Seasonality of Spiders (Araneae) in Mediterranean Ecosystems and Its Implications in the Optimum Sampling Period
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Ecological Entomology (2007), 32, 516–526 DOI: 10.1111/j.1365-2311.2007.00894.x Seasonality of spiders (Araneae) in Mediterranean ecosystems and its implications in the optimum sampling period PEDRO CARDOSO 1,2 , ISRAEL SILVA 2 , NUNO G. DE OLIVEIRA 2 a n d ARTUR R. M. SERRANO 2 1 Zoological Museum and Centre of Macroecology, University of Copenhagen, Denmark , 2Centre of Environmental Biology, Faculty of Sciences, University of Lisbon, Portugal Abstract. 1. Fields such as ecology, macroecology, and conservation biology rely on accurate and comparable data. This is especially important for mostly unknown and megadiverse taxa such as spiders and regions such as the Mediterranean. Short-term sampling programmes are increasingly seen as the best option for sampling spiders. Comparability of results, however, demands standard procedures both in methodology and in sampling period. Cost-efficiency dictates that this period should be the most species rich. 2. Pitfall trapping was conducted in 23 sites from north to south Portugal, comprising three large-scale environmental zones and many different habitat types, during 10 months in each site. The annual richness pattern, differences in this pattern between areas and habitats, the complementarity between sampling periods and possible environmental correlates of richness were studied. 3. May and June present the optimal time for collecting spiders in Mediterranean areas. Northern areas have a later peak in richness and dense tree-cover sites offer more flexibility for sampling, with a higher proportion of species present at each period throughout the year. 4. Day length is the environmental factor most correlated with species richness. Maximum daily temperature may reduce richness, especially in southernmost areas, where summer temperatures can be extremely harsh. 5. It is recommended that short-term sampling programmes, intended to give a reasonable picture of spider communities in Portugal and in the Iberian Peninsula (and possibly extending to all the Mediterranean), should be conducted during May or June, with variable flexibility according to area and habitat. The proposed suggestions should appeal to everyone working in the field, given the cost-efficiency and comparability of results by adopting a common standardised approach. Key words. Arachnids , biodiversity , complementarity , cost-effi ciency , inventory , phenology , pitfall , Portugal , Spain , standardised protocols . Introduction even misleading; however, incomplete data sets, if not exceed- ingly data deficient, can also provide good basis for different Biological surveys inevitably produce samples of the total applications ( Gaston & Rodrigues, 2003 ). Therefore, standardi- fauna, not complete inventories. If the databases used for sation and calibration of sampling methods and protocols are (macro)ecological studies, or conservation priority analyses, are fundamental for the comparison of sites or the definition of pri- incomplete or inadequate, the result will often be inaccurate or ority areas for conservation ( Stork, 1994; Cannon, 1997; Jones & Eggleton, 2000; Kitching et al. , 2001 ). Correspondence: Pedro Cardoso, Agricultural Sciences Department, Once one has chosen where to sample, there is the decision of CITA-A, University of Azores, Terra-Chã, 9701 – 851 Angra do how it is done. The sampling protocol and respective methodol- Heroísmo, Portugal. E-mail: [email protected] ogy are obviously taxon-specific. For spiders, complete protocols, © 2007 The Authors 516 Journal compilation © 2007 The Royal Entomological Society Mediterranean spiders optimum sampling period 517 or sampling packages, that gather information from a series of Materials and methods semiquantitative methods are considered the best approach for comprehensive and standardised sampling ( Coddington et al. , Study sites 1991 ). They allow optimum results with a minimum effort, al- though this is always considerable. Such methods are suitable for Four protected areas in Portugal were studied during three capturing a large number of species and simultaneously a large consecutive years, one year each area ( Fig. 1 ): (i) Douro number of specimens with replicate samples ( Coddington et al. , Internacional Nature Park (PNDI), located in north-eastern 1996; Dobyns, 1997; Toti et al. , 2000; Sørensen et al. , 2002; Portugal, sampled during 2001; (ii) Serras de Aire e Candeeiros Scharff et al. , 2003 ). Nature Park (PNSAC), located in central western Portugal, sam- Besides deciding where and how to sample the question of pled during 2002; (iii) Paúl do Boquilobo Nature Reserve when to sample must also be addressed. The answer relies (RNPB), located very close to PNSAC, sampled during 2002; fundamentally on the study of fluctuations in species rich- and (iv) Vale do Guadiana Nature Park (PNVG), located in ness of the target taxon according to the time of year, its phe- south-eastern Portugal, sampled during 2003. These areas are nology. Probably more important, it also relies on the degree located in the three environmental zones defined by Metzger to which such fluctuations are predictable year after year, its et al. (2005) as being present in Portugal – Lusitanian (PNSAC, seasonality (as defined by Wolda, 1988 ). Highly seasonal RNPB), Mediterranean North (PNDI) and Mediterranean South phenomena can allow the reliable planning of fieldwork (PNVG). Only one of the zones present in the Mediterranean activities. Many studies are devoted to seasonal changes in biome was not sampled, the aptly named Mediterranean abundance, the number of captured individuals (e.g. Chatzaki Mountains. et al. , 1998, 2005a , b ) or biomass (e.g. Pearson & Derr, 1986 ); In each protected area, most of its habitat types were sam- however, the focus of this study is on species richness, not pled, summing 23 sites. A complete data set was therefore cre- abundance or biomass, which mainly concerns individuals. ated, which allowed the comparison between many different Fluctuations in the richness of arthropods are usually much habitat types in the different geographical areas and respective higher in temperate than in tropical regions, with the latter environmental zones ( Table 1 ). having a high proportion of species that are present through- out the year ( Novotny & Basset, 1998 ). None the less, large seasonal differences in spider community composition, rich- Sampling periods ness or abundance are invariably found, even surpassing habitat differences in many cases ( Weeks & Holtzer, 2000; Although the ideal situation would have been to sample all sites Whitmore et al. , 2002 ). Given these facts, one has to con- simultaneously, this was logistically unfeasible, given that the sider the implementation of long-lasting sampling pro- objective demanded to study areas as distant from each other as grammes, of at least 1 year if not longer. Owing to the lack of possible. Logistical reasons have also made it impossible to sam- human, material, and time resources, however, this is largely ple continuously during a whole year at each area. Given that unfeasible. It is therefore imperative to optimise the duration December and January were thought a priori to be very poor in of protocols and the time of the year when they are imple- spider species, the remaining 10 sampled months were considered mented. This is best achieved by carrying out standard mini- as representative of the entire year. All sites were sampled continu- mum sampling programmes, which minimise the number and ously, from the end of January (29 – 31, depending on the year) extension of collecting periods and simultaneously giving until the beginning of December (3 – 5). Sampling periods lasted the most complete picture of diversity of any site ( Oliver & for 2 weeks, providing 22 samples each year. Short-term sampling Beattie, 1996; Duelli, 1997; Duelli et al. , 1999; Zonneveld protocols usually take between 1 and 2 weeks of fieldwork and this et al. , 2003 ). All the protocols proposed to date have tried to 2-week period was adequate for the proposed objectives. answer the most obvious question: When should one collect to guarantee maximum diversity with minimum time spent on the task? Sampling method The first objective of this work was to choose the season that consistently presented highest richness in different geographi- The main objective is to recommend the best time of year for cal regions (corresponding to broad environmental zones as de- collecting as many species as possible independently of the fined by Metzger et al. , 2005 ) and habitat types in Portugal. methods to be employed. To test all or even most of the available Second, to understand if and how do season, area, and habitat capture methods for several years and in so many sites, however, type potentially influence the sampled species richness. Third, would be a formidable challenge. Therefore, a single method was to verify the effect on accumulated richness of increasing the chosen to meet the objectives, the one that could provide the most number and duration of samples. Fourth, to assess if richness is comprehensive information with the least effort. One of the most correlated to any seasonal and easily predicted environmental frequently employed and cost-effective methods for capturing factor. Finally, to give recommendations on the best sampling spiders and other arthropods is pitfall trapping (e.g. Churchill, procedure for maximising spider species richness spending the 1993; Samu & Lövei, 1995; Brennan et al. , 1999; Churchill & least