A Handbook for the Standardised Sampling of Plant Functional Traits in Disturbance-Prone Ecosystems, with a Focus on Open Ecosystems

A Handbook for the Standardised Sampling of Plant Functional Traits in Disturbance-Prone Ecosystems, with a Focus on Open Ecosystems

CSIRO PUBLISHING Australian Journal of Botany, 2021, 69, 110 Corrigendum https://doi.org/10.1071/BT20048_CO A handbook for the standardised sampling of plant functional traits in disturbance-prone ecosystems, with a focus on open ecosystems B. J. Wigley , T. Charles-Dominique, G. P. Hempson, N. Stevens, M. te Beest , S. Archibald, W. J. Bond, K. Bunney, C. Coetsee, J. Donaldson, A. Fidelis, X. Gao, J. Gignoux, C. Lehmann, T. J. Massad, J. J. Midgley, M. Millan, D. Schwilk, F. Siebert, C. Solofondranohatra, A. C. Staver, Y. Zhou and L. M. Kruger Australian Journal of Botany 68(8) 473–531. doi:10.1071/BT20048 The authors of the above-mentioned paper regret to inform readers that one of the authors, Mariska te Beest, was incorrectly referred to in the author list and the Author contributions section as M. TeBeest. The author list should have been published as above. Please also note that Dr te Beest’s ORCID ID (https://orcid.org/0000-0003-3673-4105) has now been added to assist with connecting this work to her online profile. Journal compilation Ó CSIRO 2021 www.publish.csiro.au/journals/ajb CSIRO PUBLISHING Australian Journal of Botany, 2020, 68, 473–531 Turner Review No. 25 https://doi.org/10.1071/BT20048 A handbook for the standardised sampling of plant functional traits in disturbance-prone ecosystems, with a focus on open ecosystems B. J. Wigley A,B,X, T. Charles-Dominique C, G. P. Hempson D, N. Stevens E,F, M. TeBeestG,H,I, S. Archibald D, W. J. BondJ, K. BunneyK, C. CoetseeL,B, J. DonaldsonM, A. FidelisN, X. GaoO, J. Gignoux C, C. LehmannP,Q, T. J. MassadR, J. J. MidgleyJ, M. Millan D,F, D. SchwilkO, F. SiebertS, C. SolofondranohatraT,U, A. C. StaverV, Y. ZhouV and L. M. KrugerW,J ADepartment of Environmental Biology, Sapienza University of Rome, Piazzale Aldo Moro 5, I-00185 Rome, Italy. BSchool of Natural Resource Management, Nelson Mandela University, George, 6529, South Africa. CCNRS UMR7618, Institute of Ecology and Environmental Sciences, Paris, Sorbonne University, 4 Place Jussieu, F-75252 Paris, France. DCentre for African Ecology, School of Animal, Plant and Environmental Sciences, University of the Witwatersrand, Johannesburg, 2050, South Africa. EEnvironmental Change Institute, School of Geography and the Environment, University of Oxford, Oxford, OX1 3QY, UK. FDepartment of Botany and Zoology, Stellenbosch University, Private Bag X1, Matieland 7602, South Africa. GCopernicus Institute of Sustainable Development, Utrecht University, PO Box 80115, NL-3508 TC Utrecht, Netherlands. HCentre for African Conservation Ecology, Nelson Mandela University, PO Box 77000, Port Elizabeth, 6031, South Africa. ISouth African Environmental Observation Network (SAEON), Grasslands, Forests, Wetlands Node, Montrose, 3201, South Africa. JDepartment of Biological Sciences, University of Cape Town, Rondebosch, 7701 South Africa. KDepartment of Zoology and Entomology, University of Pretoria, Hatfield, South Africa. LScientific Services, Kruger National Park, Private Bag X402, Skukuza, 1350, South Africa. MDepartment of Biology, Wake Forest University, 049 Winston-Salem, NC 27109, USA. NUniversidade Estadual Paulista (UNESP), Instituto de Biociências, Lab of Vegetation Ecology, Avenida 24-A 1515, 13506-900, Rio Claro, Brazil. ODepartment of Biological Sciences, Texas Tech University, Lubbock, TX 79409, USA. PTropical Diversity, Royal Botanic Garden Edinburgh, Edinburgh, EH35NZ, UK. QSchool of GeoSciences, University of Edinburgh, Edinburgh, EH9 3FF, UK. RDepartment of Scientific Services, Gorongosa National Park, Sofala, Mozambique. SUnit for Environmental Sciences and Management, North-West University, Private Bag X6001, Potchefstroom 2520, South Africa. TLaboratoire de Botanique, Département de Biologie et Ecologie Végétales, Faculté des Sciences, Université d’Antananarivo BP 906, Antananarivo 101, Madagascar. UKew Madagascar Conservation Centre, Lot II J 131 B, Ambodivoanjo-Ivandry, Antananarivo 101, Madagascar. VDepartment of Ecology and Evolutionary Biology, Yale University, New Haven, CT 06511, USA. WOrganization for Tropical Studies, PO Box 33, Skukuza, 1350, South Africa. XCorresponding author. Email: [email protected] Abstract. Plant functional traits provide a valuable tool to improve our understanding of ecological processes at a range of scales. Previous handbooks on plant functional traits have highlighted the importance of standardising measurements of traits to improve our understanding of ecological and evolutionary processes. In open ecosystems (i.e. grasslands, savannas, open woodlands and shrublands), traits related to disturbance (e.g. herbivory, drought, and fire) play a central role in explaining species performance and distributions and are the focus of this handbook. We Journal compilation Ó CSIRO 2020 Open Access CC BY-NC-ND www.publish.csiro.au/journals/ajb 474 Australian Journal of Botany B. J. Wigley et al. provide brief descriptions of 34 traits and list important environmental filters and their relevance, provide detailed sampling methodologies and outline potential pitfalls for each trait. We have grouped traits according to plant functional type (grasses, forbs and woody plants) and, because demographic stages may experience different selective pressures, we have separated traits according to the different plant life stages (seedlings saplings and adults). We have attempted to not include traits that have been covered in previous handbooks except for where updates or additional information was considered beneficial. Keywords: demographic bottlenecks, disturbance, drought, fire, forbs, grasses, herbivory, plant functional traits, saplings, seedlings, woody plants. Received 6 May 2020, accepted 8 October 2020, published online 30 November 2020 Introduction and discussion (e.g. thicket or forest) is the presence of shade-intolerant fi The standardisation of methodologies used for measuring and typically C4 grasses. This de nition allowed for the fi plant functional traits at the global scale allows us to (1) reclassi cation of many systems that were previously better understand the selective forces shaping the thought of as degraded ecosystems, as ecosystems with functioning of plant communities, (2) better describe shared ecological functioning on different continents (Bond ecosystems in terms of a limited number of ecological et al. 2008;Ratnamet al. 2011; Ratnam et al. 2016), and component types, and (3) better predict species-response opened opportunities for global comparisons. Although trees rules to ecosystem perturbations (Cornelissen et al. 2003; and grasses are typically used to identify savannas, forbs Pérez-Harguindeguy et al. 2013; Díaz et al. 2016). The should not be overlooked because they often contribute the standardised measurement of traits also allows for drawing highest levels of diversity in old-growth savannas (Zaloumis comparisons among different regions of the globe (Rusch et al. and Bond 2011). Throughout this handbook, we regard the 2003). While previous trait handbooks by Cornelissen et al. functioning of grasslands, shrublands and savannas to be (2003) and Pérez-Harguindeguy et al.(2013)haveimmense similar, because of the strong parallels in their drivers. pertinence to open ecosystems (i.e. areas with climates suitable Although much of the work included in this trait handbook for forest, but dominated, instead, by grasslands, savannas, has focused on African, South American and Asian savannas open woodlands or shurblands; Bond 2019), they do not and grasslands, we consider these effective exemplars of capture the full set of traits that are relevant to the disturbance-driven ecosystems and we anticipate that the functioning of these widespread ecosystems. The selection traits covered here can be applied to other open ecosystems. pressures faced by plants in open ecosystems are largely fi driven by disturbance, often in the form of re and Demographic bottlenecks herbivory (but also seasonal flooding or drought). The plants found in these open ecosystems have, therefore, had The strong effect of disturbances on savanna structure and to evolve a diverse suite of plant traits, many of which are composition is shown both by a demographic bottleneck fi related to resisting or tolerating the dominant disturbance speci c to savannas, characterised by a high mortality of fl agent(s) (Bond and Midgley 2001; Bond 2005). woody saplings within the ame and herbivore zones (Higgins et al. 2000; Sankaran et al. 2005; Wakeling et al. 2011; Hoffmann et al. 2012), and by the distinct vertical Open ecosystems: savannas, grasslands and shrublands distribution of vegetation when fire or megaherbivores such Open ecosystems, largely dominated by shade-intolerant as elephant are present (Higgins et al. 2007;Asneret al. 2009; plants, occur in consumer-controlled (i.e. disturbance-prone) Smit et al. 2010;Midgleyet al. 2011; Asner and Levick 2012; environments that often have the potential to support closed Bond 2019). In mesic savannas, which burn frequently, tree canopy vegetation if not for the ecological processes that saplings must grow above the flame zone of grass-fuelled remove trees (Pausas and Bond 2020). The savanna biome, surface fires to escape top-kill and reach adult sizes (Trollope one of the largest terrestrial biomes (Lehmann et al. 2014), 1984). The fire trap, a concept first made explicit by Bell supports significant animal and plant biodiversity (Bond and (1984), refers to the 0–3-m zone that experiences repeated Parr 2010; Murphy et al. 2016) and provides ecosystem fires, resulting in the top-kill

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