On-line version ISSN 1806-9959
Braz. J. Bot. vol.35 no.2 São Paulo 2012
ECOLOGY AND BIOGEOGRAPHY
Pedro Higuchi*; Ana Carolina da Silva; Tiago Sousa Ferreira; Sheila Trierveiler de Souza; Juliano Pereira Gomes; Karina Montibeller da Silva; Kristiana Fiorentin do Santos
Universidade do Estado de Santa Catarina (UDESC), Centro de Ciências Agroveterinárias, Departamento de Engenharia Florestal, Av. Luiz de Camões, 2090, Bairro Conta Dinheiro, 88520-000 Lages, SC, Brazil
The present study examined the floristic composition of three fragments of Araucaria Forest (AF) in the Planalto Catarinense region of southern Brazil as well as the floristic contextualization of these areas in relation to other remnant AF sites. Three AF fragments at different altitudes were analyzed in the municipalities of Campos Novos, Lages, and Painel. Fifty 200 m2 plots were examined in each fragment and all of the trees with CBH (circumference at breast height) > 15.7 cm were identified. In order to floristically contextualize the study fragments, comparisons were made with other remnant AF sites by way of dendrograms and NMDS (Non-metric multidimensional scaling). Environmental and spatial variables were plotted on the diagram produced by the NMDS to evaluate their influence on the floristic patterns encountered. The forest fragments studied demonstrated high floristic heterogeneity, indicating that AFs cannot be considered homogeneous formations and they could be classified into 3 phytogeographical categories: i) high altitude areas influenced by cloud cover/fog, including the Painel region; ii) areas of lesser altitude and greater mean annual temperatures situated in the Paraná River basin, and iii) areas situated in the Paraná and Upper-Uruguay river basins and the smaller basins draining directly into the southern Atlantic, near Campos Novos and Lages. The environmental variables most highly correlated with species substitutions among the sites were altitude, mean annual temperature, and the mean temperature of the most humid trimester.
Key words: Araucaria Forest, arboreal species, environmental variables
Mixed Ombrophilous Forests (MOF) (IBGE 1992), also known as Araucaria Forests (AF), represent one of the principal plant formations in southern Brazil. These forests predominate on the Brazilian Meridional Plateau, principally on the southern Catarinense Plateau, between 24º and 30º S latitude at altitudes that vary between 500 and 1400 m above sea level, with disjunct occupation of sites in elevated areas of the Serra do Mar and Mantiqueira Mountains in southeastern Brazil (Klein 1960). Their largest extensions cover the hydrographic basins of the rivers that drain into the Uruguay and Paraná rivers, following the altitudinal gradients from the higher lands of the Serra do Mar and Geral Mountains towards the interior (west). AFs, like most other Brazilian forests, are currently highly fragmented due to historical anthropogenic use and occupation of the land for agriculture, cattle raising, and (more recently) large-scale monoculture plantations of Pinus spp. These activities have reduced these once extensive and continuous forests to isolated fragments with sizes and levels of disturbance that vary from one locality to another (Klein 1978, Behling & Pillar 2007).
Araucaria angustifolia is the most characteristic species of the AF phytophysiognomy as it forms the upper canopy and has a dominant character in this vegetation. In spite of the fact that A. angustifolia predominates in the upper forest canopy (Leite & Klein 1990), the middle and lower strata demonstrate large diversities of species such as Ocotea porosa (Nees & Mart.) Barroso (imbuia), Luehea divaricata Mart. (açoita-cavalo), Cedrela fissilis Vell. (cedro), Dicksonia sellowiana (xaxim), and Maytenus ilicifolia Mart. ex Reissek (espinheira-santa) (Nascimento et al. 2001, Behling & Pillar 2007).
In light of the significant environmental heterogeneity in this region, associated principally with the climate, edaphic factors, and the history of anthropogenic disturbances, it would be expected that different AF fragments would show large floristic variations. According to Klein (1978), Santa Catarina State can be divided into four distinct phytophysiognomic areas: i) Araucaria Forests in the Iguaçu-Negro basin and in the upper regions of the river basins that drain into the Uruguay River; ii) Araucaria Forests of the Pelotas-Canoas basin; iii) Araucaria Forests of the extreme west; and iv) Araucaria Forest Nuclei of the Atlantic Rainforest. According to the classification system suggested by IBGE (1992), this phytophysiognomy may be subdivided according to its altitude and its proximity to water courses into Alluvial, Sub-Montane, Montane, and High-Montane formations.
The Catarinense Plateau is considered a strategic region for forest conservation as it contains numerous headwaters that form the hydrographic basin of the Uruguay River (Sgrott 2003) and is also part of the recharging area of the Guarani aquifer (Gomes et al. 2006). Little is currently known about the floristic and phytogeographical patterns of the arboreal components of AF forest fragments (Klein 1978, Formento et al. 2004) or their interactions with environmental and spatial variables. As such, the present study was designed to: i) describe the floristic composition of three AF fragments in the Catarinense Plateau region of southern Brazil; ii) determine the floristic similarities between these fragments in relation to other AF areas in southern Brazil; and iii) evaluate the influence of environmental and spatial variables on the phytogeographical patterns encountered.
MATERIAL AND METHODS
Three AF fragments located at different altitudes in the southern Catarinense Plateau were investigated. These fragments were located in the municipalities of Campos Novos (27º35'09.69" S and 51º09'15.07" W, altitude 712 m, area = 1,551.37 ha), Lages (27º51'19.20" S and 50º10'33.39" W, altitude 993 m, area = 103.06 ha), and Painel (27º55'27.56" S and 49º56'30.14" W, altitude 1,196 m, area = 48.91 ha) (figure 1). According to the WORDCLIM databank (Hijmans et al. 2005), the mean annual precipitation levels in these fragment areas (from 1960-1990) were 1719, 1483, and 1569 mm in Campos Novos, Lages, and Painel respectively, with rainfall being well-distributed throughout the year. The mean annual temperatures (1960-1990) in these fragments were 17.1, 16.1, and 15 ºC respectively. The climates in all three study areas were classified as Cfb, following the classification system of Köppen, being characterized by rainfall uniformly distributed throughout the year and without a dry season. The fragments studied were areas within the Canoas and Pelotas River hydrological basins, and their topography is, for the most part, slightly undulating to undulating. According to the vegetation classification system of IBGE (1992), the arboreal vegetation of the Catarinense Plateau region can be classified as Montane and High-Montane Mixed Ombrophilous Forest. The fragments examined had different histories of use and disturbance, but like most of the AF remnants in the state, all of them had experienced intense harvesting of both lumber and non-lumber species during the 1960's.
The surveys of the floristic compositions of the arboreal components of the sites involved establishing 50 permanent plots (10 × 20 m = 200 m2 each), totaling 1 ha, to sample each fragment area. The permanent plots were distributed (with 30 m spacing between them) along parallel transects through the forests. The spatial arrangements of these transects were designed to adequately sample the environmental variables present in the areas, such as edge × interior and other variations related to soil types and altitudinal and topographic gradients. As such, the numbers and lengths of the transects varied among the fragments as a function of the unique characteristics of each, taking into consideration their spatial parameters (e.g. shape and total area) and environmental heterogeneity (e.g., topography). All of the arboreal individuals in each plot with CBH (circumference at breast height, measured 1.30 m above soil level) > 15.7 cm were identified and marked with aluminum tags. Plant identifications were made by comparing them with herbarium samples and through consultations with specialist and the use of specialized literature. The angiosperm species were classified into families and genera according to the APG III system (2009). The botanical material collected was incorporated into the Santa Catarina State University herbarium (LUSC).
In order to phytogeographically contextualize the study localities, previous scientific publications of floristic lists and surveys of arboreal species of remnant Araucaria forests in southern Brazil were consulted (table 1). The information concerning each species was subject to a detailed revision with respect to habitat, nomenclature, and taxonomic synonyms (which were verified on the W3 Tropicos site) (Mobot 2010). The analyses did not include plants identified only to the family or genus level.
To analyze the relationships between the floristic compositions of the areas in terms of their environmental and spatial variables, we incorporated data concerning the altitude, climate (1960-1990), and geographical coordinates of each area. The geographic coordinates of the areas were based on the publications cited earlier. Altitudinal and climatic data were obtained from the WORLDCLIM databank (Hijmans et al. 2005), including: altitude (alt), Annual mean temperature (b1), Mean diurnal range (Mean of monthly (max temp - min temp)) (b2), Isothermality (b2/ b7) (x100) (b3), Temperature seasonality (standard deviation x100) (b4), Max temperature of warmest month (b5), Min temperature of coldest month (b6), Temperature annual range (b5-b6) (b7), Mean temperature of wettest quarter (b8), Mean temperature of driest quarter (b9), Mean temperature of warmest quarter (b10), Mean temperature of coldest quarter (b11), Annual precipitation (b12), Precipitation of wettest month (b13), Precipitation of driest month (b14), Precipitation seasonality (coefficient of variation) (b15), Precipitation of wettest quarter (b16), Precipitation of driest quarter (b17), Precipitation of warmest quarter (b18), and Precipitation of coldest quarter (b19).
Both the numbers of shared and exclusive species at the Campos Novos, Lages, and Painel sites were demonstrated using a Venn diagram. In order to evaluate the floristic relationships of these areas with other remnant AF sites in southern Brazil, a dendrogram displaying the hierarchical clustering of these areas was prepared utilizing the Sorensen coefficient as a measure of distance, and the UPGMA linking method as the cluster algorithm.
In order to determine the influence of environmental and spatial variables on the floristic patterns encountered, we employed NMDS (Non-metric multidimensional scaling) multivariate analysis with two dimensions (k = 2) (Minchin 1987), and utilized the Sorensen coefficient as a measure of similarity. Although a common practice, we did not exclude rare species from these analyses as they have little influence in NMDS (Muotka et al. 2002). The adequacy of the ordination for interpretation was evaluated using a "stress" value. The variations explained by the ordination axes were calculated from nonlinear adjustments between the distances of the area as defined in the ordination as well as the original dissimilarity in values between the areas (Oksanen 2011). The significance of the "stress" value encountered was evaluated according to the methodology proposed by McCune & Grace (2002) using the Monte Carlo test with 100 permutations. The environmental and spatial variables were adjusted a posteriori and inserted in the form of vectors into the ordination produced by NMDS, using 1000 permutations and excluding the redundant variables or those with low significance in the analysis (P < 0.001).
All of the analyses were made using the R statistical program (R Development Core Team 2010). The Limma package (Gentleman et al. 2004) was used to create the Venn diagram and the Vegan (Oksanen et al. 2009) and Cluster (Maechler et al. 2011) packages were used for multivariate analyses.
Species composition and richness
A total of 147 species, 93 genera and 53 plant families were encountered in the three study fragments (table 2). Eight species were identified only to the family level (four species in Campos Novos, three species in Lages, and one species in Painel) and four only to the genus level (one species each in Campos Novos and Lages and two species in Painel); eight taxa were not identified at all due to the impossibility of making collections (two in Campos Novos, three in Lages, and three in Painel).
The families that most contributed to species richness in the three remnant areas evaluated were Myrtaceae (26 species), Fabaceae (12), Asteraceae (8), Lauraceae (8), Salicaceae (7), Solanaceae (7), and Aquifoliaceae (6). Together, these families comprised 50.68% of the total number of species identified. The richest genera were Solanum (7 species), Myrcia (7), Ilex (6), Eugenia (5), Myrceugenia (5), Maytenus (4), and Ocotea (4). Myrtaceae stood out as the family with the greatest species richness in all the three areas studied. The families that most contributed to arboreal floristic richness in Campos Novos were Myrtaceae (9 species), Fabaceae (8), Salicaceae (6), Euphorbiaceae (4), Lauraceae (4), Meliaceae (4), Rutaceae (4), and Sapindaceae (4); the species distributions among these different genera were quite homogeneous, although Myrsine stood out with three species. The families that most contributed to species richness in Lages were Myrtaceae (18 species), Lauraceae (6), Aquifoliaceae (5), Salicaceae (5), Anacardiaceae (4), Asteraceae (4), and Sapindaceae (4); the most important genera were Myrcia (5 species), Ilex (5), Eugenia (4), Myrceugenia (3), Ocotea (3), and Schinus (3). The families that most contributed to species richness in Painel were Myrtaceae (11 species), Solanaceae (7), Asteraceae (6), Aquifoliaceae (4), and Lauraceae (3); the richest genera were Solanum (7 species), Myrceugenia (5), and Ilex (4).
The species richnesses values in the remnant forest areas at Campos Novos, Lages, and Painel were 87, 91 and 49 respectively (figure 2). Only a low percentage of species were shared among all three sample areas (10.88%). The site with the greatest proportion of exclusive species was Campos Novos (45.98%), while Painel had the lowest (26.53%); the latter site shared 67.35% of its species with Lages. Of the total number species encountered in Lages, 48.35% were shared with Campos Novos, 36.26% with Painel, while 32.97% were exclusive.
Examinations of the AF fragments in southern Brazil indicated the existence of three large floristic groups (figure 3). Group 1 was composed of high-montane areas in the states of Santa Catarina and Rio Grande do Sul; Group 2 was formed by forest remnants in Paraná State in the Paraná River basin (Arapoti, Tibagi, and Irati); and Group 3 was formed by a large number of remnant areas distributed among three states located in the hydrographic basins of the Paraná and Uruguay rivers and those rivers that drain directly into the southern Atlantic. The forest remnant located in the municipality of Painel was placed in Group 1 and was the most floristically distinct but showed greater floristic similarity with Bom Jardim da Serra, SC. The fragment area in Lages (Group 3) demonstrated greater floristic similarity with another fragment from the same municipality, and both of them demonstrated high similarity with Bom Jesus, Vacaria, Caxias do Sul, Campos Novos, and São Francisco de Paula - all of which are situated in the Upper-Uruguay region.
The "stress" value in the NMDS ordination analysis (figure 4) was 15.78% (P < 0.01), which corresponds to an explained variance of 97.51% - indicating that the diagram was adequate for our interpretations. The floristic groups defined in the dendrogram are indicated in the NMDS diagram by continuous lines. The variables that demonstrated elevated significance (P < 0.001) (table 3) within the ordination are represented by vectors. It can be seen that the variables that best explained the separation of the three groups were altitude (alt), Annual mean temperature (b1), and Mean temperature of driest quarter (b9). The extremes of this gradient are represented by the areas in Group 1 (with the highest and coldest sites) and by the areas of Group 2 (with the lowest altitude and warmest sites). Internally within Group 3 (which had the largest area), the greatest species substitution occurred along an environmental gradient associated with latitude (lat), Precipitation seasonality (coefficient of variation) (b15), and Precipitation of driest month (b14).
The numbers of species encountered in each fragment in the present study were indicative of low richness (in the case of Painel) as well as elevated richness (in the case of Lages and Campos Novos) in comparison with other studies undertaken in AF fragments in southern Brazil with values that varied between 134 in Arapoti and Palmeira (PR) to 26 in Clevelândia (PR) (see table 1). Myrtaceae, the plant family that most contributed to species richness in the three study fragments, had already been cited by Mori et al. (1983) as the richest family in different Atlantic Forest formations. Additionally, a number of studies undertaken in southern Brazil have confirmed important contributions of Myrtaceae to species richness (e.g. Jurinitz & Jarenkow 2003).
The low percentages of species shared among the three fragments studied indicated that AFs are not floristically homogeneous formations. The ordination of the study areas and their relationships with the spatial and environmental variables examined demonstrated that the remnant AF areas have floristic variations that are associated with their distributions from West to East and North and South, reflecting the overall environmental heterogeneity of the region. It is worth noting that aspects related to historical perturbations in these areas also influence tree species distributions (even though this factor was not evaluated the present study) (Silva et al. 2009).
To the east, near the coast, there are important mountain chains formed by the Serra do Mar and Serra Geral mountains that constitute the highest elevations in southern Brazil. Heavy fogs frequently occur in these localities as well as snap freezes and frequent snowfalls, principally in the Serra Catarinense and Serra Gaúcha regions. This is where the areas forming floristic Group 1 are found, including the present study fragment in the municipality of Painel. A number of species encountered in this fragment were exclusive to Lages and Campos Novos, principally those belonging to the genera Ilex, Baccharis, Symphyopappus, Myrceugenia, and Solanum that contain high-altitude indicator species. The genera Drimys and Clethra which Webster (1995) identified as typical of Neotropical cloud-forest areas also occur in this fragment. The low richness observed in Painel might be a result of the high rigor and selectivity of the climatic conditions in that area.
Land elevation gradually diminishes westward as the principal watercourses in the region drain towards the Paraná and Uruguay rivers. The AF accompany the principle hydrographic basins forming these rivers, from the highest elevations to altitudes near 500 m (where the climate becomes warmer and there are stronger influences of typical components of seasonal forests). The area near Campos Novos, along the Canoas River (which together with the Pelotas River forms the Uruguay River), is an example of this situation, with a significant proportion of its species being exclusive in relation to those seen at Lages and Painel, including: Schaefferia argentinensis, Cordia americana, Myrocarpus frondosus, and Parapiptadenia rigida - all typical of Seasonal Deciduous Forests of the Upper-Uruguay river basin.
The influence of altitudinal gradients on the floristic patterns of neotropical forests has been demonstrated by a number of authors (e.g. Oliveira Filho et al. 1994, 2005, Oliveira Filho & Fontes 2000, Scudeller et al. 2001) and was shown to be (together with temperature) one of the principal variables that influenced species distributions in the three area studied here. As was noted by Oliveira Filho & Fontes (2000), various environmental changes occur along this gradient - such as temperature variations and the occurrence of freezing events - which can explain the strong changes observed in species distributions. Considering the three areas evaluated in the present study, Campos Novos is positioned at one of the extremes of this gradient at an altitude of 700 m and Painel at the other (1196 m). Both of these areas share large numbers of species with the Lages site (993 m), which is geographically situated between the two extremes and can be considered a transition area.
Passing through states of Santa Catarina to Rio Grande do Sul from north to south, there are two large hydrographic basins that together form the Platina Basin (the Paraná and Uruguay river basins). Even though this region does not experience a well-defined dry season, our results demonstrated an environmental gradient between these two basins associated principally with the precipitation regime (b14 and b15) (see figure 3). These results corroborate observations made by Klein (1978), who classified the AFs in Santa Catarina State in terms of their hydrographic basins. Authors such as Silva et al. (2006) likewise observed the tendency of areas that occur in the same hydrological basin to cluster in terms of their species compositions. One of the factors that could help explain the observed floristic compartmentalization in terms of hydrological basins would be the influence of riparian forests that function as dispersal corridors for arboreal species. This concept of tree species migration has been examined by a number of authors using different methodologies (e.g. Rambo 1951, Budke et al. 2004).
The Araucaria forests analyzed here could be classified into three distinct phytogeographical units: i) high altitude areas with lower mean annual temperatures and influences of cloud/fog cover (where the Painel site was established); ii) lower altitude areas with greater mean annual temperatures, situated in the Paraná River basin; iii) areas situated in the Paraná and Upper-Uruguay river basins as well as the smaller river basins that drain directly into the southern Atlantic (the Campos Novos and Lages sites). The environmental variables most closely correlated with the high levels of species substitutions among three sites were altitude, mean annual temperature, and mean temperature of driest quarter.
Acknowledgments - The authors thank CNPq for financial support through Edital MCT/CNPq 15/2007, which allowed us to undertake this study; the Brochmann Polis - Industrial and Florestal S/A company for allowing us access to their property at Campos Novos; and Tio Deja and Sr. Jorge Gamborgi for graciously allowing us access to their properties in Painel and Lages respectively.
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(received: March 27, 2011; accepted: April 17, 2012)