The Atlantic Forest (Portuguese: Mata Atlântica) is a South American forest that extends along the Atlantic coast of Brazil from Rio Grande do Norte state in the northeast to Rio Grande do Sul state in the south and inland as far as Paraguay and the Misiones Province of Argentina, where the region is known as Selva Misionera.

Atlantic Forest
Mata Atlântica
Area of the Atlantic Forest in Serra do Mar
Map
Map of the Atlantic Forest ecoregions as delineated by the WWF. The yellow line approximately encloses the forest's distribution.
(Satellite image from NASA)
Geography
LocationArgentina, Brazil, Paraguay
Area1,315,460 km2 (507,900 sq mi)
Discovery Coast Atlantic Forest Reserves
UNESCO World Heritage Site
Atlantic Forest in Bahia State, Brazil
LocationBrazil
IncludesUna Biological Reserve, PAU Brazil CEPLAC Experimental Station, Veracruz Station, Pau Brasil National Park, Descobrimento National Park, Monte Pascoal National Park, Linhares Forest Reserve and Sooretama Biological Reserve
CriteriaNatural: ix, x
Reference892
Inscription1999 (23rd Session)
Area111,930 ha
Atlantic Forest South-East Reserves
UNESCO World Heritage Site
View from a trail in the Salto Morato Reserve
Official nameAtlantic Forest South-East Reserves
LocationParaná and São Paulo, Brazil
Includes25 protected areas
CriteriaNatural: (vii)(ix)(x)
Reference893rev
Inscription1999 (23rd Session)
Area468,193 ha (1,807.70 sq mi)
Coordinates24°10′S 48°0′W / 24.167°S 48.000°W / -24.167; -48.000

The Atlantic Forest has ecoregions within the following biome categories: seasonal moist and dry broad-leaf tropical forests, tropical and subtropical grasslands, savannas, and shrublands, and mangrove forests. The Atlantic Forest is characterized by a high biodiversity and endemism.[1]

It was the first environment that the Portuguese colonists encountered over 500 years ago, when it was thought to have had an area of 1,000,000–1,500,000 km2 (390,000–580,000 sq mi), and stretching an unknown distance inland, making it, back then, the second largest rainforest on the planet, only behind the Amazon rainforest.[2] Over 85% of the original area has been deforested, threatening many plant and animal species with extinction.[3][4][5]

Ecology

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The Atlantic Forest region includes forests of several variations:

  • Restinga is a forest type that grows on stabilized coastal dunes. Restinga forests are generally closed canopy short forests with tree density. Open restinga is an open, savanna-like formation with scattered clumps of small trees and shrubs and an extensive layer of herbs, grasses, and sedges.[6][page needed]
  • Seasonal tropical moist forests may receive more than 2000 mm of rain a year. These include Tropical Moist: Lowland Forests, Submontane Forest, and Montane Forests.[6]
  • Tabuleiro forests are found over very moist clay soils and Tabuleiro Savannas occur over faster-draining sand soils.[6] These are humid areas that rely on water vapor from the ocean.[7]
  • Further inland are the Atlantic dry forests, which form a transition between the arid Caatinga to the northeast and the Cerrado savannas to the east. These forests are lower in stature; more open, with high abundance of deciduous trees and lower diversity when compared to tropical moist forests. These forests have between 700 and 1600 mm of precipitation annually with a distinct dry season. This includes Deciduous and Seasonal semideciduous forest each with their own lowland and montane regions.[6][7]
  • Montane forests are higher altitude wet forests across mountains and plateaus of southern Brazil. Also called Araucaria moist forests.
  • The Mussununga forests occur in southern Bahia and northern Espírito Santo states. The Mussununga ecosystem ranges from grasslands to woodlands associated with sandy spodosols. The word Mussununga is Amerindian Tupi-Guarani meaning soft and wet white sand.[8]
  • Shrubby montane savannas occur at the highest elevations, also called Campo rupestre.

The Atlantic Forest is unusual in that it extends as a true tropical rain forest to latitudes as far as 28°S. This is because the trade winds produce precipitation throughout the southern winter. In fact, the northern Zona da Mata of northeastern Brazil receives much more rainfall between May and August than during the southern summer. The geographic range of Atlantic Forest vary depending on author or institution that published them. Information on four most important boundaries as well as their union and intersection was reviewed in 2018.[9]

Geography

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The Atlantic Forest mainly covers regions of eastern Brazil (92% of the total area), but also reaches eastern Paraguay (6%) and northeastern Argentina (2%).[10]

History

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During glacial periods in the Pleistocene, the Atlantic Forest is known to have shrunk to extremely small fragmented refugia in highly sheltered gullies, being separated by areas of dry forest or semi-deserts known as caatingas.[7] Some maps even suggest the forest actually survived in moist pockets well away from the coastline where its endemic rainforest species mixed with much cooler-climate species. Unlike refugia for equatorial rainforests, the refuges for the Atlantic Forest have never been the product of detailed identification.

Biodiversity

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A jaguar

Despite having only 28% of native vegetation cover remaining,[11] the Atlantic Forest remains extraordinarily lush in biodiversity and endemic species, many of which are threatened with extinction.[12] Approximately 40 percent of its vascular plants and up to 60 percent of its vertebrates are endemic species, meaning they are found nowhere else in the world.[13] The official threatened species list of Brazil contains over 140 terrestrial mammal species found in Atlantic Forest. In Paraguay the Atlantic Forest has been heavily impacted in recent years.[14] In Paraguay there are 35 species listed as threatened, and 22 species are listed as threatened in the interior portion of the Atlantic Forest of Argentina. Nearly 250 species of amphibians, birds, and mammals have become extinct due to the result of human activity in the past 400 years. Over 11,000 species of plants and animals are considered threatened today in the Atlantic Forest.[7] Over 52% of the tree species and 92% of the amphibians are endemic to this area. The forest harbors around 20,000 species of plants, with almost 450 tree species being found in just one hectare in some locations.[15]

The Atlantic Forest is one of the best studied tropical ecosystems. For example, over 3000 tree species, 98 bat species, 94 large or medium-sized mammal species, over 2000 epiphyte species, 26 primate species, 528 amphibian species, 124 small mammal species, and over 800 bird species have been recorded in the Atlantic Forest.[16][17][18][19][20][21][22][23]

New species are continually being found in the Atlantic Forest. In fact, between 1990 and 2006 over a thousand new flowering plants were discovered.[citation needed] Furthermore, in 1990 researchers re-discovered a small population of the black-faced lion tamarin (Leontopithecus caissara), previously thought to have been extinct.[24] In 1991, the butterfly Actinote zikani was rediscovered in southern Brazil, after being declared extinct ten years earlier.[25] In southern Brazil A new species of blonde capuchin (Cebus queirozi), named for its distinguishing bright blonde hair, was discovered in northeastern Brazil at the Pernambuco Endemism Center in 2006.[26] A species of endangered three-toed sloth, named the maned sloth (Bradypus torquatus) because of its long hair, is endemic to the Atlantic Forest.[27] Hylid tree frog Dendropsophus branneri is also endemic to the Atlantic Forest.[28]

Conservation

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Human impact

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Atlantic Forest Fragmentation
 
Deforestation of Atlantic Forest in Rio de Janeiro

The incorporation of modern human societies and their needs for forest resources has greatly reduced the size of the Atlantic Forest, which has resulted in species impoverishment.[29] Almost 88% of the original forest habitat has been lost and replaced by human-modified landscapes including pastures, croplands, and urban areas. This deforestation continues at an annual rate of 0.5% and up to 2.9% in urban areas.

Agriculture: A major portion of human land use in the Atlantic Rain Forest is for agriculture. Crops include sugar-cane, coffee, tea, tobacco and more recently soybean and biofuel crops.[13]
Pasture: Even more common than using land for agriculture is the conversion of forest to cattle pastures.[30] This is commonly done by method of slash and burn which increases a forest chance of human-induced burning.
Hunting: Species in a fragmented forest are more susceptible to decline in population size because they are in a confined area that is more accessible to hunters. Larger animals make up the highest percentage of biomass. These animals are also the most rewarding to hunters and are heavily hunted in accessible fragments. This results in a change in species interactions such as seed dispersal and competition for resources.
Logging: Logging removes 10 to 80% of the canopy cover of a forest making that habitat more susceptible to natural elements such as wind and sunlight. This causes an increase in forest heating and desiccation.[31] Large amounts of organic litter and debris builds up which results in an increase in forests vulnerability to fires. Additionally, logging roads create accessibility for humans; and therefore increases the amount of human land disturbances and decreases the amount of natural forest.[31]
Fire: Human activity such as logging causes an increase in debris along forest floors that makes the Atlantic Forest more susceptible to fires. This is a forest type that is not accustomed to regular fire activity, so human induced fires dramatically affect the forest understory because plants do not have fire adaptations. In result, the forest becomes even more vulnerable to secondary fires, which are far more destructive and kill many more species including large trees.[31]

Results of human activity

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Habitat fragmentation leads to a cascade of alterations of the original forest landscape.[13] For example, the extent of human disturbances, including habitat destruction, in the Atlantic Forest has led to an extinction crisis.[32] The endemic species in this region are especially vulnerable to extinction due to fragmentation because of their small geographic ranges and low occurrence.[33] In a study of the Atlantic Forest fragments, community level biomass was reduced to 60% in plots less than 25 hectares.[34] Key ecological processes such as seed dispersal, gene flow, colonization and other processes are disturbed by fragmentation.[34] With many key vertebrate seed dispersers going extinct, it is predicted that many regional, fruit-bearing tree species in the Atlantic forest will become extinct due to failure of seedling recruitment and recolonisation.[30] With all these species already threatened, it is predicted that with the persistence of current deforestation rates the Atlantic forest will see continued extinction of species.[32]

Conservation by nongovernmental organizations

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Due to the Atlantic Forest's vast diversity of endemic plants and animals as well as the fragmentation affecting these species, many groups and organizations are working to restore this unique ecosystem. Non-governmental organizations (NGO) are huge benefactors in Brazil, providing funding as well as professional help to the Atlantic Forest due to the Brazilian Environmental Movement.[35] One organization, called BirdLife International, uses its research to preserve the area's bird biodiversity and teach people about sustainable natural resource use.[35]

Some organizations receive grants from the Critical Ecosystem Partnership Fund if they abide by its rules.[35] These include the Species Protection Program, the Program for Supporting Private Natural Heritage Reserves and the Institutional Strengthening Program.

Another strategy being implemented to maintain biodiversity within the Atlantic Forest is creating wildlife corridors. The World Bank is donating $44 million to create a corridor, which will be known as the Central Biodiversity Corridor, in the Atlantic Forest and one in the Amazon.[35] The Brazilian Development Bank has been financing, with non-reimbursable loans, 16 to 18 ecosystem restoration projects totaling 3,500 hectares and costing approximately $22 million under the so-called Iniciativa BNDES Mata Atlântica.[36] In order to preserve diversity, the state of São Paulo has created the Restinga de Bertioga State Park, a 9.3 thousand hectares park which also serves as a wildlife corridor linking the coastal regions to the Serra do Mar mountain range.[37] Some organizations, such as the Nature Conservancy, are planning to restore parts of the forest that have been lost and to build corridors that are compatible with the lifestyles of the native people.[38] The Amazon Institute is active in reforestation efforts in the northeastern state of Pernambuco, Brazil. During 2007, Joao Milanez and Joanne Stanulonis have planted 5,500 new trees in the mountains commencing with Gravata, adding to the precious little, ancient forest left.

The Pact for Atlantic Forest Restoration has assembled over 100 businesses, nongovernmental and governmental organizations around the goal of having 15 million hectares of the original ecosystem restored by 2050.[39]

The Pontifícia Universidade Católica do Rio Grande do Sul keeps a private reserve of the Araucaria moist forest ecoregion of approximately 3.100 ha called Pró-Mata, near the city of São Francisco de Paula in the state of Rio Grande do Sul. This reserve is used for research and biodiversity conservation.

Ecoregions

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Atlantic forest in Curitiba (Brazil).
 
Myiornis auricularis in the Atlantic forest of the state biological reserve of Aguaí

Terrestrial ecoregions within the Atlantic Forest Biome include:

Tropical and subtropical moist broadleaf forests
Tropical and subtropical dry broadleaf forests
Tropical and subtropical grasslands, savannas, and shrublands
Mangrove forests

See also

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References

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  1. ^ Dafonseca, G. 1985. The Vanishing Brazilian Atlantic Forest. Biological Conservation 34:17-34.
  2. ^ Por, Francis Dov. 1992. Sooretama: the Atlantic rain forest of Brazil. The Hague: SPB Academic Pub.
  3. ^ "The Atlantic Forest". The Nature Conservancy.
  4. ^ "Atlantic Forests, South America". WWF.
  5. ^ De Lima, Renato A. F.; Dauby, Gilles; De Gasper, André L.; Fernandez, Eduardo P.; Vibrans, Alexander C.; Oliveira, Alexandre A. De; Prado, Paulo I.; Souza, Vinícius C.; F. De Siqueira, Marinez; Ter Steege, Hans (2024-01-12). "Comprehensive conservation assessments reveal high extinction risks across Atlantic Forest trees". Science. 383 (6679): 219–225. doi:10.1126/science.abq5099. PMID 38207046.
  6. ^ a b c d Thomas, William Wayt, and Elizabeth G. Britton. 2008. The Atlantic coastal forest of Northeastern Brazil. Bronx, N.Y.: The New York Botanical Garden Press. ISBN 978-0-89327-498-6
  7. ^ a b c d Galindo Leal, Carlos, and Ibsen de Gusmão Câmara. 2003. The Atlantic Forest of South America: biodiversity status, threats, and outlook. Washington: Island Press.
  8. ^ Saporetti-Junior et al 2011.
  9. ^ Muylaert, Renata Lara; Vancine, Maurício Humberto; Bernardo, Rodrigo; Oshima, Júlia Emi Faria; Sobral-Souza, Thadeu; Tonetti, Vinicius Rodrigues; Niebuhr, Bernardo Brandão; Ribeiro, Milton Cezar (2018-09-11). "UMA NOTA SOBRE OS LIMITES TERRITORIAIS DA MATA ATLÂNTICA". Oecologia Australis (in Portuguese). 22 (3): 302–311. doi:10.4257/oeco.2018.2203.09. ISSN 2177-6199.
  10. ^ Fundación Vida Silvestre Argentina & WWF (2017). State of the Atlantic Forest: Three countries, 148 million people, one of the richest forests on Earth (PDF). Puerto Iguazú, Argentina.: Technical Report. p. 146.
  11. ^ Rezende, C.L.; Scarano, F.R.; Assad, E.D.; Joly, C.A.; Metzger, J.P.; Strassburg, B.B.N.; Tabarelli, M.; Fonseca, G.A.; Mittermeier, R.A. (October 2018). "From hotspot to hopespot: An opportunity for the Brazilian Atlantic Forest". Perspectives in Ecology and Conservation. 16 (4): 208–214. doi:10.1016/j.pecon.2018.10.002. ISSN 2530-0644.
  12. ^ de Lima, Renato A. F.; Oliveira, Alexandre A.; Pitta, Gregory R.; de Gasper, André L.; Vibrans, Alexander C.; Chave, Jérôme; ter Steege, Hans; Prado, Paulo I. (2020-12-11). "The erosion of biodiversity and biomass in the Atlantic Forest biodiversity hotspot". Nature Communications. 11 (1): 6347. Bibcode:2020NatCo..11.6347D. doi:10.1038/s41467-020-20217-w. ISSN 2041-1723. PMC 7733445. PMID 33311511.
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  14. ^ de la Sancha, Noé U., Sarah A. Boyle, and Nancy E. McIntyre. "Identifying structural connectivity priorities in eastern Paraguay's fragmented Atlantic Forest." Scientific reports 11, no. 1 (2021): 16129. https://doi.org/10.1038/s41598-021-95516-3
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  16. ^ de Lima, Renato A. F.; Oliveira, Alexandre A.; Pitta, Gregory R.; de Gasper, André L.; Vibrans, Alexander C.; Chave, Jérôme; ter Steege, Hans; Prado, Paulo I. (2020-12-11). "The erosion of biodiversity and biomass in the Atlantic Forest biodiversity hotspot". Nature Communications. 11 (1): 6347. Bibcode:2020NatCo..11.6347D. doi:10.1038/s41467-020-20217-w. ISSN 2041-1723. PMC 7733445. PMID 33311511.
  17. ^ Ramos, Flavio Nunes; Mortara, Sara Ribeiro; Monalisa-Francisco, Nathalia; Elias, João Pedro Costa; Neto, Luiz Menini; Freitas, Leandro; Kersten, Rodrigo; Amorim, André Márcio; Matos, Fernando Bittencourt; Nunes-Freitas, André Felippe; Alcantara, Suzana (February 2019). "ATLANTIC EPIPHYTES : a data set of vascular and non-vascular epiphyte plants and lichens from the Atlantic Forest". Ecology. 100 (2): e02541. doi:10.1002/ecy.2541. hdl:20.500.12008/30081. ISSN 0012-9658. PMID 30707454. S2CID 73437737.
  18. ^ Souza, Yuri; Gonçalves, Fernando; Lautenschlager, Laís; Akkawi, Paula; Mendes, Calebe; Carvalho, Mariana M.; Bovendorp, Ricardo S.; Fernandes-Ferreira, Hugo; Rosa, Clarissa; Graipel, Maurício Eduardo; Peroni, Nivaldo (October 2019). "ATLANTIC MAMMALS : a data set of assemblages of medium- and large-sized mammals of the Atlantic Forest of South America". Ecology. 100 (10): e02785. doi:10.1002/ecy.2785. ISSN 0012-9658. PMID 31180132. S2CID 182951440.
  19. ^ Culot, Laurence; Pereira, Lucas Augusto; Agostini, Ilaria; Almeida, Marco Antônio Barreto; Alves, Rafael Souza Cruz; Aximoff, Izar; Bager, Alex; Baldovino, María Celia; Bella, Thiago Ribas; Bicca-Marques, Júlio César; Braga, Caryne (January 2019). "ATLANTIC - PRIMATES : a dataset of communities and occurrences of primates in the Atlantic Forests of South America". Ecology. 100 (1): e02525. doi:10.1002/ecy.2525. hdl:10923/19194. ISSN 0012-9658. PMID 30317556. S2CID 52979277.
  20. ^ Vancine, Maurício Humberto; Duarte, Kauã da Silva; de Souza, Yuri Silva; Giovanelli, João Gabriel Ribeiro; Martins-Sobrinho, Paulo Mateus; López, Ariel; Bovo, Rafael Parelli; Maffei, Fábio; Lion, Marília Bruzzi; Ribeiro Júnior, José Wagner; Brassaloti, Ricardo (July 2018). "ATLANTIC AMPHIBIANS: a data set of amphibian communities from the Atlantic Forests of South America". Ecology. 99 (7): 1692. doi:10.1002/ecy.2392. hdl:11449/176529. PMID 29953585. S2CID 49589547.
  21. ^ Bovendorp, Ricardo S.; Villar, Nacho; de Abreu-Junior, Edson F.; Bello, Carolina; Regolin, André L.; Percequillo, Alexandre R.; Galetti, Mauro (August 2017). "Atlantic small-mammal: a dataset of communities of rodents and marsupials of the Atlantic forests of South America". Ecology. 98 (8): 2226. doi:10.1002/ecy.1893. hdl:11449/178972. PMID 28500789.
  22. ^ Lima, Fernando; Beca, Gabrielle; Muylaert, Renata L.; Jenkins, Clinton N.; Perilli, Miriam L. L.; Paschoal, Ana Maria O.; Massara, Rodrigo L.; Paglia, Adriano P.; Chiarello, Adriano G.; Graipel, Maurício E.; Cherem, Jorge J. (November 2017). "ATLANTIC-CAMTRAPS: a dataset of medium and large terrestrial mammal communities in the Atlantic Forest of South America". Ecology. 98 (11): 2979. doi:10.1002/ecy.1998. hdl:20.500.12219/4132. PMID 28857166. S2CID 21908829.
  23. ^ Hasui, Érica; Metzger, Jean Paul; Pimentel, Rafael G.; Silveira, Luís Fábio; Bovo, Alex A. d. A.; Martensen, Alexandre C.; Uezu, Alexandre; Regolin, André L.; Bispo de Oliveira, Arthur Â.; Gatto, Cassiano A. F. R.; Duca, Charles (February 2018). "ATLANTIC BIRDS: a data set of bird species from the Brazilian Atlantic Forest". Ecology. 99 (2): 497. doi:10.1002/ecy.2119. hdl:11449/179566. PMID 29266462.
  24. ^ Reaka-Kudla, Marjorie L, Don E. Wilson, and Edward O. Wilson. Biodiversity II: Understanding and Protecting Our Biological Resources. Washington, D.C.: Joseph Henry Press, 1997.
  25. ^ Francini, Ronaldo; Freitas, André Victor; Brown Jr., Keith Spalding (October 2005). "Rediscovery of Actinote zikani (D'Almeida) (Nymphalidae, Heliconiinae, Acraeini): natural history, population biology and conservation of an endangered butterfly in SE Brazil". ResearchGate. Retrieved August 29, 2024.
  26. ^ Pontes, A., A. Malta, and P. Asfora. 2006. A new species of capuchin monkey, genus Cebus erxleben (Cebidae, Primates): found at the very brink of extinction in the Pernambuco Endemism Centre. Zootaxa:1-12.
  27. ^ Cassano, C., M. Kierulff, and A. Chiarello. 2011. The cacao agroforests of the Brazilian Atlantic forest as habitat for the endangered maned sloth Bradypus torquatus. Mammalian Biology 76:243-250.
  28. ^ Martins-Sobrinho, Paulo Mateus; Silva, Winny Gomes de Oliveira; Santos, Elizandra Gomes dos; Moura, Geraldo Jorge Barbosa de; Oliveira, Jaqueline Bianque de (2017-07-26). "Helminths of some tree frogs of the families Hylidae and Phyllomedusidae in an Atlantic rainforest fragment, Brazil". Journal of Natural History. 51 (27–28): 1639–1648. doi:10.1080/00222933.2017.1337945. ISSN 0022-2933. S2CID 90234341.
  29. ^ Tabarelli, Marcelo; José Maria Cardoso Da Silva & Claude Gascon (June 2004). "Forest Fragmentation, Synergisms and the Impoverishment of Neotropical Forests". Biodiversity and Conservation. 13 (7): 1419–1425. doi:10.1023/B:BIOC.0000019398.36045.1b. S2CID 26608375.
  30. ^ a b IIzquierdo, Andrea E.; Carlos D. De Angelo & T. Mitchell 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. (online). doi:10.5751/ES-02377-130203. hdl:10535/3497.
  31. ^ a b c Laurance, William F., and Diana C. Useche. "Environmental Synergisms and Extinctions of Tropical Species." Conservation Biology 23.6 (2009): 1427-437.
  32. ^ a b Brooks, Thomas M., Mittermeier, Russell A., Mittermeier, Cristina G., Da Fonseca, Gustavo A. B., Konstant, William R., Flick, Penny, Pilgrim, John, Oldfield, Sara, Magin, Georgina, Hilton-Taylor, Craig. “Habitat Loss and Extinction in the Hotspots of Biodiversity.” Pérdida de Hábitat y Extinciones en Áreas Críticas para la Biodiversidad 16.4 (2002).
  33. ^ Patterson, B.D. & W. Atmar. "Analyzing species composition in fragments." Bonner Zoologische Monographen 46 (2000): Pp. 9-24.
  34. ^ a b Pütz, J. Groeneveld, L.F. Alves, J.P. Metzger, A. Huth. "Fragmentation drives tropical forest fragments to early successional states: A modelling study for Brazilian Atlantic forests." Ecological Modelling, 222. 12 (2011), pp. 1986-1997.
  35. ^ a b c d Biodiversity Hotspots - Atlantic Forest - Conservation Action Archived March 23, 2012, at the Wayback Machine. Biodiversity Hotspots - Home. 03 Oct. 2011.
  36. ^ Iniciativa BNDES Mata Atlântica
  37. ^ São Paulo Expands Atlantic Forest Protection Coverage. WWF Brasil. 03 Oct. 2011.
  38. ^ About the Atlantic Forest in Brazil. Nature Conservancy | Protecting Nature, Preserving Life. 03 Oct. 2011.
  39. ^ Pact for Atlantic Forest Restoration

Bibliography

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  • Saporetti-Junior, Amilcar Walter; Schaefer, Carlos Ernesto G. Reynaud; de Souza, Agostinho Lopes; Soares, Michellia Pereira; Araújo, Dorothy Sue Dunn; Meira-Neto, João Augusto Alves (21 September 2011). "Influence of Soil Physical Properties on Plants of the Mussununga Ecosystem, Brazil". Folia Geobotanica. 47 (1): 29–39. doi:10.1007/s12224-011-9106-9. S2CID 17406247.
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16°30′S 39°15′W / 16.500°S 39.250°W / -16.500; -39.250