Advances in Global Change Research 74

Miguel Montoro Girona

Hubert Morin
Sylvie Gauthier
Yves Bergeron Editors

Boreal Forests in
the Face of Climate
Change
Sustainable Management
Chapter 8
Ecological Classification in Forest
Ecosystem Management: Links Between
Current Practices and Future Climate
Change in a Québec Case Study

Pierre Grondin, Marie-Hélène Brice, Yan Boulanger, Claude Morneau,

Pierre-Luc Couillard, Pierre J. H. Richard, Aurélie Chalumeau,
and Véronique Poirier

Abstract Climate change is expected to profoundly impact boreal forests, ranging

from changes in forest composition and productivity to modifications in distur-
bance regimes. These climate-induced changes represent a major challenge for forest
ecosystem management, as information based on ecological classification may no
longer provide a straightforward guide for attaining management goals in the future.
In this chapter, we examine how climate change could influence the use of ecolog-
ical classification and by what means this approach can continue to be relevant for
guiding the ongoing development of management practices. We address these ques-
tions by first describing ecological classification, using the example of Québec’s
classification system, and then showing its importance in forest ecosystem manage-
ment. Using a forest landscape in Québec as a case study, we then look at how
climate change could affect boreal forest ecosystems by presenting a detailed, multi-
step analysis that considers climate analogs, habitat suitability, and changes in forest
composition. We show that at the end of the century, the vegetation of the Abies-Betula
western subdomain will not change sufficiently to resemble that of its climate analog,
currently located ~500 km to the south. Changes in fire frequency and severity could

P. Grondin (B) · A. Chalumeau · V. Poirier

Direction de la recherche forestière, ministère des Forêts, de la Faune et des Parcs du Québec,
2700 rue Einstein, Québec, QC G1P 3W8, Canada
e-mail: [email protected]
A. Chalumeau
e-mail: [email protected]
V. Poirier
e-mail: [email protected]
M.-H. Brice
Jardin botanique de Montréal, 4101 Sherbrooke Est, Montréal, QC H1X 2B2, Canada
e-mail: [email protected]
Département de sciences biologiques, Institut de recherche en biologie végétale, Université de
Montréal, 4101 Sherbrooke Est, Montréal, QC H1X 2B2, Canada
© The Author(s) 2023 219
M. M. Girona et al. (eds.), Boreal Forests in the Face of Climate Change,
Advances in Global Change Research 74,
https://doi.org/10.1007/978-3-031-15988-6_8
220 P. Grondin et al.

significantly modify forest dynamics and composition. Consequently, the potential

vegetation and the successional pathways defined under the current climate could
change and follow new successional trajectories. This possible reality forces us to
question some fundamental aspects of ecological classification. However, we argue
that ecological classification can still provide a valuable framework for future forest
management, particularly in continuing to recognize the various types of ecosys-
tems present along toposequences. Given the changes expected in forest vegetation
composition and dynamics, future variability and uncertainty must be integrated
into the current stable classification units and predictable successional trajectories
of ecological classification.

8.1 Introduction

Ecological classification provides complete descriptions of biophysical forest

features and their historical variability at various spatial scales (global, regional,
local). This ecological information is necessary to inform forest ecosystem manage-
ment, which consists of implementing silvicultural strategies that aim to ensure the
long-term maintenance of ecosystem functions and, consequently, the social and
economic benefits that forest ecosystems provide to society (Chap. 1; Gauthier
et al., 2009). However, climate change is expected to affect growth rates, distur-
bance regimes, species distributions, and, ultimately, biodiversity (Boulanger et al.,
2014, 2016; Périé & de Blois, 2016; Périé et al., 2014; Price et al., 2013). These
changes raise serious concerns for forest ecosystem management if knowledge from
ecological classification can no longer provide a reliable guide for meeting future
management objectives (Millar et al., 2007; Nagel et al., 2017; Puettmann, 2011).
In this chapter, we first introduce ecological classification and the concept of
potential vegetation (Sect. 8.2.1) and then characterize the principles of the Québec
hierarchical classification system (Sect. 8.2.2). We then consider historical variability

Y. Boulanger
Natural Resources Canada, Canadian Forest Service, Laurentian Forestry Centre, 1055 rue du
PEPS, P.O. Box 10380, Stn. Sainte-Foy, Québec, QC G1V 4C7, Canada
e-mail: [email protected]
C. Morneau · P.-L. Couillard
Direction des inventaires forestiers, ministère des Forêts, de la Faune et des Parcs du Québec,
5700 4e Avenue Ouest, Québec, QC G1H 6R1, Canada
e-mail: [email protected]
P.-L. Couillard
e-mail: [email protected]
P. J. H. Richard
Département de géographie, Université de Montréal, Complexe des sciences, P.O. Box 6128, Stn.
Centre-Ville, Montréal, QC H3C 3J7, Canada
e-mail: [email protected]
8 Ecological Classification in Forest Ecosystem Management 221

at regional and local scales (Sect. 8.2.3) and the importance of ecological classifica-
tion in forest ecosystem management (Sect. 8.2.4). Next, using a three-step analysis,
we indicate how climate change could modify boreal forest ecosystems and ecolog-
ical classification (Sect. 8.3.1). In the final section (Sect. 8.3.2), we discuss the issues
and challenges raised by climate change and illustrate how the ecological classifi-
cation framework could integrate future variability in forest dynamics in the context
of risk management. Throughout the chapter, we focus on the boreal biome, using
the case study of the Abies balsamea–Betula papyrifera western subdomain (84,567
km2 ) in Québec, Canada (hereafter the Abies-Betula w. subdomain).

8.2 Ecological Classification of Ecosystems

We define ecological classification as “an analytical process that consists of delin-

eating and defining ecosystems, at different spatial scales, on the basis of the abiotic
and biotic factors that govern their development (climate, disturbances, physical
environment, plant succession) along ecological gradients for the purposes of land
resource management” (Bailey, 2009; Barnes et al., 1982; Rowe & Sheard, 1981;
Sims et al., 1996; Whittaker, 1962). The ecological classification of ecosystems has
developed progressively in parts of the world, including in Canada, e.g., the provinces
of Québec (Fig. 8.1; MFFP 2021; Saucier et al., 2009, 2010), British Columbia
(Banner et al., 1993; MacKenzie & Meidinger, 2018; Pojar et al., 1987) and others
(Baldwin et al., 2020), as well as in the United States (Bailey, 2009), Scandinavia
(Engelmark & Hytteborn, 1999; Sjörs, 1999), and northeastern Asia (Krestov et al.,
2009).

8.2.1 Ecological Classification: Potential Vegetation

Ecological classification is based on the science of phytosociology, which originated

more than a century ago when Flahault and Schröter (1910) defined a plant asso-
ciation or community as “a vegetal grouping of determined floristic composition,
presenting a uniform physiognomy that grows in uniform site conditions.” From the
school of phytosociology emerged the concept of potential natural vegetation (here-
after potential vegetation). This concept refers to a stable state or late-successional
vegetation stage at a particular site under the existing climatic and edaphic conditions
(Baldwin et al., 2020; Blouin & Grandtner, 1971; Braun-Blanquet, 1972; Clements,
1936; Daubenmire, 1968; Küchler, 1964; Tuxen 1956 in Härdtle, 1995). Gradu-
ally, although mainly in the early 1970s, interest focused on understanding natural
disturbances, particularly fire frequency and severity, and the impacts of disturbance
on vegetation dynamics (Damman, 1964; Heinselman, 1973; Rowe & Scotter, 1973;
White, 1979). For example, in some cases, the late-successional stage is never reached
because of recurrent disturbances, such as fire, and only the early- or mid-successional
222 P. Grondin et al.

Fig. 8.1 a Circumboreal zonation modified by permission of Taylor & Francis Group from Saucier
et al. (2015) and Baldwin et al. (2020). Only the largest subdivisions are considered (1–7, dark
green). 1 North European boreal, 2 Western Siberian boreal, 3 Central Siberian boreal, 4 North-
eastern Siberian boreal, 5 Alaska–Yukon boreal, 6 West-central North American boreal, 7 Eastern
North American boreal. b–f Ecological classifications of the ministère des Forêts, de la Faune et
des Parcs du Québec (MFFP 2021 [CC-BY 4.0]; Saucier et al., 2009, 2010, with permission from
Multimondes and AgroParisTech, respectively)
8 Ecological Classification in Forest Ecosystem Management 223

stage is attained after disturbances (Bergeron et al., 2014; Cogbill, 1985; Couillard
et al. 2016; Dansereau, 1957; Frégeau et al., 2015; Frelich & Reich, 1995; Grondin &
Gosselin, 2013; Grandtner, 1966; White, 1979). Anthropogenic disturbances are also
considered an important factor that can modify the dynamics of potential vegetation
(Danneyrolles et al., 2020; Härdtle, 1995; Laflamme et al., 2016; Seastedt et al.,
2008). Québec’s ecological classification system defines potential vegetation as “a
classification unit that includes the complete set of communities associated to the
late-successional stage on a given site,” thus integrating vegetation dynamics within
the concept (Saucier et al., 2009). Potential vegetation is defined on the basis of a
particular set of tree species (early- and late-successional species) and understory
indicator species that grow together on similar site conditions. Potential vegetation is
considered permanent if soil and climatic conditions and the disturbance regime do
not change (Saucier et al., 2009). It is mainly on the basis of this predictable succes-
sional trajectory that the concept has been challenged (Loidi & Fernández-González,
2012).
The concept of potential vegetation is widely used worldwide as the basic unit
of vegetation classification systems (Härdtle, 1995; Loidi & Fernández-González,
2012; Sims et al., 1996). The distribution of potential vegetation (Fig. 8.2) can be
illustrated along a toposequence, which divides the landscape into relatively homoge-
neous sections with respect to topography (slope, elevation), microclimate, drainage,
surficial deposits, nutrient regime, soil type, disturbance regime, and forest dynamics
(Banner et al., 1993; Barnes et al., 1982; Blouin et al., 2008; Rey, 1960; Rowe &
Sheard, 1981; White 1973). This combined use of the physical environment and forest
dynamics is termed a toposequence–chronosequence approach (Damman, 1964).
Potential vegetation typical of mesic sites—zonal soils sensu Baldwin et al. (2020)
and Pojar et al. (1987)—reflects the regional climate (temperature and precipita-
tion). Other potential vegetation characterizing the toposequence is associated with
regional climate and reflects the influence of specific local environmental character-
istics (e.g., Picea-Sphagnum potential vegetation, which grows on poorly drained
sites). Toposequences present a synthetic view of the ecological information about
a landscape that can be used to produce an ecological map of potential vegetation.
Such maps present a global view of the distribution of vegetation over a specific
territory (Blouin & Grandtner, 1971; Küchler, 1964).

8.2.2 Ecological Classification: A Hierarchical Approach

Various hierarchical classifications have been developed for describing and managing
ecosystems at different spatial scales, and each has its own nomenclature and levels
(Bailey, 2009; Baldwin et al., 2020; Banner et al., 1993; Klijn & Udo de Haes, 1994;
MacKenzie and Meidinger 2017; Powell, 2000; Sims et al., 1996). The ecological
classification system developed in Québec (MFFP 2021; Saucier et al., 2009, 2010)
was influenced by European phytosociological classification (Grandtner, 1966) and
224 P. Grondin et al.

Fig. 8.2 Zonation of potential vegetation illustrated along a toposequence characterizing the
southern part of the Abies balsamea–Betula papyrifera western subdomain, Québec. The species
codes are defined in Fig. 8.7

Australian landscape classification (Jurdant et al., 1977). Québec’s ecological classi-

fication system consists of multiple hierarchy levels corresponding to various spatial
scales. At the continental scale (~1,000,000 km2 ), vegetation zones and subzones
correspond to very large territories characterized by a vegetation physiognomy and
floristic composition that reflect macroclimatic conditions controlled by latitude and
continentality. The boreal zone in Québec extends from the northern temperate to
Arctic zones and is part of the eastern North American boreal floristic subdivision
(Fig. 8.1a and b). It is subdivided from south to north into three subzones based
on vegetation structure: closed boreal forest, open boreal forest, and forest–tundra
(Fig. 8.1c). This latitudinal zonation is common in the boreal biome, which forms a
forest belt around the Northern Hemisphere (Fig. 8.1a). At the supraregional scale
(~100,000 km2 ), the bioclimatic domains and subdomains of Québec encompass
large areas characterized by relatively homogeneous mesoclimatic conditions that
are associated with the dominant potential vegetation of mesic sites and a natural
disturbance regime (Fig. 8.1d). At the regional scale (~10,000 km2 ), each ecolog-
ical region is characterized by the relative abundance of a potential vegetation and
their respective physical environments (ecological types); for example, in the Abies
balsamea–Betula papyrifera domain, ecological region 5B differs from region 5C by
having a less rugged relief and much larger areas of organic deposits. This physical
environment leads to a greater abundance of Picea-Sphagnum potential vegetation
on wet organic soils and a lower abundance of Abies-Betula potential vegetation,
which is normally typical of mesic sites in this bioclimatic domain. At the landscape
scale (~100–1,000 km2 ), the regional landscape unit is a relatively homogeneous
portion of land in terms of relief, altitude, surficial deposits, hydrography, and poten-
tial vegetation (Fig. 8.1f). The ecological district, delimited according to the same
criteria as the regional landscapes but at a finer scale, is the basic mapping unit at all
higher levels described above. The local scale (0.1–1 km2 ) is characterized by the
potential vegetation, ecological type, and forest type. The ecological type combines a
potential vegetation type and a physical environment type and is considered a stable
8 Ecological Classification in Forest Ecosystem Management 225

ecological unit. The forest type describes the observed vegetation using physiog-
nomy, overstory tree species composition, and understory indicator species of site
conditions.
In Québec, the boreal zone is subdivided into four bioclimatic domains distributed
north to south along a temperature gradient (Fig. 8.3; Grondin et al., 2007,
2014; MFFP 2021; Richard, 1987; Saucier et al., 2009). The southernmost Abies
balsamea–Betula papyrifera domain comprises mainly mixed boreal forests. The
Picea mariana–moss domain, located to the north of this southernmost domain, is
characterized by closed coniferous boreal forests. Further north, the Picea mariana–
lichen domain is characterized by open forests, whereas the northernmost forest–
tundra domain consists of a mosaic of forest and treeless communities where, with
increasing latitude, forests are increasingly scattered and confined to sheltered loca-
tions up to the Arctic tree line (MFFP 2021; Payette, 1992; Rowe & Scotter, 1973).
Most bioclimatic domains in Québec are subdivided further into two subdomains
(west and east) because of differences in precipitation, physical features, and the
dominant natural disturbance regimes, which produce differences in the dominant
vegetation (Couillard et al., 2019; Grondin et al., 2007). The climate of the western
subdomains is more continental and drier, therefore more susceptible to fire than the
eastern subdomains, which have a maritime influence, as illustrated by ombrothermal
areas (Fig. 8.4a; Rey, 1960; Richard, 1978).
At the local scale, the domain, subdomain, and ecological region are all charac-
terized by a range of potential vegetation distributed along toposequences. In the
toposequence of the southern part of the Abies-Betula w. subdomain (Fig. 8.2), the
warmer hilltops host a mixed boreal vegetation (Betula papyrifera, Abies balsamea,
Picea glauca, P. mariana) with the presence of Acer rubrum, which forms the
Abies balsamea–Acer rubrum potential vegetation (hereafter Abies-Acer). The mesic
midslopes (zonal soils) are occupied by the potential vegetation that best reflects the
climatic conditions of the Abies balsamea–Betula papyrifera bioclimatic domain,
which is the Abies-Betula potential vegetation. The lower subhydric, and often
stoniest, slopes are occupied by the Picea mariana–Abies balsamea potential vege-
tation (Picea-Abies), and Pinus banksiana may also be present. Finally, the flatter
areas host the Picea mariana–Pinus banksiana potential vegetation (Picea-Pinus)
on till or sand, whereas the wet organic soils support the Picea-Sphagnum potential
vegetation.

8.2.3 Ecological Classification: A Historical Perspective

A historical perspective (millennial scale) that considers the range of natural vari-
ability of vegetation can enhance the description of ecological classification units and
forest dynamics. The concept of natural variability was introduced in the 1990s to
incorporate an understanding of past spatial and temporal variability into ecosystem
management (Cissel et al., 1999; Morgan et al., 1994; Powell, 2000; Swanson et al.,
1994). Keane et al. (2009) defined the natural range of variability as “the variation of
226 P. Grondin et al.

Fig. 8.3 Vegetation zonation in Québec according to the level of zone, domain (1–10) and subdo-
main (W, west; E, east; N, north; S, south; C, coastal) (MFFP 2021 [CC-BY 4.0]; Saucier et al.,
2009, with permission from Multimondes). The codes (1–10) are used in Fig. 8.4 to define the
ombrothermal area of the bioclimatic subdomains. The red line is the northern limit of commercially
productive forest (MRN 2013a)

historical ecosystem characteristics and processes (vegetation, disturbance regimes,

climate) over time and space scales that are appropriate for management application.”
An approach that considers the natural range of variability assumes (1) that ecosys-
tems are complex and have a range of conditions that are self-sustaining. Beyond
that range they move into disequilibrium; and (2) historical conditions can indicate
ecosystem health (Keane et al., 2009; Kuuluvainen, 2002; Morgan et al., 1994).
8 Ecological Classification in Forest Ecosystem Management 227

Fig. 8.4 Ombrothermal area of the bioclimatic subdomains of Québec’s boreal forest (Fig. 8.3)
for a the reference horizon (1969–1990) and b the scenario under RCP8.5 for 2100. The subdo-
main codes are composed of the latitudinal gradient (1–10) and the longitudinal gradient (E:east or
W:west). Ellipses trace the 95% confidence level with a multivariate t-distribution of the data gener-
ated from global climate simulations (https://www.climateinteractive.org/tools/world-climate-sim
ulation/). Some subdomains exhibit a wide climatic range because of their large area (6–10) or the
highly variable habitat conditions (5E). The climate of the coastal forest–tundra subdomain (8C)
overlaps with the southern portion of the Picea mariana–mosses subdomain because its vegetation
is strongly influenced by rocky substrate and strong winds

Spatial variability is generally based on ecological units of the ecological classifica-

tion determined at regional (Shorohova et al., 2011) to local scales (MacLean et al.,
2021). Temporal scales define the short-, medium-, and long-term variability of forest
composition, natural disturbances (mainly fire), and climate. This variability is recon-
structed using dendrochronology and paleoecology (Chap. 2). Such information of
historical conditions helps justify and validate management strategies elaborated in
the context of climate change (Gillson & Marchant, 2014; Grondin et al., 2020;
228 P. Grondin et al.

Marcisz et al., 2018). In Québec, information regarding the long-term natural vari-
ability of ecosystems is derived from studies of postglacial vegetation, disturbance
regimes, and climate (Payette, 1993; Richard, 1993).
At the regional scale, the Holocene history of the Abies-Betula w. subdomain
was first characterized by an afforestation period, followed by the Holocene climatic
optimum, marked by the expansion of the temperate species Pinus strobus and Thuja
occidentalis (Fig. 8.5). Subsequently, the abundance of these species was reduced
as their range contracted over the following 3,000 years in response to a cooler
climate and possibly a relatively lower frequency of large fires (Ali et al., 2012).
With time, the modern-day boreal forest became established. The last millennium
has been characterized by a slight increase in P. banksiana and an opening of the
forest cover (Ali et al., 2012; Asselin et al., 2016; Bajolle, 2019; Bajolle et al., 2018;
Carcaillet et al., 2001; Fréchette et al., 2018; Hennebelle et al., 2018; Larochelle
et al., 2018; Richard et al., 2020). Today, this region experiences recurrent spruce
budworm outbreaks every 30–40 years (Saucier et al., 2009), and the fire cycle has
been approximately 275 years for the 1890–2020 period (Couillard et al., 2022).
Studies of recent forest dynamics at the local scale (toposequence, Fig. 8.2) indi-
cate that the Abies-Betula potential vegetation follows a pathway characterized by a
high abundance of B. papyrifera in the early postfire successional stages followed by
an increasing abundance of A. balsamea over time. Mature stands of A. balsamea are
subject to gap dynamics, and some stands are severely affected by spruce budworm
(Choristoneura fumiferana) outbreaks (Bergeron et al., 2014; Couillard et al., 2012;
Navarro et al., 2018; Saucier et al., 2009). The dynamics of the Picea-Abies poten-
tial vegetation can follow one of two pathways. The first pathway has fires main-
taining a stand dynamic dominated by P. mariana, with scattered A. balsamea in
regeneration. The second pathway has a sufficiently long fire interval to support a
successional dynamic toward Picea mariana–Abies balsamea stands. The Picea-
Pinus potential vegetation is controlled primarily by frequent fires that generally
favor the cycling of the same plant communities (recurrence dynamics), i.e., fires
enable the post-disturbance recovery process to produce stands resembling those
of the predisturbance state (Fig. 8.2; Couillard et al. 2016; Frégeau et al., 2015;
Martin et al., 2018). This potential vegetation can be subject to a regeneration failure
that shifts closed-canopy stands toward lichen woodlands. Various causal factors
can explain this transformation; these factors include fire intensity (severe or light),
successive fires within a short period, and the cumulative effect of an insect outbreak
or logging immediately followed by a fire. For these areas, the return of closed-
canopy stands is not possible without direct intervention, such as planting (Couillard
et al., 2021; Girard et al., 2008; Pinno et al., 2013; Schab et al., 2021; Splawinski
et al., 2019).
8 Ecological Classification in Forest Ecosystem Management 229

Fig. 8.5 Postglacial vegetation and environmental data of the Abies balsamea–Betula papyrifera
western subdomain, Québec. Pollen curves for selected coniferous (5) and broadleaf (2) tree taxa
from Lake Lili (Bajolle, 2019) are coupled with a chironomid-based reconstruction of August air
temperature at Lake Aurélie (Bajolle et al. 2018), a millennial reconstruction of annual precipitation
and summer sunshine for the entire subdomain (Fréchette et al., 2018), and a regional summary of
past climate, sequential cover type, forest density, and fires according to Bajolle (2019). Note that
some taxa of the diagram are greatly underrepresented by the pollen they shed compared to their
abundance in the field (e.g., A. balsamea, P. tremuloides, T. occidentalis), whereas others are highly
overrepresented (e.g., P. strobus, B. papyrifera). The maximum values of the P. banksiana pollen
curve at the base of the sequence (7,000–8,500 cal yr BP) reflect mainly a long-distance, northerly
transport of extraregional pollen during the afforestation stage

8.2.4 Ecological Classification: A Foundation for Forest

Ecosystem Management

Ecological classification is a framework that provides users with a straightforward

structure to practice ecosystem-based management, facilitate modeling of the spatial
distribution of ecosystems, and evaluate the impacts of changing climate and envi-
ronmental conditions on vegetation (MacKenzie & Meidinger, 2018; Sims et al.,
1996). For example, MacLean et al. (2021) present a new approach for establishing
management guidelines defined for each potential vegetation in Nova Scotia. The
forest management system developed in Québec also relies on potential vegetation
230 P. Grondin et al.

(Grondin et al., 2003; Saucier et al., 2010). More specifically, a specific silvicul-
tural scenario is developed for each ecological type or groups of ecological type
(MFFP 2020; MRN 2013b). Moreover, Québec forestry is oriented toward calcu-
lating the annual allowable cut for each management unit and for which poten-
tial vegetation plays a crucial role (Fortin & Langevin, 2010). Recently, a northern
limit of commercially productive forest was established on the basis of relation-
ships between fire regime, regeneration, and the time required for trees and forests
to reach a merchantable volume, as calculated by ecological district and considering
the potential vegetation (Fig. 8.3; Gauthier et al., 2015b; MRN 2013a).
Some management activities also consider reference conditions that serve to
ensure long-term ecosystem health. In Québec, reference conditions for defining
targets for old-growth proportions and forest composition were established at the
spatial scale of bioclimatic subdomains (Fig. 8.3; Boucher et al., 2011). These refer-
ence conditions are based on studies of natural variability, mainly related to the fire
regime of the last century or sometimes an even longer period (~200 years). The fire
regime is documented from stand origin maps—which date the last fire—and from
which a fire cycle and a proportion of old-growth forest are calculated. The long-
term range of natural variability does not determine reference conditions because
the links between long-term history and forest management must be clarified, as per
Hennebelle et al. (2018). They showed that a relatively high proportion of old-growth
forest existed throughout the Holocene in Québec. Such information would suggest
that estimates based on the recent past are similar to those based on long-term history.
By using ecological classification information to manage Québec’s forest ecosys-
tems, forest managers (1) describe the natural forest, (2) compare it to the managed
forest to highlight the differences induced by forest management, (3) translate this
information into ecological issues to be resolved and (4) integrate these issues
into management plans to reduce differences between natural and managed forests
(Harvey et al., 2009; MFFP 2017). In the Abies-Betula w. subdomain, for example,
the reference conditions indicate that even-aged mature stands (older than 80 years)
represent 30% of the subdomain compared with 31% for uneven-aged old-growth
forests (Boucher et al., 2011). These thresholds serve as guidelines to define targets
for ecosystem management plans (MFFP 2015). However, Martin et al. (2021)
showed that in the Picea mariana–mosses domain, when local features of potential
vegetations are not sufficiently considered in forest landscapes, old-growth forests are
preserved essentially on the least productive sites, such as forested wetlands (Picea-
Sphagnum potential vegetation); the most productive sites are harvested. The system
of protected areas in Québec, in which a variety of forest stands are protected, and
relatively small, protected areas (i.e., biological shelters), which protect in particular
old-growth forest, partially avoid the pattern of old-growth forests being found only
on the least productive sites. It is clear that local conditions must be better integrated
within targets set at the landscape scale, particularly with respect to complex forest
structures and their attributes (e.g., deadwood). In the context of ecosystem manage-
ment, adequate protection and adapted silviculture (partial cutting) of old-growth
forests specific to each potential vegetation are key to ensuring the resilience and
8 Ecological Classification in Forest Ecosystem Management 231

adaptive capacity of boreal forest ecosystems under global change (Martin et al.,
2018; Shorohova et al., 2011).
Overall, ecological classification is a valuable tool for summarizing spatiotem-
poral information to help guide effective conservation and forest management prac-
tices (Keane et al., 2009; Loidi & Fernández-González, 2012; MacLean et al., 2021).
However, ecological classification was developed assuming a relatively stable climate
and that potential vegetation recovers its initial structure and composition after a
disturbance (Loidi & Fernández-González, 2012; Powell, 2000; Saucier et al., 2009).
Future climate change challenges the validity of predictable successional pathways
and stable potential vegetation dynamics. This assumed predictability requires a
reanalysis of ecological classification in light of the uncertainties associated with
climate change by considering different successional trajectories for a given poten-
tial vegetation. Some mechanisms already exist in Québec’s ecological classification
system, particularly in forest mapping activities, to account for observed changes in
potential vegetation, such as shifts from closed-canopy stands (Picea-Pinus poten-
tial vegetation) to lichen woodlands (Picea mariana–lichens) caused by regeneration
failure. Over the past 15 years, several studies have attempted to improve our under-
standing of natural variability at regional and local scales (Sect. 8.2.3; e.g., Couillard
et al. 2016; Fréchette et al., 2018). These studies have greatly contributed to our
knowledge of forest dynamics and the concepts of stability and predictability associ-
ated with Québec’s ecological classification. However, vegetation changes that can
be anticipated because of ongoing climate change have yet to be considered. The
following section reflects on strategies for bringing ecological classification into the
future.

8.3 Ecological Classification and Climate Change

Future environmental conditions are expected to differ markedly from present-day

conditions. This shift will occur faster and at a greater magnitude than at any time
since the end of the last glaciation (IPCC 2021). Certain projections have ecosys-
tems falling beyond their natural range of variability (Seastedt et al., 2008). In North
America, Keane et al. (2020) projected significant changes for potential vegetation in
Montana, United States, along an altitudinal gradient. Wang et al. (2012) showed that
suitable habitats in British Columbia, Canada, for grasslands, dry forests, and moist
continental cedar–hemlock forests would expand and habitats for boreal, subalpine,
and alpine ecosystems would decrease. Moreover, in the adjacent province of Alberta,
wildfire activity is projected to accelerate the conversion of about half the province’s
upland mixed and coniferous forests to more climate-adapted deciduous woodlands
and grasslands by 2100 (Stralberg et al., 2018). In Siberia, the expansion of dark
coniferous forest—currently found in the southern boreal forest—into light conif-
erous forest (Larix laricina northern forest, often underlain by permafrost) has also
been documented (Gauthier et al., 2015a; Kharuk et al., 2007).
232 P. Grondin et al.

The consequences of climate change for Canada’s boreal forests will be numerous;
these impacts include changes in forest composition and growth rates, shifts in distur-
bance regimes, and, ultimately, altered biodiversity levels (Price et al., 2013). These
climate-induced changes in forest ecosystems thus represent a major challenge for
forest management. Climate projections must be translated into concrete information
and guidelines to prepare and assist managers in adapting forest management plans
to these changes (Forestier en chef, 2020; Gauthier et al., 2014, 2015a; Millar et al.,
2007; Nagel et al., 2017; Puettmann, 2011; Thiffault et al., 2021). In this context,
climate-induced changes could be forecast from the information and models available
at the different levels of the ecological classification system already in use in forest
ecosystem management. The following questions are thus appropriate to consider in
the discussion below:
• Question 1. Will climate change modify vegetation along the various ecological
classification scales, i.e., from bioclimatic subdomains to potential vegetation?
• Question 2. Can knowledge derived from ecological classification help respond
to the challenges inherent to climate change?

8.3.1 A Three-Step Analysis to Characterize Changes

in Climate, Species Sustainability, and Forest Dynamics

In the following section, we present an analytical approach that will address these two
questions using information generated at different levels of ecological classification.
This approach proceeds through several steps that we consider to be prerequisites for
determining silvicultural strategies in the context of climate change. Our method can
be adapted according to specific regional and local contexts. The aim is to illustrate
how climate change may impact forests at the relevant scale by focusing on the Abies
balsamea–Betula papyrifera w. subdomain in the province of Québec and comparing
current vegetation patterns with those developed under the RCP8.5 scenario from
now to 2150. As explained by the IPCC (2021),
Representative Concentration Pathways (RCP) are scenarios that include time series of emis-
sions and concentrations of the full suite of greenhouse gases (GHGs) and aerosols and
chemically active gases, as well as land use/land cover … RCPs usually refer to the portion
of the concentration pathway extending up to 2100, for which Integrated Assessment Models
produced corresponding emission scenarios.

The RCP8.5 scenario is generally considered the most severe anthropogenic

climate forcing scenario. Indeed, under this pathway, the forcing reaches 8.5 W·m−2
in 2100 and continues to increase for some time afterward. Under this scenario, the
temperature in the studied bioclimatic subdomain will increase 7–8 °C and precipi-
tation 7% by 2100 (Boulanger & Pascual Puigdevall, 2021). The Abies-Betula w.
subdomain will thus likely experience a longer fire season and more fire-prone
weather conditions, which could increase the annual area burned by 2 to 4 times
by the end of the century, as projected for various regions in Canada (Boulanger
8 Ecological Classification in Forest Ecosystem Management 233

et al., 2014). Ecological processes including tree growth, regeneration, interspe-

cific competition, and forest productivity could also be affected by these climate
changes; these altered processes could ultimately modify the forest assemblage
currently defining each potential vegetation (Boulanger & Pascual Puigdevall, 2021).
Our approach involves acquiring information about future climate analogs (Step 1),
defining potential impacts on tree species–habitat suitability at the subdomain scale
(Step 2), and finally considering local information at the potential vegetation scale
to assess changes in forest dynamics (Step 3).

8.3.1.1 Step 1. Identifying Contemporary Climate Analogs

A climate analog is a simple tool for visualizing the magnitude of climate change
at a relatively large spatial scale. This tool “identifies locations for which historical
climate is similar to the anticipated future climate at a reference location” (Grenier
et al., 2013). Here, the approach shows how the temperature and precipitation of
the Abies-Betula w. subdomain (Fig. 8.3) will be modified under RCP8.5 during the
2071–2100 period (code 5 W in Fig. 8.4b). Under this scenario, this subdomain will
experience a climate analogous to that currently experienced by the Acer saccharum–
Carya cordiformis bioclimatic subdomain (code 1 in Fig. 8.4b), located ~500 km to
the south. Northern temperate hardwoods characterize this latter area, with several
thermophilous species (e.g., Quercus, Carya, Tilia) reaching the northernmost limit
of their range and small-gap dynamics being the most common natural disturbance.
Therefore, this climatically analogous region includes very different tree species
assemblages and experiences markedly different disturbance dynamics than those
currently observed in the Abies-Betula w. subdomain.

8.3.1.2 Step 2. Assessing Future Tree Species Habitat Suitability

at the Regional Scale

Because the climate analog provides only partial insight into the impacts of climate
change on forest ecosystems, it is important to identify to what extent the projected
climate in the Abies-Betula w. subdomain will be suitable for species currently
observed in the area and those taxa that could potentially migrate into the subdo-
main. Niche models (Boisvert-Marsh et al., 2014; Périé & de Blois, 2016; Périé et al.,
2014) are used to assess the climatic vulnerability of the dominant tree species of
the subdomain (Fig. 8.6). Under the RCP8.5 climate scenario for 2071–2100, niche
models project that much of this subdomain will become a less favorable habitat for
most conifers and B. papyrifera. Populus tremuloides habitat will remain present at
a similar abundance (status quo), whereas the situation appears more critical for P.
banksiana, as the model indicates a less favorable habitat over much of the subdo-
main. Acer rubrum, a thermophilous species that favors the warmer hilltops (Fig. 8.2)
of the southern part of the western subdomain, has a distinct profile from the other
species as new suitable habitats will become available. These changes in habitat
234 P. Grondin et al.

Fig. 8.6 The impact of climate change on tree habitat suitability under scenario RCP8.5 (2071–
2100) as defined by niche models. The baseline (1969–1990) range of a species is the total area
(km2 ) of all cells where the baseline average model predicted a suitable habitat for that species in
the Abies balsamea–Betula papyrifera western subdomain and within the Abies balsamea–Betula
papyrifera potential vegetation

suitability suggest that species that have traditionally defined entire regional vegeta-
tion assemblages could become less adapted to their particular regions, resulting in
significant impacts on ecosystems.

8.3.1.3 Step 3. Assessing Local Change in Forest Composition

and Dynamics

How will the climate-induced vulnerability of these species affect the dynamics
of specific potential vegetation? Evaluating such impacts requires a more thorough
assessment of climate-induced changes on processes at the stand- (e.g., interspecific
competition, soil characteristics, productivity) and landscape- (e.g., seed dispersal,
natural disturbances) scales, which govern finer-scale forest dynamics. Differences
in soil characteristics, topographical position, and current tree species assemblages
may thus impact how specific potential vegetation will respond to climate change.
Climate-induced changes in natural disturbance regimes are another key component
to consider. Simulating changes in the disturbance regime enables assessing possible
future alterations in the natural range of variability (Fig. 8.5), which is not explic-
itly accounted for within niche models. Using a forest landscape model (LANDIS-
II; Scheller et al., 2007), we projected the future long-term compositional change
of two contrasting potential vegetation types characteristic of the Abies-Betula w.
subdomain, i.e., Abies-Betula and Picea-Pinus potential vegetation. We simulated
130 years, starting in 2020, under baseline and RCP8.5 climate scenarios. To assess
the impact of changes in the natural disturbance regime, we simulated future forest
landscapes under RCP8.5, considering the current fire regime (status quo) and a
future fire regime under climate change projections.
We found that under the RCP8.5 projected climate conditions and regardless of
the fire regime, i.e., current or projected fires (Fig. 8.7), there is a significant decrease
8 Ecological Classification in Forest Ecosystem Management 235

in total biomass after 2070 for both potential vegetation types, primarily caused by
a large decrease in the biomass of A. balsamea and/or P. mariana, the two dominant
boreal species. These decreases in total biomass are likely to be even greater when
considering the concomitant climate-induced change in fire regimes (Boulanger
et al., 2014, 2016). Our analysis distinguishes different alterations in forest dynamics
specific to each potential vegetation. For instance, A. rubrum and B. papyrifera are
projected to sharply increase in the Abies-Betula potential vegetation, whereas P.
tremuloides biomass will remain stable or increase only slightly. The increase of A.
rubrum is greater in the current fire regime than in the projected fire regime. In the
Picea-Pinus potential vegetation, however, P. tremuloides sharply increases, whereas
A. rubrum remains a minor component of the forest landscape. Still here, A. rubrum
is greater in the current fire regime and after 2100. Therefore, by 2100, the Abies-
Betula potential vegetation will be dominated mainly by conifers accompanied by
B. papyrifera, P. tremuloides, and A. rubrum. Beyond 2100, and given the projected
fires, broadleaf species (P. tremuloides, B. papyrifera, A. rubrum) will dominate. In
the Picea-Pinus potential vegetation, P. tremuloides and P. banksiana will be the
most common species. The projected dynamics for both potential vegetation types
are consistent with contemporary trends found in other studies (Boisvert-Marsh et al.,
2014; Brice et al., 2019; Fisichelli et al., 2014; Iverson et al., 2008; Jain et al., 2021).

8.3.2 Ecological Classification and Climate Change: The

Main Issues

Several conclusions can be drawn from this multistep analysis to answer both ques-
tions posed above. First, we are interested in specifying how climate change will
modify the current vegetation (Question 1). At the regional scale, the future vege-
tation of the Abies-Betula w. subdomain will not correspond to that identified by
its climate analog by the end of the century (Fig. 8.4 and Step 1). Given dispersal
limitations (Iverson et al., 2011) and forest inertia (Brice et al., 2020), it is unlikely
that the thermophilous species associated with the Acer saccharum–Carya cordi-
formis subdomain will keep pace with the >500 km northward migration of the
ombrothermal area (Boulanger & Pascual Puigdevall, 2021; Taylor et al., 2017).
With climate niches of most boreal species deteriorating within the Abies-Betula w.
subdomain, this scenario will likely introduce a climate debt (Taylor et al., 2017),
with current species assemblages being strongly maladapted to future climate condi-
tions. Under such conditions, it is improbable that regional vegetation will be at a
climate equilibrium, at least for the next several decades.
At the local scale (Fig. 8.7), our model (LANDIS-II) also suggests that increased
disturbance rates will cause a decline in the biomass of coniferous species and provide
pioneer deciduous tree species a competitive advantage because they can reproduce
vegetatively after a disturbance (Boulanger et al., 2019; Landhäusser et al., 2010).
If stands of Abies-Betula potential vegetation become dominated by P. tremuloides
236 P. Grondin et al.

Fig. 8.7 Mean stand aboveground biomass of forest species in the Abies balsamea–Betula
papyrifera western subdomain according to the LANDIS-II model and projections using the RCP8.5
scenario. The panels represent the Abies-Betula potential vegetation in midslope (left panel) and
the Picea-Pinus potential vegetation (right panel) in flat areas (Fig. 8.2). In the bottom left chart of
each panel, the current fire regime remains relatively unchanged in the future. In the bottom right
chart of each panel, the fire regime is altered in relation to projected climate change

and A. rubrum, will this community be considered (1) a mid-successional stage of

the Abies-Betula potential vegetation with a greater abundance of A. rubrum than
expected under current climate conditions, (2) a transition stage toward an Abies-
Acer potential vegetation, or (3) a new potential vegetation? Knowledge of forest
dynamics under climate change is not sufficiently advanced to project which of the
three possibilities will prevail in the future.
Furthermore, we project that warmer conditions should favor the growth and repro-
duction of thermophilous species (e.g., Acer spp.), which will then be more likely
to outcompete boreal species in the potential vegetation where they are currently
thriving. Such conditions could favor the migration of Abies-Acer potential vegeta-
tion to mid-hillside, a topographic position where conditions are not currently suitable
to support such potential vegetation in the Abies-Betula w. bioclimatic subdomain
(Fig. 8.2).
8 Ecological Classification in Forest Ecosystem Management 237

For Picea-Pinus potential vegetation, dynamic modeling (LANDIS-II) suggests

a significant amount of P. banksiana will be maintained (Fig. 8.7), even when niche
models predict a less favorable habitat for this species. This divergence relates to the
niche models relying only on climate, whereas LANDIS-II considers several other
elements, including the impacts of the fire regime.
Climate change may therefore affect vegetation as we know it today at different
scales of ecological classification, from the bioclimatic subdomain to the potential
vegetation level. At the same time, a great deal of uncertainty remains regarding future
climate and vegetation changes. The high uncertainty means that anticipated changes
in vegetation under climate change will likely be intermediate, falling between
those projected under baseline conditions (minimum change) and those of RCP8.5
(maximum change). The variability of climate scenario projections remains great
given the highly unpredictable global, political, and social conditions that will exert
a strong influence (IPCC 2021). Moreover, changes in climate conditions are likely
to evolve over several decades; therefore, it could take decades or even centuries for
highly inertial forest ecosystems, i.e., those resistant to change, to modify signif-
icantly under future climate conditions. Finally, climate and forest models each
have their predictive limitations and hence intrinsic uncertainty in addition to the
uncertainty of climate scenarios.
How can knowledge of natural variability (Sect. 8.2.3; Fig. 8.5) help us better
interpret future vegetation changes? Past patterns in vegetation illustrate previous
dynamics that can recur in the future and, therefore, help researchers formulate
hypotheses about the limits of variability that species have experienced in the past
and that may be exceeded in the future (Gillson & Marchant, 2014; Grondin et al.,
2020; Marcisz et al., 2018).
1. In the Abies-Betula w. subdomain, both types of modeling, corresponding to
the niche models (Step 2) and the LANDIS-II model (Step 3), suggest that by
2100 there will be fewer favorable habitats for conifers and a gradual decline in
their biomass. If the abundance of conifers becomes relatively low, as predicted
for RCP8.5, then there will be fewer conifers in the landscape than over the
past several thousand years (Fig. 8.5). It is difficult to compare the Holocene
abundance of coniferous species with that expected in the future; thus, we cannot
determine whether future landscapes will move beyond the range of their past
natural variability.
2. Populus tremuloides and Pinus banksiana are predicted to be abundant in the
future (Fig. 8.7), albeit within a different context compared with the early
Holocene when fires were frequent, and vegetation was in the afforestation stage
(Fig. 8.5). After the afforestation period, and mainly from 5,000 to 2,000 yr BP,
broadleaf species (B. papyrifera) remained abundant; the future Abies-Betula w.
subdomain may present a similar vegetation pattern (Fréchette et al., 2018).
3. Long-term regional natural variability indicates that P. banksiana has increased
over the last millennium in western Québec (Fig. 8.5). This trend should continue
under climate change with an increased fire incidence given the great ability of
P. banksiana to renew itself in a recurrence dynamic (Couillard et al. 2016;
238 P. Grondin et al.

Frégeau et al., 2015; Payette et al., 2012). LANDIS-II modeling also supports
this hypothesis, although there is a significant decrease in biomass, indicating
that the stands will be younger. It is difficult to comment on the spatial variation
of Pinus banksiana stands; however, one hypothesis holds that this species will
increase in geographic extent (Schab et al., 2021; Splawinski et al., 2019). All
these dynamics depend on the initial species composition at each site (greater or
lower abundance of P. banksiana), the fire cycle, and the species’ response to a
potentially shortened cycle.
4. Finally, the Holocene history of the Abies-Betula w. subdomain shows that P.
strobus and T. occidentalis were historically well represented in the landscape
during warmer climatic periods (Fig. 8.5). Both species are not considered in
contemporary modeling because of their low current abundance. On the other
hand, habitat models suggest that habitats will be suitable for the expansion of
these species (Périé et al., 2009). This migration will nonetheless depend on
multiple factors, e.g., the inertia of the vegetation.
Overall, comparing past and future scenarios allows us to better situate the poten-
tial magnitude of future vegetation change (e.g., the decreased abundance of conif-
erous species), base future projections on specific patterns observed during the
Holocene, e.g., broadleaf species such as Betula papyrifera, and propose hypotheses
regarding future vegetation change, e.g., P. banksiana, P. strobus, T. occidentalis.
Can knowledge generated by ecological classification (Sect. 8.1) be used to
address the challenges inherent to climate change (Sect. 8.2)? (Question 2). Despite
projected changes in forest ecosystem composition and dynamics at various spatial
scales, we argue that current ecological classification can still provide a useful frame-
work to inform future forest management (Loidi & Fernández-González, 2012). The
characteristics that currently define potential vegetation at the local scale (soil, surfi-
cial deposits, position, and slope) will continue to develop independently in future
ecological domains, e.g., Abies-Betula versus Picea-Pinus. The assemblages within
each potential vegetation will evolve according to the competitive abilities of its
various species. The future toposequence will continue to show a gradation of poten-
tial vegetation along a physiographic gradient. Each potential vegetation will have
a specific future successional pathway, albeit different from the current pathway
(Fig. 8.7). Moreover, each potential vegetation distributed along the toposequence
will continue to be distinct from other assemblages, and the links with the current
vegetation will likely remain for a long duration. Studies on how ecosystem inertia
could limit future vegetation dynamics must be undertaken. Overall, the hierarchy
of ecological classification (regional vs. local) and the specificity of each potential
vegetation will continue to be paramount for strategic forest planning under climate
change.
8 Ecological Classification in Forest Ecosystem Management 239

8.4 Conclusions

It is evident from our first question (Question 1) that climate change will modify
the principles of ecological classification. Given the expected modifications in forest
vegetation composition and dynamics, we must integrate the expected variability and
uncertainty of forest vegetation composition and dynamics into the current stable
and predictable potential vegetation dynamics. The results presented in this chapter,
through a three-step analysis, provide a preliminary view of the links between current
ecological classification and forest management under climate change. This infor-
mation is limited in scope and should be interpreted with caution. New knowledge,
more complete than that currently available, will be added rapidly, especially through
projects that address potential vegetation dynamics under climate change. In regard to
the possible use of ecological classification in the context of climate change (Question
2), we documented that each potential vegetation could change and be characterized
by its own successional trajectory. Thus, we must maintain and manage this diversity.
Ecological classification can and must assist forest managers in estimating poten-
tial future changes despite inherent variability and uncertainty. The challenge is to
select target species and silvicultural scenarios at the regional and local levels on the
basis of the different trajectories that current forest stands are projected to follow
under increased temperatures and precipitation amounts and changing disturbance
regimes. “Decision makers within any institution, therefore, have to find their own
way through sometimes conflicting information and face the prospect of planning
with and for uncertainty” (Gauthier et al., 2014).

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