Perspectives in Plant Ecology, Evolution and Systematics 40 (2019) 125485 Contents lists available at ScienceDirect Perspectives in Plant Ecology, Evolution and Systematics journal homepage: www.elsevier.com/locate/ppees Handbook of standardized protocols for collecting plant modularity traits T Jitka Klimešováa,b,1, Jana Martínkováa, Juli G. Pausasc, Moemy Gomes de Moraesd, Tomáš Herbenb,e, Fei-Hai Yuf,g, Javier Puntierih,i, Peter A. Veskj, Francesco de Belloa,c,k, Štěpán Janečeka,l, Jan Altmana, Beatriz Appezzato-da-Glóriam, Alena Bartuškováa, Alan Crivellaron,o, Jiři Doležala,k, Jacqueline P. Ottp, Susana Paulaq, Renáta Schnablováe,r, ⁎ Fritz H. Schweingrubers, Gianluigi Ottaviania, ,1 a Institute of Botany, The Czech Academy of Sciences, Dukelská 135, Třeboň, 37982, Czech Republic b Department of Botany, Faculty of Science, Charles University, Benátská 2, Praha, 12844, Czech Republic c CIDE-CSIC, C. Naquera Km 4.5, Montcada, Valencia, 46113, Spain d Department of Botany, Institute of Biological Sciences, Federal University of Goias, Goiânia, Goiás, 74690-900, Brazil e Institute of Botany, The Czech Academy of Sciences, Zámek 1, Průhonice, 25243, Czech Republic f Institute of Wetland Ecology & Clone Ecology, Taizhou University, Taizhou, 318000, China g Zhejiang Provincial Key Laboratory of Plant Evolutionary Ecology and Conservation, Taizhou University, Taizhou, 318000, China h Instituto de Investigaciones en Recursos Naturales, Agroecología y Desarrollo Rural, Universidad Nacional de Río Negro, Sede Andina, John O’Connor 181, San Carlos de Bariloche, 8400, Argentina i Centro Científico y Tecnológico Patagonia Norte, CONICET, San Carlos de Bariloche, 8400, Argentina j School of BioSciences, The University of Melbourne, Melbourne, Victoria, 3010, Australia k Department of Botany, Faculty of Sciences, University of South Bohemia, Na Zlaté Stoce 1, České Budějovice, 37005, Czech Republic l Department of Ecology, Faculty of Science, Charles University, Viničná 7, Prague, 12801, Czech Republic m Biological Sciences Department, ESALQ, University of São Paulo, Av Pádua Dias 11, Piracicaba, Sao Paulo, 13418-900, Brazil n Department TESAF-University of Padova, Viale dell’Università 16, Legnaro (PD), 35020, Italy o Department of Geography, University of Cambridge, Downing Place, Cambridge, CB2 3EN, United Kingdom p Forest and Grassland Research Laboratory, US Forest Service, Rocky Mountain Research Station, 8221 South Highway 16, Rapid City, South Dakota, 57702, USA q ICAEV, Universidad Austral de Chile, Campus Isla Teja, Casilla 567, Valdivia, Chile r Institute of Experimental Botany, The Czech Academy of Sciences, Rozvojová 263, Praha 6, 16502, Czech Republic s Swiss Federal Institute for Forest Snow and Landscape Research WSL, Zürcherstrasse 111, Birmensdorf, 8903, Switzerland ARTICLE INFO ABSTRACT Keywords: Plant modularity traits relevant to functions of on-spot persistence, space occupancy, resprouting after dis- Bud bank turbance, as well as resource storage, sharing, and foraging have been underrepresented in functional ecology so Carbohydrate storage far. This knowledge gap exists for multiple reasons. First, these functions and related traits have been considered Clonality less important than others (e.g., resource economics, organ-based traits). Second, collecting data for modularity Competitive ability traits can be difficult. Third, as a consequence of the previous points, there is a lack of standardized collection Ecosystem functioning protocols. We now feel the time is ripe to provide a solid conceptual and terminological framework together with Functional traits Longevity comparable protocols for plant modularity traits that can be applicable across species, regions and biomes. We Plant modules identify a suite of 14 key traits, which are assembled into five groups. We discuss the functional relevance of On-spot persistence each trait, supplying effective guidelines to assist in the use and selection of the most suitable traits inrelationto Resprouting after disturbance specific research tasks. Finally, we are convinced that the systematic study and widespread assessment ofplant Space occupancy modularity traits could bridge this knowledge gap. As a result, previously overlooked key functions could be incorporated into the functional ecology research-agenda, thus providing a more comprehensive understanding of plant and ecosystem functioning. ⁎ Corresponding author. E-mail address: [email protected] (G. Ottaviani). 1 Equal contribution. https://doi.org/10.1016/j.ppees.2019.125485 Received 8 October 2018; Received in revised form 10 July 2019; Accepted 12 August 2019 Available online 26 August 2019 1433-8319/ © 2019 Published by Elsevier GmbH. J. Klimešová, et al. Perspectives in Plant Ecology, Evolution and Systematics 40 (2019) 125485 1. Introduction primary aim of our initiative is to fill this gap, namely to present standardized collection protocols for 14 key modularity traits (Table 1) Plant functional ecology started as a discipline describing plant life potentially applicable across species and biomes. We do so by offering i) history strategies in small geographical regions (e.g., Grime, 1979) and a robust conceptual and terminological framework, ii) synthetic de- developed to broad-scale, comparative syntheses (Wright et al., 2004; scription of functional relevance of each trait, and iii) detailed protocols Chave et al., 2009). This has been facilitated by using trait-based ap- for trait assessment including field- and lab-procedures. proaches that allowed the identification of i) important axes of plant differentiation, such as size and economics spectra reflective ofallo- 1.1. Aims and structure of the handbook cation trade-offs (e.g., Westoby, 1998; Wright et al., 2004), and ii) drivers of community assembly (Götzenberger et al., 2012). At the same We identified 14 plant modularity traits arranged into five major time, functional ecology is falling short with regards to some crucial trait-groups: Anatomical features, Bud bank, Carbohydrate storage, plant functions, such as belowground nutrient acquisition, clonal mul- Clonality, and Longevity and growth (Table 1). Based on our expertise tiplication, and vegetative regeneration (Laliberté, 2017; Ottaviani working in different biomes and focused on different growth forms,we et al., 2017). This gap has been partly caused by focusing on a set of selected what we consider to be the most relevant plant traits that are functional traits that are mainly associated with resource acquisition informative on functions of on-spot persistence, space occupancy, re- and sexual reproduction (Westoby, 1998; Wright et al., 2004; Chave sprouting after disturbance, resource storage, sharing and foraging et al., 2009). These traits are, in most cases, relatively easy to collect (Weiher et al., 1999; Ottaviani et al., 2017; Klimešová et al., 2018). and measurable on aboveground organs, namely leaves, stems, and While some of these traits are fairly well-known (e.g., clonal and bud flowers/fruits, for which standardized procedures were developed bank traits) for a few biomes, community types or growth forms, others (Cornelissen et al., 2003; Pérez-Harguindeguy et al., 2013). (e.g., anatomical features, carbohydrate storage, longevity) have rarely So far, most trait-based studies and approaches rarely considered been collected for large sets of species in a comparable, meaningful way that plant organs are assembled into modules – defined as the basic for functional ecology. architectural building blocks of a plant which are accumulated during To better explain and guide readers through plant modularity, we growth (Fig. 1). The position, size and persistence of the modules i) present four sections that are not specific to any one trait. The first generate the intra- and interspecific variability of plant shapes, and ii) section on “Plant modularity” provides a general framework and voca- most importantly, affect plant functioning. Understanding plant mod- bulary for trait description and assessment. The second part reports the ularity has been a major challenge for comparative plant ecologists for “Field sampling protocol” where essential information and guidance re- quite some time (Raunkiaer, 1934; Hallé et al., 1978; White, 1979). We lated to plant collection are described (see also Appendix I now feel the time is ripe to consistently incorporate plant modularity “Standardized site description and sample labelling“). The third section on into functional ecology. “Experimental assessment of resprouting after disturbance” (included as Plant modularity traits are ecologically relevant, as they can effec- Appendix II) illustrates methods and ideas for testing functionality of tively capture functions of on-spot persistence, space occupancy, re- the proposed bud bank traits. The fourth part is the “Glossary” (included sprouting after disturbance, resource storage, sharing, and foraging (for as Appendix III) explaining specialized terminology. Plant ecologists c overviews, see Ottaviani et al., 2017 and Klimešová et al., 2018). Yet, should be aware that the assessment of plant modularity traits is time- they remain largely understudied compared to e.g., traits related to consuming and requires destructive sampling of plants. We are con- resource acquisition. We therefore consider it essential to provide vinced that combining plant modularity traits with other more widely standardized protocols for fully including plant modularity traits into studied traits is necessary