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Environment and Society: Advances in Research 15 (2024): 142–159 © e Author(s)
doi:10.3167/ares.2024.150107
Ecological Restoration, Genetics, Genomics,
and Environmental Governance
Christine Biermann and David Havlick
◾ ABSTRACT: Ecological restoration increasingly relies on genetic tools and technologies
to identify distinct populations, monitor populations, and even modify organisms to
improve tness. In this article, we review the role of genetic and genomic technologies
in restoration and conservation, using the restoration of cutthroat trout in the Western
United States as one example. Reducing restoration and conservation directives to the
molecular scale oen relies on a view of genes as discrete bits of information that pro-
duce controllable and predictable traits. is leads to life-and-death decisions about
wildlife populations, even as measures of “pure” genes for organisms are constantly
changing. We review the implications of a reductionistic approach centered on genetic
composition of organisms and consider the broader relevance of these issues to the
future of ecological restoration.
◾ KEYWORDS: biodiversity, conservation, epigenetics, genetics, genomics, hybridization,
restoration, trout
In 2019, the United Nations General Assembly adopted a resolution declaring 2021 to 2030 the
UN Decade of Ecosystem Restoration. e aim of this designation was to support and expand
“eorts to prevent, halt and reverse the degradation of ecosystems worldwide and raise aware-
ness of the importance of successful ecosystem restoration” (UN G.A. Res. 72/284). Ecological
restoration comes in many forms and can be applied to a variety of terrestrial, marine, and
aquatic contexts. From its early days, ecological restoration has included a focus on genetic
diversity and protecting and restoring ecosystems as a means of preserving genetic richness
within and across taxa. During the past three decades, as DNA testing and analyses have grown
increasingly capable of distinguishing ne dierences between organisms, ecological restoration
has turned to the molecular scale to evaluate the merits of protecting or restoring biodiversity at
a genetic level. e genetic details of organisms ranging from bison and wolves to trees, toads,
and trout have been scrutinized in recent years to advance restoration eorts and discern what
“belongs” and what does not (Merkle et al. 2007; Pröhl et al. 2021; Robinson et al. 2019; Ruther-
ford et al. 2019; Stroupe et al. 2022).
Synthesizing literature from the natural and social sciences, this article reviews and builds
upon the rise in genetic and genomic thinking, or what geographer Elizabeth Hennessy (2015)
has described as the “molecular turn” in conservation and restoration. Biodiversity has long
been conceptualized as three interconnected yet distinct levels: genetic diversity, species diver-
sity, and ecosystem diversity. With the emergence of new technologies over the past several
decades, ecological restoration and conservation have experienced a shi toward the molecular
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Ecological Restoration, Genetics, Genomics, and Environmental Governance ◾ 143
scale, with genealogical lineages, species’ genomes, and genes themselves becoming key objects
of concern and intervention (Allendorf et al. 2004; Bernos et al. 2020; Berseth 2022; Hennessy
2015; Metcalf et al. 2007, 2012; Robbins et al. 2023). When dened narrowly, genetics represents
analyses that use small numbers of molecular markers and focus on individual genes, while
genomics refers to the more recent shi to the use of large sets of molecular markers that enable
the sequencing and study of entire genomes. In this article we discuss both genetics and genom-
ics, and oen use “genetic” as a shorthand adjective to refer to both genetic and genomic data,
recognizing that the line dividing these analyses is oen fuzzy and that both types of data shape
restoration in new ways.
is article takes stock of genetic and genomic technologies and their relevance for ecologi-
cal restoration. More specically, we articulate how genetics and genomics have altered how we
understand, classify, protect, and restore nature, and how and why certain populations, taxa, and
ecosystems become identied as targets for intervention. In this review, we heed Lisa Campbell
and Matthew Godfrey (2010) in their call for social scientists to immerse themselves in genetic
science in order to study its implications for environmental governance. We empirically ground
this discussion using the restoration and management of cutthroat trout (Oncorhynchus clarkii)
populations in the Western United States as one key example, since aquatic ecosystems and
trout present specic challenges in terms of restoration and genetics (Allendorf 1988; Helfman
2007), and across the Western United States have been a particular focus of genetic-related res-
toration and conservation eorts since the late 1980s.
In the following section, we reect on key techniques and developments in the eld of res-
toration genetics, describing in brief their implications for how we think about and manage
nature. We then review existing literature that critiques a reductionist view of genes, and we
consider how advancements in biotechnology and the new science of epigenetics may com-
plicate how we think about genes and genomes in restoration. Finally, we explore alternative
frameworks for integrating genetics into restoration and conservation in a way that recognizes
other ways of knowing.
Genetics for Restoration and Conservation
Over the past half century, applications of genetics in restoration and conservation have expanded
along with the development of molecular tools and techniques. Today, molecular techniques
help to identify distinct or native populations (Amavet et al. 2023), reconstruct evolutionary
histories (Ghazi et al. 2021), monitor invasive or introgressed species (O’Donnell et al. 2023),
improve seed provenance choices (Bischo et al. 2010), and modify organisms to improve their
tness (Newhouse and Powell 2021).
In the 1970s, the study of allozymes using electrophoresis rst enabled researchers to empir-
ically test the body of population genetics theory that had been growing over the past sev-
eral decades (Allendorf 2017). Using allozymes, the amount of genetic variation within and
between populations could be quantied, allowing researchers to identify populations with
limited genetic variation or recent population bottlenecks (Leberg 1992). is was a crucial
development that had signicant implications for biodiversity conservation, making potential
genetic risks to species’ persistence visible and measurable. For example, allozyme research in
the early 1980s showed a marked lack of genetic variation in cheetah populations (O’Brien et al.
1983). is was hypothesized to be the result of a population bottleneck more than 10,000 years
ago and interpreted as placing cheetahs at greater risk of extinction. Extensive scientic debate
followed, with researchers disagreeing about the extent of the risk posed by cheetahs’ lack of
144 ◾ Christine Biermann and David Havlick
genetic variation (Caughley 1994; May 1995; Merola 1994). In both the popular media and aca-
demic literature, genetic data was used as evidence supporting more intensive research, captive
breeding, and recovery and restoration eorts focused on augmenting or maintaining genetic
variation. Conversely, some interpreted the lack of genetic variation as evidence that the species
is doomed to extinction, and therefore not warranting resources for its protection (O’Brien
1998). Each of these competing narratives emphasized cheetah genetics, diverting attention and
resources from other issues facing the species, such as habitat loss, poaching, and the illegal
wildlife trade. is emphasis on the molecular also works to reinforce the focus of conservation
and restoration eorts on single species—oen charismatic species like the cheetah—that are
privileged above others in the same ecosystem.
Alongside allozymes, researchers began to analyze mitochondrial DNA (mtDNA), which
proved particularly useful in reconstructing historical genealogical lineages and spurred the
creation of the new eld of phylogeography (Allendorf 2017). Together, allozymes and mtDNA
initiated a transition in restoration and conservation that began in earnest in the 1980s: popula-
tions were now knowable as genetic entities. Genes themselves, as well as related concepts such
as genetic divergence, gene ow, and genealogical lineages, were becoming objects of concern
and intervention (Hennessy 2015). e newfound ability to reconstruct phylogenetic relation-
ships based on mtDNA data also had implications for the concept of species and the practices
of distinguishing species, contributing to a shi toward evolutionary and lineage concepts of
species and away from morphological or biological (i.e., interbreeding) denitions of species
(de Queiroz 1998; Freudenstein et al. 2017). As science-society scholars have noted, reducing
restoration and conservation directives to the molecular scale relies on a view of genes as dis-
crete, transferable bits of information that produce controllable and predictable traits (Barnes
and Delborne 2022; Büscher et al. 2012; McAfee 2003; Rossi 2014). is leads to oen-lethal
decisions about wildlife populations, even as information about which genetics are “pure” for a
given species or subspecies may be in ux.
For cutthroat trout in the Western United States, these scientic developments reconcep-
tualized trout as genetic bodies, with the conservation value of populations now determined
in large part by their genetic composition—their purity or lack of hybridization (i.e., genetic
introgression)—but also by genetic divergence. In other words, the more genetically distinct
a population is from others in the species, the more worthy of protection it becomes. Prior to
the development of allozymes and mtDNA, patterns of genetic diversity in cutthroat trout were
poorly understood, and sheries managers and scientists struggled to determine how best to
classify, manage, and conserve populations that looked generally similar but resided in dierent
watersheds. Not only is the cutthroat trout a polytypic species, containing multiple subspecies,
but it also can hybridize in the wild with rainbow trout (Oncorhynchus mykiss), as well as among
dierent cutthroat subspecies.
Together, allozymes and mtDNA brought forward new understandings of cutthroat trout
diversity and distribution, suggesting that there were more lineages of cutthroat trout than had
been previously thought. e burgeoning molecular toolkit also enabled researchers and man-
agers to quantify the extent of hybridization using criteria other than morphological dierences.
Findings about hybridization and genetic divergence between westslope cutthroat trout (Onco-
rhynchus lewisi) and other cutthroat trout species raised new conservation questions. Should
hybridized populations receive protection under the Endangered Species Act? How should vul-
nerability be assessed for species that are widespread but oen hybridized (Allendorf et al. 2004)?
As with the cheetah, genetic threats to trout became more visible as technologies developed, and
as a result they sometimes overshadowed broader threats of habitat degradation, water manage-
ment, introduced species, climate change, and other challenges that aquatic species face.
Ecological Restoration, Genetics, Genomics, and Environmental Governance ◾ 145
e turn toward the molecular scale was hastened further by the development of polymerase
chain reaction (PCR) techniques in the 1990s. For PCR analysis, researchers amplify small seg-
ments of DNA into millions to billions of copies, which can then be examined in detail. Using
this process, researchers could measure genetic variation at the DNA level using other new tech-
niques such as amplied fragment length polymorphism, microsatellites, and single nucleotide
polymorphisms (Allendorf 2017).
More recently, next-generation sequencing (NGS) technologies have further revolution-
ized genetic approaches to restoration, enabling vast amounts of DNA to be sequenced rap-
idly and relatively inexpensively (Williams et al. 2014). An example of this is environmental
DNA (eDNA) barcoding, in which DNA in environmental samples (e.g., air or water samples)
is extracted and sequenced, and DNA sequences are then linked to taxonomic groups (Deiner
et al. 2017). Initially eDNA was used primarily for single-species detection, but through NGS
a few water samples may now essentially replace a time- and resource-intensive biodiversity
census of an entire body of water or watershed, with the added benet of minimal habitat dis-
turbance (Evans and Lamberti 2018). Still, there are shortcomings of eDNA; for example, it does
not provide a clear picture of population structure, tness, condition of organisms, or spatial
distribution. Additionally, it is particularly challenging to distinguish closely related species or
subspecies and hybridized populations using eDNA (Evans and Lamberti 2018). For species
such as cutthroat trout with unresolved taxonomies, high rates of hybridization, and local poly-
morphisms within subspecies, the deployment of eDNA is particularly challenging (Wilcox et
al. 2015). Still, eDNA is now part of a suite of technologies used in trout restoration projects,
for example by detecting survivors of sh removal projects and monitoring watersheds post-
restoration (Carim et al. 2020).
Hybridization and Gene Flow
Since the 1990s, when the establishment of molecular markers began to allow for estimations
of current and historical gene ow, a major thrust of restoration genetics has been the quanti-
cation of gene ow between species and within populations of the same species (McKay et al.
2005). Quantifying gene ow and, relatedly, measuring genetic diversity, is important for identi-
fying and selecting source populations for translocations, evaluating restoration outcomes, and
determining the appropriate scale for management (Campbell and Godfrey 2010; Mijangos et al.
2015). In some restoration contexts, gene ow between a targeted population and surrounding
populations is considered a positive outcome, increasing population persistence and enhancing
genetic diversity. In other contexts, gene ow may threaten the local adaptation of a population,
potentially undermining population persistence, as in the case of outbreeding depression, or
may result in hybridization between dierent species (McKay et al. 2005). Management center-
ing on genetics faces a tension between encouraging habitat connectivity, thereby decreasing
risks of genetic bottlenecks or vulnerability to stochastic events wiping out isolated populations,
and isolating populations in the interest of protecting genetic lineages and purity.
Hybridization between native species and non-native or introduced species is a particular
concern, as in the case of native cutthroat trout hybridizing with introduced rainbow trout in
the Western United States. is form of gene ow is oen framed as inherently negative, and in
scientic literature the use of terms such as “genetic contamination” and “genetic pollution” has
increased since the 1980s (Hirashiki et al. 2021; see also O’Brien 2006). Critiques of the use of
terms such as “genetic purity,” “genetic contamination,” “genetic pollution,” and “genetic integ-
rity” highlight their value-laden nature and/or roots in racist thinking (Biermann and Havlick
146 ◾ Christine Biermann and David Havlick
2021; Hirashiki et al. 2021; Rohwer and Marris 2015). However, ideas of “genetic purity” and
“genetic integrity” remain central to the scientic and management lexicon concerning cut-
throat trout, even if these same concepts are framed more subtly by designating populations for
conservation or recreation priorities based on thresholds of genetic purity.
Concerns about hybridization have signicant implications for how we think about and
manage nature. On one hand, the threat of hybridization has been invoked in arguments against
assisted migration and translocation, interventions that involve introducing taxa into new areas.
Concerns about gene ow are also used to caution against “willy nilly genetic intervention-
ism” (Peña-Guzmán et al. 2015: 259). On the other hand, concerns about hybridization may
be accompanied by a sense of its inevitability. For some restoration advocates, this has led to
increased support for novel forms of restoration that focus less on nativity of species and more
on ecological function (Davis et al. 2011; Hirashiki et al. 2021; Rohwer and Marris 2016).
e view of hybridization as a profound threat is countered by the idea that gene ow among
species may actually enhance evolutionary potential, reduce extinction risk, and/or improve
resilience to environmental change (Brauer et al. 2023; Vallejo-Marín and Hiscock 2016). Claire
Hirashiki and colleagues (2021) call for hybridization to be examined on a case-by-case basis.
Rather than starting from the position that hybridization threatens species survival, they con-
tend that potential positive eects on species survival or resilience should also be considered.
Even in cases where hybridization is generally considered to be a threat rather than an asset,
there are markedly dierent views about the level of risk that hybridization poses. is is the
case in scientic literature about westslope cutthroat trout (Muhlfeld et al. 2009, 2017; Young
et al. 2017), as well in responses to apparent inbreeding eects for the Bear Creek/greenback
cutthroat trout in Colorado (Fendt 2019).
In thinking about hybridization and gene ow, social scientists have analyzed how conser-
vation and restoration science function through biopolitical technologies (including genetics)
that work to dierentiate between “who or what is worthy of living—what kinds of biological
diversity are promoted in conservation projects and what kinds are not” (Biermann and Man-
seld 2014: 258). For example, Aurora Fredriksen (2015) discusses introgressive hybridization
among Scottish wildcats and domestic feral cats, exploring how taxonomy and genetic analyses
work to draw a line between unhybridized wildcats—which are cast as belonging and in need
of protection—and wild cats that are hybridized or domestic—which are presented as threats
in need of eradication. is biopolitical framework also analyzes how species-based approaches
to conservation and restoration oen run up against both care for individual organisms and
“life’s immanent, disorganising tendencies to ‘become otherwise’” (Fredriksen 2015: 690). In
Fredriksen’s research on wildcats, these “disorganizing tendencies” are the agencies of the cats
themselves. In the example of cutthroat trout restoration, these dynamics might include sh
that breach ecological barriers on their own, sh that escape death by moving to an isolated
eddy during a piscicide treatment, wildres that destroy barriers to sh passage, and so on.
is biopolitical perspective suggests that even as genetics aims to make nature knowable and
governable on the molecular scale, management eorts are constantly subverted: nature is never
fully controlled or controllable.
ese decisions about what forms of nature to protect, restore, or eradicate have spatial
implications, as genetic data are mobilized in debates about appropriate scales for managing
nature and establishment of conservation territories. In the traditional model of “fortress con-
servation,” nature is managed in bounded spaces such as national parks or wilderness areas,
within which visions of a pristine past are attempted to be re-created or maintained (Adams
2004). is vision rests upon an assumption that these places were not previously inhabited or
utilized by people, which in turn requires writing an array of Indigenous land uses and histories
Ecological Restoration, Genetics, Genomics, and Environmental Governance ◾ 147
out of view (Jacoby 2001; Spence 1996, 1999; Wilson 2020). Bringing this down to the organ-
ismic scale, Hennessy (2015: 88) suggests that genetics has enabled the geography of pristine
nature to be reimagined “as something manageable not only in Cartesian spaces, but also in the
purity of species lineages.” In such cases, a particular landscape is not sought to be protected or
restored to a pristine state, but instead the central focus becomes the “purity” or “integrity” of a
particular genealogical lineage.
At some point, prioritizing genetic purity in ecological restoration and wildlife management
will necessarily lead wildlife ocials to determine how pure is pure enough to count for conser-
vation. In the case of cutthroat trout, some states in the Western United States address this ques-
tion by setting specic thresholds of genetic purity to classify trout populations’ conservation
value. ese eorts seek to set restoration and conservation goals objectively, but there remains
some variability across states or species, and even standardized thresholds for genetic purity
reect numerical convenience (e.g., less than 1 percent or less than 10 percent introgressed)
rather than precise ecological signicance.
Besides the slightly dierent genetic standards applied to native trout management, and the
question of how these numbers are established, the genetic testing itself may be imperfect. Tech-
niques have improved over time, as we have already noted, but an inuential study from 2005
reported that only about 15 percent of genetic samples in trout could be classied with a high
degree of condence (Shepard et al. 2005). Lacking a stable and reliable means of analysis, sh
may face multiple rounds of eradication and restoration in an ongoing quest to establish truly
“pure” populations.
A restoration project in Yellowstone National Park highlights some of these issues. Aquatic
life in the park’s rivers and lakes has been heavily modied since the mid-nineteenth century,
particularly with the introduction of non-native rainbow trout, brown trout (Salmo trutta), and
brook trout (Salvelinus fontinalis). e Park Service aims to reverse this trend, identifying sites
where non-native or hybridized trout could be removed and restocked with native, unhybrid-
ized sh populations (Perkins 2020).
In 2005, the East Fork of Specimen Creek watershed was selected for Yellowstone’s rst
westslope cutthroat trout restoration project with the goal of eliminating a population that was
deemed “highly hybridized” (less than 80 percent pure) in order to restore unhybridized west-
slope cutthroat trout (Koel et al. 2006). e restoration required the construction of barriers to
isolate the watershed; the elimination of all existing sh by applying rotenone, a piscicide that
would be added in measured doses along the treatment areas of the watershed; and stocking the
now-shless waters with unhybridized westslope cutthroat trout from a combination of three
wild sources and one brood raised in a nearby hatchery managed for precisely these kinds of
restoration eorts.
In 2018 and 2019, approximately six years aer the last translocation of “genetically pure”
sh into East Fork Specimen Creek, the watershed was sampled to evaluate the success of resto-
ration eorts. Unexpectedly, of 290 successfully genotyped sh, 11.5 percent were determined
to be hybrids with a combination of westslope cutthroat trout alleles and Yellowstone cutthroat
trout (Oncorhynchus clarkii bouvieri) and/or rainbow trout alleles (Puchany 2021). is seemed
to indicate that hybridization and genetic introgression were occurring due to sh breaching a
barrier and invading from downstream, or from incomplete eradication of hybrids during the
rotenone phase. In considering next steps, Andriana Puchany (2021) concluded that complete
re-treatment of the watershed using piscicides, followed by re-stocking of “genetically pure” sh,
was likely required.
is situation raises uncomfortable questions about genetics-driven restoration decisions,
both for what these lead to intentionally and for the impacts on other organisms that can be
148 ◾ Christine Biermann and David Havlick
overlooked by managers. e lethal eects of rotenone on target sh are relatively well docu-
mented, but research less oen accounts for the impacts of this piscicide on a variety of other
organisms ranging from zooplankton and macroinvertebrates to amphibians and non-target
sh species (Beaulieu et al. 2021; Billman et al. 2011; Dalu et al. 2015). A single-species focus
in restoration projects risks disregarding the complexity and variety of organisms that may be
impacted by rotenone.
Focusing more directly on the intended impacts of rotenone treatments, it seems important
to ask, how many rounds of treatment and monitoring are required to successfully establish and
maintain unhybridized, “pure” populations? Indeed, can we expect the need for monitoring and
treatment to ever end? How many hybridized sh populations ought sheries managers be will-
ing to sacrice to establish one that is unhybridized? And more broadly, how did we arrive at
this juncture, where genes have become the target of restoration initiatives? What are the broader
implications of this shi toward the molecular scale in restoration and environmental governance?
ese and related questions rise to the fore as new genetic and genomic tools clarify the
dierences between trout lineages. is has moved the object of concern from specic rivers
or habitats toward populations with the purest (i.e., least hybridized) genes. ere are spa-
tial implications to this shi, as unhybridized populations are oen found in isolated streams
that have been disconnected from downstream waters through human modications of the
landscape, such as roads, culverts or dams (Biermann and Havlick 2021). In many restoration
projects, genetic isolation is required to protect a restored population from hybridization. is
necessitates the establishment of barriers that prevent sh from moving into the waterway from
either upstream or downstream, an intervention that again privileges a single species and de-
emphasizes or even potentially harms other taxa, such as invertebrates and amphibians.
is downscaling of restoration spurred by concerns about hybridization is potentially at
odds with longstanding ideas in conservation biology that promote landscape connectivity
through conservation corridors, mega-reserves, and networks of protected areas. In other cases,
genetic data are mobilized to support the scaling up of conservation beyond local or national
management and toward international management (Campbell and Godfrey 2010). When
genetic analyses indicated linkages among geographically disparate sea turtles in the Caribbean
Sea (e.g., linking foraging populations to distant nesting populations), researchers and conser-
vationists used this data to make the case for managing sea turtles as a common pool resource
that required conservation at an international level. Such scaling up, however, served to write
certain actors and communities out of the governance process (Campbell 2007), largely bypass-
ing the communities that were “living with, using, and conserving sea turtles” (Campbell and
Godfrey 2010: 905).
Genes as Discrete and Separable Units
One of the key questions surrounding the molecular turn in conservation and ecological res-
toration is the degree to which genes are treated as discrete, transferable bits of information
(Hennessy 2015; McAfee 2003; Rossi 2014; Valve 2011). Geographer Kathleen McAfee exam-
ined how genes are oen conceptualized as discrete entities: “functional units of information
which can be characterized precisely, counted, added or subtracted, altered, switched on and o,
or moved from one organism or one species to another” (McAfee 2003: 204). Despite scientic
evidence that supports a more complex understanding of genes and genomes, this reductionist
genetic discourse continues to circulate in various elds, from agriculture and biotechnology
to conservation and restoration. For example, the powerful eld of synthetic biology embraces
Ecological Restoration, Genetics, Genomics, and Environmental Governance ◾ 149
genes as severable or distinct units, where genetic parts, or short sequences of DNA, can be
moved, created, inserted, or modied to produce organisms with new characteristics, increased
resilience to adverse environmental factors, or even to create entirely novel forms of life. How-
ever, despite claims that biotechnology “permits precise control of life processes” (McAfee 2003:
203), the process is far messier and more complex than its nano-precision would suggest.
McAfee’s critique of operating at the molecular scale centers largely on agriculture and genetic
engineering, but many of the same concerns can be translated to other contexts such as ecolog-
ical restoration and biodiversity conservation. Genetic reductionism can abstract genes from
nature with its “spatial and temporal specicity . . . and from the environmental and social con-
texts in which [it] co-evolves” (McAfee 2003: 204). A similar critique has been made about how
genetic and genomic concepts of indigeneity reduce ancestry to genetic ancestry, thus abstract-
ing genetics from “biological and cultural kinship constituted in dynamic, long-standing rela-
tions with each other and with living landscapes” (TallBear 2013: 509). In this way, reducing our
understanding of organisms to a genetic level can be seen as a turn farther aeld from Indige-
nous or relational perspectives of socioecological connectivity (Todd 2014; Trigger et al. 2008).
With respect to environmental governance, when organisms and species are classied by and
managed primarily for their genetic composition, other aspects of their identities, such as life
histories and ecological relationships, may be obscured or overlooked (Havlick and Biermann
2021). For some species, establishing standards of genetic purity can abstract these organisms
from what are oen rich eco-social histories of particular populations. In addition, life-and-
death decisions about wildlife populations may then be predicated on an outdated and overly
deterministic view of genes. With this, the conservation value of populations is determined by
genetic composition.
According to McAfee, the reductionist view of genes-as-information also contributes to an
economic reductionism that allows genetic information to be patented and commodied down
to the molecular scale. For example, if organisms are modied through genetic engineering or
created through synthetic biology, these entities may then be subject to proprietary claims for
patenting, monetization, or the commodication of what previously were viewed as natural sys-
tems (and thus seemingly beyond the reach of complete human control). is raises concerns
about neoliberal entrainment or market capture of nature and may facilitate unconstrained
assertions of lethal control over these newly modied or hybrid ecosystems. Whether designed
with this in mind or not, genetic-focused interventions can contribute both to neoliberal biodi-
versity conservation (Büscher et al. 2012; cf Barnes and Delborne 2022) and an expansion and
deepening of human control over living organisms—what some consider a problematic exten-
sion of biopower reaching down to the building blocks of life (Lorimer 2015; Preston 2019).
is sense of control, however, may indeed be false, as it not only relies on a reductionist view
of genetics but also underestimates nonhuman agency and dynamism.
One example of how the commodication and marketization of genetic information is
deployed comes from an initiative by the International Union for the Conservation of Nature
and a technology start-up company, NatureMetrics. e eBioAtlas1 seeks to respond to the loss
of biodiversity in aquatic systems and wetlands by developing comprehensive inventories of
existing biota, utilizing tens of thousands of eDNA samples gathered across the world. Gen-
erating a global atlas of DNA found in freshwater ecosystems, the project will “target areas
threatened by climate change and development, and rapidly ll in critical gaps in knowledge to
support conservation eorts, unlock business investment to protect the natural world, and build
a rich databank to inform global policy to reverse the rapid decline in biodiversity” (NatureMet-
rics 2021). NatureMetrics promotes itself as a “global leader in using eDNA to turn nature into
data” (Cruickshanks and Czachur 2021). Businesses are expected to pay to access this data and
150 ◾ Christine Biermann and David Havlick
fund areas that are important to their operations or supply chains. In this way, genetic data are
actively monetized while also, at least prospectively, contributing to ecological restoration and
conservation eorts.
Although eBioAtlas promoters position their eort as a win-win, good for business and good
for biodiversity, critiques of this type of approach hinge not just on the commodication of
nature, but also on its reliance on genes as authoritative bit of information. McAfee argues that
while convenient, treating the gene as a unitary site conveying a genetic “code” is highly prob-
lematic (see also Kay 2000), as the gene’s “ontological status as a discrete causal unit of heredity
is increasingly in doubt” (McAfee 2003: 205). Other social analyses of genetics in restoration
have suggested that not all restoration practices rely on this same reductionism. Geographer Jai-
rus Rossi describes genes as “contextual and contingent entities” and suggests that certain kinds
of restoration practices can push against a reductionist view of genetic information and instead
lead to a view of genes as “embodied relational entities, rather than abstract information” (2014:
66). According to Rossi, this view can contribute to a decommodication of genetic knowledge
and promote more relational or socioecological restoration practices.
Epigenetics and Conservation
Another challenge of working through a strict genetic lens for restoration and conservation is
the growing understanding of epigenetics. Epigenetics is the study of heritable changes in the
ways genes work that are not associated with changes in the underlying structure or pattern of
DNA in the genetic code. In other words, epigenetics shows how environment and behavior
can aect an organism’s expression of genes in ways that can be passed down to future gen-
erations. is has contributed to what some consider a post-genomic science or a rejection of
genetic determinism (Lehrner and Yehuda 2018; Manseld and Guthman 2014). In one sense,
epigenetics counters the abstraction at work in reductionist understanding of genetics, re-
embedding genes within their biological and social environments. However, as critical analyses
of epigenetic science have shown, a non-deterministic view of genes may still reinforce eugenic
logics about biological dierences in human populations (Manseld and Guthman 2014).
Much of the epigenetic research has focused on human eects, but studies in lab animals
have contributed substantially to understanding epigenetic eects, leading to calls to incorpo-
rate conservation epigenetics in biodiversity protection (Rey et al. 2020). In recent years, a rela-
tive urry of articles has brought epigenetics into conversation with biodiversity conservation
(Amaral et al. 2020; Angers et al. 2020; Pazzaglia et al. 2021; Segelbacher et al. 2022; eissinger
et al. 2023). With epigenetic impacts such as DNA methylation likely showing greater respon-
siveness to short-term environmental changes, intraspecic epigenetic diversity may be under-
appreciated and a new front for restoration and conservation of biodiversity (Rey et al. 2020).
Fish raised in hatcheries for restoration, for example, have been shown to experience rapid
epigenetic modications that lead to reduced tness once released in the wild (Le Luyer et al.
2017). As such, epigenetics may complicate existing genetics-focused restoration strategies that
rely on raising “genetically pure” local broodstock in hatcheries.
Attending to epigenetics does not necessarily expand the focus of ecological restoration to
broader or more integrative approaches, but may facilitate more diverse ontological framings,
including blurred socio-natural constructions that lead to dierent pathways to restoring biodi-
versity (Meloni et al. 2022). With interest in epigenetics seemingly on the rise, it remains to be
seen how ready or able restoration ecologists and wildlife managers are to transfer these more
nuanced, potentially reversible features of ecosystems into practice.
Ecological Restoration, Genetics, Genomics, and Environmental Governance ◾ 151
Biotechnology, Genetic Engineering, and Synthetic Biology
With advances in biotechnology and synthetic biology, geneow between organisms is now pos-
sible not only through happenstance or painstaking programs of intentional breeding but also
through techniques such as genetic engineering and, more recently, gene editing. Gene editing via
CRISPR and similar techniques increasingly allows scientists to micromanage genetic composi-
tion and diversity within and between organisms, snipping and replacing original genetic material
with inuential bits from disparate other organisms or, in some cases, from entirely synthesized
sources (Piaggio et al. 2017; Preston 2019; Redford and Adams 2021; Redford et al. 2014).
e use of biotechnology to alter genetic material within organisms and populations con-
tinues to stir controversy; some view it as a potent new resource in restoration ecologists’ tool-
box (e.g., Barnhill-Dilling and Delborne 2019), while others raise concerns that meddling with
genetic foundations risks unforeseen consequences and destabilizes long-standing notions of
what is natural versus human-generated artifacts (Calvert 2010; Kolisis 2021). Even as debates
persist surrounding the merits of modifying and synthesizing life forms to better withstand
changing climates, degraded habitats, or other factors that imperil biodiversity, a growing array
of organisms produced through biotechnology (from older techniques such as gene transfers
through bacteria or virus vectors to newer gene editing techniques such as CRISPR) suggest that
advances in biotechnology are outpacing ethical or ecological concerns (Preston 2019).
Fully synthetic organisms have yet to be deployed in wild ecosystems, but it may only be a
matter of time before these lab creations are released, whether intentionally or not, into what
have been considered “natural” settings (Popkin 2018). US scientists created the world’s rst
organism with a synthetic genome in 2010, and less than a decade later British researchers
announced the creation of the world’s rst organism made from DNA that was entirely syn-
thetic (Sample 2019). Applications of synthetic biology currently underway range from the
de-extinction of species such as the passenger pigeon and woolly mammoth to managing (or
eradicating) malarial mosquitoes and introduced Norway rat populations in island ecosystems
(e.g., Piaggio et al. 2017).
Although synthetic biology has yet to be deployed for native trout restoration, genetic engi-
neering via bacteria or virus vectors or gene editing tools could potentially be used either to
remove undesirable material, such as rainbow trout DNA in cutthroat species, or to add new
diversity to the genes of native species in order to boost resiliency or help these organisms adapt
to changing habitat conditions, such as warmer water temperatures. e genetic makeup of
trout is already altered through less invasive, established methods as well. Since the early 1980s,
sheries scientists have heat-shocked trout to create triploid sh that develop an extra, third set
of chromosomes, rendering them sterile (Rohrer and orgaard 1986). In this way, sheries
managers have relied on genetic manipulation to stock triploid hatchery sh and reduce risks
of introgression with native trout, or to gradually eradicate non-native trout without resorting
to piscicides.
Alternative Frameworks for Using Genetics in Restoration
As we have shown, genetic science has changed not only how nature is understood but also
how it is governed, by whom, and at what scale. Genetic analyses are used to decide the value of
organisms and populations and to justify life-and-death decisions, sometimes despite incom-
plete knowledge or at the expense of other species or other forms of knowledge or values. is
is not to say that genetic techniques should be jettisoned, or that genetic knowledge is always used
152 ◾ Christine Biermann and David Havlick
in a singular fashion. Rather, a lesson we take from this review is that the science of genetics—
including the advancing elds of epigenetics and synthetic biology—provides a partial per-
spective on the world (Haraway 1988). In this nal section we explore visions of ecological
restoration and conservation that are epistemologically diverse, and that do not necessarily pri-
oritize genetic science over other values or ways of knowing. As Campbell and Godfrey (2010:
905) argue, “Any decision that this way of knowing is the best, only, or natural way, in essence
replacing or at least out-weighing other ways, is ultimately a human and political decision.”
Genetics may help us understand evolutionary patterns, interrelationships between populations
or organisms, reproductive tness, and ne distinctions between dierent types of organisms,
but it alone cannot tell us “whether or not particular outcomes are acceptable nor whether or
not they are sustainable” (Campbell and Godfrey 2010: 905). What does a culture of ecological
restoration look like when we recognize that genetics alone cannot provide an answer key for
the dicult, value-laden decisions that comprise restoration and environmental governance?
When genetic science is treated as the central arbiter of truth about a population or ecosystem,
what other opportunities, forms of knowledge, or actors are disregarded or obscured?
To begin to address these questions, it is helpful to understand the broader context of social
research on restoration and conservation. Chris Sandbrook and colleagues (2013) distinguished
between social research for conservation and social research on conservation. Research for con-
servation is driven by the mission to conserve biological diversity and seeks to improve existing
conservation policy and practice. Research on conservation may come from various other start-
ing points and generally works to “understand how conservation as a social, political practice
works” and “to situate conservation with respect to broader social and political economic issues”
(Sandbrook et al. 2013: 1488). A similar heuristic could be applied to research on and for eco-
logical restoration. Conservation and restoration professionals oen look explicitly to research
for conservation/restoration to guide management actions, and research on conservation and
restoration may be viewed as unnecessarily critical, opaque, unhelpful, or even antagonistic to
the broader mission.
A more holistic and robust approach to ecological restoration dissolves this dichotomy,
insisting that research for restoration should also reect on the context in which it is produced
and the broader social, political, ecological, and economic networks in which it is embedded.
Yet conversations about the science of restoration genetics and critical reections on genetics
in restoration and conservation have by and large proceeded separately. Natural and social sci-
entists are oen siloed in conducting research, reporting ndings, and intersecting with agency
ocials and practitioners. Rather than thinking of research on and for restoration as separate,
we envision integrating these domains to bring the insights of critical social science into the
practices of natural science and ecological restoration.
While there is a growing literature of examples of this type of boundary-crossing work (Keeve
et al. 2021; Sinner et al. 2022), critical physical geography provides one possible framework
through which to integrate genetic knowledge and restoration science with diverse approaches
such as feminist science studies, Indigenous knowledge, and queer and trans theory (Lave et al.
2018; Wöle Hazard 2022). e three tenets of critical physical geography provide a useful start-
ing point for this integration. First, we must recognize that genetics are deeply shaped by human
activities and social power relations. e genetic composition of populations, the genomic
structure of species, and the distribution of genes across space, have been fundamentally shaped
by people. We therefore can interpret genetic data in connection with human values, actions,
and social dynamics—including, but not limited to, Indigenous land uses, settler colonialism,
Western science, and conict and warfare (e.g., Darimont and Pelletier 2021). Second, the same
social dynamics and power relations that have shaped the distribution of genes across space
Ecological Restoration, Genetics, Genomics, and Environmental Governance ◾ 153
also shape who studies genetics and how they are studied. e methods and techniques pre-
viously discussed—from allozymes to epigenetics—have not emerged out of a vacuum but are
themselves shaped by social power relations in ways that aect the types of questions that are
asked, the scales at which questions are analyzed and answered, and the forms of knowledge that
are accepted in governance and decision making. Finally, this approach recognizes that genetic
knowledge has deep impacts on the organisms it purports to know as well as the other species
with whom they interact in an ecosystem.
Starting from these assumptions, it is possible to build a more expansive form of resto-
ration that is both more collaborative, interdisciplinary, community-grounded and epistem-
ically diverse. e “underows” approach of Cleo Wöle Hazard is illustrative: with Western
scientists, native communities and scientists, and community activists, Wöle Hazard (2022)
practices a collaborative river restoration science rooted in ethics of reciprocity, community
benet, and respect for multiple forms of knowledge. inking about genetics, this could mean
that research starts from a place where the authority of genetic science is not “universal or
unquestioned, but neither is science dismissed out of hand” (Wöle Hazard 2022: 26). It may
also lead researchers to follow principles of Indigenous data sovereignty in conservation and
restoration genetics (Robbins et al. 2023), and to develop opportunities for idea generation and
communication across dierence (such as the ecocultural restoration eld school cocreated by
Wöle Hazard and research partners). ese possibilities are not the focus of this article, but we
mention them here as potential pathways for incorporating both genetic data and critiques of
genetics into restoration, in ways that “see, recognize, and reach out for dierent streams—dis-
sident streams, decolonizing upwellings that rework science and governance from within but
also from below—into a lively, muddy, organic machine” (Wöle Hazard 2022: 31). Restoration
ecology has long positioned itself as an integrative eld that accommodates the contributions
of natural and physical scientists, social scientists, environmental philosophers, and contribu-
tors from the humanities. In light of the molecular turn and conservation genetics, restoration
scholars and practitioners should continue to embrace this tradition, engaging across disci-
plines and social dierences, opening to new ways of knowing, and recognizing that genetics is
one of many modes of valuing and relating to other species.
◾ ACKNOWLEDGMENTS
We appreciate the reviewers’ and editor’s helpful and constructive feedback. is material is
based upon work supported by the National Science Foundation under Grant No. SES-1922157.
◾ CHRISTINE BIERMANN is associate professor of geography and environmental studies at the
University of Colorado Colorado Springs. She is co-editor of e Palgrave Handbook of
Critical Physical Geography (with Rebecca Lave and Stuart Lane).Email: cbierman@uccs
.edu; ORCID: 0000-0002-0821-755X
◾ DAVID HAVLICK is professor of geography and environmental studies at the University of
Colorado Colorado Springs. He is the author of Bombs Away: Militarization, Conserva-
tion, and Ecological Restoration; No Place Distant; and co-editor of Restoring Layered Land-
scapes: History, Ecology, and Culture (with Marion Hourdequin). Email: dhavlick@uccs.edu;
ORCID: 0000-0002-6230-1584
154 ◾ Christine Biermann and David Havlick
◾ NOTE
1. https://ebioatlas.org/
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