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BIOTROPICA 32(4b): 786–792


Introduction: The Brazilian Atlantic Forest 1

L. Patrı´cia C. Morellato

Departamento de Botaˆnica, Plant Phenology and Seed Dispersal Research Group, Universidade Estadual Paulista, C.P. 199, 13506-900 Rio Claro, Sa˜o Paulo, Brazil


Ce´lio F. B. Haddad

Departamento de Zoologia, Universidade Estadual Paulista, C.P. 199, 13506-900 Rio Claro, Sa˜o Paulo, Brazil


The year 2000 marks 500 years of massive destruction for the Brazilian Atlantic Forest, as a consequence of the European colonization of Brazil. Today, the Atlantic Forest is restricted to ca 98,800 km 2 of remnants, or 7.6 percent

of its original extension. The Atlantic Forest continues to suffer under severe anthropogenic pressure, risking imminent

extinction of the remaining species. Our current knowledge indicates that this complex biome contains a species diversity higher than most of the Amazon forests, and also has high levels of endemism. The 13 selected articles in this special issue present data on the natural history, ecology, sustainable management, and conservation of the Atlantic Forest. These articles represent a sample of the research conducted to date in the region and suggest avenues of future research, particularly with regard to conservation alternatives for the remaining portions of the Atlantic Forest. This special issue represents one of the first general references pertaining to the Brazilian Atlantic Forest.



ano 2000 marca 500 anos de intensa destruic¸a˜o da Mata Atlaˆntica, como consequ¨eˆncia da colonizac¸a˜o europe´ia


Brasil. Hoje, a Mata Atlaˆntica esta´ restrita a aproximadamente 98.000 km 2 de remanescentes, ou 7,6% de sua

extensa˜o original, e os u´ltimos remanescentes de floresta ainda encontram-se sob intensa pressa˜o antro´pica e risco iminente de extinc¸a˜o. De acordo com o nı´vel atual de conhecimento, este bioma complexo contem maior diversidade

de espe´cies que a maioria das formac¸o˜es florestais amazoˆnicas, bem como nı´veis elevados de endemismo. Os treze

artigos selecionados para este fascı´culo especial apresentam dados sobre histo´ria natural, ecologia, manejo sustentado e conservac¸a˜o da Mata Atlaˆntica. Estes artigos representam uma amostragem das pesquisas realizadas na regia˜o, su- gerindo possibilidades de estudos futuros e discutindo alternativas para a conservac¸a˜o. Este fascı´culo especial corres- ponde a uma das primeiras refereˆncias gerais sobre a Mata Atlaˆntica brasileira.

Key words:

Atlantic Forest; biodiversity; Brazil; conservation; fragmentation; tropical forest.


lected data on the ecology and natural history of the Brazilian Atlantic Forest in the context of its endangered condition. In doing so, we hope to cat- alyze new and innovative research in the coming new century. We suggest that research on tropical forests, and especially on Brazilian Atlantic Forest, will require more coordination and direction to bring about effective conservation, rational man- agement, and public policy. We hope that this spe- cial issue will contribute to increased collaboration for future research. Before providing a brief sum- mary of the articles, we will define the Atlantic Forest as a biome, and give a brief historical ac- count of its occupation and conservation status.

1 Received 1 May 2000; revision accepted 15 September



Forest is composed of two major vegetation types:

the coastal forest or Atlantic Rain Forest (Fig. 1a, b) and the tropical seasonal forest or Atlantic Semi-deciduous forest (Fig. 1c, d). The Atlantic Rain Forest covers mostly the low to medium ele- vations ( 1000 m elev.) of the eastern slopes of the mountain chain that runs along the coastline from southern to northeastern Brazil; the Atlantic Semi-deciduous Forest extends across the plateau (usually 600 m elev.) in the center and south- eastern interior of the country (Leita˜o Filho & Mo- rellato 1997, Oliveira-Filho & Fontes 2000). The Atlantic Rain Forest experiences warm and wet cli- mate without a dry season; a seasonal climate with a relatively severe dry season (generally from April to September) predominates over the distribution of the Atlantic Semi-deciduous Forest (Morellato et al. 2000, Oliveira-Filho & Fontes 2000). The At-





Introduction 787 FIGURE 1. The two major vegetation types of the Atlantic Forest: (a) the coastal

FIGURE 1. The two major vegetation types of the Atlantic Forest: (a) the coastal Atlantic Rain Forest of the Serra do Mar mountain range, near the seashore at Praia da Fazenda, Picinguaba, Ubatuba, Sa˜o Paulo, Brazil, and (b) a view of the Atlantic Rain Forest on the Serra de Paranapiacaba mountain range, Capa˜o Bonito, Sa˜o Paulo, Brazil (Photo: M. A. Pizo); (c) the Atlantic Semi-deciduous Forest at Foz do Iguac¸u near the border of Brazil, Argentina, and Paraguay, and (d) in the Itaimbezinho, Bom Jesus, Rio Grande do Sul, represented by the temperate Araucaria forest (Photo: J. Sabino).

lantic Forest system lies almost entirely within the borders of Brazil, with small portions entering Ar- gentina and Paraguay. A detailed description of the Atlantic Forest system is presented by Oliveira-Fil- ho and Fontes in this issue.


CUPATION.—Five-hundred years ago, the Portu- guese first reached Brazil, beginning the European colonization in the east of South America (Bueno 1998, 1999). The colonization was initiated at the coast, in a land covered by luxuriant tropical forests now known as the Atlantic Forest (Dean 1995). The original area covered by the Atlantic Forest when European colonization began in A.D. 1500 was ca 1,300,000 km 2 , stretching from the state of Rio Grande do Norte at the easternmost tip of South America to as far as Rio Grande do Sul, the southernmost Brazilian state (Collins 1990). Today, 100 million Brazilians live in 3000 cities built on cleared lands of the Atlantic Forest (SOS Mata Atlaˆntica 1999). Some of these cities are among the largest in Latin America and in the world, such as

Sa˜o Paulo and Rio de Janeiro. The Atlantic Forest was cleared mainly for timber, firewood, charcoal, agriculture, cattle ranching, and the construction of cities. The remaining forests are still under severe anthropogenic pressure. This occupation, with no planning, caused a reduction of the forest to ca 98,800 km 2 , or 7.6 percent of its original extent (Ministe´rio do Meio Ambiente 1999; Fig. 2). An- other consequence was the high number of forest fragments and the various patterns of fragmenta- tion observed today (Fig. 3). The year 2000 marks 500 years of massive de- struction for this biome, which may lead to its im- minent extinction. Little is known of the Atlantic Forest, but what we do know is that this complex biome contains a species diversity higher than most of the Amazon forests, and is characterized by high levels of endemism (averaging nearly 50% overall, and as high as 95% in some groups; Brown & Brown 1992). The decimation of the Brazilian Atlantic Forest is one of the most alarming conservation problems in the world (Mori et al. 1981, Terborgh 1992,

788 Morellato and Haddad

788 Morellato and Haddad FIGURE 2. The original distribution of the Atlantic Forest in A.D. 1500

FIGURE 2. The original distribution of the Atlantic Forest in A.D. 1500 and its distribution in the 1990s. Small remnants are not represented in the map. Note that the largest remnants are concentrated between southern and southeastern Brazil.

Viana et al. 1997). Forest destruction has resulted in the elimination of many populations, and po- tentially, in the erosion of the genetic diversity of several species (Brown & Brown 1992). Based on the nearly total destruction of this biome and the high number of endemic species specialized to the forest environment, it is reasonable to suppose that many species already have gone extinct before they were described and that many others will soon van- ish.


In this special issue, we have gathered a selection of research representing varied, and occasionally disparate, approaches to the study of Atlantic For-

est biology. The common link is the biome—the Brazilian Atlantic Forest. We have tried to cover the main fields of forest structure and its dynamics, including floristic and plant phenology, plant–ani- mal interactions, and different animal groups, both insects and vertebrates. Most of the studies are at the community level, and several include a review of their subject, so we could achieve a more com- prehensive view of the Brazilian Atlantic Forest. The authors were called upon to discuss their work in the light of conservation issues. Clearly, there were a variety of ways and possibilities to choose subjects, invite contributors, and organize a collec- tion of articles. The particular selection here reflects the editors’ personal biases, but mainly covers the geographic distribution of research centers and of

the last large forest remnants of Atlantic Forest, both in southeastern Brazil; however, we acknowl- edge that research projects are being developed in other parts of the Atlantic Forest involving fields of knowledge not included in this special issue. The first article, by Oliveira-Filho and Fontes (2000), focuses on the definition of the Atlantic Forest based on floristic composition. Which forest vegetation can be considered as being part of the Atlantic Forest? What is the distribution of the At- lantic Forest? These and other questions are ad- dressed through an extensive analysis of the floristic composition of rain and semi-deciduous forests (103 sites, 2400 species), taking into account geographic and climatic variables. They suggest a broader definition of Atlantic Forest than is usually considered, including both semi-deciduous season- al forest and the rain forest. Differences in species composition occur between these forest types and these differences in turn are associated with rainfall seasonality (Oliveira-Filho & Fontes 2000). The phenological patterns analyzed by Morel- lato et al. (2000), involving 200 tree species, support the differences between the two Atlantic Forests (Morellato et al. 2000). This is the first comprehensive study to analyze the phenological patterns of Atlantic Rain Forest trees. Increasing climatic seasonality, especially variation in rainfall and the length of the dry season, influences the floristic composition and the seasonal patterns of both forests. While light and temperature are the key factors regulating seasonal patterns of Atlantic Rain Forest trees (Morellato et al. 2000), rainfall seasonality and low temperatures are suggested as the main factors affecting the phenology of trees in the Atlantic Semi-deciduous Forest (Morellato et al. 1989). Floristic diversity and phenology of the hum- mingbird-pollinated flora from Atlantic Rain Forest are discussed by Buzato et al. (2000). The plant populations bloom annually for the most part, and the community has staggered blooming times. Her- mit hummingbirds are the major pollinators of hummingbird-pollinated floras in the Atlantic Rain Forests of southeastern Brazil. Considering the or- nithophilous species, Bromeliaceae is the most im- portant family (Buzato et al. 2000). Two other pa- pers also deal with plant–animal interactions in the Atlantic Forest. Galetti et al. (2000) compare the frugivory by four of the five toucan species that occur in Atlantic Rain Forest. The diet of toucans is richer in the lowlands than in the highlands. The two species of the genus Ramphastos do not occur sympatrically in the highlands and their sympatry



in the lowlands occurs only at the peak of fruit availability, suggesting competitive exclusion (Gal- etti et al. 2000). Pizo and Oliveira (2000) discuss

the diverse interactions between litter-foraging ants and fallen fruits (non-myrmecochorous diaspores) on the forest floor of the Atlantic Rain Forest. This latter study suggests that this kind of interaction may be especially common in lowland rain forests, where the abundance of ants is dependent on the abundance and year-round availability of fruits (Pizo and Oliveira 2000). The three articles provide

a glimpse of the richness of plant–animal interac-

tions in the Atlantic Forest. The diversity of reproductive modes in hylid

frogs and a description of a new mode in anurans

is discussed by Haddad and Sawaya (2000). Eleven

of 12 reproductive modes known for hylid frogs around the world occur in the Atlantic Forest. The high diversity of reproductive modes observed for Atlantic Forest hylid frogs is the result of success- fully utilizing the diversified and humid microhab- itats in this biome (Haddad & Sawaya 2000). Di- versity and centers of endemism for South America mammals are investigated by Costa et al. (2000) using the parsimony analysis of endemicity. The patterns of distribution for marsupials, rodents, and primates of South America are compared with

a more restricted data set covering only the Atlantic Forest. Among 904 species of South American mammals, 229 are found in Atlantic Forest, 73 of those endemic to this biome (Costa et al. 2000). They confirm the idea of a fauna structured in the form of centers of endemism. The last six articles address questions related to Atlantic Forest conservation. Trajano (2000) pro- vides a synthesis of the knowledge on the cave fau- na in the Atlantic Forest. She concludes that the fauna of Atlantic Forest caves is not uniquely dif- ferent than the cave faunas of Cerrado and Caatin- ga. She discusses past and present conservation

problems of Atlantic Forest cave ecosystems, in the light of its intrinsic fragility and other distinctive features such as high degree of endemism and the morphological, ecological, and behavioral differ- entiation of these endemic species. The main threats to the cave fauna are deforestation (even on

a small scale), mining and the pollution resulting

from mining and other activities, overcollection, and unsupervised visitation. Due to their peculiar- ities, the presence of caves should be a strong pri- ority for surrounding land preservation (Trajano


A case study focused on one species, the ‘‘pal- miteiro’’ (palm heart tree) or Euterpe edulis, gives

790 Morellato and Haddad

790 Morellato and Haddad FIGURE 3. Variously fragmented forest landscapes in the Brazilian Atlantic Forest (black

FIGURE 3. Variously fragmented forest landscapes in the Brazilian Atlantic Forest (black forest; inclined hatching high second growth; vertical hatching tree plantation; dots Cerrados in a, coastal plain shrub restinga in e– f; larger dots Araucaria Forest in f). (a) Northwest of Brası´lia, DF (15 30 S, 48 W), headwater woods, and gallery forest in a hilly Cerrado landscape; (b) Campinas (CP), SP (22 50 S, 47 W), scattered Semi-deciduous Forest frag- ments including Santa Genebra Reserve (sg) and Ribeira˜o Cachoeira (rc) in an agricultural landscape; (c) Caitetus Ecological Station (ct), SP (22 20 S, 49 40 W), large and small Semi-deciduous Forest fragments in a mixed cattle ranch landscape; (d) Serra do Japi, Jundiaı´ (JD), SP (23 20 S, 47 W), montane forest of various ages in a

us an example of a native species that can be har- vested, but within the context of forest conserva- tion (Reis et al. 2000). Palmiteiro is a native palm tree having an economic importance that relies upon its edible product, the palm heart. Reis et al. (2000) propose a sustainable management system for natural populations of palmiteiro, conserving the forest in which it occurs. The fragmentation of Atlantic Forest and its conservation are the subject of the four remaining studies (Strier 2000, Machado & Fonseca 2000, Tabanez & Viana 2000, Brown & Freitas 2000). Some common ideas underlying these articles are:

the need for preserving even small forest frag- ments; the importance of connecting the frag- ments either to ensure the conservation of nearby fragments or to preserve or to restore ‘‘corridors’’ of vegetation, allowing the fauna to move from one fragment to another; and alternative manage- ment procedures, such as the control of edge ef- fects, which are particularly a problem in small fragments. All authors agree that forest fragments contain a high diversity of species. In turn, these fragments, through management, reveal the po- tential to maintain healthy populations of even large animals. Strier studied the impact of fragmentation and conservation on the ‘‘muriquis,’’ Brachyteles arachnoides, which is the largest nonhuman American primate endemic to the Atlantic Forest of southeastern Brazil. This endangered primate has a surprisingly low extinction probability, considering a high density population inhabiting a small, disturbed forest fragment, with no evi- dence of deleterious inbreeding effects. A high extinction probability is predicted for low den- sity populations living in large, undisturbed for- ests, but comparative data are needed to evaluate this prediction (Strier 2000). The effects of At- lantic forest fragmentation in the avifauna are discussed by Machado and Fonseca (2000). They find a positive correlation between number of species and fragment size, and great variation in species composition among seasons and across elevational gradients. Conservation of protected fragments, the protection of a larger number of



forest fragments, and the establishment of ‘‘cor- ridors’’ to connect fragments are suggested as management procedures needed to achieve the conservation of this rich and unique avifauna. Tabanez and Viana (2000) analyze the use of ‘‘eco-units,’’ or forest types, to determine the de- gree of disturbance for fragments of Atlantic Semi-deciduous Forest in southeastern Brazil, based on tree density, basal area, and species di- versity. They conclude that the eco-unit concept may be a useful tool to identify forest fragments that correspond to samples of local and regional diversity. The authors discuss the importance of conserving forest fragments and the utilization of some management procedures. Finally, Brown and Freitas (2000) discuss the use of butterflies as indicators for conservation of the Atlantic For- est. The authors base their analysis on 21 thor- oughly studied sites and 2100 species of but- terflies quantitatively censused over 35 years. They recognize six distinct subregions along the Atlantic Forest, each with relatively unique fau- nal compositions. They conclude that species richness is well predicted by landscape connec- tivity alone, or by composite indices of environ- mental heterogeneity, natural disturbance, and (negatively) anthropogenic disturbance. We believe that the special issue has accom- plished the intended goals: (1) to provide a repre- sentative sampling of studies that have been carried out in the Brazilian Atlantic Forest; (2) to show new approaches for future research; (3) to offer suggestions and support for conservation; and (4) to provide a general reference on the Brazilian At- lantic Forest.


The publication of the Brazilian Atlantic Forest special issue was made possible by a generous grant from FA-


PESP (Fundac¸a˜o de Amparo A Pesquisa do Estado de Sa˜o Paulo, proc. 98/12690-0) and additional support by FMB. We would like to thank Carlos A. Joly and Bob Marquis for the encouragement to produce the special issue, the latter for suggestions and corrections to this introductory article, all the authors for their contribu- tions, Jaime R. Somera for producing the distributional map of the Atlantic Forest, and Marco A. Pizo and Jose´

tree plantation landscape; (e) west of Caraguatatuba, SP (23 40 S, 45 40 W), steep mountains with nearly continuous heavy rain forest, restinga on coastal plain (center right, lower left); (f) southwest of Joinville (JV), SC (26 20 S, 48 50 W), a ‘‘conservation landscape’’ of small agriculture in a hilly forested matrix, with shrub restinga near the ocean and Araucaria Forest (larger dots) to the west. Scale bar (at top of b) 2 km in a; 5 km in b, c, d, and e; and 14 km in f. (Redrawn by A. V. L. Freitas; Brown and Freitas 2000; b–e from Kronka et al. [1993]).

792 Morellato and Haddad

Sabino for the photographs. L. Patrı´cia C. Morellato and Ce´lio F. B. Haddad are grateful to CNPq (Conselho Na- cional de Desenvolvimento Cientı´fico e Tecnolo´gico) for

the research productivity fellowships and thank FAPESP (grants 95/9626-0 and 95/9269-2, respectively) for sup- porting current research in the Atlantic Forest.


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