The northern hemisphere forestry literature

suggests that latitudinal or altitudinal gradients, or both, can be important for

detecting the
scale of local adaptation, but that other factors
such as habitat, rainfall and topographical differences can also be signifcant
(Ennos, Worrell
and Malcolm, 1998). There is evidence for adaptive variation over reasonably short
distances in a
number of conifer tree species in western North
America, owing to features such as aspect and altitude (e.g. Adams and Campbell,
1981; Sorensen,
1994). This is particularly marked in areas with
oceanic climates, where environmental gradients
are much steeper than in more continental sites.
Field, greenhouse and laboratory studies on conifer species in the northwestern
United States
show that a signifcant proportion (typically 25�
45 percent) of the genetic variation within populations is accounted for by
climatic (e.g. rainfall
and temperature) or location (e.g. latitude, altitude, slope aspect, distance from
ocean) variables
that re?ect environmental factors specifc to each
location. There are often differences between
provenances from warmer and colder climates,
the former showing adaptation to the longer
growing season in lower latitudes but suffering
from early or late frosts when moved too far into
higher latitudes. The degree of risk in transplanting across a species�
distribution is correlated
more with environmental changes than with
the geographical distance moved (Adams and
Campbell, 1981; see Insight 5). This suggests that
habitat matching may be a more useful means of
determining where seed should be sourced than
would be an arbitrary distance from the site to be
restored. Provenance trials of a number of tropical tree species show that most
morphological genetic variation occurs within rather than between
provenances. In most of the species studied, ranking reversals (adaptation) or
signifcant genotype � environment interactions only occur with
large environmental site differences (e.g. dry vs
wet zones, alkaline vs acidic soils). Unfortunately,
almost nothing is currently known about local
adaptation in temperate southern hemisphere
species.
Currently there are too few studies from too
few regions of the world to allow for predictions
regarding the scale and importance of local adaptation for the myriad of life-
history traits and
evolutionary histories of tree species that require restoration. For example, Leimu
and Fischer
(2008) used only 32 species in their local adaptation meta-analysis, none of which
were tree species. There is also a need for reciprocal transplant
experiments (RTEs), which test the ftness of
�home� and �away� genotypes within the sites
from which the genotypes originate (Primack and
Kang, 1989) and can mimic natural regeneration
by establishing seedlings in a forest at close spacings to encourage early
competition and with
minimal intervention (e.g. little or no weeding).
Germplasm selected and tested in forestry trials
or plantations for growth, form and other commercial criteria may be less suited to
the more
competitive environment of semi-natural forests
and restoration.
The scale over which species show adaptation
to their environment depends on the degree of
habitat heterogeneity, in particular the specifc
habitat characteristics that affect a species, and
the interaction with gene ?ow. Dispersal levels
may be a useful high-level predictor of the importance of local adaptation, under
the premise
that species with long-range gene ?ow are less
likely to generate strong local adaptation, whereas restricted gene ?ow is more
likely to generate
genotypes adapted to their local environment.
Extensive gene ?ow in widely distributed tree
species suggests that local adaptation over a small
geographic scale is unlikely unless selection forces
are very strong.
2.5. Are non-local seed sources
ever appropriate?
In highly modifed or degraded landscapes, using non-local seed may be entirely
appropriate
or indeed the only option for restoration. Mitigating the negative physical effects
associated
with vegetation removal, such as loss of topsoil,GENETIC CONSIDERATIONS IN
ECOSySTEM RESTORATION USING NATIvE TREE SPECIES
31
altered hydrological ?ows or increased nutrient
loads, may require specifc germplasm that is able
to cope with these conditions. For example, saline scalds in southern Australia
that developed
following the removal of deep-rooted perennials
are not generally amenable to restoration using
local species, let alone local seed. In these cases,
planting saline-tolerant varieties of other species
may be the only option to prevent further degradation of valuable agricultural
land. The loss of
diversity at genes of major effect may also require
sourcing of seed from non-local populations. For
example, small populations of self-incompatible
plants can be mate-limited if diversity in the incompatibility locus is low,
requiring seed from
beyond the local area to introduce new mating
types. However, impacts on local species and
communities that may arise from using non-local
seed need to be considered carefully, preferably
prior to restoration and using an appropriate risk
management framework (Byrne, Stone and Millar, 2011). This should also include
analysis of the
risk of not undertaking restoration and allowing
landscape degradation and biodiversity loss to
continue.
2.6. Local seed sources may not
produce restoration-quality
seed
Habitat fragmentation remains a major threat
to biodiversity worldwide through the loss of
populations and consequent altered biotic and
abiotic processes (Bakker and Berendse, 1999;
Eriksson and Ehrlen, 2001; Hobbs and Yates,
2003; Lienert, 2004). Unfortunately, some regions of the world have now reached a
tipping
point, such that whole biomes may be in danger
of collapse (Hoekstra et al., 2005). The most immediate consequence of
fragmentation for use
of native species in restoration and farm systems
is limitations to seed supply following the loss of
individuals and populations. But several negative genetic and demographic effects
associated
with fragmentation can also have an impact on
the quantity and quality of seed available. The removal of trees and populations
from landscapes
directly reduces genetic diversity, most of which
is irreplaceable since genetic mutations accumulate slowly over long evolutionary
periods (i.e.
tens of thousands to millions of generations).
Diversity is further eroded in small populations
by drift resulting from random sampling within
populations, as well as inbreeding as a result of
trees in remnant populations often being more
highly related than those in larger populations
(Barrett and Kohn, 1991; Ellstrand and Elam,
1993). Reduced ftness and productivity are commonly documented effects associated
with genetic erosion and inbreeding, both of which can
have an impact on a population�s ability to persist
in stressful situations or changing environments
(Frankham, Ballou and Briscoe, 2002; Hughes et
al., 2008). Other negative outcomes include poor
reproductive success, smaller, poor-quality plants
and increased susceptibility to pests and pathogens (Lienert, 2004 and references
therein). Over
time, this exposes small populations to decline
through recruitment failure (Figure 2.1) and limits their utility as appropriate
seed sources for
restoration. Limited seed supply and poor-quality
seed are two major impediments to the successful
planting of native species and restoration of native vegetation, especially at the
landscape level.
Worldwide analyses of fragmentation impacts
on plant reproduction indicate that some species are shifting towards selfng
(Aguilar et al.,
2006; Aguilar et al., 2008; Eckert et al., 2010), but
how this translates to seed production depends
largely on reproductive strategy. For example,
species that cannot self or mate with close relatives (self-incompatible) will not
produce seed
unless pollinated by distantly- or non-related
trees and small, self-incompatible populations
are often characterized by reduced seed production, severely limiting quantities
available for
restoration. In contrast, species that can self and
mate with close relatives (self-compatible) continue to produce seed but this is
often less ft, being smaller, slower to germinate and with poorer
survival (Buza, Young and Thrall, 2000; YoungTHE STATE OF THE WORLD�S FOREST
GENETIC RESOURCES � THEMATIC STUDy
PART 2
32
et al., 2000; Mathiasen, Rovere and Premoli,
2007). Restoration using this seed is therefore
likely to produce poorer results than expected
and over the long term is less likely to develop
into a self-sustaining population. A requirement
that only local seed be used for restoration can
drive practitioners to use seed from small, inbred
populations that are unlikely to produce positive long-term restoration outcomes,
but rather
create more small, inbred populations, with limited long-term persistence. One
consideration is
that populations restored with a narrow genetic
base may be limited in their ability to respond
to the rapid predicted shifts in climatic variables
(Helenurm, 1998).
2.7. Adaptation and climate
change
There are theoretical reasons that underlie observed patterns of adaptive
variation; these also
suggest that many tree species over large areas
may fail to show local adaptation at a very narrow
scale. The prevalence of extensive gene ?ow may
counteract selection, while the temporal variation in selective forces that trees
experience (e.g.
yearly variation in temperature, frosts or rainfall)
is likely to have a stabilizing effect rather than the
directional selection that would lead to highly localized adaptation. Given the
long life of trees,
the environment is also likely to have altered over
the lifespan of a tree or only a few generations,
such that a particular site no longer experiences
the same conditions under which the trees originally evolved. These factors explain
the relative
lack of adaptation over short distances in many
tree species. Temporal variation in environment
is particularly important for trees, not only with
respect to past adaptation but also in the context
of predicted climate change (e.g. Broadmeadow,
Ray and Samuel, 2005), and thus undue emphasis
on local seed sources may also cause problems.
Figure 2.1.
Simplifed representation of how low genetic
diversity and inbreeding can impact on plant
population persistence and seed production in
small populations of plantsGENETIC CONSIDERATIONS IN ECOSySTEM RESTORATION USING
NATIvE TREE SPECIES
33
2.8. benefts of using larger but
more distant seed sources
Using large populations as primary seed sources
for restoration not only ensures that seed quality
will be higher but also that larger quantities are
available. In many cases a population of 100�200
plants would be large enough to provide goodquality seed, but more than 400 plants
may be
needed for some species. A good restoration outcome is also more likely if the
habitat of the site to
be restored is matched as closely as possible with
that of the nearest large population. Seed from
these large populations could be augmented
with that collected from small populations closer
to the restoration site to capture any useful genetic diversity they may contain
(Broadhurst et
al., 2008). To capture as much genetic diversity as
possible from large populations, as many plants
as practically possible should be sampled broadly
across the site, collecting from a range of cohorts,
from various sides of plant canopies without disrupting biotic associations that
also rely on this
seed. Breed et al. (2012) reviewed such strategies for sourcing restoration seed
(Box 2.1) and
summarized their suitability for mitigating climate change and habitat
fragmentation impacts
(Table 2.1).
The mixing of introduced and native germplasm
raises the issue of outbreeding depression; the
potential problem of reduced vigour as adapted
gene complexes are broken up or the proportion
of locally adapted alleles is reduced. As with local
adaptation, evidence for outbreeding depression
comes from herbaceous species that show highly
localized adaptation (see Hufford and Mazer,
2003) and there is little evidence for its occurrence in trees at distances of less
than hundreds
of kilometres (e.g. Hardner et al., 1998, Boshier
and Billingham, 2000). For example large-scale
importation of cheap seed from Eastern Europe
has shown problems of maladaptation in Britain.
But it seems unlikely that use of material from
maritime France that matches future climate predictions (Broadmeadow et al., 2005)
and of similar
phylogeographic origins will face such problems,
nor lead to outbreeding depression problems on
introgression with British material.
2.9. Conclusions
Any genetic conservation policy for native trees
should aim at conserving the evolutionary potential of their populations, rather
than at preserving a particular genetic structure and status.
The extent and scale of local adaptation in many
tree populations, and thus its practical importance to restoration efforts, remain
in doubt.
While there is a need for more feld trials, both
of the traditional provenance or progeny and
RTE types, to provide more information on the
scale of adaptation, planting of native trees continues apace and demand for seed
from certifed
sources increases. There is good evidence to suggest that emphasis on a very
restricted view of
what is �local� will not lead to better-adapted
tree populations and is more likely to lead to use
of stock of limited genetic diversity than would a
broader approach.
It has been argued that, given the lack of extensive trials investigating adaptive
variation
in native tree populations, the precautionary
principle should be adopted in sourcing germplasm for planting trees (e.g. Flora
Locale, 1999;
UKWAS, 2007). This is expressed as the use of local seed, although the subsequent
view of what
constitutes the local population varies from a
particular forest to large seed zones. However,
given current evidence for trees, i.e. clear dangers from inbreeding and loss of
genetic diversity, with extensive gene ?ow and adaptation at
a broad scale, it seems more logical to apply the
precautionary principle in terms of ensuring the
use of genetically diverse material with the capacity to adapt to current and
future conditions.THE STATE OF THE WORLD�S FOREST GENETIC RESOURCES � THEMATIC
STUDy
PART 2
34
Strict local provenancing: collecting seeds
from plants that are located physically very close
to the revegetation site (e.g. Natural England,
United Kingdom: 5 miles; Western Australian Forest
Management Plan 2004�2014: 15 km).
Relaxed local provenancing: collecting seeds
with a bias towards certain ecological criteria, and
avoiding small population fragments (e.g. Australian
FloraBank: soil type, altitude and climate).
Predictive provenancing (Sgr�, Lowe and
Hoffmann, 2011): use of naturally occurring
genotypes experimentally determined to be adapted
to projected conditions. This technique requires
data on local adaptation of target species (e.g. by
reciprocal transplant experiments), as well as climate
projections for these species at a revegetation site
(e.g. by bioclimatic modelling).
Composite provenancing (Broadhurst et al.,
2008): collecting a mixture of seed that attempts
to mimic natural gene-?ow dynamics. For example,
recommended proportions of seed collected from
local, intermediate and distant distance-classes could
be determined by estimating the pollen dispersal
kernel for target species.
Admixture provenancing (Breed et al., 2012):
collecting seed only from large populations, focusing
on capturing a wide selection of genotypes from a
diversity of environments with no spatial bias towards
the revegetation site. These seeds are then admixed
for sowing or planting, generating a population
with a mixture of genotypes from a wide array of
provenances.
References
Breed, M.F., Stead, M.G., Ottewell, K.M., Gardner,
M.G. & Lowe, A.J. 2012. Which provenance and where?
Seed sourcing strategies for revegetation in a changing
environment. Conserv. Genet., November 2012. doi:
10.1007/s10592-012-0425-z.
Broadhurst, L.M., Lowe, A., Coates, D.J., Cunningham,
S.A., McDonald, M., Vesk, P.A. & Yates, C. 2008. Seed
supply for broadscale restoration: maximising evolutionary
potential. Evol. Appl., 1: 587�597.
Sgr�, CM, Lowe, A.J. & Hoffmann, A.A. 2011. Building
evolutionary resilience for conserving biodiversity under
climate change. Evol. Appl., 4: 326�337.
Source: Breed, M.F., Stead, M.G., Ottewell, K.M., Gardner, M.G.
& Lowe, A.J. 2012. Which provenance and where? Seed sourcing
strategies for revegetation in a changing environment. Conserv.
Genet., November 2012. doi: 10.1007/s10592-012-0425-z.
Box 2.1.
Summary of alternative strategies for sourcing seed for restoration
TAbLE 2.1.
Suitability of provenancing techniques under climate change with habitat
fragmentation
Provenancing
technique
Adaptive
potential
benefts
Genetic rescue
benefts
Low genetic
load
Suitable
with high
uncertainty
Economically
effcient
Likely
population
success
Strict local x*
Relaxed local x*
Predictive x x x�
Composite x x x x
Admixture x x x x x
* May experience high failure rates, negating the economic beneft.
� Beneft rests on successfully matching genotype ftness with future conditions.
Source: Breed et al. (2012).GENETIC CONSIDERATIONS IN ECOSySTEM RESTORATION USING
NATIvE TREE SPECIES
35
Current threats to the maintenance of genetic
diversity come principally from poor practice in
seed collection; undue emphasis on restricting
the area of collection or poor instruction of collectors can limit the number of
trees and hence
genetic diversity sampled, leading to the establishment of trees with restricted
genetic diversity
and limited future adaptive potential. A study
of the few remnant ash and rowan trees in the
denuded Carrifran valley in southern Scotland
showed that large amounts of genetic diversity
are maintained, making them suitable for use in
restoration despite their highly fragmented nature (Bacles, Lowe and Ennos, 2004).
In contrast,
some of the locally sourced material planted as
part of the Carrifran wildwood restoration project was shown to be low in genetic
diversity
(Kettle, 2001), presumably because of poor collection practices, which impose
limitations on the
future potential of the population.
It is disturbing to contemplate that some of
the poorest seed sources exist in the very regions
where restoration is most needed and that continued requirements for using local
seed simply
perpetuate the problem. In many regions of the
world with fragmented forest populations, being
able to reliably source large volumes of quality
seed of native species can be challenging. Not
only are there fewer populations from which seed
can be collected, but fragmentation has split continuous populations into much
smaller and more
isolated remnants, which can impact the quality
and quantity of seed (e.g. Lowe et al., 2005). The
implications from this are that (i) remnant vegetation contains all of the
diversity that is left that
is extremely valuable, and (ii) it is important that
most of the diversity that does remain is used
for restoration (i.e. avoid over-collection from a
few populations). In regions where fragmentation is high, should the rules for
using local seed
change? Can we afford the luxury of being too
restrictive about seed sources? Are small, fragmented and probably inbred
populations so precious that we cannot source seed from beyond
our comfort zone?
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Vander Mijnsbrugge, K., Bischoff, A. & Smith, B. 2010.
A question of origin: where and how to collect seed for
ecological restoration. Basic Appl. Ecol., 11: 300�311.
Wilkinson, D.M. 2001. Is local provenance important in
habitat creation? J. Appl. Ecol., 38: 1371�1373.
Williams, G.C. 1966. Adaptation and natural selection.
Princeton, NJ, USA, Princeton University Press.
Young, A.G., Brown, A.H.D, Murray, B.G., Thrall, P.H.
& Miller, C. 2000. Genetic erosion, restricted mating
and reduced viability in fragmented populations of
the endangered grassland herb Rutidosis leptorrhynchoides. In A.G. Young & G.M.
Clarke, eds. Genetics,
demography and viability of fragmented populations,
pp. 335�359. Cambridge, UK, Cambridge University
Press.GENETIC CONSIDERATIONS IN ECOSySTEM RESTORATION USING NATIvE TREE SPECIES
39
A population of trees or shrubs is autochthonous
if it has regenerated naturally since its arrival after the last glaciation; any
human intervention in
breeding should have occurred with strictly local
material only. For long-lived species such as trees,
autochthony assumes a continuous presence at a
given site since post-glacial immigration (Kleinschmit, Kownatzki and Gegorius,
2004). This implies a continuity of local genetic diversity after
thousands of years of natural selection. Trees and
shrubs that belong to native species but are imported from other climatic zones or
geographic
regions are not autochthonous.
After many years of neglect, the use of native species in afforestation and
landscape programmes is gaining importance all over the
world, based on the basic underlying ecological
principle that native species and genotypes will
be well adapted to local conditions and will have
co-evolved with other components of local forest
ecosystems. This has led to massive plantations
of indigenous tree and shrub species in Western
Europe, not only in forestry but also for native
woodland restoration and other landscape plantings, such as thickets, wooded banks
and hedgerows. A major challenge is to ensure that planting material used
represents the genetic variation
and diversity within native species. Several initiatives have been developed in
various European
countries to promote the use of locally sourced
seeds for the production of planting stock (e.g.
Belgium: Vander Mijnsbrugge, Cox and Van
Slycken, 2005; Germany: Kleinschmit, Leinemann
and Hosius, 2008; Denmark: Kjaer et al., 2009).
Here we describe in detail the programme on the
production of autochthonous planting stock in
Flanders, Belgium.
3.1. Why should autochthonous
diversity be protected?
There is a high demand for �native� planting
stock in Flanders, Belgium, and to a broader extent in many Western European
countries. The
use of native planting material is promoted by
a wide range of public organizations. However,
planting stock of native material in commercial
nurseries is largely not autochthonous. Seeds of
native species are often imported, originating
from foreign provenances, often in Eastern European countries. This is especially
true for shrub
species. For trees, in the European Union, Council Directive 1999/105/EC of 22
December 1999
(Council of the European Union, 2000) regulates
the marketing and transport of forest reproductive material through an obligatory
certifcation
system indicating the origin of the material (although control in practice is not
perfect). However, certifcation is not obligatory for shrubs, and
Chapter 3
Continuity of local genetic diversity
as an alternative to importing
foreign provenances
Kristine Vander Mijnsbrugge1,2
1 Research Institute for Nature and Forest, Belgium
2 Agency for Nature and Forest, BelgiumTHE STATE OF THE WORLD�S FOREST GENETIC
RESOURCES � THEMATIC STUDy
PART 2
40
shrub germplasm is commonly imported from
Eastern and Southern Europe where cheaper
seed is available. Nursery managers often do not
know or are not interested in the exact origin of
the seed they obtain. Tree seed may also be imported when seed is not available
from offcially
approved sources or supplies are too limited to
meet requirements.
Introduction of non-local material can
have numerous negative consequences. Nonautochthonous planting stock may be poorly
adapted to local growing conditions, which can
lead to negative consequences such as lower
ftness (e.g. McKay et al., 2005; Krauss and He,
2006; Edmands, 2007; Laikre et al., 2010; Vander
Mijnsbrugge, Bischoff and Smith, 2010). Problems
may only become evident many years after seemingly successful establishment.
Intraspecifc hybridization of local and introduced genotypes
may result in outbreeding depression, i.e. reduced ftness in subsequent
generations, loss
of genetic diversity and loss of adaptation, and
less adapted characteristics can introgress into
the autochthonous populations. The introduction of non-local material may also have
negative
effects on associated plant and animal species.
Imported hawthorn (Crataegus monogyna) has
been shown to ?ower several weeks earlier than
native hawthorn, potentially threatening the insects and birds whose reproductive
cycles are synchronized with this event (Hubert and Cottrell,
2007). In addition, purity of the species can be
problematic in commercial planting stock. A genetic study on commercially available
hawthorn
in Flanders, grown from seeds imported from
Hungary, showed that it comprised a mixture of
C. monogyna and C. monogyna � C. rhipidophylla
(Debeer, 2006).
3.2. Inventory of autochthonous
woody plants
A simple way to ensure the continuity of local genetic diversity is the production
of autochthonous
planting stock. For this an overview is needed of
the remnant autochthonous populations still present. A survey was conducted to
locate remaining
autochthonous populations in Flanders, Belgium,
from 1997 to 2008. The evaluation of autochthony in the feld was conducted
following the
methodology presented by Maes (1993). In short,
areas of woody vegetation that are indicated as
forest on historical maps are identifed. Information on ?ora, soil conditions and
geomorphology
further refne the selection of potentially relevant sites. In the feld, the woody
vegetation is
evaluated according to a set of criteria. The tree
or shrub must be a wild variety and old. No evidence must be seen of plantation
(e.g. trees in
lines). The site must be located within the natural
geographic range of the species, and the growth
conditions correspond to the ecological requirements of the species. The tree or
shrub must be
present on similar sites in the surrounding area. A
variety of plants in the tree, shrub or herb layer is
indicative of undisturbed woodland and ancient
forests. If hedges or wooded banks have been
planted with locally sourced material the plants
can be considered autochthonous.
The fndings show that autochthonous woody
plants have become seriously endangered in
Flanders, with only about 6 percent of the current forest cover holding
autochthonous woody
plants. Several causes for this loss of autochthonous material are evident. Only 11
percent
of Flanders is now forested and what there is is
highly fragmented as a result of centuries of intensive forest use. Small felds
have been replaced
by large, open expanses of farmland, with the
consequent disappearance of wooded banks, old
hedges and small forests on farmland.
The inventory data (in Flemish) are accessible
on the internet (www.natuurenbos.be).