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Acta Botanica Brasilica

Print version ISSN 0102-3306

Acta Bot. Bras. vol.27 no.2 Feira de Santana Apr./June 2013 



Community dynamics in a species-rich patch of old-growth forest in a global changing scenario



Markus GastauerI,II; João Augusto Alves Meira NetoI,II,*

IUniversidade Federal de Viçosa, Departamento de Biologia Vegetal, Laboratório de Ecologia e Evolução das Plantas, Viçosa, MG, Brazil
IICentro de Ciências Ambientais Floresta-Escola, Frutal, MG, Brazil




Ecological theory predicts that, in mature ecosystems, species richness, the number of individuals and the biomass of individuals will remain in a relatively stable state of equilibrium. The aim of this study was to test that theory. In 2001 and 2010, we conducted censuses of all trees with a circumference at breast height > 10 cm in a one-hectare plot in a seasonal semideciduous old-growth forest in southeastern Brazil. We compared the two censuses in terms of species richness and diversity, computing growth, recruitment and mortality rates, as well as gains and losses of basal area. Between 2001 and 2010, species richness declined from 224 to 218 species and the basal area increased from 37.86 to 40.16 m2 ha-1. Overall turnover (the mean difference between mortality and recruitment) was lower than would be expected for a seasonal semideciduous forest, indicating stability and slight successional advance. This interpretation is supported by the observation that pioneer species and canopy species both showed higher mortality than recruitment. However, uncommon species (< 10 individuals in the 2001 census) showed higher mortality than recruitment and became rarer, whereas most species that were abundant in 2001 became more abundant by 2010. These observations, as well as the decline in species richness, although statistically not significant, match the predictions of ecological theory for scenarios in which formerly contiguous ecosystems become fragmented and the remnants become isolated within the landscape. Nevertheless, further censuses are needed in order to test the idea that the observed patterns are not explained by natural oscillations but are consequences of environmental changes related to human activity.

Key words: Community dynamics, habitat loss, immigration rate, landscape fragmentation, species richness.




Climate change (IPCC 2007; Matesanz et al. 2010), habitat destruction and fragmentation (Wright 2010) threaten species richness and diversity worldwide. Because tropical forests account for the major part of terrestrial biodiversity (Millennium Ecosystem Assessment 2005), climate change is expected to have severe effects on such forests. To outline and understand the influences of global changes on forest communities, their species richness and diversity, it is necessary to conduct long term monitoring studies known as community dynamics studies (Losos & Leigh Jr. 2004).

Species composition within a given community undoubtedly varies over time (Rosenzweig 1995; Hubbell 2001; Magurran 2011). These variations depend on the disturbance regime (Connell 1978; Molino & Sabatier 2001; Machado & Oliveira-Filho 2010), ecological drift and stochasticity (Hubbell 2001).

The division of formerly contiguous forests into fragments of variable size and shape also alters the ecological processes of the natural dynamics of the forest (Tabarelli et al. 2004). In addition, changes in resource availability influence growth, mortality and recruitment of trees within forest communities (Ernest et al. 2009).

Among tropical forests, the Atlantic Forest of Brazil is one of the most diverse ecosystems (Stehmann et al. 2009). Due to its high degree of endemism and endangered status, it is considered a biodiversity hotspot (Myers et al. 2000). Once covering up to 1,500,000 km2 (Câmara 2005), only 11% of the original Atlantic Forest remains, most of it as small secondary forest patches (Ribeiro et al. 2009). Species-rich old-growth forests are extremely rare.

The aim of this study is to test whether the mature tree community of the Seu Nico Forest (SNF) shows stability in the form of dynamic equilibrium over time, in terms of community dynamics, such as mortality and recruitment rates (Oliveira-Filho et al. 2007); species richness (MacArthur & Wilson 1967); the number of individuals or zero-sum dynamics (Hubbell 2001); and basal area per individual (Ernest et al. 2009).



Study site

The SNF is a 35 ha patch of species-rich old-growth forest (20º47'44"S; 42º50'50"W) on the Bom Sucesso Farm, in the town of Viçosa, which is in the state of Minas Gerais, Brazil. The owners of the SNF state that it has never been logged (Campos et al. 2006). According to the Köppen climate classification system (Köppen 1948), the climate of the region is type Cwb (Peel et al. 2007), a humid mesothermal climate with mild, rainy summers and dry winters. The predominant soils are deeply weathered oxisols, which are found within a small-scale mosaic of inceptisols on slopes and neosols in sedimentation areas of valley bottoms. The predominant vegetation is characterized as submontane seasonal semideciduous forest (Veloso et al. 1991).

Data collection

Within the SNF, a 100 × 100 m (1-ha) area was marked off and divided into 100 plots of 10 × 10 m. In an initial census, conducted for all of the plots between October 2000 and March 2001 (Irsigler 2001), all trees with a circumference at breast height > 10 cm-corresponding to a diameter (DBH) > 3.2 cm-were tagged and identified. Because it is conventional to use diameter rather than circumference in the international literature, we will hereafter refer to the DBH. During a second census, carried out between December 2009 and February 2010, we measured the circumference of all surviving individuals within the 100 plots. Recruits (individuals that had not been tagged but met the inclusion criteria) were also tagged and identified. Specimens were collected, identified and deposited in the Herbarium of the Federal University of Viçosa (code, VIC). Species nomenclature follows the database compiled by Forzza et al. (2012). Species classification follows the Angiosperm Phylogeny Group III guidelines (APG III 2009).

Data analysis

To compare species richness between the two censuses, we calculated the species-area relationship using the power model proposed by Arrhenius (1921) to fit the species-accumulation curve:

where S is the number of species; A is the area; and c and z are constants corresponding to the intercept with the y-axis (c) and the slope (z) of the linearized species-area relationship after log transformation of S and A. For the fitting, we assessed the average number of species within groups of 2, 5, 10, 25 and 50 plots, as well as within the study site as a whole (all 100 plots). Only neighboring plots were grouped.

Linearized in logarithmic space, the slope of the species-area relationship describes the difference between plots and is therefore interpreted as spatial turnover, or beta diversity (Condit et al. 1996). For statistical comparison of the values, we used Microsoft ExcelTM to perform linear least squares regression.

For both censuses, the biodiversity indices of Simpson and Shannon-Wiener, as well as Fisher's alpha, were computed with the software EstimateS (Colwell & Coddington 1994; Colwell 2005). Statistical differences in diversity as well as in species richness, basal area and number of individuals were checked for significance by a two-sample t-test after testing for normal distribution by a Shapiro-Wilk test with STATISTCA software, version 7.0 (StatSoft Inc., Tulsa, OK, USA). Parameters were calculated for each plot.

The community dynamics were calculated as proposed by Losos & Lao (2004). To calculate the average growth rate between censuses, the increase in diameter for each tree was divided by the interval (in years) between the two censuses. The average growth rate is the average of the growth rates of all trees meeting the inclusion criteria.

The difference in the mortality rate of trees between the censuses is the natural logarithm of the proportional relationship between the number of trees tagged in the first census and that of those surviving to the second census, divided by the time between the two:

where MR is the mortality rate, NS is the number of survivors (trees that were alive in both censuses), N1 is the NS minus the number of undetected individuals or individuals that died between the two censuses, and T is the average interval between censuses (in years). The rate of tree recruitment was calculated as follows:

where RR is the recruitment rate, and NR is the number of new trees appearing between censuses (i.e., recruits).

For any given plot, a loss in basal area (defined as the area below the 3.2 cm DBH level) resulted from mortality and from stem breakages. To calculate the loss in basal area between the two censuses (the total basal area of trees tagged in the first census that had died or whose basal area had otherwise been lost by the time of the second census), we employed the following equation:

where BAloss is the total basal area lost, and ABM is the difference in basal area between the two censuses. Similarly, a gain in basal area was calculated as follows:

where BAgain is the total basal area gained, BAS2 is the total basal area of the trees surviving from the first to the second census, and BAS1 is the total basal area of those same individuals in the first census. For the calculation of demographic dynamics, all stems of multi-stemmed individuals were included.

We calculated demographic dynamics not only for the community as a whole but also for different size classes, as proposed by Losos & Lao (2004): class I (DBH = 3.2-9.9 cm); class II (DBH = 10-29.9 cm); and class III (DBH > 30 cm).

Species were classified according to their regeneration, stratification and dispersal strategies. Community dynamics were calculated separately for all pioneer and all non-pioneer species following the classification of Swaine & Whitmore (1988): for understory species, which typically do not reach heights above 15 m; for all canopy species, which typically reach heights above 15 m (Liebsch et al. 2008); and for animal-dispersed and non-animal-dispersed species (Ingle 2003). Information about successional, dispersal and dispersal strategies was drawn from the following studies (in alphabetical order, not sorted by relevance): Appolinário et al. (2005), Aquino & Barbosa (2009), Araújo et al. (2005, 2006), Araujo et al. (2010), Brandão et al. (2009), Cappelatti & Schmitt (2009), Carvalho et al. (2006, 2007), Carvalho & Nascimento (2009), Chagas et al. (2001), Colonetti et al. (2009), Higuchi et al. (2008a), Leite & Rodrigues (2008), Lemos (2008), Lima et al. (2010), Lopes et al. (2002), Marangon et al. (2007), Metzger et al. (1997), Norden et al. (2009), Nunes et al. (2003), Oliveira-Filho et al. (2004, 2007), Paula et al. (2004), Peixoto et al. (2004), Pinto et al. (2005), Rolim et al. (1999), Silva et al. (2004), Stranghetti et al. (2003), and Yamamoto et al. (2007). In cases of contradictory information, species were allocated to the strategy indicated in the greatest number of references. Community dynamics were calculated separately for common species (< 9 individuals) and uncommon species (> 10 individuals).



Plot census history

Overall, the number of individuals increased in our plot within the SNF. However, of the 224 species surveyed in 2001, only 214 remained in 2010. Both changes were less than significant (p=0.89 and p=0.81 for the number of individuals and number of species, respectively). Within the community examined, only four new species were recruited between the two censuses (Table 1). Although this net loss of six species reduced the z slope of the species-area relationship (Fig. 1), the differences were not significant (z=0.545 ± 0.032 in 2001 and z=0.531 ± 0.030 in 2010). The various diversity indices indicated an insignificant loss of tree diversity within the plot (Table 2). The basal area increased (from 37.86 m2 ha-1 in 2001 to 40.16 m2 ha-1 in 2010), albeit insignificantly (p=0.44).





Community dynamics

Between the first and second censuses, the number of individuals increased because recruitment exceeded mortality (Table 3). In terms of basal area, gains surpassed losses, resulting in a net gain from 2001 to 2010. Among the common species, mortality and recruitment rates were nearly identical. However, among the uncommon species, the mortality rate was higher than was the recruitment rate. In comparison with that observed for the community as a whole, mortality was disproportionately high among the pioneer and non-animal dispersed species. Among the understory species, recruitment exceeded mortality.

Population dynamics

Of the nine species that were the most abundant in 2001, eight increased in abundance, only Bathysa nicholsonii being less represented in 2010 than in 2001 (Table 4). Of the 20 species with the highest basal areas, 10 increased their basal area by more than 10%. In five species, there was a moderate to high increase in basal area, which decreased in another five species.

Species ranks from both censuses showed an s-shaped curve (Fig. 2). Due to lower species richness and higher abundance of common species in 2010, the curve trended downward, although the slope decreased only slightly from 2001 to 2010 (from -0.0089 ± 0.0009 to -0.0083 ± 0.0008).



Mortality rates were high for some of the uncommon species, such as Vernonanthura diffusa (15.4% yr-1) and Astronium fraxinifolium (13.1% yr-1), whose populations declined. The populations of other species increased considerably, with above-average recruitment rates. Examples are Siparuna reginae (23.1% yr-1) or Eugenia florida (13.4% yr-1). Among the most abundant species, recruitment rates were above average for Siparuna guianensis (3.1% yr-1), Euterpe edulis (4.1% yr-1), Virola gardneri (3.3% yr-1) and Eugenia cf. lambertiana (3.9% yr-1), whereas mortality rates were above average for Bathysa nicholsonii (2.6% yr-1), S. guianensis (2.4% yr-1), Pourouma guianensis and (2.5% yr-1) and Casearia ulmifolia (2.4% yr-1).



Within the 1 ha plot of the SNF evaluated, we obtained low values for the key features of community dynamics, such as recruitment, mortality and growth rates, as well as losses and gains of basal area (Leigh et al. 2004; Thompson et al. 2004). Within the Atlantic Forest and the Cerrado domains, mortality and recruitment rates are generally higher for flooded gallery forests (Lopes & Schiavini 2007; Higuchi et al. 2008b; Fontes & Walter 2011) than for non-flooded gallery forests (Pinto 2002; Oliveira-Filho & Felfili 2008), for seasonal deciduous forests (Carvalho & Felfili, 2011), for seasonal semideciduous forests (Apollinario et al. 2005; Oliveira-Filho et al. 1997, 2007; Silva & Araújo 2009; Machado & Oliveira-Filho 2010) and for evergreen forests (Rolim et al. 1999; Saiter et al. 2011).

The mortality and recruitment rates reported in the literature are higher than those observed for our study site, indicating that there was high stability in the SNF. This stability is congruent with the findings reported by Saiter et al. (2011) for other old-growth, mature forests.

For the SNF plot evaluated, size class was not found to correlate with mortality or recruitment. In disturbed forests, mortality is typically higher among the smaller size classes usually because of the effect known as self-thinning (Oliveira-Filho et al. 2007; Saiter et al. 2011). However, the observations that mortality exceeded recruitment among pioneer species and recruitment exceeded mortality among animal-dispersed species, as well as among understory species, indicates a slight successional advance of the SNF as a mature forest, making it even more representative of an old-growth forest (Liebsch et al. 2008).

Turnover rates were high only for some of the uncommon species (data not shown). Due to a low number of individuals, even small alterations within these populations result in high rates. Among the twenty most abundant species, turnover rates were above average for only a few, including S. guianensis, E. edulis, V. gardneri, and E. cf. lambertiana, which are non-pioneer, animal-dispersed species, the occurrence of which indicates forest maturation (Liebsch et al. 2008). Several of the most abundant species showed high mortality rates, the highest being for P. guianensis. Because P. guianensis is a pioneer species, a decline in its population is also indicative of the ongoing successional change in the SNF.

Although the recruitment and mortality rates, as well as the gains and losses of basal area, indicate stability and slight successional advance for the tree community of the SNF, our comparison of the two censuses revealed some tendencies that merit discussion. As species richness and diversity declined, common species became more common, while rare species became rarer. In addition, the number of individuals and the total basal area both increased. The fact that those tendencies were not significant might be due to the small sample size. However, it is also possible that they were stochastic, describing only the natural oscillations within the dynamic equilibrium of the seasonal semideciduous Atlantic Forest (Rees et al. 2001; Lopes & Schiavini 2007; Paiva et al. 2007). Nevertheless, an increase in the number of individuals contradicts the zero-sum assumptions made by Hubbell in a theoretical work (2001). In addition, because the amount of biomass supported within an ecosystem depends on energy and resource inputs (Ernest et al. 2009), the increase in basal area might be explained by increased resource availability (Lewis et al. 2004), perhaps due to climate change (IPCC 2007). This is alarming, because the same tendencies have recently been observed in other regions of the Atlantic Forest (Higuchi et al. 2008b; Carvalho & Felfili 2011; Saiter et al. 2011).

Regarding the decline in species richness, Felfili et al. (2000) stressed the difficulty in interpreting what they referred to as "pseudoextinction", because species might still be present within the community in form of seeds, seedlings or treelets that have not yet met the inclusion criterion. However, if a net loss of six species represents a natural oscillation of the tree community, a net gain of six species in nine years, perhaps in the next census or in similar studies, could be expected. We are not aware of any studies reporting such net gains in primary or late secondary forests; on the contrary, most dynamic studies conducted to date have also reported declining numbers of species (Oliveira-Filho et al. 1997; Bunyavejchewin et al. 2004; Lee et al. 2004; Leigh et al. 2004; Thompson et al. 2004; Werneck & Franceschinelli 2004; Higuchi et al. 2008a, 2008b; Machado & Oliveira-Filho 2010).

Because these observations have been made within a landscape that was logged recently (about 150 years ago) and is now fragmented, they should be discussed from a metacommunity perspective. According to the species-area relationship, one of the most widely studied patterns in ecology (Tjørve 2003; Martin & Goldenfeld 2006; Colwell et al. 2012), habitat area loss causes species loss (Pimm & Raven 2000; Ney-Nifle & Mangel 2000; Fischer 2000; He & Hubbell 2011). However, decades of research on species-area relationships have also revealed that species richness is lower on islands than on contiguous land masses of the same area (Rosenzweig 1995; Lomolino 2001). Because landscape fragmentation might be interpreted as the transformation of former contiguous habitats into small patches, forming continental islands, theory predicts that a secondary, delayed loss of species should occur after landscape fragmentation (Rosenzweig 1995).

According to the theory of biogeography, species richness at a given site is achieved by maintaining equilibrium among the rates of immigration, speciation (Hubbell 2001) and extinction (MacArthur & Wilson 1967). In the present study more species disappeared from the community than were recruited. Therefore, we might assume that there was a disturbance of this equilibrium, indicating problems with the immigration of propagules of new species to the SNF. Against this conclusion stands the observation that recruitment rates were higher for animal-dispersed species than for non-animal-dispersed species (Table 3), and that the four new species were animal-dispersed. This indicates that an adapted fauna is actually dispersing seeds and fruits within the fragment and should guarantee genetic exchange between the SNF and neighboring fragments. However, as research activities in the last decades have shown, these secondary fragments are less species-rich (Lopes et al. 2002; Meira-Neto & Martins 2002; Paula et al. 2004; Ribas et al. 2004; Ferreira Jr. et al. 2007). This reduces the richness of arriving seeds and therefore the immigration rate of propagules from new species. Nevertheless, natural extinction, in our case measured as pseudoextinction (Felfili et al. 2000; Carvalho & Felfili 2011), continues unchanged, which causes ongoing species loss. In this scenario, species loss in the SNF will continue until a new equilibrium is reached between extinction and reduced immigration rate. These theoretical explications of the species loss observed in the SNF are congruent with the findings of Hubbell (2001), who ran simulations of the distribution of species abundance in local communities. In isolated stands (communities with reduced immigration rates), common species become more common and rare species become rarer, as was observed in the SNF plot evaluated here. This is a vicious cycle: due to isolation, common species become more common in the local community but also increase their abundance in the metacommunity. Because abundance in the local community, even in isolation, still depends on species abundance in the metacommunity, this commonness of already common species further increases metacommunity abundance. Increased mortality of uncommon species supports the categorization of the species loss observed in the SNF as time-delayed species loss after landscape fragmentation.

As shown above, our tree community dynamics study of the SNF indicated stability and low dynamics in this unique patch of primary forest. The slight, insignificant decline in species richness and the increase in basal area might be due to natural oscillations around an equilibrium value but might also be interpreted as consequences of environmental changes that are currently affecting vegetation and will do so into the future. Future censuses in the same SNF plot showing similar tendencies will support our hypothesis that the changes observed in this, one of the last old-growth forest remnants in the region, are due to human activity worldwide.



We are grateful to David Teixeira Irsigler for carrying out the first census in the SNF. This study received financial support from Suzano Pulp and Paper in the form of a doctoral scholarship grant to M.G. Authors also thank CNPq, FAPEMIG, MCTI and SECTES-MG for financial support. J.A.A.M.N. has research scholarship of CNPq.



APG (Angiosperm Phylogeny Group) III. 2009. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III. Botanical Journal of the Linnean Society 161:105-121.         [ Links ]

Appolinário, V.; Oliveira Filho, A.T. & Guilherme F.A.G. 2005. Tree population and community dynamics in a Brazilian tropical semideciduous forest. Revista Brasileira de Botânica 28(2):347-360.         [ Links ]

Aquino, C. & Barbosa, L.M. 2009. Classes sucessionais e síndromes de dispersão de espécies arbóreas e arbustivas existentes em vegetação ciliar remanescente (Conchal, SP), como subsídio para avaliar o potencial do fragmento como fonte de propágulos para enriquecimento de áreas revegetadas no Rio Mogi-Guaçu, SP. Revista Árvore 33(2):349-358.         [ Links ]

Araújo, F.S.; Martins, S.V.; Meira Neto, J.A.A.; Lani, J.L. & Pires, I.E. 2005. Florística da vegetação arbustivo-arbórea colonizadora de uma área degradada por mineração de caulim, em Brás Pires, MG. Revista Árvore 29(6):983-992.         [ Links ]

Araújo, F.S.; Martins, S.V.; Meira Neto, J.A.A.; Lani, J.L. & Pires, I.E. 2006. Estrutura da vegetação arbustivo-arbórea colonizadora de uma área degradada por mineração de caulim, em Brás Pires, MG. Revista Árvore 30(1):107-116.         [ Links ]

Araujo, M.M.; Chami, L.; Longhi, S.J.; Avila, A.L. & Brena, D.A. 2010. Análise de agrupamento em remanescente de Floresta Ombrófila Mista. Ciência Florestal 20(1):1-18.         [ Links ]

Arrhenius, O. 1921. Species and area. Journal of Ecology 9:95-99.         [ Links ]

Brandão, C.F.L.S.; Marangon, L.C.; Ferreira, R.L.C.; Lins & Silva, A.C.B. 2009. Estrutura fitossociológica e classificação sucessional do componente arbóreo em um fragmento de floresta atlântica em Igarassu - Pernambuco. Revista Brasileira de Ciências Agrárias 4(1):55-61.         [ Links ]

Bunyavejchewin, S.; Baker, P.J.; LaFrankie, J.V. & Ashton, P.S. 2004. Huai Kha Khaeng Forest Dynamics Plot, Thailand. Pp. 482-491. In: Losos, E.C. & Leigh Jr., E.G. (Eds.). Tropical Forest Diversity and Dynamism - Findings from a Large-Scale Plot Network. London, The University of Chicago Press.         [ Links ]

Câmara, I.G. 2005. Breve história da conservação da Mata Atlântica. Pp. 31-42. In: Galindo-Leal, C. & Câmara, I.G. (Eds.). Mata Atlântica: biodiversidade, ameaças e perspectivas. São Paulo/Belo Horizonte, Fundação SOS Mata Atlântica/Conservação Internacional.         [ Links ]

Campos, E.P.; Silva, A.F.; Meira Neto, J.A.A. & Martins, S.V. 2006. Florística e estrutura horizontal da vegetação arbórea de uma ravina em um fragmento florestal no município de Viçosa, MG. Revista Árvore 30:1045-1054.         [ Links ]

Cappelatti, L. & Schmitt, J.L. 2009. Caracterização da flora arbórea de um fragmento urbano de Floresta Estacional Semidecidual no Rio Grande do Sul, Brasil. Pesquisas, Botânica 60:341-354.         [ Links ]

Carvalho, F.A.; Nascimento, M.T. & Braga, J.M.A. 2006. Composição e riqueza florística da Floresta Atlântica submontana na região de Imbaú, Município de Silva Jardim, RJ. Acta Botanica Brasilica 20(3):727-740.         [ Links ]

Carvalho, F.A.; Nascimento, M.T. & Braga, J.M.A. 2007. Estrutura e composição florística do estrato arbóreo de um remanescente de Mata Atlântica submontana no município de Rio Bonito, RJ, Brasil (Mata Rio Vermelho). Revista Árvore 31(4):717-730.         [ Links ]

Carvalho, F.A. & Felfili, J.M. 2011. Variações temporais na comunidade arbórea de uma floresta decidual sobre afl oramentos calcários no Brasil Central: composição, estrutura e diversidade florística. Acta Botanica Brasilica 25(1):203-214.         [ Links ]

Carvalho, F.A. & Nascimento, M.T. 2009. Estrutura diamétrica da comunidade e das principais populações arbóreas de um remanescente de Floresta Atlântica Submontana (Silva Jardim-RJ, Brasil). Revista Árvore 33(2):327-337.         [ Links ]

Chagas, R.K.; Oliveira-Filho, A.T.; van den Berg, E. & Scolforo, J.R.S. 2001. Dinâmica de populações arbóreas em um fragmento de Floresta Estacional Semidecidual Montana em Lavras, Minas Gerais. Revista Árvore 25(1):39-57.         [ Links ]

Colonetti, S.; Citadini-Zanette, V.; Martins, R.; Santos, R.; Rocha, E. & Jarenkow, J.A. 2009. Florística e estrutura fitossociológica em floresta ombrófila densa submontana na barragem do rio São Bento, Siderópolis, Estado de Santa Catarina. Maringá 31(4):387-405.         [ Links ]

Condit, R.; Hubbell, S.P.; LaFrankie, J.V.; Sukumar, R.; Manokaran, N.; Foster, R.B. & Ashton, P.S. 1996. Species-Area and Species-Individual Relationships for Tropical Trees: A Comparison of Three 50-ha Plots. Journal of Ecology 84:549-562.         [ Links ]

Connell, J.H. 1978. Diversity in tropical rain forests and coral reefs. Science 199:1302-1310.         [ Links ]

Colwell, R.K. & Coddington, J.A. 1994. Estimating terrestrial biodiversity through extrapolation. Philosophical Transactions of the Royal Society London B 345:101-108.         [ Links ]

Colwell, R.K. 2005. EstimateS: Statistical estimation of species richness and shared species from samples. Version 8.2. User's Guide and application published at:, retrieved October 2012.         [ Links ]

Colwell, R.K.; Chao, A.; Gotelli, N.J.; Lin, S.-Y.; Mao, C.X.; Chazdon, L.R. & Longino, J.T. 2012. Models and estimators linking individual-based and sample-based rarefaction, extrapolation, and comparison of assemblages. Journal of Plant Ecology 5:3-21.         [ Links ]

Ernest, K.M.; White, E.P. & Brown, J.H. 2009. Changes in a tropical forest support metaboliczero-sum dynamics. Ecology Letters 12:507-515.         [ Links ]

Felfili, J.M.; Rezende, A.V.; Silva-Junior, M.C. & Silva, M.A. 2000. Changes in fl oristic composition of cerrado sensu stricto in Brazil over a nine year period. Journal of Tropical Ecology 16:579-590.         [ Links ]

Fischer, M. 2000. Species loss after habitat fragmentation. Tree 15:396.         [ Links ]

Ferreira Jr., W.G.; Silva, A.F.; Meira Neto, J.A.A.; Schaefer, C.E.G.R.; Dias, A.S.; Ignácio, M. & Medeiros, M.C.M.P. 2007. Composição florística da vegetação arórea de um trecho de Floresta Estacional Semidecídua em Viçosa, Minas Gerais, e espécies de maior ocorrência na região. Revista Árvore 31(6):1121-1130.         [ Links ]

Fontes, C.G. & Walter, B.M.T. 2011. Dinâmica do componente arbóreo de uma mata de galeria inundável (Brasília, Distrito Federal) em um período de oito anos. Revista Brasileira de Botânica 34(2):145-158.         [ Links ]

Forzza, R.C. (coord.) 2012. Lista de Espécies da Flora do Brasil. Online access at, retrieved October 2012.         [ Links ]

He, F. & Hubbell, S.P. 2011. Species-area relationships always overestimate extinction rates from habitat loss. Nature 473:368-371.         [ Links ]

Higuchi, P.; Oliveiro-Filho, A.T.; Silva, A.C.; Machado, E.L.M.; Santos, R.M. & Pifano, D.S. 2008a. Dinâmica da comunidade arbórea em um fragmento de floresta estacional semidecidual montana em lavras, minas gerais, em diferentes classes de solos. Revista Árvore 32:417-426.         [ Links ]

Higuchi, P.; Oliveiro-Filho, A.T.; Bebber, D.P.; Brown, N.D.; Silva, A.C. & Machado, E.L.M. 2008b. Spatio-temporal patterns of tree community dynamics in a tropical forest fragment in South-east Brazil. Plant Ecology 199:125-135.         [ Links ]

Hubbell, S.P. 2001. The Unified Neutral Theory of Biodiversity and Biogeography. Princeton/Oxford, Princeton University Press.         [ Links ]

Ingle, N.R. 2003. Seed dispersal by wind, birds, and bats between Philippine montane rainforest and sucessional vegetation. Oecologia 134(2):251-261.         [ Links ]

IPCC (Intergovernmental Panel on Climate Change) 2007. Climate Change 2007 - The Physical Science Basis. Contribution of Working Group I to the Forth Assessent Report of the IPCC. Cambridge, Cambridge University Press.         [ Links ]

Irsigler, D. T. 2001. Composição florística e estrutura de um trecho primitivo de Floresta Estacional Semidecídua em Viçosa, MG. Master thesis, Universidade Federal de Viçosa.         [ Links ]

Lee, H.S.; Tan, S.; Davies, S.J.; LaFrankie, J.V.; Ashton, P.S.; Yamakura, T.; Itoh, A.; Ohkubo, T. & Harrison, R. 2004. Lambir Forest Dynamics Plot, Sarawak, Malaysia. Pp. 527-539. In: Losos, E.C. & Leigh Jr., E.G. (Eds.). Tropical Forest Diversity and Dynamism - Findings from a Large-Scale Plot Network. London, University of Chicago Press.         [ Links ]

Leigh Jr., E.G.; Lao, S.L.; Condit, R.; Hubbell, S.P.; Foster, R.B. & Pérez, R. 2004. Barro Colorado Island Forest Dynamics Plot, Panama. Pp. 451-463. In: Losos, E.C. & Leigh Jr., E.G. (Eds.). Tropical Forest Diversity and Dynamism - Findings from a Large-Scale Plot Network. London, University of Chicago Press.         [ Links ]

Leite, E.C. & Rodrigues, R.R. 2008. Fitossociologia e caracterização sucessional de um fragmento de Floresta Estacional no suedeste do Brasil. Revista Árvore 32(3):583-595.         [ Links ]

Lemos, P.H.D. 2008. Efeito de borda no componente arbórea de um fragmento de floresta semidecídua, Viçosa, MG. Master thesis, Universidade Federal de Viçosa, Brasil.         [ Links ]

Lewis, S.L.; Phillips, O.L.; Baker, T.R. et al. 2004. Concerted changes in tropical forest structure and dynamics: evidence from 50 South American long-term plots. Philosophical Transactions of the Royal Society London B 359:421-436.         [ Links ]

Liebsch, D.; Marques, M.C.M. & Goldenberg, R. 2008. How long does the Atlantic Rain Forest take to recover after a disturbance? Changes in species composition and ecological features during secondary succession. Biological Conservation 141:1717-1725.         [ Links ]

Lima, A.L.S.; Zanella, F. & Castro, L.D.M. 2010. Crescimento de Hymenaea courbaril L. var. stilbocarpa (Hayne) Lee et Lang. e Enterolobium contortisiliquum (Vell.) Morong (Leguminosae) sob diferentes níveis de sombreamento. Acta Amazonica 40(1):43-48.         [ Links ]

Lomolino, M.V. 2001. The species-area relationship: new challenges for an old pattern. Progress in Physical Geography 25:1-21.         [ Links ]

Lopes, W.P.; Paula, A.; Sevilha, A.C. & Silva, A.F. 2002. Composição da flora arbórea de um trecho de Floresta Estacional no Jardim Botânica da Universidade Federal de Viçosa (Face Sudoeste), Viçosa, Minas Gerais. Revista Árvore 26(3):339-347.         [ Links ]

Lopes, S.F. & Schiavini, I. 2007. Dinâmica da comunidade arbórea de mata de galeria da Estação Ecológica do Panga, Minas Gerais, Brasil. Acta Botanica Brasilica 21:249-261.         [ Links ]

Losos, E.C. & Leigh Jr., E.G. 2004. The Growth of a Tree Plot Network. Pp 3-7. In: Losos, E.C. & Leigh Jr., E.G. (Eds.). Tropical Forest Diversity and Dynamism - Findings from a Large-Scale Plot Network. London, The University of Chicago Press.         [ Links ]

Losos, E.C. & Lao, S.L. 2004. Forest Dynamics Plots. Introduction. Pp 433-450. In: Losos, E.C. & Leigh Jr., E.G. (Eds.). Tropical Forest Diversity and Dynamism - Findings from a Large-Scale Plot Network. London, The University of Chicago Press.         [ Links ]

MacArthur, R. & Wilson, E.O. 1967. The Theory of Island Biogeography. Princeton, Princeton University Press.         [ Links ]

Machado, E.L.M. & Oliveira-Filho, A.T. 2010. Spatial patterns of tree community dynamics are detectable in a small (4 ha) and disturbed fragment of the Brazilian Atlantic forest. Acta Botanica Brasilica 24:250-261.         [ Links ]

Magurran, A.E. 2011. Measuring biological diversity in time (and space). Pp. 85-94. In: Magurran, A.E. & McGill, B.J. Biological Diversity - frontiers in measurement and assessment. New York, Oxford University Press.         [ Links ]

Marangon, L.C.; Soares, J.J.; Feliciano, A.L.P.; Lins, C.F. & Brandão, S. 2007. Estrutura fitossociológica e classificação sucessional do componente arbóreo de um fragmento de Floresta Estacional Semidecidual, no município de Viçosa, Minas Gerais. Cerne, Lavras 13(2):208-221.         [ Links ]

Martin, H.G. & Goldenfield, N. 2006. On the origin and robustness of power-law species-area relationships in ecology. Proceedings of the National Academy of Sciences 103:10310-10315.         [ Links ]

Matesanz, S., Escudero, A. & Valladares, F. 2009. Impact on three global change drivers on a Mediterranean shrub. Ecology 90:2609-2621.         [ Links ]

Meira Neto, J.A.A. & Martins, F.R. 2002. Composição florística de uma Floresta Estacional Semidecidual Montana no município de Viçosa-MG. Revista Árvore 26(4):437-446.         [ Links ]

Metzger, J.P.; Bernacci, L.C. & Goldenberg, R. 1997. Pattern of tree species diversity in riparian forest fragments of different widths (SE Brazil). Plant Ecology 133:135-152.         [ Links ]

Millennium Ecosystem Assessment. 2005. Biodiversity: What is it, where is it, and whyis it important? Pp 18-29. In: Ecosystems and Human Well-Being: Biodiversity Synthesis. World. Washington DC, Resources Institute.         [ Links ]

Molino, J.F. & Sabatier, D. 2001. Tree Diversity in Tropical Rain Forests: A Validation of the Intermediate Disturbance Hypothesis. Science 294:1702-1704.         [ Links ]

Myers, N.; Mittermeier, R.A.; Mittermeier, C.G.; Fonseca, G.A.B. & Kent, J. 2000. Biodiversity hotspots for conservation priorities. Nature 403:853-858.         [ Links ]

Ney-Nifle, M. & Mangel, M. 2000. Habitat loss and changes in the species-area relationship. Conservation Biology 14:893-898.         [ Links ]

Norden, N.; Chave, J.; Belbenoit, P.; Caubère, A.; Châtelet, P.; Forget, P.-M.; Piéra, B.; Viers, J. & Thébaud, C. 2009. Interspecific variation in seedling responses to seed limitation and habitat conditions for 14 Neotropical woody species. Journal of Ecology 97:186-197.         [ Links ]

Nunes, Y.R.F.; Mendonça, A.V.R.; Botezelli, L.; Machado, E.L.M. & Oliveira-Filho, A.T. 2003. Variações da fisionomia, diversidade e composição de guildas da comunidade arbórea em um fragmento de floresta semidecidual em Lavras, MG. Acta Botanica Brasilica 17(2):213-229.         [ Links ]

Oliveira-Filho, A.T.; Mello, J.M. & Scolforo, J.R.S. 1997. Effects of past disturbances and edges on tree community strucutre and dynamics within a fragment of tropical semideciduous forest in south eastern Brazil over a five-year period (1987-1992). Plant Ecology 131:45-66.         [ Links ]

Oliveira-Filho, A.T.; Carvalho, D.A.; Vilela, E.A.; Curi, N. & Fontes, M.A.L. 2004. Diversity and structure of the tree community of a fragment of tropical secondary forest of the Brazilian Atlantic Forest domain 15 and 40 years after logging. Revista Brasileira de Botânica 27(4):685-701.         [ Links ]

Oliveira-Filho, A.T.; Carvalho, W.A.C.; Machado, E.L.M.; Higuchi, P., Appolinário, V.; Castro, G.C.; Silva, A.C.; Santos, R.M.; Borges, L.F., Corrêa, B.S. & Alves, J.M. 2007. Dinâmica da comunidade e populações arbóreas da borda e interior de um remanescente florestal na Serra da Mantiqueira, Minas Gerais, em um intervalo de cinco anos (1999-2004). Revista Brasileira de Botanica 30:149-161.         [ Links ]

Oliveira, A.P. & Felfi li, J.M. 2008. Dinâmica da comunidade arbórea de uma mata de galeria do Brasil Central em um período de 19 anos (1985-2004). Revista Brasileira de Botânica 31:597-610.         [ Links ]

Paiva, L.V.; Araújo, G.M. & Pedroni, F. 2007. Structure and dynamics of a woody plant community of a tropical semi-deciduous seasonal forest. Revista Brasileira de Botanica 30:365-373.         [ Links ]

Paula, A.; Silva, A.F.; Marco Jr., P.; Santos, F.A.M. & Souza, A.L. 2004. Sucessão ecológica da vegetação arbóres em uma Floresta Estacional Semidecidual, Viçosa, MG, Brasil. Acta Botanica Brasilica 18(3):407-423.         [ Links ]

Peel, M.C.; Finlayson, B.L. & McMahon, T.A. 2007. Updated world map of the Köppen-Geiger climate classification. Hydrology and Earth System Science 11:1633-1644.         [ Links ]

Peixoto, G.L.; Martins, S.V.; Silva, A.F. & Silva, S. 2004. Composição florística do componente arbórea de um tracho de Floresta Atlântica na Área de Proteção Ambiental da Serra da Capoeira Grande, Rio de Janeiro, RJ, Brasil. Acta Botanica Brasilica 18(1):151-160.         [ Links ]

Pimm, S.L. & Raven, P. 2000. Extinction by numbers. Nature 403:483-483.         [ Links ]

Pinto, J.R.R. 2002. Dinâmica da comunidade arbóreo-arbustiva em uma floresta de vale no Parque Nacional da Chapada dos Guimarães, Mato Grosso. Ph.D. thesis, Universidade de Brasília, Brasília.         [ Links ]

Pinto, L.V.A.; Botelho, S.A.; Oliveira-Filho, A.T. & Davide, A.C. 2005. Estudo da vegetação como subsídio para propostas de recuperação das nascentes da Bacia hidrográfica do Riberão Santa Cruz, Lavras, MG. Revista Árvore 29(5):775-793.         [ Links ]

Rees, M.; Condit, R.; Crawley, M.; Pacala, S. & Tilman, D. 2001. Long-term studies of vegetation dynamics. Science 293:650-658.         [ Links ]

Ribas, R.F.; Meira Neto, J.A.A.; Silva, A.F. & Souza, A.L. 2004. Composição florística da dois trechos em diferentes etapas serais de uma Floresta Estacional Semidecidual em Viçosa, Minas Gerais. Revista Árvore 27(6):821-830.         [ Links ]

Ribeiro, M.C.; Metzger, J.P.; Martensen, A.C.; Ponzoni, F.J. & Hirota, M.M. 2009. The Brazilian Atlantic Forest: How much is left, and how is the remaining forest distributed? Implications for conservation. Biological Conservation 142:1141-1153.         [ Links ]

Rolim, S.G.; Couto, H.T.Z. & Jesus, R.M. 1999. Mortalidade e recrutamento de árvores na Floresta Atlântica em Linhares (ES). Scientia forestalis 55:49-69.         [ Links ]

Rosenzweig, M.L. 1995. Species diversity in space and time. Cambridge, Cambridge University Press.         [ Links ]

Saiter, F.Z.; Guilherme, F.A.G.; Thomaz, L.D. & Wendt, T. 2011. Tree changes in a mature rainforest with high diversity and endemism on the Brazilian coast. Biodiversity and Conservation 20:1921-1949.         [ Links ]

Silva, C.T.; Reis, G.C.; Reis, M.G.F.; Silva, E. & Chaves, R.A. 2004. Avaliação temporal da florística arbórea de uma Floresta Secundária no município de Viçosa, Minas Gerais. Revista Árvore 28(3):429-441.         [ Links ]

Silva, M.R. & Araújo, G.M. 2009. Dinâmica da comunidade arbórea de uma floresta semidecidual em Uberlândia, MG, Brasil. Acta Botanica Brasilica 23:49-56.         [ Links ]

Stehmann, J.R.; Forzza, R.C.; Salino, A.; Sobral, M.; Costa, D.P. & Kamino, L.H.Y. (org.) 2009. Plantas da Floresta Atlântica. Rio de Janeiro, Instituto de Pesquisas Jardim Botânico do Rio de Janeiro.         [ Links ]

Stranghetti, V.; Ituralde, R.B.; Gimenez, L.R. & Almella, D. 2003. Florística de um fragmento florestal do sítio São Pedro, município de Potirendaba, Estado de São Paulo. Maringá 25(1):167-172.         [ Links ]

Swaine, M.D. & Whitmore, T.C. 1988. On the definition of ecological species groups in tropical rain forests. Vegetatio 75:81-86.         [ Links ]

Tabarelli, M.; Silva, J.M.C. & Gascon, C. 2004. Forest fragmentation, synergisms and the impoverishment of neotropical forests. Biodiversity and Conservation 13:1419-1425.         [ Links ]

Thompson, J.; Brokaw, N.; Zimmermann, J.K.; Waide, R.B.; Everham III, E.M. & Schaefer, D.A. 2004. Luquillo Forest Dynamics Plot, Puerto Rico, United States. Pp 540-550. In: Losos, E.C. & Leigh Jr., E.G. (Eds.) Tropical Forest Diversity and Dynamism - Findings from a Large-Scale Plot Network. London, University of Chicago Press.         [ Links ]

Tjørve, E. 2003. Shapes and functions of species-area curves: a review of possible models. Journal of Biogeography 30:827-835.         [ Links ]

Veloso, H.P.; Rangel Filho, A.L.R. & Lima, J.C.A. 1991. Classificação da Vegetação Brasileira, adaptada a um Sistema Universal. Rio de Janeiro, Departamento de Recursos Naturais e Estudos Ambientais do Intituto Brasileiro de Geografia e Estatística - IBGE.         [ Links ]

Werneck, M.S. & Franceschinelli, E.V. 2004. Dynamics of a dry forest fragment after the exclusion of human disturbance in southeastern Brazil. Plant Ecology 174:337-346.         [ Links ]

Wright, S.J. 2010. The future of tropical forests. Annals of the New York Academy of Sciences 1195:1-27.         [ Links ]

Yamamoto, L.F.; Kinoshita, L.S. & Martins, F.R. 2007. Síndromes de polinização e de dispersão em fragmentos da Floresta Estacional Semidecídua Montana, SP, Brasil. Acta Botanica Brasilica 21(3):553-573.         [ Links ]



Submitted: 26 April, 2012
Accepted: 16 December, 2012



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