**5. Beta diversity of Atlantic Forest formations**

124 The Dynamical Processes of Biodiversity – Case Studies of Evolution and Spatial Distribution

Table 5. (Continued)

**Alluvial\*** 

According to the Venn diagrams (Figure 4), the two Rainforest types share at least 99 tree species, most of them very common in the Montane formations of both forest types, like *Cinnamodendrum dinisii, Ocotea porosa, Drimys brasiliensis* and *Ocotea odorifera*. The surveys carried out in the Dense Rainforest shared 94 species with the Semideciduous Forest, which in turn shared 88 species with the Araucaria Rainforest. At least 50 arboreal species (8% of the species measured) occur in the three Atlantic Forest types, such as *Campomanesia xanthocarpa, Casearia sylvestris, Alchornea triplinervia, Nectandra megapotamica, Sloanea guianensis, Cupania vernalis, Casearia decandra, Syagrus romanzoffiana, Blepharocalyx salicifolius, Myrsine umbellata, Ocotea pulchella* and *Ilex paraguariensis*, among the most abundant.

A total of 272 species were found exclusively in the Dense Rainforest, e.g. *Tibouchina trichopoda*, *Andira anthelmia*, *Handroanthus umbellatus*, *Tabebuia cassinoides*, *Clusia criuva*, *Pera glabrata*, *Ternstroemia brasiliensis*, *Virola bicuhyba*, *Marlierea tomentosa*, *Cupania oblongifolia*, *Protium kleinii*, *Quiina glazovii*, *Aspidosperma pyriccolum*, *Myrcia freyreissiana*, *Siphoneugena reitzii*, *Drimys angustifolia* and *Handroanthus catarinensis*.

At least 51 tree species were found only in Araucaria Rainforest surveys, such as *Podocarpus lambertii, Nectandra grandiflora, Lithraea brasiliensis, Guettarda uruguensis, Curitiba prismatica, Symplocos celastrinea, Myrrhinium atropurpureum, Myrcianthes pungens, Ocotea nutans, Erythroxylum deciduum, Cinnamomum amoenum and Zanthoxylum kleinii.* 

Among the 127 species found only in the Semideciduous Seasonal Forest (Figure 4), the most abundant are *Chrysophyllum gonocarpum, Metrodorea nigra, Astronium graveolens, Parapiptadenia rigida, Plinia rivularis, Casearia gossypiosperma, Triplaris americana, Balfourodendron riedelianum, Peltophorum dubium, Holocalyx balansae, Aspidosperma polyneuron and Gallesia integrifolia.* 

The dendrogram using Sorensen's similarity coefficients for tree species (Figure 5) shows two major distinct groups. The first one comprises the five Dense Rainforest formations, and the second comprises the Araucaria Rainforest and the Semideciduous Seasonal Forest, reinforcing the data given in Venn diagram (Figure 4). These two types of Atlantic Forest

ATLANTIC FORESTS

Fig. 4. Venn diagrams presenting the number of measured tree species shared in the 58 sites of the three types of Atlantic Forest in the state of Paraná, Southern Brazil.

Arboreal Diversity of the Atlantic Forest

for these variables in studies in Southeastern Brazil.

and diversities.

Cumulative percentage variance:

presented in Table 1.

of Southern Brazil: From the Beach Ridges to the Paraná River 127

The dendrogram based on abundance data (Figure 6) shows some different results as compared with the dendrogram based on the Sorensen's similarity. In the first group are the two Semideciduous Forests, while in the second are all studied Rainforests. This group is divided in two subgroups: one presenting the highest tree species richness and diversities (Shannon-Wiener index) among the rainforests analyzed and other with the lowest richness

The first axis resulting from the CCA (eigenvalue = λ = 0.637) showed a gradient associated with altitude on one hand and annual temperature an rainfall on the other, separating Montane Rainforests (on the left) and Lowland and Alluvial Rainforests (on the right) (Figure 7). Oliveira-Filho & Fontes (2000) and Scudeller et al. (2001) found similar patterns

Eigenvalues (species) 0.637 0.425 0.311 0.212

 of species data 4.4 7.3 9.4 10.8 of species-environment relation 40.2 67.0 86.7 100.0 Species-environment correlations 0.899 0.860 0.826 0.212 Significance of species-environment correlation (Monte Carlo test) 0.001 0.001 0.001 0.001 Fig. 7. Biplot of a Canonical Correspondence (CCA) applied to species found in 58 sites in Atlantic Forests in the state of Paraná, Southern Brazil. Numbers are related to sites

**Axis 1 Axis 2 Axis 3 Axis 4** 

share 24.5% of the total species. The two Rainforests share 19.6% of the species and the Semideciduous Seasonal Forest share 16.2% of the species with the Dense Rainforest. These values differ from those found by Oliveira-Filho & Fontes (2000), in which Dense Rainforest and Semideciduous Forest of Southeastern Brazil shared a high proportion of tree species in their checklists: 50% and 66% respectively.

The comparatively high floristic similarity within the Southeastern region than in the Southern region, can be related to the absence of Araucaria Rainforests between these two tropical forests further north.

In the group of the Dense Rainforest, "slope forests" were separated from the "coastal and alluvial plain" forests (Figure 5). The second main group divided the Araucaria Rainforest and the Semideciduous Seasonal Forest.

Pair wise comparisons of the Sorensen's similarity coefficients show values ranging from 0.09 between the Alluvial Semideciduous Forest and the Upper Montane Dense Rainforests to 0.61 between the two formations of the Semideciduous Forest.

Fig. 5. Dendrogram showing the similarity between tree species measured in nine main formations of the Atlantic Forest of the state of Paraná, Southern Brazil. The cluster analysis was carried out using Sorensen's similarity coefficients, squared euclidean distances and the Ward's method of agglomeration (DRF – Dense Rainforest; ARF – Araucaria Rainforest; SF – Seasonal Forest; A – Alluvial; L – Lowland; S – Submontane; M – Montane; U – Upper Montane).

Fig. 6. Dendrogram clustering all Atlantic Forest sites through abundance similarities for tree species with squared euclidean distances and the Ward's method of agglomeration (DRF – Dense Rainforest; ARF – Araucaria Rainforest; SF – Seasonal Forest; A – Alluvial; L – Lowland; S – Submontane; M – Montane; U – Upper Montane).

share 24.5% of the total species. The two Rainforests share 19.6% of the species and the Semideciduous Seasonal Forest share 16.2% of the species with the Dense Rainforest. These values differ from those found by Oliveira-Filho & Fontes (2000), in which Dense Rainforest and Semideciduous Forest of Southeastern Brazil shared a high proportion of tree species in

The comparatively high floristic similarity within the Southeastern region than in the Southern region, can be related to the absence of Araucaria Rainforests between these two

In the group of the Dense Rainforest, "slope forests" were separated from the "coastal and alluvial plain" forests (Figure 5). The second main group divided the Araucaria Rainforest

Pair wise comparisons of the Sorensen's similarity coefficients show values ranging from 0.09 between the Alluvial Semideciduous Forest and the Upper Montane Dense Rainforests

their checklists: 50% and 66% respectively.

and the Semideciduous Seasonal Forest.

Distance

Montane).

Distance

0

ASF

SSF

– Lowland; S – Submontane; M – Montane; U – Upper Montane).

LDRF

UDRF

Fig. 6. Dendrogram clustering all Atlantic Forest sites through abundance similarities for tree species with squared euclidean distances and the Ward's method of agglomeration (DRF – Dense Rainforest; ARF – Araucaria Rainforest; SF – Seasonal Forest; A – Alluvial; L

ADRF

AARF

SDRF

MDRF

MARF

2 4 6

8

ADRF

LDRF

SDRF

MDRF

Fig. 5. Dendrogram showing the similarity between tree species measured in nine main formations of the Atlantic Forest of the state of Paraná, Southern Brazil. The cluster analysis was carried out using Sorensen's similarity coefficients, squared euclidean distances and the Ward's method of agglomeration (DRF – Dense Rainforest; ARF – Araucaria Rainforest; SF – Seasonal Forest; A – Alluvial; L – Lowland; S – Submontane; M – Montane; U – Upper

UDRF

AARF

MARF

ASF

SSF

to 0.61 between the two formations of the Semideciduous Forest.

tropical forests further north.

The dendrogram based on abundance data (Figure 6) shows some different results as compared with the dendrogram based on the Sorensen's similarity. In the first group are the two Semideciduous Forests, while in the second are all studied Rainforests. This group is divided in two subgroups: one presenting the highest tree species richness and diversities (Shannon-Wiener index) among the rainforests analyzed and other with the lowest richness and diversities.

The first axis resulting from the CCA (eigenvalue = λ = 0.637) showed a gradient associated with altitude on one hand and annual temperature an rainfall on the other, separating Montane Rainforests (on the left) and Lowland and Alluvial Rainforests (on the right) (Figure 7). Oliveira-Filho & Fontes (2000) and Scudeller et al. (2001) found similar patterns for these variables in studies in Southeastern Brazil.

Fig. 7. Biplot of a Canonical Correspondence (CCA) applied to species found in 58 sites in Atlantic Forests in the state of Paraná, Southern Brazil. Numbers are related to sites presented in Table 1.

Significance of species-environment correlation (Monte Carlo test) 0.001 0.001 0.001 0.001

Arboreal Diversity of the Atlantic Forest

of Southern Brazil: From the Beach Ridges to the Paraná River 129

Barddal, M.L., Roderjan, C.V., Galvão, F. & Curcio, G.R. (2004). Caracterização florística e

Bianchini, E., Silveira, R.P., Dias, M.C. & Pimenta, J.A. (2003). Diversidade e estrutura de

Blum, C.T., Santos, E.P., Hoffmann, P.M. & Socher, L.G. (2001). Análise Florística e

Blum, C.T., Silva, D.A.T., Hase, L.M. & Miranda, D.L.C. (2003). Caracterização Florística e

Blum, C.T. (2006). A Floresta Ombrófila Densa na Serra da Prata, Parque Nacional Saint-

Blum, C.T. & Roderjan, C.V. (2007). Espécies indicadoras em um gradiente da Floresta

Blum, C.T. & Petean, M.P. (2008). Flora. In: *Estudo de Impacto Ambiental da BR-487, trecho* 

Blum, C.T. (2010). Os componentes epifítico vascular e herbáceo terrícola da Floresta

Borghi, W.A., Martins, S.S., Del Quiqui, E.M. & Nanni, M.R. (2004). Caracterização e

Campos, J.B., Romagnolo, M.B. & Souza, M.C. (2000). Structure, composition and spatial

Castella, P.R., Britez, R.M. (2004). *A Floresta com Araucária no Paraná: conservação e diagnóstico* 

Caviglione, J.H., Kiihl, L.R.B., Caramori, P.H. & Oliveira, D. (2000). *Cartas climáticas do* 

Cordeiro, J. & Rodrigues, W.A. (2007). Caracterização Fitossociológica de um remanescente

Cordeiro, J. (2010). Compartimentação pedológico-ambiental e sua influência sobre a

de Floresta Ombrófila Mista em Guarapuava, PR. *Revista Árvore*, Vol. 31, No. 3, pp.

florística e estrutura de um remanescente de Floresta Ombrófila Mista na região de

*and Technology*, Vol. 43, No. 2, pp. 185-194, ISSN 1516-8913

*dos remanescentes florestais*, FUPEF/PROBIO/MMA, Brasília

www.iapar.br/modules/conteudo/conteudo.php?conteudo=677

*Paraná*, IAPAR – Instituto Agronômico do Paraná, Retrieved from:

*Cruzeiro do Oeste – Porto Camargo, PR.* ENGEMIN / DNIT, Curitiba.

Araucária, PR. *Ciência Florestal*, Vol. 14, No. 2, ISSN 0103-9954

*Botanica Brasilica*, Vol. 17, No. 3, pp. 405-419, ISSN 0102-3306

Universidade Federal do Paraná. Curitiba, Paraná, Brasil

*Paraná e Santa Catarina*, Curitiba, november 2001

*Social*, Foz do Iguaçú, november 2003

Vol. 5, No. 2, pp. 873-875, ISSN 1679-2343

Paraná. Curitiba, Paraná, Brasil

ISSN 1415-9112

545-554, ISSN 0100-6762

fitossociológica de um trecho sazonalmente inundável de floresta aluvial, em

espécies arbóreas em área alagável do município de Londrina, sul do Brasil. *Acta* 

Estrutural de um Trecho de Floresta Ombrófila Densa Montana no Morro dos Perdidos, Serra de Araçatuba, PR. *Proceedings of VI Encontro Regional de Botânicos do* 

Ecológica de Remanescentes Florestais no Rio das Cinzas, Norte Pioneiro/PR. *Proceedings of Seminário Nacional Degradação e Recuperação Ambiental - Perspectiva* 

Hilaire/Lange, PR - caracterização florística, fitossociológica e ambiental de um gradiente altitudinal. M.Sc Dissertation. Pós-graduação em Engenharia Florestal,

Ombrófila Densa na Serra da Prata, Paraná, Brasil. *Revista Brasileira de Biociências*,

Ombrófila Densa ao longo de um gradiente altitudinal na Serra da Prata, Paraná. Ph.D Thesis. Pós-graduação em Engenharia Florestal, Universidade Federal do

avaliação da mata ciliar à montante da hidrelétrica de Rosana, na Estação Ecológica do Caiuá, Diamante do Norte, PR. *Cadernos da Biodiversidade*, Vol. 4, No. 2, pp. 9-18,

distribution and dynamics of tree species in a remnant of Semideciduous Seasonal Alluvial Forest of the Upper Paraná River floodplain. *Brazilian Archives of Biology* 

The second axis (λ = 0.425) shows the Semideciduous Seasonal Forest tree species and their abundance change with increasing the distance from the ocean, and decreasing rainfall (and probably increasing rainfall seasonality). The opposite occurred with Submontane, Alluvial and Lowland Dense Rainforests.

The relatively high eigenvalues found (> 0.3), indicate considerable abundance and species turnover along gradients mainly in axis 1 and 2. As also found by Scudeller et al. (2001), the low variance explained by the first axis indicates that other variables not investigated or methodological restrictions in this study probably influenced the abundance distribution.
