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Does Basin Wildrye (Leymus Cinereus) Show Local Adaptation When Deployed According to Generalized Provisional Seed Zones in the Central Basin and Range Ecoregion?

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Does Basin Wildrye (Leymus Cinereus) Show Local Adaptation When Deployed According to Generalized Provisional Seed Zones in the Central Basin and Range Ecoregion? 112 A typical wildland basin wildrye (Leymus cinereus) population. NATIVEPLANTS | 22 | 2 | SUMMER 2021 REFEREED RESEARCH Does basin wildrye (Leymus cinereus) show local adaptation when deployed according to generalized provisional seed zones in the Central Basin and Range ecoregion? Scott Jensen, Val Jo Anderson, William Christensen, Bruce Roundy, Stanley Kitchen, and Loreen Allphin ABSTRACT Identifying genetically appropriate plant materials for seed-based restoration relies on the principle of local adaptation: the intent to match adaptive genetic characteristics to variation in ecological clines pertinent to plant establishment and persistence. Seed zone maps delin- eate some of these relationships. Generalized provisional seed zones (GPSZ) were developed for use where species-specific seed zones are lacking, as was the case for basin wildrye at the initiation of this study. In this study, basin wildrye (Leymus cinereus (Scribn. & Merr.) Á. Löve [Poaceae]) sources from 25 wild populations and 4 commercial varieties were planted at 4 test sites representing the species distribution across GPSZ in the central basin and range ecoregion. Sources were seeded separately into 5 blocks for each of 2 treatments and data were grouped by tetraploid or octoploid cytotype and local or non-local origin for compari- son. Treatments included coverage with row cover or uncovered controls. We assessed initial and short-term survival. Plantings failed at 2 sites in both 2013 and 2014, with too few plants to quantify differences. At the remaining 2 sites we found no differences in initial survival or short-term persistence between pooled local sources compared to pooled non-local sources. Among commercial sources, the cultivar ‘Magnar’ initially outperformed local pooled materi- als at Fountain Green but not at Nephi. This difference was not evident 3 and 4 y later. Initial establishment under row cover was dramatically better than uncovered controls but persisted only through years 3 and 4 at the Fountain Green site. Jensen S, Anderson VJ, Christensen W, Roundy B, Kitchen S, Allphin L. 2021. Does basin wildrye (Leymus cinereus) show local adaptation when deployed according to generalized provisional seed zones in the Central Basin and Range ecoregion? Native Plants Journal 22(2):112–122. KEY WORDS local adaptation, basin wildrye, Leymus cinereus, Great Basin, provisional seed zones, row cover, Poaceae NOMENCLATURE USDA NRCS (2020) Photos by Scott Jensen This article was prepared by a U.S. government employee as part of the employee’s official duties and is in the public domain in the United States. This open access article is distributed under the terms of the CC-BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0) and is freely available online A typical wildland basin wildrye (Leymus cinereus) population. at: http://npj.uwpress.org. 113 NATIVEPLANTS | 22 | 2 | SUMMER 2021 n recent years, plant material programs and policies have Range (CBR) ecoregion, its known distribution ranges over emphasized the development and deployment of geneti- 1100 m (3630 ft) in elevation and across 9, level 4 ecoregions cally appropriate native plant materials (Richards and (Omernik 1987). A substantial array of climatic and ecologi- IChambers 1996; Rogers and Montalvo 2004; Shaw and others cal variation occurs throughout this species distribution that 2008; Oldfield and Olwell 2015; Wood and others 2015). The may lead to the development of localized adaptions between foundation of genetic suitability is the principle of local adap- spatially or ecologically divergent populations (Hereford tation with the intent of exploiting genetic variation to both 2009). Across the western portion of the species distribu- preserve and capitalize on spatially diverse functional traits to tion, Culumber and others (2013) genetically distinguished 3 improve restoration outcomes (Leimu and Fischer 2008). In metapopulation races corresponding to the Columbia, Rocky application, using genetically appropriate plant materials re- Mountain, and Great Basin regions. Octoploid cytotypes were quires a suitable pairing of restoration sites with seed sources, more abundant in the western portion of the species range, so that the developing plant community provides desired eco- and tetraploid cytotypes were more abundant in the eastern system services and persists through environmental challenges portion. In the Great Basin, where the Great Basin race is fully (Johnson and others 2010b; Jones 2013). encompassed by the more broadly distributed Rocky Moun- Genecological studies are a common first step in under- tain race, both cytotypes are common. In a recently completed standing morphological and phenological variation that can be common-garden study (Johnson and Vance-Borland 2016), used to identify adaptive genetic differences between popula- genetic variation for both cytotypes was linked to source cli- tions (Campbell 1986; Erickson and others 2004; Johnson and mates. That information was used to delineate 15 genecologi- others 2015). In genecological studies, numerous populations cal seed zones for basin wildrye. of the same species are planted in one or more common en- To test the principle of local adaptation, we hypothesized vironments, and many traits that may have adaptive signifi- that wild basin wildrye sources planted into test sites matching cance are evaluated. Resulting data, when paired with climate their home GPSZ would demonstrate local adaptation through records from source population locations, are used to develop higher initial establishment and better short-term persistence. seed transfer guidelines, which are often displayed as seed zone We also hypothesized local GPSZ pooled sources would simi- maps. For a number of prominent Great Basin restoration larly outperform the cultivars ‘Magnar’, ‘Trailhead’, and ‘Con- grasses, genecological work is underway (Johnson and oth- tinental’, which originate from outside the Great Basin, and ers 2010a; Johnson and others 2012; St Clair and others 2013; Great Basin ‘Tetra’, which is a composite of 31 sources from Johnson and others 2015) and was recently completed for basin multiple GPSZ’s and ecoregions. From experience using fabric wildrye (Leymus cinereus (Scribn. & Merr.) Á. Löve [Poaceae]) row cover to improve seedling establishment in nursery beds, (Johnson and Vance-Borland 2016). Yet, where genecological we included row cover as a treatment option in this experiment work is absent, many species will, of necessity, be planted with- and hypothesized basin wildrye establishment would be better out species-specific seed zone recommendations (Bower and under row cover. others 2014). In these cases, surrogate approaches have been suggested (Omernik 1987; Milberg and Andersson 1998; Har- METHODS grove and Hoffman 2005; Vogel and others 2005; Johnson and others 2010b; Miller and others 2011; USDA 2013). Within the Great Basin, known populations of basin wild- In this study, we evaluated a leading surrogate, general- rye (n = 107) were mapped to 9 of the 20 Great Basin GPSZ’s ized provisional seed zones (GPSZ) (Bower and others 2014; (WWETAC 2021) using ArcMap 10.1 (ESRI 2012). Five zones Kramer and others 2015), as a method of matching basin were poorly represented, containing 1 to 3 populations within wild rye source populations to representative restoration sites. their boundaries, while 4 zones accounted for the remaining GPSZ delineate areas of similar winter minimum temperature 91.6% of populations. Bower and others (2014) demonstrated and annual aridity, characteristics important to plant adapta- GPSZ’s to be most effective when nested within Omernik’s level tion. When paired with ecoregions (Omernik 1987), they parti- III ecoregions. Consequently, for this study, source populations tion areas into climatically and ecologically distinct units and were restricted to origins within the CBR ecoregion. Basin are recommended as a starting point for developing seed trans- wildrye population locations were obtained from a US Forest fer guidelines. Service database developed over the course of several decades Basin wildrye was selected as the test species because it of plant material work and were originally located by travel is a common restoration species (Krabacker 2013) in the In- throughout the area, consultation with federal and state agency termountain region, yet its genecological seed zones had not personnel, or university and online herbarium sources. been developed at the time this study was initiated. Its dis- Basin wildrye occurs in 2 cytotypes, tetraploid (2n = 28) and tribution extends from New Mexico to Saskatchewan and octoploid (2n = 56) (Ogle and others 2012). Crossing cytotypes 114 then west to the Pacific Coast. Within the Central Basin and results in sterile seed or unstable hexaploids (Young and others NATIVEPLANTS | 22 | 2 | SUMMER 2021 BASIN WILDRYE IN GENERALIZED PROVISIONAL SEED ZONES 2013). Consequently, cytotypes should not be combined when Leaves from 2 plants per site were processed to determine each assembling multi-origin seed sources for grow out or on res- population’s cytotype. Across the western portion of the spe- toration projects. As our interest, in this study, was to evalu- cies distribution, Culumber and others (2013) found no cases ate only establishment and short-term persistence, sources of mixed ploidy within populations of basin wildrye. (Table 1) were not isolated by cytotype but data were grouped When developing genetically appropriate stock seed sup- by cytotype
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