Multiscale Patterns of Riparian Plant Diversity and Implications for Restoration
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RESEARCH ARTICLE Multiscale Patterns of Riparian Plant Diversity and Implications for Restoration Joshua H. Viers,1,2 Alexander K. Fremier,3 Rachel A. Hutchinson,1 James F. Quinn,1 James H. Thorne,1 and Mehrey G. Vaghti4 Abstract herbaceous species, whereas woody species were largely 2 Planning riparian restoration to resemble historic refer- cosmopolitan across the nearly 38,000 km study area. At ence conditions requires an understanding of both local the floodplain scale, riparian floras reflected species rich- and regional patterns of plant species diversity. Thus, ness and dissimilarity patterns related to hydrological and understanding species distributions at multiple spatial disturbance-driven successional sequences. These findings scales is essential to improve restoration planting success, reinforce the importance of concurrently evaluating both to enhance long-term ecosystem functioning, and to match local and regional processes that promote species diver- restoration planting designs with historic biogeographic sity and distribution of native riparian flora. Furthermore, distributions. To inform restoration planning, we exam- as restoration activities become more prevalent across the ined the biogeographic patterns of riparian plant diver- landscape, strategies for restoration outcomes should emu- sity at local and regional scales within a major western late the patterns of species diversity and biogeographic U.S.A. drainage, California’s Sacramento—San Joaquin distributions found at regional scales. Valley. We analyzed patterns of species richness and com- plementarity (β-diversity) across two scales: the watershed scale and the floodplain scale. At the watershed scale, spa- Key words: biodiversity, biogeography, multiscale analysis, tial patterns of native riparian richness were driven by restoration, riparian, Sacramento, San Joaquin. Introduction are often intended to revegetate degraded and disturbed natural Conservation of biodiversity is a critical component of ecolog- areas with native plant species, are conducted on a site-by-site ical restoration worldwide (Gann & Lamb 2006) and is artic- basis without strategic planning aimed at promoting ecosys- ulated as a unifying principle for riparian restoration (Ward tem complexity or enhancing biodiversity. Thus, horticultural &Tockner2001).Incertainregions,andespeciallyincer- restoration is often reactive to local conditions and opportuni- tain ecosystems, restoration efforts are common and cover ties rather than regional strategies, in part because of a desire sufficiently large areas such that large-scale biogeographic to promote ecosystem functioning and biophysical processes. patterns of species diversity should be described and repre- Short of defining and implementing regional restoration strate- sented in restoration planning. For example, large coordinated gies to meet multiscale objectives (Hobbs & Norton 1996), we ecosystem restoration efforts, which are intended to restore suggest that a basic understanding of local and regional pat- ecosystem functions and processes, are ongoing for the Chesa- terns of plant diversity has the potential to improve how prac- peake Bay (Hassett et al. 2005), the Florida Everglades (Sklar titioners conduct horticultural restoration, specifically species et al. 2005), the Colorado River (Patten & Stevens 2001), selection during the design phase and evaluation of restoration and the San Francisco Bay-Delta (Kondolf et al. 2008). Over success. large areas such as these, it is clear that no single restoration Riparian plant restoration efforts typically focus on a small design can emulate the inherent complexity of these ecosys- number of large, overstory species rather than explicitly tar- tems. However, many horticultural restoration efforts, which geting overall species diversity (Sweeney & Czapka 2004). It is assumed that certain species have a large impact on the 1 Department of Environmental Science & Policy, University of California, Davis, ecosystem function (e.g. bank stability, nutrient processing, One Shields Avenue, Davis, CA 95616, U.S.A. wildlife habitat, etc.) and that, if present, a host of other species 2 Address correspondence to J. H. Viers, email [email protected] will colonize after initial targeted species are established. This 3 Department of Fish and Wildlife Resources, College of Natural Resources, University of Idaho, Moscow, ID 83844-1136, U.S.A. horticultural approach often achieves initial aims of targeted 4 Placer Land Trust, 11661 Blocker Drive, Suite 110, Auburn, CA 95603, U.S.A. species establishment, yet it is unclear if a native herbaceous understory component can subsequently equal remnant ripar- 2011 Society for Ecological Restoration International doi: 10.1111/j.1526-100X.2011.00787.x ian forests in native species richness (Holl & Crone 2004). Restoration Ecology 1 Patterns of Riparian Diversity for Restoration Further, there is little evidence that the cumulative result of knowledge rather than conducting wide-ranging studies (Wohl many localized restoration projects captures the inherent pat- et al. 2005; Young et al. 2005). Although many practition- terns of biodiversity found over large areas (Bedford 1999) ers likely possess an intuitive sense of how local actions fit nor that the current spatial distribution of restoration projects into broader ecosystem restoration efforts, there are too few across large areas is strategically sound if overall ecosystem examples of how baseline assessments can be utilized to ben- function or watershed-scale patterns of historic species distri- efit their projects, such as analyzing biogeographic maps of bution is a desired outcome (Kershner 1997; Teal & Peterson species diversity to determine if local projects are comple- 2005). With the recent profusion of horticultural restoration mentary in their scope or expected outcome, or alternately efforts (Bernhardt et al. 2007; Kondolf et al. 2007; Palmer evaluating field-based measures of species complementarity to et al. 2007), it is increasingly likely that uncoordinated activ- determine if a restoration project planting list is too homo- ities—coupled with narrowly defined objectives and limited geneous in its representation. Not only could the adoption implementation budgets—will influence the overall composi- of these principles early in the objective setting phase of tion and future trajectory of biodiversity in riparian ecosystems restoration help improve broader ecosystem outcomes, but also at a regional scale. provide practitioners with scientifically supported benchmarks For example, many riparian and floodplain restoration from which to objectively evaluate restoration outcomes. efforts in California’s San Francisco Bay-Delta and Central Our objective was to provide some context to the premise Valley are large in size, widely distributed, and could clearly that understanding biogeographical patterns of riparian plant impact the regional character of plant diversity (Alpert et al. diversity at local and regional scales is critical to developing 1999) and associated wildlife (Gardali et al. 2006; Golet contemporary restoration objectives. We performed a multi- et al. 2009). The widely implemented horticultural approach scale study on species distributional patterns within a riparian in this region is to establish native woody species with the restoration context, and provided critical analyses of diver- expectation that native understory species will colonize in sity patterns found in a large geographic region undergoing time (Palmer et al. 1997; Hilderbrand et al. 2005). The long- multiple, and largely uncoordinated, restoration projects as a term successes, and failures, of these projects have been well framework for restoration planning and evaluation. We cata- studied (Holl & Crone 2004) and ultimately shown to not logued multiple scale-specific studies of plant species presence adequately restore native species richness and composition to assess the patterns of species richness and turnover across when compared to remnant forest stands (McClain et al. 2011). ahydrologicallyandclimaticallycomplexregionofCalifor- Although the current approach is likely to improve certain nia. We used these data to understand patterns of diversity ecosystem functions on a localized basis (Dufour & Piegay in relation to watershed position and floodplain structure. We 2009), it is also likely to influence longer-term successional asked, over large areas, what is the number of native riparian trends and composition of regional floras. species and how variable is species richness? And, what is We believe that a regional understanding of species dis- the expected dissimilarity of herbaceous and woody species tributions is necessary if local efforts are to complement in a watershed and floodplain context (e.g. the dissimilarity in larger watershed-scale strategies aimed at ecosystem restora- species composition that would result from species represen- tion. Biogeographic patterns can help define regional restora- tation along environmental gradients or regionally from one tion objectives while also improving restoration evaluation and watershed to another)? At the watershed scale, we expected identifying gaps in habitat type representation. Although many native species richness to be uniformly distributed through- restoration projects value the concept of biodiversity (Mayer out the Central Valley due to its physical geography (i.e. 2006) from the perspective of providing localized genotypic it is a large alluvial basin with