**4.1 Analysis of the sampled sites in the Atlantic Forests**

Among the 58 sites, comprising 29 ha of plots and 36627 sampled individuals, selected from 39 studies, the Dense Rainforest has the highest number of sampled sites (29 out of a total of 58) and the highest number of tree individuals found (Table 2). This great amount of data available is due to the high number of Upper Montane Dense Rainforest sampled sites, which present relatively many areas with primary and well preserved vegetation. In spite of the relatively smaller sampled area, small-sized trees present in high abundance make up for a high number of individuals found.

On the other hand, the Alluvial formation of the Dense Rainforest is the least sampled one (Table 2). The lack of phytosociological studies conducted in this formation may be related to its actual small cover area (representing only 0.89% of the remnants of Dense Rain Forest) (Pires et al., 2005), and also to its level of degradation, which make the search for typical and representative remnants difficult. According to this data, the studied area is almost six times smaller than the area of the neighbor Lowland formation. Therefore, this understudied vegetation needs more studies.

The low number of studied sites in the Semideciduous Seasonal Forest in the state of Paraná is also remarkable, especially when the great covering area (Figure 1) and the considerable latitudinal extension (22º 30' - 26º 30' S) of this ecosystem are considered. The lack of studies is related to the advanced stage of degradation of this forest type. In a few decades this ecosystem was reduced to scarce and fragmented remnants, generally in bad conservation conditions. We did not find phytosociological surveys in the Southwestern region of the state of Paraná, where the Semideciduous Forest is also found (Figure 1). Moreover, forest structure studies are not available even in the most representative remnant of the Submontane Semideciduous Forest of Southern Brazil, located in the Iguaçu National Park, a protected area of approximately 1852.6 km2. This fact points out to the urgency of knowing better this important ecosystem and its resources.

Besides the similar sampled areas shown by the three Atlantic Forest types (Table 2), and highest number of studied sites for the Dense Rainforest, the sum of the 10 sites of the Semideciduous Forest comprises the largest sampled area (11.8 ha). The two most extensively sampled formations are the Montane Araucaria Rainforest and the Submontane Semideciduous Forest. However, these sampling areas are very small compared to the sampling area of other surveys in different ecosystems. As an example, a single study in the Amazonian Dense Forest, could easily cover an area of 20 ha (Pitman et al., 2002; Laurance et al., 2010).

#### **4.2 Alfa diversity**

In the 58 selected phytosociological surveys, 700 species, 256 genera and 83 families were sampled (Table 2). Of the total number of species, 10 *taxa* represent groups with

The emergent irregular canopy can reach around 35-40 m height. In the upper stratum are common *Aspidosperma polyneuron*, *Handroanthus heptaphyllus*, *Gallesia integrifolia*, *Balfourodendron riedelianum*, *Peltophorum dubium*, *Astronium graveolens*, *Diatenopteryx sorbifolia, Parapitadenia rigida, Cariniana estrellensis, Cedrela fissilis, Albizia hasslerii, Lonchocarpus guilleminianus*, *Machaerium stipitatum*, *Holocalyx balansae*, *Rauvolfia sellowii*, and *Nectandra megapotamica*, among others. The dominated strata are characterized by *Guarea macrophylla*, *Actinostemon concolor*, *Metrodorea nigra*, *Sorocea bomplandii* and *Pilocarpus pennatifolius* (Hueck, 1972; Leite et al., 1986; Leite & Klein, 1990; Maack, 2002; Roderjan et al., 2002; SEMA, 2002).

Among the 58 sites, comprising 29 ha of plots and 36627 sampled individuals, selected from 39 studies, the Dense Rainforest has the highest number of sampled sites (29 out of a total of 58) and the highest number of tree individuals found (Table 2). This great amount of data available is due to the high number of Upper Montane Dense Rainforest sampled sites, which present relatively many areas with primary and well preserved vegetation. In spite of the relatively smaller sampled area, small-sized trees present in high abundance make up

On the other hand, the Alluvial formation of the Dense Rainforest is the least sampled one (Table 2). The lack of phytosociological studies conducted in this formation may be related to its actual small cover area (representing only 0.89% of the remnants of Dense Rain Forest) (Pires et al., 2005), and also to its level of degradation, which make the search for typical and representative remnants difficult. According to this data, the studied area is almost six times smaller than the area of the neighbor Lowland formation. Therefore, this understudied

The low number of studied sites in the Semideciduous Seasonal Forest in the state of Paraná is also remarkable, especially when the great covering area (Figure 1) and the considerable latitudinal extension (22º 30' - 26º 30' S) of this ecosystem are considered. The lack of studies is related to the advanced stage of degradation of this forest type. In a few decades this ecosystem was reduced to scarce and fragmented remnants, generally in bad conservation conditions. We did not find phytosociological surveys in the Southwestern region of the state of Paraná, where the Semideciduous Forest is also found (Figure 1). Moreover, forest structure studies are not available even in the most representative remnant of the Submontane Semideciduous Forest of Southern Brazil, located in the Iguaçu National Park, a protected area of approximately 1852.6 km2. This fact points out to the urgency of knowing

Besides the similar sampled areas shown by the three Atlantic Forest types (Table 2), and highest number of studied sites for the Dense Rainforest, the sum of the 10 sites of the Semideciduous Forest comprises the largest sampled area (11.8 ha). The two most extensively sampled formations are the Montane Araucaria Rainforest and the Submontane Semideciduous Forest. However, these sampling areas are very small compared to the sampling area of other surveys in different ecosystems. As an example, a single study in the Amazonian Dense Forest,

In the 58 selected phytosociological surveys, 700 species, 256 genera and 83 families were sampled (Table 2). Of the total number of species, 10 *taxa* represent groups with

could easily cover an area of 20 ha (Pitman et al., 2002; Laurance et al., 2010).

**4. The studied Atlantic Forest formations** 

for a high number of individuals found.

better this important ecosystem and its resources.

vegetation needs more studies.

**4.2 Alfa diversity** 

**4.1 Analysis of the sampled sites in the Atlantic Forests** 

undetermined species at the family level and 58 *taxa* represent groups with undetermined species at the genus level. It is important to mention that the Atlantic Forest is habitat for many other tree species that were not found due to the criterion of inclusion, or due do the area needed to sample, that neither checklists can cover completely.


Table 2. Richness and other diversity parameters of the analyzed Atlantic Forest formations in the state of Paraná, Southern Brazil.

Among the Atlantic Forest types analyzed in this study, the Dense Rainforest is the richest in tree species (469). These species are distributed in 174 genera and 72 families. This tropical forest presents the highest tree diversity according to the Shannon-Wiener index, being its Montane and Submontane formations the most diverse (Table 2; Figure 3). Even though the analysis shows the highest diversity value in the Montane formation, according to many studies, the Submontane formation shows a tendency of being the most diverse (Guapyassú, 1994; Roderjan, 1994; Jaster, 1995; Athayde, 1997; Blum, 2006).

Tabarelli & Mantovani (1999) compiled phytosociological studies on the Dense Rainforest in Southeastern Brazil, which comprised 432 species measured in 2.3 ha and considered 2640 trees measured through the quarter-plot method. According to these authors the richness of these forests are low when compared to other Neotropical Forests of South America. However, more studies considering similar conditions (*e.g.* larger plots – 1 ha – with homogeneous sites) are needed to corroborate these results.

However, some diversity indexes such as Eveness presented for the Atlantic Forest types can be influenced by the disproportionate sampling among their formations.

According to Table 2, the Araucaria Rainforest presents less than half of the Dense Rainforest tree species richness (220), in part due to its lower environmental heterogeneity

Arboreal Diversity of the Atlantic Forest

formations of both rainforests (Table 4).

Semideciduous Forest (Tables 3 and 4).

and Asteraceae richness.

of Southern Brazil: From the Beach Ridges to the Paraná River 121

Forest. Some tree families, such as Fabaceae, Moraceae, Sapotaceae and Meliaceae, show a tendency of preferring warmer environments. On the other hand, Aquifoliaceae, Asteraceae and Myrsinaceae present comparatively high floristic expression in the Rainforest environments. In addition, Melastomataceae presents high species richness only in the Dense Rainforest. According to Gentry (1995) and Oliveira-Filho et al. (2000), with increasing altitude there is a decrease in Fabaceae richness and an increase in Aquifoliaceae

At the genus level, *Eugenia* is the richest in the tropical forests (mainly in the Dense Rainforest), even though the Araucaria Rainforest (subtropical) presents many species of this taxon. Oliveira-Filho et al. (2000) observed the same tendency in an extensive survey on floristic differentiation patterns among Atlantic Forests in Southeastern Brazil. The richness of *Eugenia* is relatively higher in slope rain forests, and its relative position is reversed with *Myrcia* in lowland (including alluvial) and plateau (Araucaria Rainforest) formations. *Marlierea* has a considerable number of species in Lowland and Submontane formations of the Dense Rainforest, whilst *Myrceugenia* has many species in Upper Montane and Montane

*Ocotea* is among the three genera with the highest number of tree species in the three Atlantic Forest types in the state of Paraná. *Ilex* and *Myrceugenia* have a considerable number of species in the two rainforests, whilst *Trichilia* and *Cordia* have many species in the

**Dense Rainforest (S=469; N=29) Araucaria Rainforest (S=220; N=19) Semideciduous Forest (S=282; N=10) Family S Genus S Family S Genus S Family S Genus S**  Myrtaceae 112 *Eugenia* 39 Myrtaceae 49 *Myrcia* 14 Fabaceae 47 *Eugenia* 13 Lauraceae 39 *Myrcia* 25 Lauraceae 23 *Ocotea* 11 Myrtaceae 32 *Nectandra* 8 Fabaceae 35 *Ocotea* 18 Fabaceae 18 *Eugenia* 10 Lauraceae 19 *Ocotea* 8 Rubiaceae 24 *Miconia* 12 Asteraceae 10 *Myrceugenia* 8 Meliaceae 13 *Trichilia* 8 Melastomataceae 21 *Ilex* 11 Rubiaceae 10 *Ilex* 6 Rubiaceae 11 *Cordia* 7 Moraceae 12 *Myrceugenia* 11 Salicaceae 8 *Myrsine* 6 Solanaceae 11 *Solanum* 7 Sapotaceae 12 *Calyptranthes* 10 Solanaceae 8 *Solanum* 6 Salicaceae 10 *Inga* 6 Aquifoliaceae 11 *Inga* 9 Aquifoliaceae 6 *Symplocos* 6 Euphorbiaceae 9 *Myrcia* 6 Euphorbiaceae 11 *Nectandra* 9 Myrsinaceae 6 *Casearia* 5 Rutaceae 9 *Casearia* 5 Asteraceae 9 *Marlierea* 8 Sapindaceae 6 *Maytenus* 5 Boraginaceae 7 *Ficus* 5 Myrsinaceae 9 *Myrsine* 8 Symplocaceae 6 *Allophylus* 4 Moraceae 7 *Lonchocarpus* 5 Annonaceae 8 *Symplocos* 8 Annonaceae 5 *Lonchocarpus* 4 Sapotaceae 7 *Machaerium* 5 Meliaceae 8 *Coccoloba* 7 Celastraceae 5 *Machaerium* 4 Annonaceae 6 *Pouteria* 5 Salicaceae 8 *Psychotria* 7 Euphorbiaceae 4 *Nectandra* 4 Malvaceae 6 *Sloanea* 4 Sapindaceae 8 *Ficus* 6 Rutaceae 4 *Piptocarpha* 4 Sapindaceae 6 *Zanthoxylum* 4 Table 3. The 15 best represented families and genera measured in the three Atlantic Forest types of the state of Paraná, Southern Brazil. S = number of species, N = number of sites.

The most abundant species found in phytosociological studies in the Atlantic Forest formations in the state of Paraná are given in Table 5. Except for the Alluvial, the sampling for all the other formations can be considered very representative of what can be found in these forests. This compilation obtained data for at least 2800 individuals and four sites for each formation, whereas phytosociological studies for the Semideciduous Seasonal forest in

the Southwest of the state of Paraná cannot be found.

and altitudinal range. In addition, the colder climate in the Araucaria Rainforest region probably restricts the occurrence of a substantial number of species.

The Semideciduous Seasonal Forest shows intermediate values of species richness (282) and diversity when compared with the two types of Rainforests. (Table 2; Figure 3).

Fig. 3. Tree richness and Shannon-Wiener diversity indexes of the Atlantic Forest formations in the state of Paraná, Southern Brazil.

The higher richness when compared with the Araucaria Rainforest is due to some distinct environmental conditions, such as lower altitude, which result in higher temperatures. On the other hand, the restriction imposed by the seasonal climate, especially by the unfavorable dry season, prevents the Seasonal Forest to reach values of species richness similar to those observed in the Dense Rainforest (i.e. 66% higher). A lower difference (31%) in this parameter between these two types of forest was observed by Oliveira Filho & Fontes (2000) in floristic surveys in Southeastern Brazil.

The Submontane formations present the highest richness of tree species, followed by the Montane formations (Figure 3). The Lowland, Upper Montane and Alluvial formations present lower richness. From the data available, the Alluvial Dense Rainforest has the lowest number of species. However, this is certainly related to the lack of inventories in this type of forest. Due its proximity and environmental similarity with the coastal Lowland formation, it is probable that the real values of tree species richness of both forests are similar.

Therefore, among the Atlantic Forest formations, the Alluvial Araucaria Rainforest can be considered the least rich in tree species. This is due to two important environmental factors: the milder climate, with frequent frosts; and the hydromorphic feature of its soils, with high susceptibility to flooding during the rainiest periods.

According to Table 3, Myrtaceae, Fabaceae and Lauraceae are the families with the highest tree species richness of the Atlantic Forest, making up to 40% of the measured Rainforest species and 35% of the Semideciduous Seasonal Forest species.

Myrtaceae presents the highest number of tree species in the two Rainforests, reaching 112 species in the Dense Rainforest, while Fabaceae is the richest family of the Semideciduous

and altitudinal range. In addition, the colder climate in the Araucaria Rainforest region

The Semideciduous Seasonal Forest shows intermediate values of species richness (282) and

3,93

3,20

157 148

Fig. 3. Tree richness and Shannon-Wiener diversity indexes of the Atlantic Forest formations

The higher richness when compared with the Araucaria Rainforest is due to some distinct environmental conditions, such as lower altitude, which result in higher temperatures. On the other hand, the restriction imposed by the seasonal climate, especially by the unfavorable dry season, prevents the Seasonal Forest to reach values of species richness similar to those observed in the Dense Rainforest (i.e. 66% higher). A lower difference (31%) in this parameter between these two types of forest was observed by Oliveira Filho & Fontes

The Submontane formations present the highest richness of tree species, followed by the Montane formations (Figure 3). The Lowland, Upper Montane and Alluvial formations present lower richness. From the data available, the Alluvial Dense Rainforest has the lowest number of species. However, this is certainly related to the lack of inventories in this type of forest. Due its proximity and environmental similarity with the coastal Lowland formation,

Therefore, among the Atlantic Forest formations, the Alluvial Araucaria Rainforest can be considered the least rich in tree species. This is due to two important environmental factors: the milder climate, with frequent frosts; and the hydromorphic feature of its soils, with high

According to Table 3, Myrtaceae, Fabaceae and Lauraceae are the families with the highest tree species richness of the Atlantic Forest, making up to 40% of the measured Rainforest

Myrtaceae presents the highest number of tree species in the two Rainforests, reaching 112 species in the Dense Rainforest, while Fabaceae is the richest family of the Semideciduous

it is probable that the real values of tree species richness of both forests are similar.

<sup>88</sup> <sup>79</sup> <sup>78</sup>

2,48

3,38

Richness Shannon-Wiener H'

probably restricts the occurrence of a substantial number of species.

211 210

<sup>265</sup> <sup>252</sup>

in the state of Paraná, Southern Brazil.

(2000) in floristic surveys in Southeastern Brazil.

susceptibility to flooding during the rainiest periods.

species and 35% of the Semideciduous Seasonal Forest species.

4,61 4,46 4,38 4,64

diversity when compared with the two types of Rainforests. (Table 2; Figure 3).

4,26

Forest. Some tree families, such as Fabaceae, Moraceae, Sapotaceae and Meliaceae, show a tendency of preferring warmer environments. On the other hand, Aquifoliaceae, Asteraceae and Myrsinaceae present comparatively high floristic expression in the Rainforest environments. In addition, Melastomataceae presents high species richness only in the Dense Rainforest. According to Gentry (1995) and Oliveira-Filho et al. (2000), with increasing altitude there is a decrease in Fabaceae richness and an increase in Aquifoliaceae and Asteraceae richness.

At the genus level, *Eugenia* is the richest in the tropical forests (mainly in the Dense Rainforest), even though the Araucaria Rainforest (subtropical) presents many species of this taxon. Oliveira-Filho et al. (2000) observed the same tendency in an extensive survey on floristic differentiation patterns among Atlantic Forests in Southeastern Brazil. The richness of *Eugenia* is relatively higher in slope rain forests, and its relative position is reversed with *Myrcia* in lowland (including alluvial) and plateau (Araucaria Rainforest) formations. *Marlierea* has a considerable number of species in Lowland and Submontane formations of the Dense Rainforest, whilst *Myrceugenia* has many species in Upper Montane and Montane formations of both rainforests (Table 4).

*Ocotea* is among the three genera with the highest number of tree species in the three Atlantic Forest types in the state of Paraná. *Ilex* and *Myrceugenia* have a considerable number of species in the two rainforests, whilst *Trichilia* and *Cordia* have many species in the Semideciduous Forest (Tables 3 and 4).


Table 3. The 15 best represented families and genera measured in the three Atlantic Forest types of the state of Paraná, Southern Brazil. S = number of species, N = number of sites.

The most abundant species found in phytosociological studies in the Atlantic Forest formations in the state of Paraná are given in Table 5. Except for the Alluvial, the sampling for all the other formations can be considered very representative of what can be found in these forests. This compilation obtained data for at least 2800 individuals and four sites for each formation, whereas phytosociological studies for the Semideciduous Seasonal forest in the Southwest of the state of Paraná cannot be found.

Table 4. The 15 best represented families and genera measured in the nine Atlantic Forest formations of the state of Paraná, Southern Brazil. S = number of species, N = number of sites. \*Data from Zacarias (2008).

Arboreal Diversity of the Atlantic Forest

of sites. \*Data from Zacarias (2008).

**Alluvial\*** 

of Southern Brazil: From the Beach Ridges to the Paraná River 123

Table 5. Species with the highest number of individuals measured for each formation of Atlantic Forest in the state of Paraná, Southern Brazil. I = number of sampled individuals, N = number


Table 5. Species with the highest number of individuals measured for each formation of Atlantic Forest in the state of Paraná, Southern Brazil. I = number of sampled individuals, N = number of sites. \*Data from Zacarias (2008).

Table 4. The 15 best represented families and genera measured in the nine Atlantic Forest formations of the state of Paraná, Southern Brazil. S = number of species, N = number of sites.

\*Data from Zacarias (2008).

**Alluvial\*** 


Table 5. (Continued)

Arboreal Diversity of the Atlantic Forest

N = 29 S = 469

Dense Rainforest

Dense Rainforest

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

*reitzii*, *Drimys angustifolia* and *Handroanthus catarinensis*.

*Erythroxylum deciduum, Cinnamomum amoenum and Zanthoxylum kleinii.* 

of Southern Brazil: From the Beach Ridges to the Paraná River 125

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* 

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,* 

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,* 

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

Rainforest Semideciduous

*Araucaria* Rainforest

**S = 504 S = 359**

Dense Rainforest

**S = 580 S = 631**

Fig. 4. Venn diagrams presenting the number of measured tree species shared in the 58 sites

**<sup>321</sup> <sup>94</sup> <sup>165</sup> <sup>272</sup> <sup>127</sup>**

*Peltophorum dubium, Holocalyx balansae, Aspidosperma polyneuron and Gallesia integrifolia.* 

N = 19 S = 220

**316 99 89 100 88 171**

*Araucaria*

Semideciduous Forest

of the three types of Atlantic Forest in the state of Paraná, Southern Brazil.

N = 10 S = 282

Semideciduous Forest

Forest

*Araucaria* Rainforest

**50**

**44**

**51 <sup>38</sup> <sup>49</sup>**
