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A Role for Assisted Evolution in Designing Native Plant Materials for Domesticated Landscapes TA Jones* and TA Monaco

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A Role for Assisted Evolution in Designing Native Plant Materials for Domesticated Landscapes TA Jones* and TA Monaco CONCEPTS AND QUESTIONS A role for assisted evolution in designing native plant materials for domesticated landscapes TA Jones* and TA Monaco Developers of native plant propagation materials for wildland restoration may emphasize naturally occur- ring genetic patterns or, in contrast, the material’s empirical performance in comparative field trials. We contend that both approaches have value and need not be mutually exclusive. Anthropogenic influences have pushed many ecosystems across ecological thresholds, to less desirable states, so that actively manag- ing for “domesticated nature” – nature as modified, either intentionally or inadvertently, by humans – is more realistic and more likely to succeed than recreating the original ecosystem. Furthermore, when domes- ticated nature is the most reasonable objective, empirical performance, together with geographical origin, are plausible criteria for choosing restoration plant material. For altered ecosystems, we suggest that evolu- tion should be assisted by the inclusion of plants that (1) reflect general historical evolutionary patterns, (2) are particularly suited to the modified environment, (3) are able to adapt to contemporary selection pres- sures, and (4) contribute to the restoration of ecosystem structure and function. Front Ecol Environ 2009; doi:10.1890/080028 s a matter of policy, scientists and land managers return to their previous state is not feasible. Kareiva et al. A sometimes emphasize protection and sustainability (2007) challenge scientists to manage these landscapes of lands that have been only minimally altered by anthro- by balancing trade-offs between ecosystem services. pogenic influences. The goal, with these less altered Ultimately, to be successful, management objectives must ecosystems, is to maintain endemic genotypes. On the be based on pragmatism (Hobbs et al. 2006) and sound other hand, many ecosystems have been drastically scientific principles (Gallagher and Carpenter 1997). altered (ie domesticated), leaving very few truly wild Here, we consider issues surrounding the development places on Earth (Kareiva et al. 2007). Previous genera- and choice of plant propagation materials for restoring tions of conservationists have written off lands impacted domesticated ecosystems. First, as an example of a domes- by humans as “lost causes”, yet the discipline of restora- ticated landscape, we describe the modification of west- tion ecology has emerged, with the goal of restoring ern North America’s sagebrush steppe by an invasive ecosystem processes to such lands (MacMahon 1997). plant–wildfire cycle. Next, we make the argument for Many scientists believe that these domesticated systems plants that restore evolutionary and ecological processes must be managed at some point (Gallagher and under such dramatic circumstances. Finally, we contend Carpenter 1997). Hobbs et al. (2006) suggest managing that human-assisted evolution offers the best hope for highly modified systems for utilitarian purposes when a repairing ecosystem structure and function when land- scapes have become domesticated. We use the term “eco- logical restoration” in the broad sense – that is, “the In a nutshell: process of assisting the recovery of an ecosystem that has • Indigenous genetic material may no longer be adapted in mod- been degraded, damaged, or destroyed”, as defined by the ified ecosystems that have crossed ecological thresholds Society for Ecological Restoration International Science • Genetically manipulated plant materials developed to over- and Policy Working Group (SER 2004). come common biotic and abiotic stresses are useful for restoring ecosystem structure, function, and biodiversity in modified ecosystems A controversial topic: options for restoration plant • Considering both empirical performance and geographic origin materials is important when fashioning plant materials to assist evolution along a desirable trajectory Two paradigms have long vied for the allegiance of scien- tists who develop plant materials for restoration. The “evolutionary” paradigm seeks to restore putative natural USDA-Agricultural Research Service, Forage and Range Research patterns of genetic variation, in order to generate an evo- Laboratory, Utah State University, Logan, UT *(thomas.jones@ lutionary trajectory as similar as possible to that which ars.usda.gov) prevailed before the advent of any anthropogenic distur- © The Ecological Society of America www.frontiersinecology.org Assisted evolution in domesticated landscapes TA Jones and TA Monaco bance, such as weed invasion or soil degradation. Great When are genetically manipulated plants an importance is therefore placed on genetic identity. In appropriate choice? contrast, the “resource” paradigm depends on the delivery Anthropogenic disturbances may cause plant communi- of economic products and ecosystem services, with mini- ties to cross one or more ecological thresholds, which will mal regard for species or genotype. These two models be difficult or impossible to reverse. We suggest that the have arisen within scientific traditions with contrasting decline of local genotypes under such conditions indicates attitudes toward the conservation of biological resources that the primary RGP is no longer as well adapted to the and with distinct philosophical beliefs as to how the local environment as it once was, and that this decline is world should be. In the US, this distinction corresponds a consequence of natural selection, a process that under to the historic division between the biocentric philoso- “natural” circumstances is considered desirable (Meffe phy of John Muir and the anthropocentric philosophy of and Carroll 1997). The anthropogenic perturbation of the Gifford Pinchot. These contrasting viewpoints influence sagebrush–steppe ecosystem is unnatural, but the biologi- the choice of seeds used in restoration projects; indeed, cal response to the perturbation is in accordance with the there has long been a healthy diversity of opinions normal workings of nature (Botkin 1990). regarding standards for the genetic identity of plant mate- Although the “local is best” assumption (Johnson et al. rials with the ecosystem restoration community. 2004) that underpins the general preference for the pri- But what is the best approach in choosing plants for the mary RGP is arguably valid when thresholds have not yet restoration of modified environments where domesti- been crossed, we contend that, for altered ecosystems, cated nature is the most viable remaining option? Vast this debate may be moot. Instead, the appropriate ques- landscapes in the North American Intermountain West, tion becomes, “is local still good enough to confer for example, are increasingly being modified by plant acceptable ecological function now that novel evolution- invasions, degradation of soil and hydrological processes, ary forces are at work?”. We believe that local may no and loss of biodiversity. Under such conditions, the evo- longer be good enough, based on simple observation; that lutionary paradigm alone will not be able to overcome is, if local was still good enough, we would not have an the problems encountered when modified conditions ren- invasive plant problem. The essential question then der local populations less adapted than before. Here, we becomes, “can we design better plant materials for the suggest that both paradigms have an important role in the monumental challenge sitting on our doorstep?”. restoration of damaged wildlands. Wise land managers will try to combine principles from both traditions to Costs and risks effectively realize their restoration objectives. We developed the Restoration Gene Pool (RGP) con- If we are to pursue a comprehensive plant materials cept (Jones 2003; Jones and Monaco 2007) to clarify approach, such as that described above, the associated plant options for restoration. The four RGPs – primary, costs and risks must be considered. The first cost results secondary, tertiary, and quaternary – are placed in from the research infrastructure required to develop the descending order of preference to the restoration practi- plants. A multidisciplinary, problem-solving approach tioner. The primary RGP includes materials that are and a long-term commitment are essential. Specific goals indigenous to the restoration site and to ecologically sim- for improving ecological function in western North ilar sites, as well as material from sites that are genetically America include controlling wildfire, inhibiting invasive connected to the site. Typically, seeds are either collected plant populations, and restoring soil structure and nutri- directly from the site or propagated under cultivation ent dynamics. To be successful, plant ecologists, physiolo- from wildland seed collections. The secondary RGP con- gists, soil scientists, rangeland scientists, seed scientists, sists of target-species material that is disqualified from the geneticists, plant breeders, weed scientists, and engineers primary RGP, based on the definition given above. must all work together to confront the problem at various Typically, secondary RGP materials are intended for ecological scales. The research objective must be to broader geographical use than are primary RGP materials; develop seeding technologies, weed management proto- they are often “releases”, either cultivars or pre-variety cols, and plant materials that may be prescribed for reha- germplasms (propagating material that has not yet been bilitating the land and restoring ecological
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