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Tree-Fall Gaps Effects on Spider (Araneae) Assemblages in an Atlantic Forest Landscape in Northeastern Brazil

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Tree-Fall Gaps Effects on Spider (Araneae) Assemblages in an Atlantic Forest Landscape in Northeastern Brazil Open Journal of Animal Sciences, 2014, 4, 118-133 Published Online June 2014 in SciRes. http://www.scirp.org/journal/ojas http://dx.doi.org/10.4236/ojas.2014.43016 Tree-Fall Gaps Effects on Spider (Araneae) Assemblages in an Atlantic Forest Landscape in Northeastern Brazil Marcelo Cesar Lima Peres1,2,3*, Kátia Regina Benati2,3, Alessandra Rodrigues S. de Andrade2,3, Marcos Vinicius Alvim Guimarães2, Tércio da Silva Melo2, Antonio Domingo Brescovit4, Jacques Hubert Charles Delabie3,5 1Territorial Planning and Social Development, Catholic University of Salvador, Salvador, Brazil 2Ecology Center and Animal Conservation, Universidade Católica do Salvador, Salvador, Brazil 3Federal University of Bahia, Salvador, Brazil 4Butantan Institute, Collections Zoological Laboratory, São Paulo, Brazil 5Laboratory of Myrmecology, UESC/CEPLAC, Centre for Cocoa Research, Ilhéus, Brazil Email: *[email protected] Received 15 April 2014; revised 20 May 2014; accepted 27 May 2014 Copyright © 2014 by authors and Scientific Research Publishing Inc. This work is licensed under the Creative Commons Attribution International License (CC BY). http://creativecommons.org/licenses/by/4.0/ Abstract The study investigates the effect of spatial and temporal tree-fall gaps structure on spiders’ as- semblage in an Atlantic forest fragmented area in Brazil. It was conducted in the Michelin Ecologi- cal Preserve-REM (Bahia), 190 ha forest remnant. Samples were collected on leaf-litter (50 × 50 cm) at five tree-fall gaps formations (<150 m2), within five adjacent primary forest and five inner edge parcels. During 16 months (between May 2009 and October 2012), 480 m2 leaf-litter samples were collected, from which spiders were extracted using mini-Winkler traps. The observed and estimated richness of spider’s species was higher at the edge (p < 0.01). The habitat structures differ significantly among the three habitat types (MRPP, p < 0.01) and also during the tree-fall gaps aging gradient (MRBP, p < 0.01). There were significant differences on spiders’ species com- position, comparing the three habitats (MRPP, p < 0.05). The composition of spider’s species changed as tree-fall gaps aged (MRBP, p < 0.05). We argue that the tree-fall gaps play, on a local scale, an important role in acting on the time-space distribution dynamics of spider’s species as- semblages, although the time effect should still be evaluated. Keywords Atlantic Forest, Edge Effect, Habitat Selection, Spatiotemporal Effect, Neotropical Fauna *Corresponding author. How to cite this paper: Peres, M.C.L., et al. (2014) Tree-Fall Gaps Effects on Spider (Araneae) Assemblages in an Atlantic Forest Landscape in Northeastern Brazil. Open Journal of Animal Sciences, 4, 118-133. http://dx.doi.org/10.4236/ojas.2014.43016 M. C. L. Peres et al. 1. Introduction Natural disturbances role in shaping the tropical forests biological communities’ structure is widely known [1]. Natural disturbances increase habitat heterogeneity by promoting specialization and resource partitioning among species, whereas at the same time, they prevent from competitive exclusion fostering diversity [2]-[4]. Gaps formed by the fall of one or more trees are, apparently, the most common disturbance in forest environments, and these forms an opening in the canopy structure [5]. However, tree-fall gaps are likewise considered one of the main contributors for tropical forests’ regeneration, significantly adding to its floristic diversity [6] [7], as they alter the habitat structure. The regeneration witnessed on tree-fall gaps promotes the development of primary and pioneer species, and the secondary one promotes remarkable changes on trees’ population’s dynamics, species composition and growth rate [8] [9]. Therefore, it has been considered as an essential mechanism for the forests diversity’s main- tenance [3] [10] [11]. Tree-fall gap regeneration can be divided into three phases: tree-fall gaps formation, building phase, and mature phase [12]. These phases duration vary, and depend on factors such as size [6] [13], alleviation [14], severity of disturbance [13], soil condition [7] and pre-existing vegetation [15] [16]. The tree-fall gap phase is considered the most important to determine floristic composition [17], since it is characterized by an intense gap, changing microclimatic conditions and plant community structures [15]. It is estimated to last from four to six years in tropical forests [9]. Tree-fall gaps influence the spatial and temporal distribution of plants, therefore, affect these organisms’ in- teractions to other animals [18] [19]. However, few studies have focused on animals, whilst most of them have pointed out structural aspects, such as area, age, geometry and type of fall, as well as vegetal organisms rather than their interactions [6] [10] [13]. This knowledge bulk has found significant tree-fall gaps influence on plant community structure. Following a thorough literature review we can refer to: studies on trees [3], pioneer plants [6] [14], bamboos [20], seedling growth [21], and gap and grazing areas interactions with the establishment, growth and survival of pioneer species [22]. Previous studies of animal related tree-fall gaps include: birds [1], ants [23]-[25] and spiders [26] [27]. No significant differences were found, on the aforementioned publications, between tree-fall gaps and mature fo- rests on birds and orb-weaver spiders’ studies. However, studies on ants have revealed controversial results. In a forest in India, species richness differed between forest tree-fall formations [23], whilst in a study conducted in a forest in Panama, no difference was found on species’ abundance, richness and composition [24]. More recently, a study in a forest in Costa Rica found that the estimated richness differ between mature forest and adjacent tree-fall gap, although species composition did not show significant difference [25]. Most studies tend to com- pare the fauna in young tree-fall gaps to intact forest, ignoring that tree-fall gaps may change over time [25]. Thus, time scale approaches are essential to understand the tree-fall gaps dynamics, especially when related to certain animal groups such as arachnids. Spiders (Arachnida: Araneae) are among the most abundant and diverse animal group, with 44,540 species currently described in the World [28]. They act as secondary predators on the trophic structure where they have a role [29]. They then, directly or indirectly regulate important taxa’ abundance of ecosystem processes, such as predation, pollination, decomposition and dispersion [30]. Spiders’ distribution and density are related to envi- ronmental factors such as temperature, relative humidity and luminosity [31] [32], habitat physiognomy, plant richness and composition [33]. In tropical forests such as any other natural landscape, habitat structure influ- ences the richness and composition of spider’s assemblages species [34]). Leaf litter structure also affects spid- ers’ abundance [35], richness and composition [36] [37]. Therefore, natural disturbance caused by tree fall gaps may also influence the spider’s assemblages [38] [39]. This study investigates whether tree-fall gaps show time-space scale effect on composition of spider’s assem- blages species in an Atlantic forest remnant at the Michelin Ecological Reserve, Igrapiúna, Bahia, northeastern Brazil. We addressed four questions: 1) Does the habitat structure of tree-fall gap (microclimate and microhabi- tat) significantly differ from that found in adjacent mature forest and inner edge? 2) Does the richness and com- position of tree-fall gap species significantly differ from those found in adjacent mature forest and inner edge? 3) Does the habitat structure of natural tree-fall gaps (microclimate and microhabitat) significantly change dur- ing vegetation regeneration process? 4) Does the spider’s species’ assemblage composition of tree-fall gaps sig- nificantly change throughout the regeneration process? 119 M. C. L. Peres et al. 2. Material & Methods 2.1. Study Area The study was conducted at the Ecological Reserve Michelin (REM) (13˚50'S, 39˚10'W, 90 to 383 m above sea level), situated in the city of Igrapiúna and Ituberá (state of Bahia, Brazil). It is located 18 km away from the coast in a region known as “BaixoSul”, 200 km south from the State capital city of Salvador. The 3096 ha REM area consists of a vegetation mosaic on different successional stages. Landscape id modified by different types of human disturbance, agriculture and pasture converted. And other forms of anthropogenic pressure (such as logging, hunting and exploitation of the palm). Approximately, 25% of the reserve is intended to rubber tree monoculture plantations [40]. The mean annual rainfall is 2000 mm and the temperature ranges from 21˚C to 28˚C. [40]. The reserve contains 1800 ha of lowland evergreen hill forest distributed in three main fragments: Vila 5/Pancada Grande fragment with 625 ha, the 140 ha Luis Inácio forest, and the 550 ha Pacangê forest, which is contiguous with a 13,000 ha forest [40]. This study was conducted in Mata da Vila Cinco with 190 ha, which is one of the remnants. The northern access to this fragment is constituted by a land mosaic dominated by mature and young primary forest, where the canopy reaches uniformly from 18 - 25 m. The southern access consists of mature second-growth forest, with trees ranging from eight to 13 m height, abundant vines and thick leaf litter. At higher slopes in the landscape there is a group of mature primary forest with several old trees. The local jus- sarapalm, bromeliads, vines and other epiphytes are abundant and well distributed in this remnant, which is crossed by a small river, two streams and several springs. The forest is surrounded by cocoa, rubber and banana plantations and is limited to the north by the “Mata do Rio” forest, another section of the reserve (K. Flesher, pers. comm.). 2.2. Sampling Design Eight field surveys were conducted every two months, from July 2009 to October 2010. On the northern sector of Mata Vila Cinco we defined the tree-fall gaps and forest sampling points, both in the mature forest sections.
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