are often trapped within small fragments and unable to migrate, as discussed in Tabarelli, et al. 2010. Amphibians are a taxon of particular concern due to their high endemicity and the increasing threat posed by chytrid fungus, as shown in Carnaval, et al. 2006.

  • Carnaval, A. C. O. Q., R. Puschendorf, O. L. Peixoto, V. K. Verdade, and M. T. Rodrigues. 2006. Amphibian chytrid fungus broadly distributed in the Brazilian Atlantic Rain Forest. EcoHealth 3.1: 41–48.

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    This research reports the results of histological screenings on nearly one hundred anurans from the Atlantic rain forest, showing a wide spread of chytrid fungus, with an infection record coinciding with the first observed declines in amphibians.

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  • Lapenta, M. J., and P. Procópio-de-Oliveira. 2008. Some aspects of seed dispersal effectiveness of golden lion tamarins (Leontopithecus rosalia) in a Brazilian Atlantic forest. Tropical Conservation Science 1.2: 122–139.

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    This article examines the role of golden lion tamarins in the dispersal of seeds by analyzing the seeds found in fecal deposits. This evidence shows that the golden lion tamarin may be a keystone species, as it provides ecosystem services for so many other species.

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  • Martini, A. M. Z., P. Fiaschi, A. M. Amorim, and J. L. da Paixão. 2007. A hot-point within a hot-spot: A high diversity site in Brazil’s Atlantic Forest. Biodiversity and Conservation 16.11: 3111–3128.

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    This paper describes an area of exceptionally high biodiversity within the Atlantic Forest, located within southern Bahia, revealing it to be the second most diverse area in the world.

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  • Mittermeier, R. A., P. R. Gill, M. Hoffmann, et al. 2005. Hotspots revisited: Earth’s biologically richest and most endangered terrestrial ecoregions. Washington, DC: CEMEX.

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    This book takes a detailed look at the world’s biodiversity hotspots, looking into levels of endemism for many taxonomic groups.

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  • Myers, N., R. A. Mittermeier, C. G. Mittermeier, G. A. Da Fonseca, and J. Kent. 2000. Biodiversity hotspots for conservation priorities. Nature 403.6772: 853.

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    This is a key paper on the subject of biodiversity hotspots. It identifies areas of the world that are most concentrated in biodiversity and puts them forward as a “silver bullet” conservation strategy to conserve the highest level of biodiversity within the smallest area.

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  • Tabarelli, M., A. V. Aguiar, M. C. Ribeiro, J. P. Metzger, and C. A. Peres. 2010. Prospects for biodiversity conservation in the Atlantic Forest: Lessons from aging human-modified landscapes. Biological Conservation 143.10: 2328–2340.

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    This review provides a good introduction to the history of Atlantic Forest disturbance and discusses in detail various potential conservation options, highlighting the value of protected areas and old-growth forests. It also contains a useful table of studies that have examined the effects of habitat loss on different taxa and the conservation insights they provide.

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Biodiversity Data

There have been large-scale efforts to gather data on Atlantic Forest biodiversity. There are several public data sets known as the Atlantic data papers, which represent a high proportion of diversity present in the region, including: 94 species of mammal (see Souza, et al. 2019), 26 primate species (see Culot, et al. 2019), 745 bird species (see Hasui, et al. 2018), 2,095 epiphyte species (see Ramos, et al. 2019), 279 butterfly species (see Santos, et al. 2018), 528 amphibian species (see Vancine, et al. 2018), and 98 bat species (see Muylaert, et al. 2017). Several of these data sets monitor assemblages (e.g. Culot, et al. 2019) over time. As well as simple occurrence records, some of the data sets collate information on species traits and interactions, such as plant–frugivore interactions in Bello, et al. 2017, and bird traits including body mass and wing length in Rodrigues, et al. 2019. Going beyond simple species occurrence data allows researchers to investigate patterns over time such as demographics and make inferences about how anthropogenic stressors affect morphology. Interaction data is particularly valuable as it can uncover species mutualisms and the extent to which certain species depend on others. This contributes to the deeper understanding of how ecological communities work, which can be useful to inform better conservation policies. For the state of São Paulo, there is also information available on the SinBiota 2.1 platform, as described in Mira, et al. 2011. This was created by the Biota/Fapesp program to integrate information generated by all researchers funded by this program.

  • Bello, C., M. Galetti, D. Montan, et al. 2017. Atlantic frugivory: A plant–frugivore interaction data set for the Atlantic Forest. Ecology 98.6: 1729.

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    This data set contains pairwise observations between 331 vertebrate species and 788 plant species from the Atlantic Forest.

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  • Culot, L., L. A. Pereira, I. Agostini, et al. 2019. ATLANTIC‐PRIMATES: A dataset of communities and occurrences of primates in the Atlantic Forests of South America. Ecology 100.1: e02525.

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    This detailed data set contains georeferenced locations of all twenty-six primate species found in the Atlantic Forest as well as one introduced species. It describes 700 primate communities made up of over 8,000 individual single-species occurrences and covers the entire Atlantic Forest range including Brazil, Argentina, and Paraguay.

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  • Hasui, É., J. P. Metzger, R. G. Pimentel, et al. 2018. ATLANTIC BIRDS: A data set of bird species from the Brazilian Atlantic Forest. Ecology 99.2: 497.

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    This is a large data set comprising over 33,000 individual birds of 832 species from the Atlantic Forest in Brazil. It includes location, date, sampling mode, altitude, and type of habitat.

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  • Mira, C., P. Feijao, T. Duque-Estrada, J. Meidanis, and C. A. Joly. 2011. The SinBiota 2.0 biodiversity information system. In Proceedings 2011 Seventh IEEE International Conference on e-Science. 5–8 December 2011. pp. 142–149. Los Alamitos, CA: IEEE.

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    This conference paper discusses the history of the SinBiota platform and future plans to include new technologies.

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  • Muylaert, R. d. L., R. D. Stevens, C. E. L. Esbérard, et al. 2017. ATLANTIC BATS: A data set of bat communities from the Atlantic Forests of South America. Ecology 98.12: 3227.

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    This data set comprises information on over 90,000 individual bat captures totaling ninety-eight species. The data reported were collected in 205 sites from 135 studies.

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  • Ramos, F. N., S. R. Mortara, N. Monalisa-Francisco, et al. 2019. ATLANTIC EPIPHYTES: A data set of vascular and non‐vascular epiphyte plants and lichens from the Atlantic Forest. Ecology 100.2: e02541.

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    An extensive data set containing 2,095 species of epiphytes from the Atlantic Forest including 89,270 individual records. The data set includes records from 1824 to early 2018.

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  • Rodrigues, R. C., É. Hasui, J. C. Assis, et al. 2019. ATLANTIC BIRD TRAITS: A data set of bird morphological traits from the Atlantic forests of South America. Ecology 100.6: e02647.

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    A large data set of bird traits from 67,197 individuals of 711 species. The traits recorded include sex, body mass, body length, and reproductive stage.

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  • Santos, J. P. dos, A. V. L. Freitas, K. S. Brown, et al. 2018. Atlantic butterflies: A data set of fruit‐feeding butterfly communities from the Atlantic forests. Ecology 99.12: 2875.

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    A large data set of butterfly occurrence records from the Atlantic Forest including 7,062 presence records for 279 species dating from 1949 to 2018.

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  • Souza, Y., F. Gonçalves, L. Lautenschlager, et al. 2019. ATLANTIC MAMMALS: A dataset of assemblages of medium and large-sized mammals of the Atlantic Forest of South America. Ecology 100.10: e02785.

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    This data set compiles 129 studies resulting in occurrence data of ninety-four species of mammal in the Atlantic Forest across 244 assemblages.

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  • Vancine, M. H., K. da Silva Duarte, Y. S. de Souza, et al. 2018. ATLANTIC AMPHIBIANS: A data set of amphibian communities from the Atlantic Forests of South America. Ecology 99.7: 1692.

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    This data set of Atlantic Forest amphibians includes 17,619 records of 528 species. The records date between 1940 and 2017.

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Habitat Transformation

Habitat loss, fragmentation, and degradation are the major threats to biodiversity in the Atlantic Forest, thus there is a substantial body of literature on this subject. Lôbo, et al. 2011 showed that habitat transformation has over time modified diverse communities within five physiognomic subtypes (including evergreen, semideciduous, and open forest) of the Atlantic Forest into a homogenized set of disturbance-specialist species. Indeed, studying in southern Bahia (world’s second highest concentration of tree species), the authors of Benchimol, et al. 2017 found that forest loss leads to nonrandom floristic shifts, such that shade-intolerant species (e.g., pioneers) become more common than shade-tolerant species below 30 percent of forest cover. The loss of species also drives evolutionary changes in seed size, as shown by Galetti, et al. 2013. Within the evergreen and semideciduous forests, Santos, et al. 2008 demonstrated that fragmentation and the resulting creation of more edges have severely reduced functional traits of tree assemblages. But habitat loss is not the only driver of species loss. Using a multi-taxa database with over 2,200 community-level estimates from 1,097 sites, Püttker, et al. 2020 showed that forest-dependent species respond negatively to habitat loss and fragmentation, and that in areas with over 30 percent of forest cover, habitat fragmentation was as important as or more important than habitat loss in driving changes in species richness. This was not the first time that it has been shown that the effects of fragmentation on species are dependent on the amount of forest cover. Pardini, et al. 2010 demonstrated that within the ombrophilous dense forests, the size of a forest fragment only positively affects biodiversity when the landscape level forest cover is intermediate (around 30 percent of forest cover). The impacts of edge effects have been shown for a variety of abiotic and biotic conditions. Magnago, et al. 2015 showed that forest edges are drier and warmer, and these abiotic changes affect forest structure. Changes to habitat structure can then have knock-on effects on other taxa. For instance, Filgueiras, et al. 2011 found that dung beetle diversity was impacted by the impoverished flora of small patches. Banks-Leite, et al. 2010 found that edge effects likely drive the patch area effects on birds in the Atlantic Forest. This is because large patches experience a weaker influence of edge effects than small patches, which have higher edge-to-area ratio.

  • Banks-Leite, C., R. M. Ewers, and J. P. Metzger. 2010. Edge effects as the principal cause of area effects on birds in fragmented secondary forest. Oikos 119.6: 918–926.

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    This article demonstrates that edge and area effects are intrinsically confounded in fragmented landscapes, but that the magnitude of edge-to-interior differences increases in larger patches. When controlling for edge effects, the authors show that patch size does not affect birds.

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  • Benchimol, M., E. Mariano-Neto, D. Faria, et al. 2017. Translating plant community responses to habitat loss into conservation practices: Forest cover matters. Biological Conservation 209:499–507.

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    This study shows that species richness of plants declines drastically below 30 percent of forest cover, and that in deforested areas the assemblages of seedlings and saplings is very different from that of mature trees. These results suggest that the incredibly speciose forest of southern Bahia will progressively lose species as mature trees are replaced by younger individuals.

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  • Filgueiras, B. K., L. Iannuzzi, and I. R. Leal. 2011. Habitat fragmentation alters the structure of dung beetle communities in the Atlantic Forest. Biological Conservation 144.1: 362–369.

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    This research is based on field work performed in nineteen forest fragments of varying size in the Atlantic Forest in northeastern Brazil. It demonstrates not only that fragment area directly affects dung beetle diversity, but that low tree species diversity and lower levels of shade-tolerant plants also lowered dung beetle diversity.

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  • Galetti, M., R. Guevara, M. C. Côrtes, et al. 2013. Functional extinction of birds drives rapid evolutionary changes in seed size. Science 340.6136: 1086–1090.

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    This article makes the link between the reduction in seed size of a keystone palm species and the functional extinction of large-gape seed dispersers in the Brazilian Atlantic Forest. This shows a short-term adaptation to loss of large birds.

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  • Lôbo, D., T. Leão, F. P. Melo, A. M. Santos, and M. Tabarelli. 2011. Forest fragmentation drives Atlantic forest of northeastern Brazil to biotic homogenization. Diversity and Distributions 17.2: 287–296.

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    This article makes the important observation that land cover change is altering tree flora communities and that since 1980 similarity in species composition has increased by 20–40 percent.

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  • Magnago, L. F. S., M. F. Rocha, L. Meyer, S. V. Martins, and J. A. A. Meira-Neto. 2015. Microclimatic conditions at forest edges have significant impacts on vegetation structure in large Atlantic forest fragments. Biodiversity and Conservation 24.9: 2305–2318.

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    This study provides a good working example of how edge effects impact microclimatic conditions and how these changes affect vegetation structure at edges.

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  • Pardini, R., A. A. Bueno, T. A. Gardner, P. I. Prado, and J. P. Metzger. 2010. Beyond the fragmentation threshold hypothesis: Regime shifts in biodiversity across fragmented landscapes. PloS One 5.10: e13666.

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    This article presents a model that describes mechanisms and consequences of changes in biodiversity within fragmented landscapes, taking factors such as local extinction risk and immigration rates. They show that patch size only positively affects biodiversity in landscapes with intermediate amounts of cover (30 percent). In highly forested (50 percent) or deforested (10 percent) landscapes, species richness is not affected by patch size.

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  • Püttker, T., R. Crouzeilles, M. Almeida-Gomes, et al. 2020. Indirect effects of habitat loss via habitat fragmentation: A cross-taxa analysis of forest-dependent species. Biological Conservation 241:108368.

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    This article analyzes data from the Synthesis in Atlantic Forest Ecology and Sustainability Group. This group has collated data from various projects, putting together a database on amphibians, reptiles, birds, mammals, spiders, harvestmen, beetles, butterflies, termites, bees, ants, and other insects, as well as bryophytes, pteridophytes, and higher plants. Here they report their first results on the relative effects of habitat loss and fragmentation on forest specialists.

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  • Santos, B. A., C. A. Peres, M. A. Oliveira, A. Grillo, C. P. Alves-Costa, and M. Tabarelli. 2008. Drastic erosion in functional attributes of tree assemblages in Atlantic forest fragments of northeastern Brazil. Biological Conservation 141.1: 249–260.

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    This article examines data of tree assemblages in a hyperfragmented part of the Atlantic Forest in northeastern Brazil, showing that smaller fragments and higher edge length within the evergreen and semideciduous physiognomies have drastic effects on functional traits from seed size to proportion of pioneer and emergent species.

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Vertebrates and Habitat Transformation

Birds are one of the most commonly studied taxa in the Atlantic Forest due to their diversity and sensitivity. Responses of birds to fragmentation have been well documented: Zurita and Bellocq 2010 found forest cover to be the main driver of differences in bird communities in Argentina, while Morante-Filho, et al. 2015 showed the number of bird species in southern Bahia abruptly changed at a threshold of 50 percent forest cover. Banks-Leite, et al. 2012 showed that around São Paulo responses of bird communities to fragmentation do not to conform to the classical ecological species–area relationship. Instead, the main drivers of changes in bird communities across a gradient of disturbance are purported to be individual species reaching their extinction threshold. Despite conservation efforts, many species remain at high risk of extinction (as demonstrated in Canale, et al. 2012), particularly large mammals such as the jaguar, which, as shown by Paviolo, et al. 2016, have undergone high rates of extirpation. Umetsu and Pardini 2007 found that small mammals, particularly endemic species, have been found to be sensitive to land use change. The response of bats is less clear, as Gorresen and Willig 2004 found the highest levels of bat diversity in moderately fragmented landscapes. With regards to the response of amphibians to fragmentation, Becker, et al. 2007 found habitat loss to be a key driver of amphibian declines, particularly for forest species; and amphibians are particularly vulnerable to the coupled effects of fragmentation and climate change, as shown by Loyola, et al. 2014. The effects of fragmentation on reptiles are less studied, but Lion, et al. 2016 demonstrated that reptiles can benefit from even small forest fragments.

  • Banks-Leite, C., R. M. Ewers, and J. P. Metzger. 2012. Unraveling the drivers of community dissimilarity and species extinction in fragmented landscapes. Ecology 93.12: 2560–2569.

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    This article demonstrates that using species–area relationship in fragmentation studies does not hold for bird communities in the Atlantic Forest and that changes in community composition are primarily driven by species-level extinction thresholds.

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  • Becker, C. G., C. R. Fonseca, C. F. B. Haddad, R. F. Batista, and P. I. Prado. 2007. Habitat split and the global decline of amphibians. Science 318.5857: 1775–1777.

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    This article shows that as well as habitat loss and the rapid spread of chytrid fungus driving the losses of amphibians globally, fragmentation of habitat is also a key driver, particularly in species that migrate to forests from an aquatic larval stage.

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  • Canale, G. R., C. A. Peres, C. E. Guidorizzi, C. A. F. Gatto, and M. C. M. Kierulff. 2012. Pervasive defaunation of forest remnants in a tropical biodiversity hotspot. PloS One 7.8.

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    This paper examines local extinctions of 18 mammal species in 196 patches of the Brazilian Atlantic Forest, and finds that of a possible 3,528 populations that could have persisted, only 767 have, and each forest patch only retains an average of 3.9 species.

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  • Gorresen, P. M., and M. R. Willig. 2004. Landscape responses of bats to habitat fragmentation in Atlantic forest of Paraguay. Journal of Mammalogy 85.4: 688–697.

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    This article reports the outcome of an extensive survey effort on different landscape within the Atlantic Forest of Paraguay, finding the highest species richness in partly forested fragments.

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  • Lion, M. B., A. A. Garda, D. J. Santana, and C. R. Fonseca. 2016. The conservation value of small fragments for Atlantic Forest reptiles. Biotropica 48.2: 265–275.

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    This fragmentation study focuses on reptile communities in the Brazilian Atlantic Forest; though the main predictor of reptile species richness and abundance is fragment size, both the matrix quality and shape of the fragment contribute to the communities present.

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  • Loyola, R. D., P. Lemes, F. T. Brum, D. B. Provete, and L. D. Duarte. 2014. Clade‐specific consequences of climate change to amphibians in Atlantic Forest protected areas. Ecography 37.1: 65–72.

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    This paper shows that the suitable range for most amphibian species would contract under climate change and that the responses are clade-specific. It starts to piece together how understanding the changes in the phylogenetic pool may lead to a more comprehensive idea of the effect of climate change on assembly-related processes.

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  • Morante-Filho, J. C., D. Faria, E. Mariano-Neto, and J. Rhodes. 2015. Birds in anthropogenic landscapes: The responses of ecological groups to forest loss in the Brazilian Atlantic Forest. PLoS One 10.6.

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    The authors examined bird species grouped by whether they were forest specialists or habitat generalists and ran models to investigate the effect of fragmentation. The results find that all ecological groups show a similar forest cover threshold value of 50 percent, where species numbers abruptly change.

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  • Paviolo, A., C. De Angelo, K. M. Ferraz, et al. 2016. A biodiversity hotspot losing its top predator: The challenge of jaguar conservation in the Atlantic Forest of South America. Scientific Reports 6.1: 1–16.

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    A thorough review of the status of the jaguar’s habitat in the Atlantic Forest, combining information from fourteen research groups. It shows that jaguars only persist in 2.8 percent of the Atlantic Forest in low densities.

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  • Umetsu, F., and R. Pardini. 2007. Small mammals in a mosaic of forest remnants and anthropogenic habitats—evaluating matrix quality in an Atlantic forest landscape. Landscape Ecology 22.4: 517–530.

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    This paper investigates matrix use by small mammals of the Atlantic Forest, showing that assemblages of small mammals are highly dissimilar between native vegetation and anthropogenic land use. Endemic small mammals typically occupy native vegetation.

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  • Zurita, G. A., and M. I. Bellocq. 2010. Spatial patterns of bird community similarity: Bird responses to landscape composition and configuration in the Atlantic forest. Landscape Ecology 25.1: 147–158.

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    This paper examines how continuous and fragmented forest landscapes affect the similarity of bird communities, finding that forest cover explains most of the variation. It also finds that native bird communities are more resilient to forest loss in landscapes dominated by planted trees.

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Conservation and Policy

The Atlantic Forest is protected by the Forest Code, which is an environmental law created in Brazil in 1965, when most of the deforestation had already taken place. By law, landowners are required to set aside 20 percent of their land for native habitat, as well as protect riparian forests, hilltops, and other environmentally sensitive areas. The Forest Code was revised recently, weakening the protection of the Atlantic Forest, as discussed by Soares-Filho, et al. 2014. This revision is particularly problematic given that Banks-Leite, et al. 2014 has shown that at least 30 percent of native habitat is required to protect biodiversity within the Atlantic Forest. Due to the pressing need to preserve its unique yet endangered biota, a group of academics, NGOs, industry, and government formed the Atlantic Forest Restoration Pact, an initiative which aims to restore 15 Mha of habitat in the Atlantic Forest by 2050. This pledge comes as part of Brazil’s commitment to the Bonn Challenge. Crouzeilles, et al. 2019 shows that the Atlantic Forest Restoration Pact has already facilitated the restoration of roughly 700,000 ha, estimating that by 2020 there will be 1.5 Mha under restoration. Rezende, et al. 2018 has estimated that if landowners comply with the new Forest Code to restore riparian forest (i.e., forest strip along rivers), by 2038 the vegetation cover in the Atlantic Forest will be close to 35 percent, bringing hope to the preservation of this charismatic and species-rich system.

  • Banks-Leite, C., R. Pardini, L. R. Tambosi, et al. 2014. Using ecological thresholds to evaluate the costs and benefits of set-asides in a biodiversity hotspot. Science 345.6200: 1041–1045.

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    This study analyzes the responses of mammals, birds, and amphibians to habitat loss to show that the minimum amount of area required to maintain biodiversity is 30 percent of forest cover. The authors used these results to plan a domain-wide restoration strategy and demonstrate that with 6.5 percent of Brazil’s annual expenditure on agricultural subsidies, it would be possible to restore priority areas back to 30 percent of cover.

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  • Crouzeilles, R., E. Santiami, M. Rosa, et al. 2019. There is hope for achieving ambitious Atlantic Forest restoration commitments. Perspectives in Ecology and Conservation 17.2: 80–83.

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    The authors document the amount of forest that has been restored between 2011 and 2015 and how much restoration they expect to be under way by 2020. They discuss how this progress is due to the activities promoted by the Atlantic Forest Restoration Pact. They discuss how these activities could help restoration elsewhere.

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  • Rezende, C. L., F. R. Scarano, E. D. Assad, et al. 2018. From hotspot to hopespot: An opportunity for the Brazilian Atlantic Forest. Perspectives in Ecology and Conservation 16:208–214.

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    This study presents the most updated estimate of extent of forest cover within the Atlantic Forest. It also discusses that landowners are legally required to restore 5.2 Mha of currently degraded riparian area. Through adequate enforcement, restoring these areas could cause native vegetation cover to increase to 35 percent of forest cover.

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  • Soares-Filho, B., R. Rajão, M. Macedo, et al. 2014. Cracking Brazil’s Forest Code. Science 344.6182: 363–364.

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    This policy forum discusses the origins of Brazil’s Forest Code, the changes that were implemented recently, and their impacts on forest cover and carbon storage.

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