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Assessing Spider Species Richness and Composition in Mediterranean Cork Oak Forests

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Assessing Spider Species Richness and Composition in Mediterranean Cork Oak Forests acta oecologica 33 (2008) 114–127 available at www.sciencedirect.com journal homepage: www.elsevier.com/locate/actoec Original article Assessing spider species richness and composition in Mediterranean cork oak forests Pedro Cardosoa,b,c,*, Clara Gasparc,d, Luis C. Pereirae, Israel Silvab, Se´rgio S. Henriquese, Ricardo R. da Silvae, Pedro Sousaf aNatural History Museum of Denmark, Zoological Museum and Centre for Macroecology, University of Copenhagen, Universitetsparken 15, DK-2100 Copenhagen, Denmark bCentre of Environmental Biology, Faculty of Sciences, University of Lisbon, Rua Ernesto de Vasconcelos Ed. C2, Campo Grande, 1749-016 Lisboa, Portugal cAgricultural Sciences Department – CITA-A, University of Azores, Terra-Cha˜, 9701-851 Angra do Heroı´smo, Portugal dBiodiversity and Macroecology Group, Department of Animal and Plant Sciences, University of Sheffield, Sheffield S10 2TN, UK eDepartment of Biology, University of E´vora, Nu´cleo da Mitra, 7002-554 E´vora, Portugal fCIBIO, Research Centre on Biodiversity and Genetic Resources, University of Oporto, Campus Agra´rio de Vaira˜o, 4485-661 Vaira˜o, Portugal article info abstract Article history: Semi-quantitative sampling protocols have been proposed as the most cost-effective and Received 8 January 2007 comprehensive way of sampling spiders in many regions of the world. In the present study, Accepted 3 October 2007 a balanced sampling design with the same number of samples per day, time of day, collec- Published online 19 November 2007 tor and method, was used to assess the species richness and composition of a Quercus suber woodland in Central Portugal. A total of 475 samples, each corresponding to one hour of Keywords: effective fieldwork, were taken. One hundred sixty eight species were captured, of which Araneae 150 were recorded inside a delimited one-hectare plot; this number corresponds to around Arra´bida 90% of the estimated species richness. We tested the effect of applying different sampling Biodiversity assessment approaches (sampling day, time of day, collector experience and method) on species rich- Iberian Peninsula ness, abundance, and composition. Most sampling approaches were found to influence the Methodology species measures, of which method, time of day and the respective interaction had the Portugal strongest influence. The data indicated that fauna depletion of the sampled area possibly Quercus suber occurred and that the inventory was reaching a plateau by the end of the sampling process. Richness estimators We advocate the use of the Chao estimators as best for intensive protocols limited in space Semi-quantitative sampling and time and the use of the asymptotic properties of the Michaelis–Menten curve as a Stop-rules stopping or reliability rule, as it allows the investigator to know when a close-to-complete inventory has been obtained and when reliable non-parametric estimators have been achieved. ª 2007 Elsevier Masson SAS. All rights reserved. * Corresponding author. Universidade dos Ac¸ores, Dept. Cieˆncias Agra´rias, Terra-Cha˜, 9701-851 Angra do Heroı´smo, Portugal. Tel.: þ351 295 402 206; fax: þ351 295 332 605. E-mail address: [email protected] (P. Cardoso). 1146-609X/$ – see front matter ª 2007 Elsevier Masson SAS. All rights reserved. doi:10.1016/j.actao.2007.10.003 acta oecologica 33 (2008) 114–127 115 1. Introduction et al., 2003). However, they were never fully standardized and optimized as the effort concerning the various variables, Despite their fundamental roles in natural ecosystems, eco- e.g. time of day, was mostly related to the available resources system services and potential use in identifying conservation and personal experience, instead of being a consequence of priority areas, arthropods have largely been ignored in conser- the thorough analysis of previous results. In this sense, the vation studies (Franklin, 1993; Kremen et al., 1993; New, work now presented is also not fully standardized but it is 1999a,b). When corrected for knowledge bias, data on arthro- a first step in a series of similar studies that intend to reach pods show that risk of extinction is as real for them as it is for a truly standardized and optimized protocol. vertebrates (Thomas and Morris, 1994; MacKinney, 1999; The Mediterranean region is considered one of the most Dunn, 2005). As a consequence of the current data and knowl- important global biodiversity hotspots, the only one partly lo- edge deficit, nowadays most conservation studies and deci- cated in Europe (Cowling and Samways, 1994; Mittermeier sions necessarily rely on data predominately from plants, et al., 1998; Me´dail and Que´zel, 1999; Myers et al., 2000; Brooks birds and mammals, but their function as good indicators et al., 2002). Shortcuts for the rapid assessment of spider rich- for conservation priorities which ought to be relevant for all ness in the Mediterranean have already been proposed by us- other living beings still needs to be proved. When this as- ing higher taxa surrogates (Cardoso et al., 2004a) or indicator sumption is tested on comparable datasets, at least birds taxa (Cardoso et al., 2004b), but no field protocol has been pro- and mammals appear to be rather ineffective in relation to posed for the habitats in the region (but see Jime´nez-Valverde the use of various arthropod taxa (e.g. Lund and Rahbek, and Lobo, 2005, 2006). 2000, 2002; Lund, 2002). The objectives of this study were to determine: (1) the fea- Standardization and calibration of sampling methods and sibility of sampling the entire community over a limited time- protocols are fundamental issues for the comparison of sites frame; (2) the factors (faunal depletion, time of day, collector and the definition of priority areas for conservation (Stork, experience or method) which most strongly affect collecting 1994; Cannon, 1997; Jones and Eggleton, 2000; Kitching et al., productivity and sample composition; and (3) the richness es- 2001; Pereira and Cooper, 2006). The non-existence of such timators which present the best results for intensive semi- protocols is one of the reasons why arthropods are usually rel- quantitative sampling of spiders. This work is part of a larger egated to a secondary place or even not considered in conser- project that intends to create a standardized and optimized vation programs (New, 1999a,b). Nevertheless, standardized sampling protocol for Mediterranean spiders; ultimately it protocols have already been proposed and used for some ar- will be applicable in most habitats of this region. thropods like ants (Agosti and Alonso, 2000) and carabid bee- tles (Niemela et al., 2000). Spiders (order Araneae) are one of the most speciose ar- 2. Materials and methods thropod orders. It has been estimated that one hectare of trop- ical forest may support between 300 and 800 species of spiders 2.1. Study site at any given time (Coddington et al., 1991). They are among the most numerous arthropods in many samples in all kinds The study was undertaken in a cork oak (Quercus suber L.) of habitats (e.g. Basset, 1991; Borges and Brown, 2004). woodland of about 10 ha, located in the Vale da Rasca, Arra´- Spiders have been mostly sampled by methods originally bida Nature Park (PNA), in central Portugal, at an altitude of intended for other arthropods, which allows for comparable 60 m (N 3830.700, W 00858.800). The canopy density was rel- samples between sites or dates. However, only a very limited atively sparse, with trees varying from 4 to 8 m in height. The number of methods are simultaneously applied in most stud- understorey was equally sparse although continuous, and ies, leaving many species undetected due to the failure of cov- dominated by rockrose (Cistus ladanifer L.), heather (Erica ering most microhabitats. Spiders are very diverse in their arborea L.) and other herbaceous plant species typical of Med- ways of life, and sampling them requires a combination of iterranean habitats. The ground was mostly bare and leaf lit- methods. Also, many sampling programs neglect to consider ter cover was sparse. the time of day as an important variable. Neither do they pay any attention to the collectors’ experience nor their influ- ence on the composition of the assemblage collected. For spi- 2.2. Sampling procedures ders, complete protocols or sampling packages that gather information from a series of semi-quantitative methods A single square sampling plot, 100 m wide and hereafter called have proved to be the most effective (Coddington et al., ‘‘sampling plot’’ or just ‘‘plot’’, was established in the centre of 1991). They achieve the best results with a minimum of effort. the habitat, at least 100 m from the edge, in order to reduce Such methods are adequate for capturing a large number of spatial edge effects. The majority of the sampling was concen- species as well as large numbers of specimens with replicate trated in this plot with some additional sampling done outside samples. Exhaustive sampling protocols directed at spiders, the area yet still inside the habitat. mainly based on semi-quantitative methods, have been Sampling followed a balanced design, which means that experimented with in several parts of the world, i.e. Bolivia the same effort was applied to each factor, i.e., sampling (Coddington et al., 1991), Peru (Silva and Coddington, 1996), day, time of day, collector and method. These independent the USA (Coddington et al., 1996; Dobyns, 1997; Toti et al., variables were tested for their effect on the richness, abun- 2000), Tanzania (Sørensen et al., 2002) and Denmark (Scharff dance and composition of samples. 116 acta oecologica 33 (2008) 114–127 A semi-quantitative sampling design was defined based on Brush – Brushing the tree bark with a soft large brush was the time spent collecting each sample. One person-hour of ef- used to dislodge mainly cryptic spiders (8 samples, none from fective fieldwork was used as a sample unit.
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