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\begin{document}

\title{The eco-evolutionary risks of not changing seed provenancing practices
in changing environments}



\author[1]{Rebecca Jordan}%
\author[2]{Peter Harrison}%
\author[3]{Martin Breed}%
\affil[1]{CSIRO}%
\affil[2]{University of Tasmania}%
\affil[3]{Flinders University}%


\vspace{-1em}



  
  \date{May 19, 2024}


\begingroup
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\selectlanguage{english}
\begin{abstract}
Rapidly changing environments combined with increasing global
restoration initiatives require improved seed sourcing strategies for
native revegetation. Sourcing seed from local populations (local
provenancing) has been the long-standing default for native revegetation
for numerous eco-evolutionary reasons including local adaptation and
species co-evolution. However, the evidence-base has shifted, revealing
risks for both non-local and local provenancing in changing
environments. As alternative strategies gain interest, we argue for
effective decision-making that weighs the risks of changing and not
changing seed sourcing strategies in a changing environment that
transcends a default position and the polarising local vs. non-local
debate.%
\end{abstract}%



\sloppy


\textbf{The eco-evolutionary risks of not changing seed provenancing
practices in changing environments}

Rebecca Jordan \textsuperscript{1}, Peter A.
Harrison\textsuperscript{2}, Martin Breed \textsuperscript{3}

\textsuperscript{1} CSIRO, Environment, Sandy Bay, Tasmania 7005,
Australia. rebecca.jordan@csiro.au

\textsuperscript{2} Australian Research Council Centre for Forest Value
\& School of Natural Sciences, University of Tasmania, Sandy Bay,
Tasmania 7005, Australia. paharrison@utas.edu.au

\textsuperscript{3} College of Science and Engineering, Flinders
University, Bedford Park, SA 5042, Australia.
martin.breed@flinders.edu.au

Statement of authorship: All authors conceived this viewpoint and
contributed to the writing and editing of the manuscript.

Running title: Risks of not changing provenancing practices

Article type: Viewpoint

`Abstract' (first paragraph) word count: 96

Main text (incl. `abstract') word count: 1963

Number of references: 44

Number of figures: 0

Number of tables: 1

Data accessibility statement: No data associated with this study

Corresponding author: Rebecca Jordan. 15 College Rd, Sandy Bay, Tasmania
7005, Australia. Tel: +61 3 6237 5641. rebecca.jordan@csiro.au

\textbf{CONTEXT}

Rapidly changing environments combined with increasing global
restoration initiatives require improved seed sourcing strategies for
native revegetation. Sourcing seed from local populations (local
provenancing) has been the long-standing default for native revegetation
for numerous eco-evolutionary reasons including local adaptation and
species co-evolution. However, the evidence-base has shifted, revealing
risks for both non-local and local provenancing in changing
environments. As alternative strategies gain interest, we argue for
effective decision-making that weighs the risks of changing \emph{and
not changing} seed sourcing strategies in a changing environment that
transcends a default position and the polarising local vs. non-local
debate.

Revegetation aims to restore ecosystems by reintroducing biodiverse
plant communities that support key services and functions that humans
rely upon (e.g., water filtration, carbon sequestration). How to best
source seed to achieve resilient, long-term, self-sustaining plant
populations has received substantial attention (Jones 2013;
Prober\emph{et al.} 2015; Bucharova 2017). Local provenancing targets
adaptations that have evolved to maintain a fitness advantage to the
local environment (Leimu \& Fischer 2008; Hereford 2009), enhancing the
establishment and long-term success of the revegetation plantings. Local
provenancing also maintains important co-evolved biotic interactions
such as plant-fungi and plant-pollinator relationships (Grady \emph{et
al.} 2017; Bucharova \emph{et al.} 2021) whilst avoiding negative
genetic effects of introducing non-local genotypes, such as outbreeding
depression and swamping of local genotypes (Byrne \emph{et al.} 2011;
Bucharova \emph{et al.} 2021). For these reasons, and others, local
provenancing has been the long-standing default strategy for ecosystem
restoration.

However, environmental change and associated impacts on plant fitness
have raised concerns for the long-term viability of the default use of
local provenancing. Changes to local environmental conditions (e.g.,
direct and indirect effects anthropogenic change, such as climate change
and land-use) may decouple adaptation and fitness, resulting in greater
risk of local seed being maladapted (Etterson \& Shaw 2001; Anderson
2016). For example, seed sourced from small, fragmented local remnants
can result in genetically depauperate, inbred progeny (Breed \emph{et
al.} 2015; Aguilar \emph{et al.} 2019) that are less fit and more
vulnerable to climate change (Nickolas \emph{et al.} 2019). Thus, the
perception that local provenancing is a `safe' option may be misleading
in a rapidly changing world (Frankham \emph{et al.} 2011; Ralls \emph{et
al.} 2018).

Proposed alternative provenancing strategies that mix seed from local
and non-local provenances aim to address these environmental
change-induced issues by increasing genetic diversity or introducing
putatively pre-adapted genotypes that enhance resilience and long-term
adaptability to environmental change. Though there is a shift in
perceptions toward these alternative strategies (e.g Australia,
Hancock\emph{et al.} 2023), concerns remain about the risk of using
non-local provenances, with some sectors continuing to recommend only
local provenancing (Hancock \& Encinas-Viso 2021). Such hesitance
possibly reflects a paucity of evidence on outcomes of using non-local
provenances (Twardek \emph{et al.} 2023). While risks associated with
introducing non-local provenances in a revegetation context are valid,
equally valid are the risks of using local provenances under current and
future conditions.

Here, we argue that the risks of changing \emph{and not changing} the
local provenancing default in a changing environment needs to be weighed
when determining the most appropriate provenancing strategy. We focus on
the eco-evolutionary risks associated with provenance choice, including
the short- and long-term fitness of plantings and their resilience and
future adaptability, rather than whether provenances are local or
non-local \emph{per se} . We aim to help move the discussion away from a
polarised local vs. non-local debate, and towards a risk-based,
context-dependent rethinking of how to source seed for revegetation.

\textbf{WHEN LOCAL MAY NOT BE BEST}

The risks associated with maintaining the default local provenancing
strategy may be greater from an eco-evolutionary perspective than using
alternative strategies that include both local and non-local provenances
(summarised in Table 1). While this is not an exhaustive list, we
present four case studies, with the aim of highlighting the transition
from a state of known risks for non-local provenances to a state of
known risks for both non-local and local provenances.

\emph{Climate change shifting local advantage}

As climates change, there is a risk of local adaptations becoming
decoupled from environmental selective pressures, resulting in the
maladaptation of local genotypes. Evidence of such decoupling was
demonstrated in \emph{Eucalyptus gunnii} , a subalpine foundation tree
species endemic to Tasmania, Australia. Consistent with local
adaptation, local provenances initially showed higher fitness in
reciprocal common garden trials (Potts 1985). However, following
two-decades of declining precipitation and increasing temperatures
(Sanger \emph{et al.} 2011), local provenance fitness declined
(\textless{}30\% survival), with the lower elevation, non-local
provenance showing higher fitness than the local provenance
(\textgreater{}70\% survival; Prober \emph{et al.} 2016). Similar
impacts of changing environments have been observed in other forest
species (Jump \emph{et al.} 2009) and natural systems more broadly
(Parmesan 2006; Scheffers \emph{et al.} 2016).

While future change may decouple local adaptations from home
environments, climate change to date may already be outpacing adaptation
-- an outcome known as adaptation lag. Evidence for adaptation lag was
shown in \emph{Quercus lobata} (valley oak), where two, early-age
provenance trials showed greater relative growth of provenances when
planted into cooler climates than their current homesite climate
(\textasciitilde{}10\% greater growth; Browne \emph{et al.} 2019). Using
a genomics-informed selection model, Browne et al. (2019) predicted
gains of up to 25\% in relative growth of individuals pre-adapted to
current environments of the test site (Browne \emph{et al.} 2019).
Optimal growth associated with historic rather than current climate is
consistent with adaptation lag. While growth is a plastic trait and
species may persist in suboptimal environments, similar evidence of
adaptation lag has been shown in several tree species (Carter 1996;
Aitken \emph{et al.} 2008; Fr\selectlanguage{ngerman}éjaville \emph{et al.} 2020).

\emph{Demographic and evolutionary history}

Demographic history plays an important role in structuring the genetic
and adaptive variation within and among populations (Hewitt 2000). The
historical expansion from refugia or genetic bottlenecks can, for
example, often leave evolutionary fingerprints that have fitness effects
on modern-day populations. Such demographic processes have been argued
to explain the differential survival and fecundity of two natural
populations of \emph{Arabidopsis thaliana} (Thale cress) from opposing
latitudinal ends of its native European range (Sweden and Italy). While
local provenances showed higher fitness on average in reciprocal
transplant trials (Ågren \& Schemske 2012), inter-provenance crosses
between the two populations tended to have higher fecundity than the
local provenance when planted in Sweden (Ågren \emph{et al.} 2013).
Furthermore, several Italian alleles provided a fitness advantage over
the Swedish alleles when plants were grown in Sweden (Ågren \emph{et
al.} 2013). Demographic history, in particular postglacial expansion in
northern Europe reducing genetic variation most likely led to increased
fixation of these maladaptive alleles.

\emph{Novel environments from land use change}

In some situation, environmental change, such as mining, agriculture,
and deforestation, has resulted in novel environments. This presents a
risk that the local provenances may no longer be fit for these novel,
local conditions. Evidence of this is shown in two metallophyte
legumes,\emph{Mimosa acutistipula} var. \emph{ferrea} and \emph{Dioclea
apurensis} , which are promising candidates for the restoration of
degraded areas of mineral rich soils in the Amazon, Brazil. Adaptive
genomic variation showed that local provenances would perform best in
moderately disturbed sites (e.g., sites altered by fire)
(Carvalho\emph{et al.} 2020). However, no provenance in the sampled area
matched predictions for highly disturbed sites with substantially
altered conditions (i.e., ex-mining sites; (Carvalho \emph{et al.}
2020). With no suitable local provenance, regional mixing across
multiple adaptive units was recommended to capture genetic variation and
enhance adaptability, thereby de-risking revegetation plantings.
Together with previous findings of rapid, microgeographic adaptation
post-mining (e.g. Antonovics and Bradshaw 1970; Antonovics 2006), these
results highlight that provenance choice may vary dependant on site
history, with environmental changes having the potential to
significantly influence provenance fitness in novel environments.

Together, these examples highlight that current and future environmental
changes may have reduced local provenance fitness, and thus increased
the risk associated with using local provenances in particular
revegetation contexts. Further, they highlight the need to consider
current and future environmental change, climate and anthropogenic, as
well as effects of evolutionary history (e.g. demographic history) when
selecting provenances, to ensure both the short- and long-term fitness
and productivity of revegetation plantings. One approach could be mixing
local seed with seed from non-local provenances already occupying
current or future climates, which may de-risk the long-term resilience
of revegetation plantings.

\textbf{MOVING AWAY FROM A DEFAULT}

Reframing provenancing strategies on eco-evolutionary risks highlights
that all strategies come with risks, even local provenancing. We
recommend moving the discussion beyond the local vs. non-local debate
and towards context-dependant choices that enhance resilience and future
adaptability. Growing evidence suggests that no single strategy will be
appropriate for all contexts. Rather, the most appropriate strategy in
any situation will be influenced by a range of context-dependant
factors.

To create resilient plantings, provenancing must consider the dynamic
conditions imposed by environmental change (Aitken \& Bemmels 2016).
Genetic variation is a key factor in long-term evolutionary resilience
and adaptability (Sgrò \emph{et al.} 2011; Kardos \emph{et al.} 2021).
Provenancing strategies that mix local and non-local provenances aim to
boost resilience by increasing genetic diversity, including adaptive
variation. Considering eco-evolutionary genetics can therefore help
guide whether local, non-local or a combination of both may be most
appropriate to enhance genetic variation and thus adaptability in any
given situation (Hoffmann \emph{et al.} 2020). While we focussed on the
eco-evolutionary risks associated with provenance choice, other factors
may influence provenance choice, including ecological (e.g., biotic
interactions; (Bucharova \emph{et al.} 2021), logistical (e.g.,
availability of seed; (Broadhurst \emph{et al.} 2016) and socio-cultural
(e.g., significance of local genetic identity of plants; Maurice\emph{et
al.} 2013). However, these factors need to be weighed against the
eco-evolutionary risks associated with not changing provenancing
strategy, especially in changing environments.

Moving forward, it will be important to address barriers and knowledge
gaps that currently inhibit context-dependant provenancing decisions.
These include better understanding of the long-term performance, and
risks associated with, both local and non-local provenances for
different species in different restoration contexts and deriving
practical guidance on provenancing choice (Breed \emph{et al.} 2018).
Greater direct evidence on the long-term performance of not only
different provenances (e.g., local and non-local provenances) but also
different provenancing strategies will also assist decision-making
(Bailey \emph{et al.} 2021). Determining whether generalized
recommendations can be derived based on plant life history, functional
traits and/or genomic information across species will also help guide
recommendations for species without empirical data.

Several initiatives are working to address these knowledge gaps. For
example, there are already several practitioner guides that incorporate
climate uncertainty, evolutionary genetics, and landscape fragmentation
into seed sourcing decisions with the aim of increasing genetic
diversity and adaptability (Harrison \emph{et al.} 2017;
Rossetto\emph{et al.} 2019). Establishing long-term provenance trials
(as per example 1 above), especially those that are embedded within
mixed species revegetation plantings (e.g. Bailey \emph{et al.} 2021),
will help generate long-term fitness data on provenances in a relevant
revegetation context and thus help enable adaptive seed sourcing
decisions into the future.

\textbf{CONCLUDING REMARKS}

With a focus on eco-evolutionary factors, our viewpoint is that no
provenancing strategy is risk free. The risk of changing from the
default local provenancing strategy needs to be weighed against the risk
of not changing. We do not suggest local provenancing is no longer
appropriate, nor that provenancing strategies must change from local.
There are clear situations where local will be most appropriate and
where factors not discussed in depth here (e.g., logistics and cultural)
may influence provenance choice more than eco-evolutionary factors.
Rather, the choice of provenancing strategy should consider all risks
when determining the most appropriate provenancing strategy for a given
species and context. As such, the intention of our paper is to move the
discussion of provenancing away from a default strategy and whether
local provenances are more-or-less appropriate than non-local
provenances, and towards conscious and context-dependent provenancing
decisions.

\textbf{ACKNOWLEDGEMENTS}

We thank Linda Broadhurst, Ary Hoffmann and Suzanne Prober for comments
on an earlier draft of this manuscript. This work was supported by the
Australian Research Council (grant numbers LP190100051, LP190100484,
DP180100668, DP210101932 to M.F.B; and P.A.H was supported by
IC150100004). All authors declare that, to the best of their knowledge,
they have no conflicts of interest.

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Table 1: Examples of where only local provenancing may not be best and
pose a risk to the long-term success of restoration plantings.

\emph{FIGURE REPRINT PERMISSIONS REQUIRED}\selectlanguage{english}
\begin{longtable}[]{@{}lllll@{}}
\toprule
\begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
ISSUE\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
Long term performance\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
Adaptation lag\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
Demographic history\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
Altered environments\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
Short term fitness not indicative of long term or life-time fitness,
especially for long-lived organisms\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
Adaptation not keeping pace with climate change, resulting in local
being adapted to historic not current conditions.\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
Demographic history, such as recent expansion and bottlenecks, may
reduce local adaptation.\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
Significant environmental changes, especially anthropogenic change,
resulting in novel local conditions\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
Species\strut
\end{minipage} & \begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
Cider gum (\emph{Eucalyptus gunnii-archeri})\strut
\end{minipage} & \begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
Valley oak (\emph{Quercus lobata})\strut
\end{minipage} & \begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
Thale cress \emph{(Arabidopsis thaliana)}\strut
\end{minipage} & \begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
Metallophyte legumes (\emph{Mimosa acutistipula} var \emph{ferrea} and
\emph{Dioclea apurensis})\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
Location\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
Tasmania, Australia \textsuperscript{1}\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
California, USA \textsuperscript{2}\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
Sweden and Italy \textsuperscript{3}\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
Amazon, Brazil \textsuperscript{4}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
Overview\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
35 year old provenance trial. Survival, growth and frost damage used to
assess provenance performance.\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
Combined common garden trials and genomic analysis; assessing relative
growth and associated genomic loci\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
Multi-year reciprocal transplant trials and genomic analysis assessing
performance of both wild and F1 recombinant inbred lines.\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
Genomic analysis; predicting performance based on genotype-climate and
genotype-phenotype associations.\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
Key finding\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
Performance of local declined over time, with non-local outperforming
local after 25 years.\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
Predicted greater growth using non-local seed sources accounting for
adaptation lag.\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
Swedish population was most fit in Sweden in only 3 of 5 years; Swedish
alleles in Sweden were often maladaptive.\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
Local genotypes predicted not to match highly disturbed (ex-mining)
sites.\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
\textbf{Take home}\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
\textbf{Local not best in long-term, despite initial short-term
outcomes}\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
\textbf{Local already not best due to adaptation lag behind recent
climate changes}\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
\textbf{Local not most fit due to demographic history impacts on local
populations}\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
\textbf{Local no longer suitable where site conditions have
significantly changed}\strut
\end{minipage}\tabularnewline
\bottomrule
\end{longtable}

\textsuperscript{1}Prober et al. 2016; \textsuperscript{2} Browne et al.
2019;\textsuperscript{3} \selectlanguage{ngerman}Ågren \emph{et al.} 2013; Ågren \& Schemske
2012;\textsuperscript{4} Carvalho et al. 2020

\selectlanguage{english}
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