*Dirección para correspondencia
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Abstract

The forest understory is made up of resident and transitory species and can be much richer than the canopy. With the purpose to describe the contribution of these groups to the woody understory, five Atlantic Forest fragments were selected and ]]>
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Key words: height classes, floristic composition, density, natural regeneration, richness, functional groups.


Resumen
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El sotobosque forestal está compuesto por especies residentes y transitorias. Con el objetivo de describir la contribución de esos grupos en el sotobosque leñoso, cinco fragmentos de Bosque Atlántico fueron seleccionados en el nordeste de Brasil. El muestreo incluyó individuos con circunferencia a la altura del pecho (CPA)<15cm y con circunferencia a la altura del suelo (CAS)≥3cm. Las especies fueron cuantificadas y clasificadas en residentes o transitorias y la ]]>

Palabras clave: clases de altura, composición florística, densidad, regeneración natural, riqueza de especies, Bosque Atlántico, grupo funcional.

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Several life forms coexist in the forest understory, which contributes to high species richness (Gentry & Dodson 1987, Schnitzer & Carson 2000). There are two main approaches to view the regeneration dynamics of this stratum: either as a physiognomic component defined by a maximum height or diameter, including young trees, seedlings, saplings, and shrubs, regardless of ]]> et al. 2002, Rayol et al. 2006, Silva et al. 2007, Marangon et al. 2008, Gomes et al. 2009, Lü et al. 2010, Sansevero et al. 2011); or as functional groups of plants that ]]> et al. 2004, Araújo et al. 2006, Santos et al. 2008, Aquino & Barbosa 2009, Onofre et al. 2010).

The resident group ]]> et al. 1994, 1995). The transitory group is generally made up of tree individuals with dendrometric values lower than those defined for the tree stratum (normally diameter at breast ]]> et al. 1995), the understory’s composition may include much more species than the canopy (Galeano et al. 1998, Lü et al. 2010). Future canopy composition, in turn, is dependent on the density of transitory species in the understory, as well as on the recruitment of these species from the lower to the highest height classes (Clark & Clark 1992, Volpato 1994, Kobe 1999). Species that occur in all ]]> et al. 2000), caused by physical damage, leaf litter, vertebrates (Scariot 2000, Ickes et al. 2001, Santos & Válio 2002) or by biological damage such as predation and parasitism (Cadenasso & Picket 2000).

The physiognomic and ]]> et al. 2003) and allow us to: better estimate the richness and state of conservation (Richards 1996), diagnose the dynamics of natural forest fragments (Finol 1971), measure responses to soil and climatic variations and to environmental stress (Harms et al. 2004), plan management actions, forestry practices and vegetation ]]> et al. 2003, Silva et al. 2007, Sansevero et al. 2011).

In a mature tropical forest, the floristic composition of the canopy and understory is expected to be distinct from each other (Jardim & Hosokawa 1986) due to the low ]]> et al. (2008) who recorded an increase in the similarity between the canopy and the understory from a more disturbed area (0.45; edge) to a more conserved one (0.59; forest interior) in a 300ha mature Atlantic forest fragment.
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In fragmented landscapes, analyzing the understory’s structure and identifying its functional components has been crucial to diagnose the effects of disturbances (Martins & Rodrigues 2002, Ceccon et al. 2006), fragmentation and edge effects (Benitez-Malvido 1998, Laurance et al. 2007, Bouroncle & Finegan 2011). In the Brazilian Atlantic Forest, where most areas have been ]]> et al. 2009). Yet, despite its status as a world conservation hotspot (Mittermeier et al. 2005), and the fact that many plants spend all or a considerable part of their lives in the understory (Harms et al. 2004), few studies have analyzed the understory vertical structure (e.g. Silva et al. 2007, Marangon et al. 2008), described who is who in this stratum regarding plant habit (Souza et al. 2009) or successional category (Tabarelli & Mantovani 1999, Aquino & Barbosa 2009,Onofre et al. 2010, Sansevero et al. 2011) or compared understory composition and structure among forests (Gomes et al. 2009).
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Only 4.6% of the original vegetation remains in the extreme North of the Atlantic Forest’s distribution area (Lima 1998), in the form of small, irregular fragments, most less than 500ha in size (Trindade et al. 2008). Such forests are located mainly at the bottom of deep valleys - areas where small springs and watercourses can be found - and have closed canopies. They are, thus, riparian forests (Naiman et al. 2005) made up of assemblages that border watercourses in areas where the interfluvial vegetation is also forest (Metzger et al. 1997). In addition to the typically riparian species, species from the upper forests can also be found in these habitats, since the riparian vegetation serves as an important source of diaspores for forest remnants’ natural regeneration and recolonization processes (Triquet et al. 1990).

]]> With the purpose of describing the physiognomic and functional structure of the understory in Atlantic Forest fragments, five differently sized remnants were selected from a highly fragmented landscape. The study had five goals: 1) to uncover species richness and composition; 2) to discover how many and which canopy species are found in the understory; 3) to functionally classify species as resident or transitory; 4) to investigate the existence of a pattern in the understory’s structure among the different fragments; and 5) to discover the present and potential regenerative capacity of ]]>

Material and methods

Study area: Five fragments were studied, all from the extreme North of the state of Pernambuco, Northeastern Brazil (07º41’05” - 07º54’17” S and ]]> et al. 2008). Following Köppen’s classification, climate is of type As’ (tropical wet), with an average precipitation of 1 689mm and rain concentrated from April-August; the average temperature is 25.1ºC (data from the São José sugarcane processing plant for the period 1998-2006). The fragments are located on the Barreiras geological formation, which dates back to the Plio-Pleistocene and consists of unconsolidated sandy-clay sediments of continental origin. The terrain is made up of flattened tableland cut by deep, narrow ]]> et al. 2005).

Composition and structure data sampling: In order to analyze the understory, 20 non-contiguous 5x5m ]]>

To sample to tree stratum, we ]]> et al. 2006).

The botanical material collected was added to the Herbarium Geraldo Mariz (UFP) of the Federal University of Pernambuco/Usina São José Collection. When possible, samples were identified up to the species level with the aid of specific literature, comparison with material from the UFP, PEUFR and IPA herbaria that had been previously identified by specialists. ]]>

Analysis of data structure and ]]>

The subdivision of typical species into the low understory and upper understory categories was based on the works of Vilela et al. (1995), Oliveira & Amaral (2005), Gomes et al. (2009), Souza et al. (2009) and Onofre et al. (2010). Upper ]]>

Studies on natural regeneration ]]> et al. (1994) and Marangon et al. (2008). Marangon et al. (2008) used the minimum 1m height limit and justified that individuals with such a height have better morphological characteristics and, thus, allow for a more reliable identification. In this study, we also included plants that were less than 1m tall, as Oliveira-Filho et al. (1994) have stated that this makes it possible to sample more individuals. Individuals were grouped into the following height classes: class 1-height<1m; class 2-height≥1m and <2m; class 3-height≥2m and <3m; and class 4-height ≥3m with CBH less than 15cm.

Species importance in the understory was analyzed based on the Natural Regeneration Index by ]]> et al. (2007), ]]> et al. (2008) and Santos et al. (2008) used Volpato’s modified version for describing natural regeneration in Brazilian forests. This latter incorporates a more detailed analysis per height class when calculating the index, which considers the relative density of the species in each class.

In order to calculate the Natural Regeneration Index, absolute and relative density and frequency ]]> ij=(RD_ij+RF_ij)/2 where NRC_ij is the natural regeneration estimate of the i^th species of the j^_th plant size class, as a percentage, RD_ij is the relative density for the i^th species of the j^th natural regeneration size class, RF_ij is the relative frequency for ]]> th species of the j^th natural regeneration size class, i is the 1, 2, 3... p^th species sampled, and j is the 1, 2, 3, and 4 classes.

The next step was to calculate the Total Natural Regeneration estimate (TNR) for each species by adding the natural regeneration indexes of each class according to the formula ]]> i=ΣNRC_ij, where TNR_i is the total natural regeneration estimate of the i^th species, NRC_ij is the natural regeneration estimate of the i^th species of the j^th plant size class, i is the 1, 2, 3... p^th species sampled, and j is the 1, 2, 3 and 4 classes.

]]> Results

Species richness and composition: A total of 163 species distributed among 44 families were sampled from the understory of the five fragments analyzed (fragment median=63.8 species, SD=21.72, n=5), varying from 32 species in F3 to 93 species in F1 (Table 1). One ]]> Table 1). Fragment F3 had the highest similarity (0.68), while fragments F4 and F5 were the least similar (0.46 and 0.45, respectively).

Families Myrtaceae (17 species/227 ]]>

]]> In addition to the families cited above, despite its lower richness, the family Lecythidaceae (3 species/447 individuals) was high in density. Anaxagorea dolichocarpa Sprague & Sandwith (Annonaceae), Eschweilera ovata (Cambess.) Miers. (Lecythidaceae) and Symphonia globulifera L.f. (Clusiaceae) stood out due to the number of individuals (24.10% of the total) and ]]>

Species classification into stratification categories: The percentage of transitory species was greater than that of typical understory species (Fig. 1A). An average of 67.01% (SD=3.76) transitory species were recorded, while lower and ]]>
Transitory species ]]> Fig. 1B). Conversely, the lower and upper understory species represented 11.52% (11.33) and 27.06% (14.50) of the individuals, respectively. Fragment F5 differed from the others because it had a greater percentage of individuals in the lower understory (30.79%) and a smaller percentage of individuals in the upper understory (12.22%) in relation to fragments F1 (G=29.30, p<0.05), F2 (G=19.50, p<0.05) and F3 (G=54.82, p<0.05). Fragment F4 was different due to its higher percentage transitory ]]>

]]> Among the 163 species recorded in the five fragments’ understory, looking at the 10 with highest regeneration indexes (TNR) in each fragment, 34 species are listed (Table 2). These species include the most important in at least one fragment, in which they added up to 10.75-31.25% of the total natural regeneration. Transitory species were the majority, with 21 recorded species (61.76%), while the lower and upper understory had six (17.65%) and seven species (20.59%), respectively.

Regenerative capacity and vertical structure of the understory: Most species with high total natural regeneration indices (TNR) were also well distributed among height classes and had the highest densities (Fig. 2). Only 37 species - less than 21% of the total richness - had densities equal to or higher than ]]> Table 1); the highest percentage was recorded for F3, which stood out for housing 31.25% of the species with 15 or more individuals. Species such as Inga thibaudiana DC., Myrcia racemosa (O. Berg) Kiaersk., Miconia prasina (Sw.) DC. (F2) and Pilocarpus cf. giganteus Engl. (F3) all had less than 15 individuals each. Nevertheless, they were among those with the highest TNR, as they were recorded in all of the height classes (the exceptions were M. prasina and P. cf. giganteus, which were found in three classes, yet had high relative frequencies in relation to the others). Tapirira ]]> Aubl. (F1), E. ovata (F3), Henriettea succosa (Aubl.) DC., M. racemosa, Erythroxylum citrifolium A. St.-Hil.,Coccoloba sp16 (F4), Calyptranthes brasiliensis Spreng. and Piper caldense C.DC. (F5), although not being among the species with the highest regeneration, ]]> T. guianensis and M. racemosa - were absent from one of the height classes.

The percentages of ]]> Fig. 3A). The transitory species were the best represented in all of the classes, followed by those from the upper and lower understory ( Fig. 4). The average number of typical lower and upper understory species was similar for classes 1 and 2.

In all of the forest remnants, the species that occurred in only one class were concentrated in the second height class (except for F3, where most occurred solely in class 4) (Fig. 3B). The distribution of transitory and typical understory ]]>
Few species occurred in all of the height classes - percentages of such species varied from 16.67% in F2 to 28.13% in F3 (Fig. 3C). Most species only occurred in one class, with percentages varying from 25% in F3 to 45.16% in F1. Transitory ]]>

Discussion

Species richness and composition: ]]> et al. 1998, Schnitzer & Carson 2000, Lü et al. 2010), as the lower stratum encompasses transitory and resident species. In these forests, the understory flora may represent 50% or more of the total species (Schnitzer & Carson 2000) and augment the list of tree species up to 30% (Lins-e-Silva 2010).

According to Jardim & Hosokawa (1986), the floristic composition of tropical forests is very different for the upper stratum and understory. Nevertheless, the 54% average similarity among the understory and tree stratum of the different fragments studied here was higher than what is commonly found in the literature (Alves & Metzger 2006). It is important to point out that the herbaceous understory was not sampled and the upper understory ]]>

With the exception of Family ]]> et al. 2009, Onofre et al. 2010, Lü et al.2010), as well as in seasonal forests and savannah (Vilela et al. 1995, Cardoso-Leite et al. 2004, ]]>

Species classification into stratification categories: In a manner similar to that already recorded by other authors (Vilela et al. ]]> et al. 2009), the understory of the studied fragments is also composed primarily by species of the canopy which are passing through the lower strata. Due to the different methodologies used for sampling this stratum, widely varied percentages for tree species have been recorded in the understory- ranging, for example, from 54.54% (Vilela et al. 1995) to 89.18% (Gomes et al. 2009). Despite this variation, all ]]> et al. 2004).

Few species were abundant and most ]]> et al. 2010) and for the tree stratum (Richards 1996). These species, classified as rare or locally rare, generally occur in abundance percentages above 25% (Nappo et al. 2004, Oliveira & Amaral 2005); differently from abundant species, these are more likely to be substituted by others as the forest develops, either for natural reasons or due to disturbances in the area (Campos & Landgraf 2001).
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Regenerative capacity and vertical structure of the understory: Regarding species’ vertical distribution, most were between 1-2m high, similar to what was found by Silva et al. (2007) and Marangon et al. (2008). The species that were found in all height classes - despite their lower occurrence percentages - must have their growth accompanied and their successional characteristics ]]> et al.2007). The exceptions are the species that – due to particular characteristics - are resident and never surpass the lower stratum (Finol 1971).

Species that scored ]]> et al. 2007, Marangon et al. 2008), with few exceptions. Some species remained among those with high TNR, despite low relative density, because of their homogeneous vertical distribution and higher relative frequency in relation to the others. Nevertheless, some species did not have a high regeneration rate ]]>

]]> The results of this study reinforced the idea that the understory of forest fragments is made up mainly of transitory species that are well distributed among the height classes and have higher densities, which favors their future presence in the structure and composition of the canopy. Typical understory species, however, occur in two strata: the lower understory (which includes the species that don’t generally surpass four meters, such as those from families Piperaceae, Rubiaceae and Melastomataceae) and the upper understory (intermediate between the low understory and the ]]>

]]> Acknowledgments

This study is part of the Fragments Project/Phase II - “Sustainability of Atlantic Forest Remnants in Pernambuco and their Implications for Local Conservation and Development”, a Brazil-Germany scientific collaboration program (“Science and Technology for the Atlantic Forest”) financed by CNPq ]]>
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*Correspondencia:
Juliana Silva Gomes-Westphalen:Universidade Federal do Ceará, Campus do Pici, Programa de Pós-Graduação em Ecologia e Recursos Naturais, Bloco 906, 60455-760, Fortaleza, CE, Brasil. juli.ufrpe@gmail.com,
Ana Carolina Borges Lins-e-Silva: Universidade Federal Rural de Pernambuco, Departamento de Biologia, Área de Ecologia, 52171-900, Recife, PE, Brasil. anacarol@db.ufrpe.br
Francisca Soares de Araújo: Universidade Federal do Ceará, Campus do Pici, Programa de Pós-Graduação em Ecologia e Recursos Naturais, Bloco 906, 60455-760, Fortaleza, CE, Brasil. tchesca@ufc.br
1. Universidade Federal do Ceará, Campus do Pici, Programa de Pós-Graduação em Ecologia e Recursos Naturais, Bloco 906, 60455-760, Fortaleza, CE, Brasil; juli.ufrpe@gmail.com, tchesca@ufc.br
2. Universidade Federal Rural de Pernambuco, Departamento de Biologia, Área de Ecologia, 52171-900, Recife, PE, Brasil; anacarol@db.ufrpe.br

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Received 11-VIII-2011. Corrected 06-II-2012. Accepted 05-III-2012.