Received: 14 August 2019 | Revised: 20 October 2019 | Accepted: 23 October 2019 DOI: 10.1111/btp.12737 ORIGINAL ARTICLE Guidelines for including bamboos in tropical ecosystem monitoring Belén Fadrique1 | Joseph W. Veldman2,3 | James W. Dalling4,5 | Lynn G. Clark6 | Lia Montti7,8 | Eduardo Ruiz-Sanchez9 | Débora C. Rother10,11 | Francisca Ely12 | William Farfan-Ríos13,14,15 | Paul Gagnon16 | Cecilia M. Prada4 | Juan Carlos Camargo García17 | Sonali Saha18 | Thomas T. Veblen19 | Ximena Londoño20 | Kenneth J. Feeley1,21 | Cara A. Rockwell22 1Department of Biology, University of Miami, Coral Gables, FL, USA 2Department of Ecology and Conservation Biology, Texas A&M University, College Station, TX, USA 3Instituto Boliviano de Investigación Forestal, Santa Cruz, Bolivia 4Department of Plant Biology, University of Illinois at Urbana-Champaign, Urbana, IL, USA 5Smithsonian Tropical Research Institute, Balboa, Republic of Panama 6Department of Ecology, Evolution, and Organismal Biology, Iowa State University, Ames, IA, USA 7Instituto de Investigaciones Marinas y Costeras-CONICET, Instituto de Geología de Costas y del Cuaternario-Universidad Nacional de Mar del Plata, Buenos Aires, Argentina 8Instituto de Biología Subtropical, Universidad Nacional de Misiones-CONICET, Puerto Iguazú, Argentina 9Departamento de Botánica y Zoología, Centro Universitario de Ciencias Biológicas y Agropecuarias, Universidad de Guadalajara, Zapopan, Mexico 10Departamento de Biologia Vegetal, Instituto de Biociências, Universidade Estadual de Campinas, Cidade Universitária, São Paulo, Brasil 11Departamento de Ciências Florestais, Universidade de São Paulo, Escola Superior de Agricultura “Luiz de Queiroz”, Piracicaba, Brasil 12Facultad de Ciencias, Instituto Jardín Botánico de Mérida, Universidad de los Andes, Mérida, Venezuela 13Living Earth Collaborative, Washington University in Saint Louis, St. Louis, MO, USA 14Center for Conservation and Sustainable Development, Missouri Botanical Garden, St. Louis, MO, USA 15Herbario Vargas (CUZ), Escuela Profesional de Biología, Universidad Nacional de San Antonio Abad del Cusco, Cusco, Peru 16Institute for Water Resources, U.S. Army Corps of Engineers, Alexandria, VA, USA 17Universidad Tecnológica de Pereira, Pereira, Colombia 18Miami Dade College, Miami, FL, USA 19Department of Geography, University of Colorado Boulder, Boulder, CO, USA 20Sociedad Colombiana del Bambú, Montenegro, Quindío, , Colombia 21Fairchild Tropical Botanical Garden, Coral Gables, FL, USA 22Department of Earth and Environment, International Center for Tropical Botany, Florida International University, Miami, FL, USA Correspondence Belén Fadrique, Department of Biology, Abstract University of Miami, Coral Gables, FL 33146, Bamboos are a diverse and ecologically important group of plants that have the po- USA. Email: [email protected] tential to modulate the structure, composition, and function of forests. With the aim of increasing the visibility and representation of bamboo in forest surveys, and Funding information National Science Foundation, Grant/Award to standardize techniques across ecosystems, we present a protocol for bamboo Number: DEB 1743249; Programa Bosques monitoring in permanent research plots. A bamboo protocol is necessary because Andinos Biotropica. 2020;00:1–17. wileyonlinelibrary.com/journal/btp © 2020 The Association for Tropical Biology and | 1 Conservation 2 | FADRIQUE ET AL. Associate Editor: Jennifer Powers Handling Editor: Ferry Slik measurements and sampling schemes that are well-suited to trees are inadequate for monitoring most bamboo species and populations. Our protocol suggests counting all bamboo culms (stems) in the study plot and determining bamboo dimensions based on two different approaches: (a) measuring a random subset of 60 culms and calculat- ing the average dimensions or (b) measuring all culms. With data from 1-ha plots in the Peruvian Andes, we show that both approaches provide very similar estimates of bamboo basal area. We suggest including all mature culms rooted inside change the to each plot from all woody bamboo species with maximum diameters ≥1 cm. We also present recommendations on how to collect vouchers of bamboo species for identification. Data collected according to our proposed protocols will increase our understanding of regional and global patterns in bamboo diversity and the role of bamboo in forest dynamics. Abstract in Spanish is available with online material. KEYWORDS Bambusoideae, forest monitoring, Peru, protocol, tropical ecology 1 | INTRODUCTION consensus view among a group of tropical bamboo experts and for- est ecologists on the current state of bamboo ecology. To that end, To fully understand the ecology of tropical forests, and their role in we present a protocol for bamboo monitoring that should serve as global climate dynamics, we must embrace their complexity. Forests a guide to standardize aboveground bamboo measurements across include a diversity of taxa with different life forms—not just trees— a wide diversity of ecosystems. Finally, to inform our guidelines and all of which have unique natural histories, physiologies, and inter- demonstrate the utility of the protocol, we include a section on actions. While short- and long-term forest monitoring efforts have Protocol justification using data collected in the Peruvian Andes. traditionally focused on tree demographic data (Bastin et al., 2018; Phillips et al., 2009), in the last few decades, research on palms, li- anas, epiphytes, and understory plants has revealed the important 1.1 | Bamboo life history, diversity, and distribution influence of these non-tree life forms on forest communities (Farris- Lopez, Denslow, Moser, & Passmore, 2014; Phillips et al., 2002; Ter Bamboos are the largest grasses on Earth and exhibit some of the Steege et al., 2013). For example, during most of the 20th century, li- fastest growth rates of all plants (Pearson, Pearson, & Gomez, 1994). anas (woody vines) were excluded from forest censuses and their ef- For example, Guadua weberbaueri (tropical South America) can fects on forest dynamics were largely ignored (Schnitzer & Bongers, grow 3 m/month in height during the rainy season (Silveira, 2001) 2002; Schnitzer & Carson, 2000). After three decades of extensive and Bambusa bambos (South-East Asia) can grow up to 30 cm/day research (Putz, 1984; Schnitzer, Putz, Bongers, & Kroening, 2015), we (Shanmughavel & Francis, 1996). Bamboos have great plasticity in now know that lianas are fierce competitors that reduce tree growth, their physiological traits and can present combinations of traits increase tree mortality, and limit tree regeneration (Gerwing & Lopes typical of both early successional (high photosynthesis capacity, Farias, 2000; Ingwell, Wright, Becklund, Hubbell, & Schnitzer, 2010; fast growth rate) and late-successional species (shade-tolerance, Schnitzer & Carson, 2010), and in turn alter ecosystem processes long leaf life span) (Lei & Koike, 1998; Montti, Villagra, Campanello, such as forest transpiration and carbon sequestration (Schnitzer & Gatti, & Goldstein, 2014); as such, many bamboos are adapted to live Bongers, 2002). We also know that liana abundances are increasing in open environments as well as the forest understory (Judziewicz, in many forests due to global climate change, including increased Clark, Londoño, & Stern, 1999). frequency of disturbances (Schnitzer & Bongers, 2011). Belowground, bamboos produce extensive rhizome networks Like lianas, bamboos (family Poaceae, subfamily Bambusoideae) that allow for clonal growth and a rapid resprouting response after include many ecologically important taxa that are ignored or avoided disturbances (Griscom & Ashton, 2006; Smith & Nelson, 2011; in most studies of forest dynamics, diversity, and ecophysiology, pre- Stapleton, 1998). There are three main rhizome morphologies that sumably because bamboos are assumed to have minimal influence lead to different patterns of culm spacing, from tightly aggregated on large-scale forest dynamics (but see section on Impacts of bamboo (i.e., clumping) to distantly spaced (i.e., running). Pachymorph rhi- on Neotropical forest dynamics). With the aim of increasing the vis- zomes have short, thick segments with a sympodial branching pat- ibility of this important but understudied subfamily, we present a tern that usually lead to tightly clumped culms. Leptomorph rhizomes FADRIQUE ET AL. | 3 have long, slender segments with monopodial branching pattern 2000) and microsatellites (Abreu et al., 2011) such as SSRs and ISSRs (running), or in some cases, with tillering culms (clumped). However, (Ely, Rada, Fermin, & Clark, 2019; Kitamura & Kawahara, 2009; Yang, some pachymorph taxa have elongated necks, producing more or An, Gu, & Tian, 2012) have been used successfully to identify clones less diffuse clumps or distantly spaced culms, the latter mimicking within bamboo populations. New techniques such as RAD-seq have the pattern produced by leptomorph rhizomes. Amphimorph rhi- shown promising results to separate species and individuals (Wang, zomes are a combination of both pachy- and lepto- morphologies and Zhao, Eaton, Li, & Guo, 2013) and to establish phylogenetic rela- show amphipodial branching (see figure 2 in Judziewicz et al., 1999; tionships between genera (Wang et al., 2017). Because of these McClure, 1966). The fibrous roots produced by bamboo rhizomes limitations to the use of molecular tools for identifying bamboos enhance and facilitate water uptake (Cochard, Ewers, & Tyree,