EDINB. J. BOT. 52(2): 141-194(1995) 141 A STUDY OF THE ORIGIN OF CENTRAL BRAZILIAN FORESTS BY THE ANALYSIS OF PLANT SPECIES DISTRIBUTION PATTERNS A. T. OLIVEIRA-FILHO* & J. A. RATTERf The floristic nature of central Brazilian forests, as well as their links to other main forest formations of eastern tropical South America, is assessed by means of multivariate analyses of 106 existing floristic checklists and by the analysis of a series of dot-maps showing the distribution of 55 woody species. Most species of central Brazilian forests seem to conform to two main distribution patterns: (1) species of deciduous and semideciduous forests are dependent essentially on the occurrence of patches of soils of intermediate to high fertility within the cerrado domain and tend to be distributed mostly along a northeast-southwest arch connecting the caatingas to the chaco boundaries; (2) considerable numbers of gallery forest species are dependent on high soil moisture and many appear to link the Amazonian and Atlantic rainforests by crossing the cerrado region in a northwest-southeast route via the dendritic net of gallery forests. Many gallery forest species are habitat generalists, some even occurring in the cerrado vegetation. Nevertheless, significant numbers of gallery endemics are also present. Galleries of the west and north of the Cerrado Province show stronger floristic links to the Amazonian rainforests, while those of the centre and south show stronger affinity with the montane semideciduous forests of southeastern Brazil. Present-day distribution patterns are discussed in the light of the current knowledge of palaeoenvironmental changes in the Neotropics. A natureza floristica das florestas do Brasil Central bem como seus lacos com outras grandes formacoes florestais da America do Sul tropico-oriental e avaliada por meio de analises multivariadas de 106 listagens floristicas existentes e pela analise de uma serie de mapas de distribuicao geografica de 55 especies lenhosas. A maioria das especies das florestas do Brasil Central parece ajustar-se a dois grandes padroes de distribuicao: (1) especies de florestas deciduas e semideciduas dependem essencialmente da ocorrencia de manchas de solos de media a alta fertilidade dentro do dominio dos cerrados e tendem a se distribuir principalmente ao longo de um arco nordeste- sudoeste que conecta as caatingas as fronteiras do chaco; (2) um consideravel numero de especies de florestas de galeria depende basicamente de alta umidade no solo e muitas delas parecem distribuir-se das florestas pluviais da Amazonia ate as Atlanticas, cruzando a regiao do cerrado numa rota noroeste-sudeste atraves da rede dendritica de florestas de galeria. Muitas especies de floresta de galeria sao generalistas quanto ao habitat, sendo inclusive compartilhadas com os cerrados. Um significativo numero, * Depto. Ciencias Florestais, Universidade Federal de Agricultura de Lavras, Lavras 37200-000, MG, Brazil. t Royal Botanic Garden Edinburgh, Inverleith Row, Edinburgh EH3 5LR, UK. This paper is one of the results of a postdoctoral collaborative study carried out at the Royal Botanic Garden Edinburgh from September 1992 to March 1994, under the sponsorship of the CNPq, Brazilian Government's Council for Research (grant n° 201995/91-3). Support also came from the European Community for the project 'Biodiversity of the Brazilian cerrado' (financial contribution B92/4-3040/9304) which also funded some of the fieldwork. 142 A. T. OLIVEIRA-FILHO & J. A. RATTER entretanto, e endemico das florestas de galena. As galerias do oeste e norte da Provincia do Cerrado mostram uma ligacao floristica mais forte com as florestas pluviais da Amazonia, ao passo que as do centro e sul mostram uma estreita afinidade com as florestas semideciduas montanas do sudeste brasileiro. Os padroes de distribuicao geografica atuais que emergiram das analises sao discutidos so ba luz do presente conhecimento sobre as mudancas paleoambientais nos Neotropicos. INTRODUCTION The Cerrado Biogeographic Province of tropical South America covers an area of about 2 million km2 in Brazil, corresponding to one quarter of the land surface of the country, plus small areas in northwestern Paraguay and eastern Bolivia; it is surpassed in area only by the Amazonian Province (Cabrera & Willink, 1973). The predominant vegetation prior to the main human disturbances of this century was of a woody savanna with a physiognomy varying from closed woodland to open grassland. The Cerrado Province extends mainly over the plateaux of central Brazil and is part of the so-called 'diagonal of open formations' (Vanzolini, 1963) or 'corridor of xeric vegetation' (Bucher, 1982), that includes the much drier Caatinga Province, in northeastern Brazil, and Chaco Province, in Argentina-Paraguay- Bolivia. This dry corridor runs between the two main regions of moist forest of tropical South America, the Amazonian Province, in the northwest, and the Atlantic- Paranense Provinces, in the east and southeast (Fig. 1). Most of the recent and controversial discussion on the palaeoenvironmental changes of tropical South America addresses the waxing and waning of both the open seasonal formations and humid forests during the late Quaternary climatic fluctuations. Geoecological data derived from soils, landforms and palynology, plus present-day distribution patterns of plants and animals, have given substantial evi- dence for the expansion of open vegetation formations and accompanying forest contraction during the glacial maxima as a result of drier, cooler macroclimates (Brown & Ab'Saber, 1979; Van der Hammen, 1982; Dickinson & Virji, 1987). Most studies indicate two main dry, cool periods of forest contraction, the first and more severe during the Pleistocene, coinciding with the Northern Hemisphere Wurm- Wisconsin glaciation, and the second during the Holocene (Ab'Saber, 1979, 1980, 1982; Bigarella & Andrade-Lima, 1982). Although most authors now agree that major glacial cycle-related climatic and vegetation shifts did occur, a great deal of controversy still focuses on the role that possible fragmentation of forest into isolated refugia played in speciation and on present-day distribution patterns (Haffer, 1969, 1982; Prance, 1973; Brown & Ab'Saber, 1979; Benson, 1982; Endler, 1982; Forero & Gentry, 1987; Beven et al., 1984). The postulated geographical location of some refugia has also been seriously challenged (Liu & Colinvaux, 1985; Colinvaux, 1989). One of the most concrete sources of evidence for palaeoenvironmental changes at any latitude is the pollen record. However, palaeopalynological data for the Neotropics are very scarce and most of the studies relating pollen record to palaeo- climatic changes are centred in the Andean and Amazonian regions or in Central America (Van der Hammen, 1974; Absy & Van der Hammen, 1976; Absy, 1982; Liu ORIGIN OF CENTRAL BRAZILIAN FORESTS 143 Guayana Province Amazonian Province Atlantic Province Paranense Province Caatinga Province Chaco Province Brazilian savannas FIG. 1. Map showing the distribution of Brazilian savannas (cerrados), according to IBGE (1993), superimposed on the seven phytogeographic provinces of eastern tropical South America, according to Cabrera & Willink (1973). & Colinvaux, 1985; Colinvaux, 1989; Markgraf, 1989; Busch & Colinvaux, 1990; Busch et al., 1990; Absy et al., 1991). It was only recently that Ledru (1991, 1993) contributed the first palaeopalynological study for the Cerrado Province of central Brazil, carried out at Salitre in western Minas Gerais. She concluded from the analy- sis of pollen distribution and modern vegetation and climate that between 33,000 and 25,000 BP the region was moister than today and was covered by rainforests (Ledru, 1993). During the Pleistocene glacial maximum, from 17,000 to 13,000 BP, a drier climate predominated and the proportion of trees in the vegetation declined, although only in the final term did arboreal cover reduce to the extent of a forest retreat. In another recent study, based on the analysis of plant species distribution, Prado & Gibbs (1993) proposed that during the Pleistocene glacial maximum the present-day corridor of xeric vegetation was covered mostly by an extensive seasonal woodland formation. This agrees with the findings of Ledru (1993) at Salitre (right in the middle of the corridor) who suggested the predominance of a seasonal arboreal 144 A. T. OLIVEIRA-FILHO & J. A. RATTER vegetation during most of the Pleistocene, when the climate would be drier than now but not severely arid as previously thought (Brown & Ab'Saber, 1979; Bigarella & Andrade-Lima, 1982). According to Ledru (1993), the period following the last Pleistocene glacial maxi- mum was characterized by the expansion of the Araucaria forest, indicating the predominance of a cooler and moister climate until 8500 BP. This period suffered a sudden and short-term interruption between 11,000 and 10,000 BP, corresponding to a dry interval accompanied by forest retreat. After 8500 BP the Araucaria forest was replaced by mesophytic forest, indicating an episode of moist climate with higher temperatures lasting until 5500 BP. The mid-Holocene dry interval which ensued at about 5000 BP caused a severe reduction in the forest area; since then forest expansion has once more occurred but not enough to recover the whole extent of the previous mesophytic forest period. Some authors believe that disturbances, both anthropo- genic and natural (fire in particular), helped retard forest expansion within the Cerrado Province during the late Holocene, despite the fact that climate and soils certainly favour the establishment of forests in most of the region (Reis, 1971; Klein, 1975). The isolated areas of cerrado (savanna) that are found enclosed by the Amazonian forest (see Fig. 1) provide evidence both of the forest expansion through a previously cerrado-dominated landscape and of the maintenance of cerrado vegetation in areas where the climatic conditions could support forest vegetation (Bigarella et al., 1975). It is now clear that the effects of the late Quaternary climatic fluctuations on the open vegetation formations of central Brazil were not restricted to a simple expansion and contraction of cerrado vs. rainforest, but also included complex changes within the dry corridor region itself. Cerrados, seasonal forests, rainforests, and even Araucaria forests appear to have thrived in various epochs at least in certain areas of the region. In addition to climatic variation, other factors such as soils, biotic pressure and disturbance could also have had profound palaeoenvironmental effects, and further data are necessary to provide a reliable description of the past history of the vegetation cover of central Brazil. The dry corridor of open formations has been considered by many authors as an important impediment to species migration between the two main South American forest regions, thus explaining many of the differences in floristic composition between them. On the other hand, the disjunct distribution patterns of a considerable number of species in the coastal Atlantic and Amazonian rainforests suggest the possibility of past forest links (Rambo, 1961; Rizzini, 1963; Andrade-Lima, 1966). Andrade-Lima (1964) and Rizzini (1963) refer to a possible migration route through a postulated mesophytic forest corridor that would have traversed the present-day Caatinga Province of northeastern Brazil at certain periods since the late Tertiary. Andrade-Lima (1982) suggested that the montane forests ('brejos') that presently exist isolated within the semi-arid region are relics of an ancient and wider forest cover. Bigarella et al. (1975) argued that, in addition to this Northeast Bridge, the floristic similarity between the eastern Amazonian and southeastern Atlantic rain- forests demonstrates that there must have been a second migration route, which they called the Southeast-Northwest Bridge. ORIGIN OF CENTRAL BRAZILIAN FORESTS 145 Although Bigarella et al. (1975) did not themselves suggest how the Southeast- Northwest Bridge connected the two forest regions, one could postulate that such a connection could have run through central Brazil, either as a continuous forest corridor or as a series of forest patches between which 'island-hopping' occurred. The idea of the Cerrado Province acting as an effective obstacle to the migration of forest species seems rather doubtful when a detailed map of present-day vegetation is examined. The map of Brazilian savannas (cerrados) (Fig. 1), extracted from IBGE (1993), clearly shows that the distribution of cerrados in central Brazil is far from continuous. When this is superimposed on the Cerrado Province of Cabrera & Willink (1973) a great proportion of gaps is shown (Fig. 1). These correspond essen- tially to patches of semideciduous forests, deciduous forests and to the transitional mosaics between these forests and the cerrado, called 'areas of ecological tension' by IBGE (1993). In addition to this, the savanna (cerrado) vegetation formation in the IBGE's classification system includes gallery forests, which are ubiquitous throughout the cerrado region. Therefore, forests are a far from negligible component of central Brazilian vegetation and their role as migration routes for forest species cannot be neglected, especially when evidence exists of a wider forest cover during a number of episodes in the late Quaternary. In fact, even now, the occurrence of mesophytic forest (deciduous and semideciduous forests) within the cerrado biome is very extensive and generally underestimated; during a recent journey by road from the Distrito Federal to Estreito in Maranhao, crossing c. 1400km of the states of Goias and Tocantins, one of us (J.A.R.) estimated that probably nearly 50% of the vegetation traversed was degraded mesophytic forest or the closely related meso- trophic cerradao. The deciduous forests that occur within the Cerrado Province on patches of calcium-rich soils have already been regarded by Prado & Gibbs (1993) as an important 'island-hopping' migration route connecting the semi-arid caatingas to the semideciduous forests of the Parana basin and of the Andean flanks. If such a NE-SW route existed in central Brazil for deciduous calciphilous species, the exist- ence of a NW-SE route for species associated with higher soil moisture is also conceivable through gallery forests. A number of authors have already suggested that the central Brazilian gallery forests represent floristic intrusions of the Amazonian and/or Atlantic forests into the cerrado domain. Cabrera & Willink (1973) mentioned that the galleries are actually a net connecting the forests of the Amazonian and Paranense Provinces. Based on floristic similarities, Rizzini (1979) stated that the galleries, together with other hinterland forests, are 'mediterranean extensions of the great Atlantic forest'. He also added that in Mato Grosso the continuous link to the Hylaea would give the galleries an Amazonian floristic nature. In fact, a considerable number of Amazonian species are known to penetrate the cerrados via gallery forests in that state, including species of rubber-tree (Hevea), a genus which is often reported to define the borders of the Amazonian forests (Veloso, 1966; Pires & Prance, 1977; Daly & Prance, 1989). Pires (1984) argued that if forest refuges did exist in the Amazon during the glacial ages they would not have had the postulated island 146 A. T. OLIVEIRA-FILHO & J. A. RATTER configuration, but a dendritic pattern following river drainage. This author also suggested that present-day galleries within the cerrado region constitute hodiernal forest refuges. In the present contribution we investigate the floristic nature of central Brazilian forests by the analysis of present-day distribution patterns. Our work was based essentially on multivariate analyses of existing floristic checklists and on the analysis of a series of dot-maps showing the distribution of woody species. The links to other main forest formations were assessed, with special attention to the connections to the Amazonian and Atlantic forests. Although we are aware of the many risks of drawing conclusions from the still poor floristic record, we believe that major patterns did emerge from our analyses and that these added an important contribution to the discussion of the past evolution of central Brazilian ecological spaces .In addition, we believe that the results contribute valuable information on the pattern of bio- diversity in the region and that this type of information is essential in the planning of conservation areas. MATERIALS AND METHODS Selection of floristic checklists One of the main purposes of this study was to assess the patterns of floristic differen- tiation of central Brazilian forests, both among the many forest types of the region and in relation to the forests of neighbouring vegetation provinces. To that end, a total of 106 floristic checklists were obtained from literature and other sources avail- able in the Royal Botanic Gardens of Edinburgh and Kew in 1992-94. These were selected from floristic surveys of the woody flora of forests of eastern tropical South America, with special emphasis on central Brazilian forests. To ensure accuracy, only surveys with at least 85% of the taxa identified to species level were used. Unfortunately, standardization by area size proved impossible as this varied widely. However, 'regional lists' (those for areas of thousands of hectares) were included only for non-central Brazilian forests, which were incorporated in the dataset as paradigms. The 106 floristic checklists comprised only areas which could be classified physiognomically as forests or dense woodlands, i.e. predominantly arboreal, although sometimes with an incomplete canopy. Therefore, the array of vegetation types ranged from dense Amazonian rainforests to cerradoes (forested savannas). The 106 forest areas chosen extended from the state of Roraima, in northern Brazil, to Chaco and Santa Fe Provinces, in northeastern Argentina, and from the state of Pernambuco, in eastern Brazil, to Pando Province, in northern Bolivia. Their location is shown in Fig. 2 while Table 1 gives summarized information for each area. Classification of the vegetation types The environmental information available for the 106 forest areas was either very poor or hardly comparable due to the miscellaneous methods used (especially for ORIGIN OF CENTRAL BRAZILIAN FORESTS 147 FIG. 2. Map of Brazil and neighbouring countries showing the location of the 106 forest areas used in the analyses. The forest areas are identified by their number in Table 1, followed by the two-letter code corresponding to their classification according to the modified IBGE's classification system for Brazilian vegetation (see text, pp. 154-5). soil data). However, an environmental background was necessary to make sense of the results emerging from the multivariate analyses. So, we decided to apply to the forest areas a vegetation classification system highly dependent on both physiognomy and environmental conditions and to assess the classification categories in the multi- variate analyses. The choice of a single well-defined classification system which suited our needs also allowed skirting the byzantine discussion about the best nomenclature for Brazilian vegetation. We chose the IBGE classification system for Brazilian vegetation (Veloso et al., 148 A. T. OLIVEIRA-FILHO & J. A. RATTER TABLE 1. Vegetation formation, locality, geographical coordinates, altitude, climatic type and source for each of the 106 forest areas used in the analyses. The two-letter codes after the identification numbers correspond to the classification of the forest areas according to the modified IBGE classification system for Brazilian vegetation (see text, pp. 154-5), which is also given in full in the following column. The double capitals after the localities are the official abbreviations of Brazilian states. Latitudes are South unless indicated. Altitudes are medians where a range was given in the source. An HS after the reference indicates that herbarium specimens were also used in the preparation of the checklist. Ident. Vegetation formation no. (modified IBGE system) Locality 1 Gf Gallery forest Serra do Roncador (MT) 2 As Submontane open rainforest Serra do Roncador (MT) 3Sd Dystrophic cerradao Serra do Roncador (MT) 4Cs Submontane deciduous forest Serra do Roncador (MT) 5 Sm Mesotrophic cerradao Serra do Roncador (MT) 6Gf Gallery forest Rio Suia-Missu (MT) 7Cs Submontane deciduous forest Vale dos Sonhos (MT) 8Cs Submontane deciduous forest Torixoreu (MT) 9 Sm Mesotrophic cerradao Torixoreu (MT) 10 Sd Dystrophic cerradao Chapada dos Guimaraes (MT) 11 Gf Gallery forest Chapada dos Guimaraes (MT) 12 Sm Mesotrophic cerradao Rio Claro, Cuiaba (MT) 13 Sm Mesotrophic cerradao Pocone, Pantanal (MT) 14 Sm Mesotrophic cerradao Faz. Acurizal, Pantanal (MT) 15 Cb Lowland deciduous forest Faz. Acurizal, Pantanal (MT) 16 Fa Alluvial semideciduous forest Faz. Acurizal, Pantanal (MT) 17 Cb Lowland deciduous forest Pocone, Pantanal (MT) 18 Fa Alluvial semideciduous forest Pocone, Pantanal (MT) 19 Sm Mesotrophic cerradao Faz. Nhumirim, Pantanal (MS) 20 Cb Lowland deciduous forest Faz. Nhumirim, Pantanal (MS) 21 Cb Lowland deciduous forest Rio Abobral, Pantanal (MS) 22 Cb Lowland deciduous forest Faz. Salina, Pantanal (MS) 23 Sm Mesotrophic cerradao Faz. Salina, Pantanal (MS) 24 Cs Submontane deciduous forest Corumba (MS) 25 Cb Lowland deciduous forest Porto Murtinho (MS) 26 Fa Alluvial semideciduous forest Rio Ivinheima (MS) 27 Sd Dystrophic cerradao Ilha do Bananal (TO) 28 As Submontane open rainforest Ilha do Bananal (TO) 29 Da Alluvial dense rainforest Ilha do Bananal (TO) 30 Cs Submontane deciduous forest Ilha do Bananal (TO) 31 Sm Mesotrophic cerradao Barrolandia (TO) 32 Sm Mesotrophic cerradao Figueiropolis (TO) 33 Cm Montane deciduous forest Padre Bernardo (GO) 34 Gf Gallery forest Silvania (GO) 35 Cm Montane deciduous forest FERCAL(DF) ORIGIN OF CENTRAL BRAZILIAN FORESTS 149 TABLE 1. (cont.) Lat. Long. Alt. Climate S* W (m) (Koppen) Source 12°49' 51°46' 380 Aw/Am Ratter et al. 1973, HS 12°49' 51°46' 400 Aw/Am Ratter et al. 1973, HS 12°49' 51°46' 400 Aw/Am Ratter et al. 1973 12°49' 51°46' 400 Aw/Am Ratter et al. 1973, 1977, 1978b 12°49' 51°46' 400 Aw/Am Ratter et al. 1973, 1977 12°46' 52°03' 350 Aw/Am Ratter et al. 1978a, HS 15°00' 54°30' 450 Aw Ratter et al. 1977, 1978b 15°53' 52°15' 490 Aw Furley et al. 1988, Ratter et al. 1978b 15°53' 52° 15' 490 Aw Furley et al. 1988 15°21' 55°49' 350 Aw Oliveira-Filho & Martins 1986, Oliveira-Filho et al. 1990 15°21' 55°49' 350 Aw Oliveira-Filho 1989, Oliveira-Filho et al. 1990 15°20' 55°51' 270 Aw Oliveira-Filho & Martins 1986, 1991 16°25' 56°50' 95 Aw Cunha 1990, Ratter et al. 1988 17°45' 57°37' 85 Aw Prance & Schaller 1982 17°45' 57°37' 85 Aw Prance & Schaller 1982 17°45' 57°37' 85 Aw Prance & Schaller 1982 17°05' 56°57' 85 Aw Cunha 1990 17°10' 56°57' 85 Aw Cunha 1990 18°59' 56°39' 89 Aw Ratter et al. 1988 18°59' 56°39' 89 Aw Ratter et al. 1988 19°45' 56°45' 95 Aw Cunha et al. 1986, Ratter et al. 1988, HS 19°30' 56°10' 95 Aw Dubs 1992 19°30' 56°10' 95 Aw Dubs 1992 19°01' 57°39' 120 Aw/Bs Ratter et al. 1988, Prado et al. 1992, HS 21°15' 57°40' 100 Aw/Bs Prado et al. 1992 22°47' 53°32' 250 Cw Assis 1991 10°25' 50°20' 240 Aw/Am Ratter 1987 10°25' 50°20' 240 Aw/Am Ratter 1987 10°25' 50°20' 240 Aw/Am Ratter 1987 10°25' 50°20' 240 Aw/Am Ratter 1987 10°05' 48°55' 250 Aw Ratter et al., unpubl. data 12°04' 49°10' 400 Aw Ratter et al., unpubl. data 15°15' 48°30' 650 Aw Ratter et al. 1977, 1978b 16°40' 48°37' 1050 Aw Felfili 1990, 1993b, HS 15°40' 47°56' 1100 Aw Ramos 1989 150 A. T. OLIVEIRA-FILHO & J. A. RATTER TABLE 1. (cont.) Went. Vegetation formation no. (modified IBGE system) Locality 36 Gf Gallery forest PN Brasilia (DF) 37 Gf Gallery forest APA Gama-C. do Veado (DF) 38 Sd Dystrophic cerradao Faz. Agua Limpa (DF) 39 Fm Montane semideciduous forest 40 Gf Gallery forest Brasilia Botanic Garden (DF) 41 Cs Submontane deciduous forest APA Rio S. Bartolomeu (DF) 42 Cm Montane deciduous forest Pandeiros, Januaria (MG) 43 Sm Mesotrophic cerradao Sagarana (MG) 44 Gf Gallery forest Sagarana (MG) 45 Gf Gallery forest Paracatu (MG) 46 Gf Gallery forest Patrocinio (MG) 47 Gf Gallery forest EE Panga, Uberlandia (MG) 48 Fm Montane semideciduous forest PN Serra da Canastra (MG) 49 Fm Montane semideciduous forest ESAL, Lavras (MG) RB Poco Bonito, Lavras (MG) 50 Fm Montane semideciduous forest Bom Sucesso (MG) 51 Fm Montane semideciduous forest Camargos, Itutinga (MG) 52 Fm Montane semideciduous forest Madre de Deus de Minas (MG) 53 Fm Montane semideciduous forest 54 Ds Submontane dense rainforest Serra S. Jose, Tiradentes (MG) 55 Db Lowland dense rainforest RF Rio Doce (MG) 56 Db Lowland dense rainforest Southern Bahia (BA) 57 Fs Submontane semideciduous forest RF CVRD, Linhares (ES) 58 Fm Montane semideciduous forest Anhembi (SP) 59 Fs Submontane semideciduous forest PM Grota Funda, Atibaia (SP) 60 Fs Submontane semideciduous forest RE Bauru (SP) APA Corumbatai, Ipeiina (SP) 61 Fs Submontane semideciduous forest UNESP, Jaboticabal (SP) 62 Fm Montane semideciduous forest Serra do Japi, Jundiai (SP) 63 Fs Submontane semideciduous forest RB Mogi Guacu (SP) 64 Fs Submontane semideciduous forest RE Porto Ferreira (SP) 65 Fs Submontane semideciduous forest Rio Claro (SP) 66 Fs Submontane semideciduous forest PE S. Rita Passa Quatro (SP) 67 Fs Submontane semideciduous forest PE Teodoro Sampaio (SP) 68 Fm Montane semideciduous forest FR S. Jose dos Campos (SP) 69 Db Lowland dense rainforest EE Ubatuba (SP) 70 Db Lowland dense rainforest PE Ilha do Cardoso (SP)