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Roeder, M. et al. Traits

Roeder, M. et al. Traits and growth of liana regeneration in primary and secondary forests functional diversity of growth forms or species in a community. Analyses of traits have gained importance in ecological research over the last few decades within biomes (e.g. Chapin et al. 1996; Garwood 1996; Ackerly et al. 2002; Poorter & Bongers 2006) and across biomes (e.g. Wright et al. 2004). Performance of seedlings may be closely related to several leaf traits (e.g. Reich et al. 1992). The most obvious difference of performance in plant regeneration is the trade-off between growth and survival, resulting in persistent, slow-growing species with high survival rates versus fast-growing species with low survival rates (Wright 2002). However, a continuum exists between these extremes in trade-offs (Gilbert et al. 2006). Trait data also offer a possibility to describe the main features of a plant community without the need for species identification. This can be a useful approach in tropical environments, on account of their high diversity and the scarcity of knowledge on some groups such as lianas. With the increase of disturbed areas in tropical regions (secondary forest or fragmented forests), the density of lianas is expected to increase. However, only species with trait combinations resembling pioneers might benefit from these areas, and liana infestation will also be influenced by land-use history, such as burning. Studies on the regeneration of lianas, including performance and traits of seedlings and sprouts, in secondary and primary forest are scarce. One study of seedling and sapling performance in primary and secondary forest of Panama indicated that lianas and tree seedlings had widely overlapping ranges in growth rates and survival (Gilbert et al. 2006). We investigated the following main questions: Do traits of liana regeneration differ in secondary forest types of varying land-use history and primary forest? Is the variability of traits and growth different in the forest types? To answer these questions, we used traits of liana seedlings, saplings and sprouts related to whole plant habit and leaf morphology, as well as herbivory and relative growth rate during 1 year. Traits at community and species level were investigated in primary forest in the Central Amazon, Brazil, and in the two main types of secondary forest found in the region, which resulted from different intensities of former land use. Methods Study site The study was conducted in the area of the Biological Dynamics of Forest Fragments Project (BDFFP) (Lovejoy & Bierregaard 1990), north of Manaus, Brazil (2120-250S, 601W). The annual rainfall in Manaus averages 2285 mm (INMET 2010), with a June–October dry season. The mean annual temperature is 26.7 1C (Lovejoy & Bierregaard 1990; Laurance 2001). Soils were classified as nutrient-poor acidic xanthic ferralsols (Ranzani 1980; FAO 2003) with a high clay content and good drainage (Chauvel et al. 1987; Laurance et al. 1999). Elevation ranges from 50 to 100 m a.s.l. and topography consists of plateaux and deeply dissected valleys. Mature vegetation is evergreen, non-inundated (terra firme) lowland tropical forest, with trees 35–40-m tall (Rankin-de Mérona et al. 1992). Density of trees (Z10 cm diameter at breast height [DBH]) is on average 613 stems ha 1 (Laurance 2001). Tree diversity is high, with more than 1000 species in the area (Rankin-de Mérona et al. 1992; Ribeiro et al. 1999; Laurance 2001) and 280 species ha 1 Z10 cm DBH (De Oliveira & Mori 1999). The area of the BDFFP includes primary forest and, at the time of the study, 17- to 25-year-old secondary forest which established after clear-cutting. Two main types of secondary forests (Vismia and Cecropia forest) developed in the area (Mesquita et al. 2001): Vismia forest established when areas were used as cattle pasture for about 4 years and experienced repeated intense fires. At the beginning of our study (2007), the secondary forest in these areas was 17 to 19 years old (T. Vizcarra, pers. com.) and dominated by the genus Vismia (Clusiaceae) and species of the Melastomataceae family. In areas that experienced few or no fires and no cattle ranching, initially a forest dominated by Cecropia (Urticaceae) developed. This forest type was about 25 years old in 2007 and was characterized by greater tree species diversity than the Vismia forest, although few living Cecropia trees remained (nonetheless it is referred to as Cecropia forest). Study plots Plots were established at four sites ranging from 8 to 40 km from one another. Initially 18 secondary forest plots (ten Vismia and eight Cecropia), and nine primary forest plots were studied. All plots of the same forest type were always distributed over three sites and, within a site, were located 0.5–3.0 km from each other (Fig. 1). Secondary forest plots were 25 to 1200 m from primary forest. A plot consisted of three long subplots (1.5 m 30 m), 1.5 m apart, resulting in a total plot area of 135 m 2 . Stand structural characteristics and soil properties of each forest type are provided in Table 1. Methods of soil analysis are described elsewhere (Roeder et al. 2010). A detrended correspondence analysis (DCA) of log-transformed tree family data from stems Z5 cm DBH confirmed the separation of the secondary forests into two groups (details in Roeder et al. 2010). Vismia forest had less canopy cover (0.74) than Cecropia or primary forest (both 0.87) (Table 1). Later, three (in growth analysis four) plots were excluded (two Cecropia and one or two Applied Vegetation Science Doi: 10.1111/j.1654-109X.2011.01152.x © 2011 International Association for Vegetation Science 109

Traits and growth of liana regeneration in primary and secondary forests Roeder, M. et al. Fig. 1. Location of the study site and the 27 study plots. Encircled plots were excluded from the analyses of traits or growth rates at plot level. Source: BDFFP, 1990 &1995. Vismia forest plots) from analysis due to small plant sample size (see below). Study species During the survey of liana regeneration in 27 plots, 2319 plants comprising more than 160 species or morphospecies, including many single- and doubletons, were found (Roeder et al. 2010). Species richness as well as regeneration density per plot was highest in primary forest (Table 1). For the present study, the 26 most important species or morphospecies were chosen, which included 72% of all measured plants. The modified importance value index, IVI (Kershaw & Looney 1985) was used, calculated as relative abundance1relative frequency. Species with an IVIZ2 were included in this study (Appendix S1). Thus singletons, doubletons and rare species were excluded. The sample size of the 26 species varied between 17 and 286 plants per species; in total 1542 plants were used. Many species were spatially clumped and occurred in only a few plots. Species were assigned to a main habitat (primary, Vismia or Cecropia forest) when two-thirds of plants occurred in one habitat type; otherwise species were regarded as co-occurring. When traits were examined per plot, only plots that contained at least 15 plants of the 26 most abundant species were included in the analysis, resulting in 24 plots with 15 to 209 plants. Therefore, in 17 plots more than 64% of all liana individuals were included and in seven plots 37–49% of the liana regeneration. Number of plots per forest type was eight for Vismia, seven for Cecropia and nine for primary forest. For each forest type, plots were still distributed over three sites. Species richness and regeneration density was still significantly different between forest types with the reduced data set. Applied Vegetation Science 110 Doi: 10.1111/j.1654-109X.2011.01152.x © 2011 International Association for Vegetation Science

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