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How Plot Shape and Spatial Arrangement Affect Plant Species How plot shape and spatial arrangement affect plant species richness counts: implications for sampling design and rarefaction analyses Behlül Guler, Anke Jentsch, Iva Apostolova, Sandor Bartha, Juliette Bloor, Giandiego Campetella, Roberto Canullo, Judit Hazi, Jurgen Kreyling, Julien Pottier, et al. To cite this version: Behlül Guler, Anke Jentsch, Iva Apostolova, Sandor Bartha, Juliette Bloor, et al.. How plot shape and spatial arrangement affect plant species richness counts: implications for sampling design andrar- efaction analyses. Journal of Vegetation Science, Wiley, 2016, 27 (4), pp.692-703. 10.1111/jvs.12411. hal-01555265 HAL Id: hal-01555265 https://hal.archives-ouvertes.fr/hal-01555265 Submitted on 3 Jul 2017 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Journal of Vegetation Science 27 (2016) 692–703 How plot shape and spatial arrangement affect plant species richness counts: implications for sampling design and rarefaction analyses Behlul€ Guler,€ Anke Jentsch, Iva Apostolova, Sandor Bartha, Juliette M.G. Bloor, Giandiego Campetella, Roberto Canullo, Judit Hazi, Jurgen€ Kreyling, Julien Pottier, Gabor Szabo, Tsvetelina Terziyska, Emin Ugurlu, Camilla Wellstein, Zita Zimmermann & Jurgen€ Dengler Keywords Abstract Biodiversity; Discontiguous; Effective area; Grassland; Sampling unit; Scale dependence; Questions: How does the spatial configuration of sampling units influence Spatial autocorrelation; Spatial extent; Spatial recorded plant species richness values at small spatial scales? What are the con- grain; Species–area relationship; Species– sequences of these findings for sampling methodology and rarefaction analyses? extent relationship; Vegetation plot Location: Six semi-natural grasslands in Western Eurasia (France, Germany, Abbreviations Bulgaria, Hungary, Italy, Turkey). Ae = effective area; SAR = species–area 9 relationship; SAER = species–area–extent Methods: In each site we established six blocks of 40 cm 280 cm, subdivided relationship; SER = species–extent relationship; into 5 cm 9 5 cm micro-quadrats, on which we recorded vascular plant species STAR = species–time–area relationship; presence with the rooted (all sites) and shoot (four sites) presence method. Data STR = species–time relationship. of these micro-quadrats were then combined to achieve larger sampling units of 0.01, 0.04 and 0.16 m² grain size with six different spatial configurations Received 11 October 2015 Accepted 29 February 2016 (square, 4:1 rectangle, 16:1 rectangle, three variants of discontiguous randomly Co-ordinating Editor: Janos Podani placed micro-quadrats). The effect of the spatial configurations on species rich- ness was quantified as relative richness compared to the mean richness of the square of the same surface area. Guler,€ B. ([email protected])1,2, Jentsch, A. ([email protected])2,3, Results: Square sampling units had significantly lower species richness than Apostolova, I. ([email protected])4, other spatial configurations in all countries. For 4:1 and 16:1 rectangles, the Bartha, S. increase of rooted richness was on average about 2% and 8%, respectively. In 5,6 ([email protected]) , contrast, the average richness increase for discontiguous configurations was 7%, Bloor, J.M. ([email protected])7, 17% and 40%. In general, increases were higher with shoot presence than with Campetella, G. ([email protected])8, rooted presence. Overall, the patterns of richness increase were highly consis- Canullo, R. ([email protected])8, tent across six countries, three grain sizes and two recording methods. Hazi, J. ([email protected])9, Conclusions: Our findings suggest that the shape of sampling units has negligi- Kreyling, J. (juergen.kreyling@ – uni-greifswald.de)10, ble effects on species richness values when the length width ratio is up to 4:1, Pottier, J. ([email protected])7, and the effects remain small even for more elongated contiguous configurations. Szabo, G. ([email protected])5, In contrast, results from discontiguous sampling units are not directly compara- Terziyska, T. ([email protected])4, ble with those of contiguous sampling units, and are strongly confounded by Ugurlu, E. ([email protected])1,11, spatial extent. This is particularly problematic for rarefaction studies where spa- 12 Wellstein, C. ([email protected]) , tial extent is often not controlled for. We suggest that the concept of effective Zimmermann, Z. 5 area is a useful tool to report effects of spatial configuration on richness values, ([email protected]) , – Dengler, J. (corresponding author, and introduce species extent relationships (SERs) to describe richness increases [email protected])3,13,14 of different spatial configurations of sampling units. 1Biology, Faculty of Science & Letters, Celal Bayar University, Muradiye Campus, 45040 Manisa, Turkey; 2Disturbance Ecology, University of Bayreuth, Universit€atsstr. 30, 95447 Bayreuth, Germany; 3Bayreuth Center of Ecology and Environmental Research (BayCEER), University Journal of Vegetation Science 692 Doi: 10.1111/jvs.12411 © 2016 International Association for Vegetation Science B. Guler€ et al. Species richness vs plot shape and arrangement of Bayreuth, Universit€atsstr. 30, 95447 7Grassland Ecosystem Research Team, UR874, 11Forest Engineering, Faculty of Forestry, Bursa Bayreuth, Germany; INRA, 5 Chemin de Beaulieu, 63039 Clermont- Technical University, 152 Evler Str. 2/10, 16330 4Institute of Biodiversity and Ecosystem Ferrand, France; Yıldırım, Bursa, Turkey; Research, Bulgarian Academy of Sciences, 23 8Plant Diversity and Ecosystems Management 12Faculty of Science and Technology, Free Acad. G. Bonchev Str., 11113 Sofia Sofia, Unit, School of Biosciences & Veterinary University of Bozen, Piazza Universita 5, 39100 Bulgaria; Medicine, University of Camerino, Via Pontoni 5, Bozen, Italy; 5Institute of Ecology and Botany, MTA Centre for 62032 Camerino (MC), Italy; 13Plant Ecology, University of Bayreuth, Ecological Research, Alkotmany u. 2–4, 2163 9Institute of Botany and Ecophysiology, Szent Universit€atsstr. 30, 95447 Bayreuth, Germany; Vacr atot, Hungary; Istvan University, Pater K. u. 1, 2100 God€ oll€ o,} 14German Centre for Integrative, Biodiversity 6School of Plant Biology, The University of Hungary; Research (iDiv) Halle-Jena-Leipzig, Deutscher Western Australia, 35 Stirling Highway, Crawley, 10Experimental Plant Ecology, Institute of Botany Platz 5e, 04103 Leipzig, Germany WA 6009, Australia; and Landscape Ecology, University of Greifswald, Soldmannstr. 15, 17487 Greifswald, Germany; has two main drivers (Nekola & White 1999): first, the Introduction distance decay in climate, soil, topography, composition In ecology and conservation, species richness is probably of species of other trophic levels, as well as of human the most frequently used measure of diversity because it is land-use patterns creates environmental filters that easily measurable in a multitude of different situations and become, on average, more and more dissimilar with dis- comprehensible even for non-specialists. Accurate quan- tance, thus selecting for increasingly different plant spe- tification of species richness requires appropriate sampling cies composition. Second, biological processes of the decisions regarding sample size, the selection and configu- plant species themselves are strongly distance-depen- ration of sampling units (in vegetation science called quad- dent, such as lateral spread, dispersal, gene flow and rats, vegetation plots or just plots), as well as their size and species–species interactions, including facilitation or par- shape (Kenkel et al. 1989; Bacaro et al. 2015). Given that asitism. Such biological processes can even overrule – to species richness on average increases with area (Arrhenius some extent – environmental filtering, leading to the 1921; Connor & McCoy 1979; Dengler 2009), comparisons occurrence of species in ecologically suboptimal habitats of species richness counts are usually only meaningful which are spatially close to ecologically optimal source between sampling units of the same grain size. However, habitats. This phenomenon occurs both at local and at there are at least three other factors that can distort com- biogeographic distances and has been termed ‘mass parisons of species richness for a given area: (1) the shape effect’ (Shmida & Wilson 1985) or ‘vicinism’ (Zonneveld of the sampling unit used to assess species richness (elon- 1994). Generally, distance decay in compositional simi- gated vs compact); (2) the dispersion or contingency of larity of plant assemblages should be relatively low subplots that constitute the overall area to be quantified when there are less pronounced environmental gradi- (spatial arrangement; contiguous vs discontiguous); and ents and/or when species with good dispersal ability of (3) in the case of plants and other sessile organisms, the diaspores and genes are considered, and vice versa.Ifwe method by which an individual
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