Fire is a stronger driver of forest than

logging in the boreal forest of eastern

Canada
Yan Boucher,Isabelle Auger,Jean Noël,Pierre Grondin,Dominique Arseneault,
First published: 14 October 2016

https://doi.org/10.1111/jvs.12466

Abstract
Aims
Our study aimed to: (1) document the preindustrial (1925) forest composition prior to
extensive logging; (2) document the magnitude of changes from 1925 to 2005; and (3)
identify the relative influence of logging and natural disturbances as drivers of the present-
day forest composition.

Location
Boreal forest in central Quebec, eastern Canada.

Methods
We used a dense network of georeferenced historical (~1925) forest plots (n = 30 033) to
document preindustrial forest composition. We evaluated the magnitude of changes with
the present-day using modern plots (1980s to 2000s). We reconstructed a long-term,
spatially explicit history of logging, spruce budworm outbreaks (Choristoneura
fumiferana [Clem.], SBO), and fire using historical maps and field surveys.

Results
In the preindustrial period, late successional coniferous taxa (Abies
balsamea and Picea spp.) dominated the landscape, whereas early successional deciduous
taxa (Betula spp. and Populus spp.) were confined to recently burned areas. In the present-
day landscape, large areas dominated by late successional coniferous taxa have been
replaced by early successional deciduous taxa. Forest communities dominated by early
successional deciduous taxa increased sharply throughout the study area. Logging has been
a minor driver of these changes compared to fire and SBOs.
Conclusions
This study demonstrates the importance of documenting the long-term history of both
anthropogenic and natural disturbances in order to assess their relative contributions to the
development of the present-day forest ecosystems. Natural disturbances have remained the
main drivers of forest composition during the 20th century, whereas logging played a less
important role. In the current context of global change, long-term experimental research is
required to help forecast impacts of natural disturbances and forest management on boreal
forest composition.

Introduction
Land-use change is one of the most important agents of global change, and has important
consequences for forest ecosystems (Ellis 2015; Gauthier et al. 2015). Over the last century,
management for timber production has extensively affected ecosystems of North America
and Eurasia (Spies et al. 1994; Östlund et al. 1997; Boucher et al. 2014). In most areas of
the northeastern United States and Canada, logging causes more tree mortality than all
other factors combined (Cyr et al. 2009; Canham et al. 2013). Such an additive and
relatively periodic human-induced disturbance in forest ecosystems has altered the natural
disturbance regime by increasing the frequency and modifying the nature of disturbances
(Turner & Gardner 2015). Clear-cut logging creates large and open environments that are
suitable for colonization by early successional species (Keenan & Kimmins 1993) and may
thus drive forest composition outside of its natural range of variability (Landres
et al. 1999).

From a biodiversity conservation perspective, it is fundamental to evaluate the long-term

impact of logging on tree species composition and to adjust forestry practices if major
changes are observed (Lindenmayer et al. 2006). Yet, several factors complicate this
evaluation. First, natural disturbances interact with logging and may mask or alter its long-
term influence. Second, the forest composition prior to the early 20th century is unknown in
many areas, making it difficult to detect changes. Third, it is challenging to draw a spatially
explicit, long-term (>80 yr) regional picture of natural and anthropogenic disturbances (but
see Weir & Johnson 1998; Boucher & Grondin 2012).

However, in eastern Canada, Price Brothers & Company evaluated wood volumes within
its timber harvest limits by conducting extensive and detailed forest inventories from 1924
to 1926. Tree stems were identified to the species level and measured at breast height
(DBH) in a very large number of plots (Delisle-Boulianne et al. 2011; Terrail et al. 2014).
Because a large part (~60%) of the forests then surveyed had remained unlogged, these
inventories provide a unique opportunity to document the ‘preindustrial’ forest
composition. Moreover, exhaustive maps of logged areas (1925–2005), fires (1820–2005)
and spruce budworm (SBO) (Choristoneura fumiferana [Clem.]) outbreaks (1938–1985)
can be used to statistically assess the relative contribution of these agents on the ensuing
compositional changes. Accordingly, this study aims to: (1) document preindustrial boreal
forest composition at the regional scale, at the start of the industrial logging period in the
early 20th century (~1925); (2) document the magnitude of changes from 1925 to 2005; and
(3) identify the relative influence of logging and natural disturbances as driver of the
present-day forest composition. To our knowledge, this is the first study to combine
extensive forest inventories from the early 20th century with detailed, spatially explicit
logging and natural disturbance histories in order to disentangle the relative contributions of
these agents on the long-term forest composition.

Methods
Study area
The study area is located in south-central Quebec, in eastern Canada, at the southeastern
range limit of the boreal forest (Rowe 1972). It extends from 48°21′°N to 47°37′°N and
from 72°07′°W to 69°43′°W (Fig. 1a), in the Laurentian Highlands of Quebec, within the
Canadian Shield's Grenville Province. The topography consists of low hills (100–350 m)
along the northern section bordered by the Saguenay and the Saint-Lawrence Rivers, and of
high hills (550–1000 m) in the central and southern sectors (Fig. 1b). Mean annual
precipitation varies from 800 to 1600 mm, of which 30% to 40% falls as snow. Mean
annual temperatures vary from −2.5 to +2.5 °C (Robitaille & Saucier 1998). The study area
contains the ‘South Kenogami’ (3650 km2) and ‘Lower Saguenay’ (2400 km2) timber
harvest limits (Fig. 1b), which were originally owned by Price Brothers & Company from
the late 19th century to the early 1960s. The Price Brothers & Company archives contain
forest inventory plots, forest maps and extensive information about past forest
management, dating back from the early 20th century until ca the 1960s (Delisle-Boulianne
et al. 2011).
Figure 1

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Location of the study area in the boreal forest of south-central Quebec, eastern Canada (a); timber
harvest limits, elevation gradient, present-day agricultural areas and location of cities, villages and
hamlets (b); timeline of major historical events in the study area (c). Modified from Boucher et al.
(2014).

Currently, the major forest types are comprised of balsam fir (Abies balsamea [L.] Mill.),
black spruce (Picea mariana [Mill.] B.S.P.), paper birch (Betula papyrifera Marsh.) and
trembling aspen (Populus tremuloïdes L.). Research conducted within the study area
(Blais 1965; Boucher et al. 2014) indicates that the natural disturbance regime is
characterized by stand-replacing fires and SBO. Despite the fact that the Innu First Nation
occupied the area well before arrival of the first Europeans, nothing in the ethnographic
literature confirms their use of fire, contrary to other parts of northeastern North America
(Tulowiecki & Larsen 2015). Although Europeans began colonizing the surrounding
landscape near Québec City at the end of the 17th century, settlement only started around
the mid-19th century (Fig. 1c). This coincided with the onset of selective logging for saw
timber of large-diameter trees, mainly pines (Pinus spp.) and white spruce (Picea glauca
Moench [Voss]), along main watercourses. Industrial clear-cut logging targeting conifer
trees began in the early 20th century with the rise of the pulp and paper industry (Girard &
Perron 1989). Throughout the 20th century, natural regeneration was the main process that
allowed forest re-establishment after logging. By comparison, conifer plantation was a less
common practice.

Data sources – preindustrial forest (1925)

The main data used to reconstruct forest composition prior to logging are forest plots
(n = 30 033) derived from an early inventory conducted by Price Brothers & Company
from 1924 to 1926, preserved in the Archives nationales du Québec. In the rest of the
article, we will refer to this period as 1925, and associate it with the preindustrial forest.
From the original, larger data set (63 751 plots), we removed all plots located in areas
disturbed by logging prior to 1925. The methodology used by the company was described
in its management plans (Price Brothers & Company, Ltd. 1940; Delisle-Boulianne
et al. 2011). Half-chain-wide (~10 m) inventory transects were systematically placed
perpendicular to the main rivers at ~800-m (0.5 mile) intervals and divided into contiguous
rectangular plots (2 chains × 0.5 chain, i.e. ~400 m2) over the entire study area. Each plot
was georeferenced from its position on a 1925 forestry map, with the aid of ArcGis 9.1
software (Environmental Systems Research Institute, Redlands, CA, US). The species and
DBH (in 2-inch or ~5 cm classes) of all trees with a DBH >3 inches (~7.6 cm) in each plot
was recorded on a tally sheet during the original survey. Inventoried taxa or species (in
order of importance) were balsam fir, black spruce, white spruce, all spruces (Picea spp.),
paper birch, aspens (Populus spp.), yellow birch (Betula alleghaniensis Britt.), eastern
white cedar (Thuja occidentalis L.), white pine (Pinus strobus L.), red pine (Pinus
resinosa Aiton), jack pine (Pinus banksiana Lamb.), all pines (Pinus spp.), tamarack
(Larix laricina K. Koch), maples (Acer spp.), ashes (Fraxinus spp.) and red oak (Quercus
rubra L.). In order to avoid possible inconsistencies in species identification, we grouped
the following species at the genus level: Picea spp. (P. mariana,
P. glauca), Betula spp. (B. papyrifera, B. alleghaniensis), Populus spp. (P. tremuloides,
P. balsamifera L., P. grandidentata Michx.) and Pinus spp. (P. banksiana, P. strobus,
P. resinosa). Less common tree species (maples, ashes, red oak and others) were grouped
under ‘other spp.’.
Data sources – contemporary forest (2005)
The present-day forest composition was described from plots (n = 3 321) surveyed during
the three most recent forest inventories (1980s, 1990s, 2000s) conducted by the
Government of Quebec. For the rest of the article, we will refer to this period as 2005, and
associate it with the present-day forest. The 400-m2 circular plots are distributed along
transects that vary from 500 to 1500 m in length and are arranged according to a stratified
random sampling plan in order to represent the main stand types. Tree species and DBH
(>7 cm) in 2-cm classes were recorded in each plot (MRNQ 2008). To allow comparisons
between the preindustrial and present-day forests, we converted the diameters taken in 1925
using the imperial system to current standard metric units. The difference of 0.6 cm
between the lower DBH thresholds of 7.6 cm in 1925 vs 7 cm in 2005 was compensated for
by the fact that the two periods are compared on the basis of the relative abundance of the
taxa. Species and taxa inventoried in 2005 were reclassified according to the 1925
classification.

Data sources – Explanatory variables and disturbance

history
Explanatory variables were divided into three data sets that correspond to a
priori important vegetation drivers: logging, natural disturbances (fire and SBOs) and
physiographical variables (elevation a.s.l. and drainage class). Logging and fire histories
were obtained from the study of Boucher et al. (2014), who used old forestry maps and
field survey to reconstruct the spatially explicit history of clear-cut logging before
mechanization (1925–1965 [hereafter referred to as the early logging period]) and during
the modern mechanized period (1966–2005 [the modern logging period]), as well as the
history of stand-replacing fires >200 ha from 1820 to 2005. Because fires are harder to
detect as time passes, 200 ha is usually the minimum size used to reconstruct fire history
over the last two centuries. Moreover, several fire history reconstruction studies in eastern
Canada have shown that small fires play only a minor role in the total area burned
(Bergeron et al. 2004). We also used the number of years of defoliation caused by 20th-
century SBOs (1938–1985) as a proxy to estimate the influence of the insect. We calculated
this value as the number of years for which >35% of the landscape cover was affected by
moderate or severe defoliation (mean 24.2 yr; range 17–34 yr), using the historical survey
grid (5ʹ latitude by 5ʹ longitude) produced by the Quebec Government (Gray et al. 2000 for
details). Mean elevation a.s.l. (precision: 10 m; range: 100–1000 m; mean: 518 m) of each
cell was calculated using a DEM provided by the MRNQ (2000). Soil drainage data,
indicating the type of drainage percentage of the cell occupied by low, moderate and
excessive drainage founded within a soil profile, was obtained from provincial maps
(1:50 000 scale) derived from aerial photographs and field sampling (MRNQ 2008; see
Appendix S1 and Fig. 1 for a representation of disturbance-related explanatory variables).

Analysis
To allow spatio-temporal comparisons of forest composition and to compensate for
different sampling densities between the 1925 and the 2005 periods, we divided the entire
study area into 165 sampling cells (5 km × 5 km each). After eliminating cells with less
than ten plots from each period (1925 and 2005), we retained 130 cells containing a total of
33 354 plots (n 1925 = 30 033; n 2005 = 3 321). Thereby, statistical relationships between
each explanatory variable and forest composition were made at the cell scale.

The relative basal area (i.e. the percentage of total basal area calculated from all available
plots in each cell) of each taxon was first calculated for 1925 and 2005, and then mapped
using seven classes (Class 1: 0–2.5%, Class 2: 2.5–5%, Class 3: 5–15%, Class 4: 15–25%,
Class 5: 25–35%, Class 6: 35–45%, Class 7: >45%). For each taxon, we subtracted the
relative basal area of 1925 from that of 2005 to map the changes in each cell. A linear
model of the change in relative basal area from 1925 to 2005 was fitted for each taxon to
statistically assess the change between the two periods. Spatial autocorrelation was
accounted for by modeling the residuals with a spatial correlation structure (SAS mixed
procedure, SAS/STAT 12.3; SAS, Cary, NC, US).

We used a K-means cluster analysis to assign community types (groups) based on the
relative basal area of taxa within 25-km2 cells and to measure changes in forest composition
from 1925 to 2005. The K-means analysis, which maximizes extra-group variance and
minimizes inter-group variance, was conducted after subjecting the basal area to a
Hellinger transformation using the vegan v. 2.3-1 package in R (v 0.98.978; R Foundation
for Statistical Computing, Vienna, AT). We clustered data of each of the 130 cells from
both periods in a single run to allow an objective classification. We identified the optimal
number of groups using a simple structure index (SSI) criterion. A community type was
then attributed a posteriori to each cell for each period (1925 and 2005). A cross-tabulation
procedure (transition matrix) using the gmodels package in R was used to document
changes in community types from 1925 to 2005. The kappa statistic (Cohen 1960) was
calculated for each community type to assess the level of change from 1925 to 2005 using
R package irr v 0.84.
We used RDA to identify the most important explanatory variables correlated with the
relative basal area of taxa in 1925 and 2005 for each cell. We also considered the
explanatory variables that were correlated with the observed changes in relative basal area
of taxa from 1925 to 2005. The RDA generates a Y matrix response formed by the relative
basal area of each taxon, constrained by an X matrix (Legendre & Gallagher 2001)
comprised of three broad classes of explanatory variables: logging, natural disturbances and
physiography. Relative basal area was standardized using the Hellinger transformation.
Before conducting the RDA, multicollinearity between the explanatory variables of the X
matrix was verified. We retained the variables with variance inflation factors (VIF) <10
(Borcard et al. 2011). Finally, to quantify the variance of taxa composition explained by
each group of explanatory variables, we partitioned the variation with a series of partial
RDAs between the Y and X matrices (Peres-Neto et al. 2006). This analysis determines the
unique, shared and total explained variation contributed by each set of explanatory
variables. The unique explained variation is the portion of the total variation that is
explained by a given variable, after controlling for all others. All these analyses were
conducted using the package vegan v 2.3-1, following the steps proposed by Borcard et al.
(2011).

Results
In 1925, late successional coniferous taxa (Abies and Picea spp.) composed most (>81.6%)
of the total basal area over the entire study area; the early successional deciduous
taxon Betula spp. occupied 16.6% (Table 1, Fig. 2) and other taxa accounted for <2%. In
2005, the relative basal area of Picea spp. and Abies dropped by 8.8% and 7.4%,
respectively, in favour of early successional deciduous taxa such as Populus spp.
(+6.7%), Betula spp. (+4.7%) and the ‘other species’ class, currently dominated by red
maple (Acer rubrum) (+4.0%; Table 1, Fig. 2).

Table 1. Relative basal area of taxa contributing at least 0.5% of the basal area in the 1925 and
2005 forest surveys over the entire study area

Taxa 1925 (% Basal Area) 2005 (% Basal Area) Difference (2005–1925)

Coniferous Taxa

Abies 43.4 35.9 –7.4

Taxa 1925 (% Basal Area) 2005 (% Basal Area) Difference (2005–1925)

Picea spp. 38.2 29.4 –8.8

Thuja 0.4 1.0 0.5

Pinus spp. 0.4 0.7 0.3

Deciduous Taxa

Betula spp. 16.6 21.3 4.7

Populus spp. 0.8 7.5 6.7

Other spp. 0.2 4.2 4.0

Figure 2

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Spatial distribution pattern of Abies, Picea spp., Betula spp., Populus spp., Thuja, Pinus spp. and
the ‘other spp.’ group in 1925 and 2005. The present-day composition of the ‘other spp.’ group is
dominated by red maple (Acer rubrum).

In 1925, Picea spp. and Abies were the dominant or co-dominant taxa in 87.7% and 80.8%
of all cells; by 2005, these values had dropped by 21.3% and 5.2%, respectively. In
contrast, the largest increase was observed for the early successional deciduous
taxa Betula spp. and Populus spp. (+16.6% and +8.4%, respectively; Table 2). A significant
change between the two periods, as shown by a 95% CI that excludes the value of 1
(Appendix S2), was observed for all taxa, but was especially evident for Abies, Picea spp.
and Betula spp. For the less common taxa, the high frequency of 0 values led to a poor
statistical fit. Nonetheless, the change over time seems obvious for Populus spp. and the
‘other spp.’ group, since the reference line (no change) is almost completely outside the
scatter plot (Appendix S3).

Table 2. Percentage of cells where each taxon was dominant or co-dominant across the study area
in 1925 and 2005. The dominance status was determined by ranking the relative basal area of taxa
within each of the 130 cells, and identifying those that ranked first and second

Taxa 1925 (% of Cells) 2005 (% of Cells)

Dominant Dominant + Co-Dominant Dominant Dominant + Co-Dominant

Coniferous Taxa

Abies 50.0 80.8 49.6 75.6

Picea spp. 34.6 87.7 26.7 66.4

Thuja 0.0 0.8 0.0 0.0

Pinus spp. 0.0 0.0 0.0 0.0

Deciduous Taxa

Betula spp. 15.4 30.0 17.6 46.6

Populus spp. 0.0 0.8 6.1 9.2

Other spp. 0.0 0.0 0.0 2.3

The K-means cluster analysis allowed the identification of five forest communities both in
1925 and 2005, which we named on the basis of the dominant taxa (Fig. 3a). In 1925, the
C5 community, dominated by Abies, was the most abundant type, covering 35.4% of the
cells. Together, the C1 and C2 communities, dominated by Picea spp., covered 43.1% of
the cells. In 2005, the situation reversed: communities dominated by early successional
deciduous taxa (C3 and C4) had become 2.5 times more abundant and covered more than
53.9% of the study area. Overall, the C1 and C2 communities, dominated by Picea spp.,
were 1.8 times less abundant in 2005 (23.8%) than in 1925 (43.1%; Table 3, Fig. 3b). The
transition matrix between time periods shows that 54.7% of the cells dominated by
coniferous taxa (C2 and C5 communities), which were the most abundant in 1925, evolved
toward C3 and C4 communities now dominated by early successional deciduous taxa.
Cohens’ kappa coefficient was 0.18, indicating a very dynamic landscape.

Figure 3

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Forest communities identified using K-means clustering (a), and community transitions (1925–
2005) (b). The communities were named according to the dominant taxon.

Table 3. Transition matrix calculated from the observed changes in forest community types (C1 to
C5)a for each cell from 1925 to 2005. The composition of each community type is detailed in
Fig. 3a
 1925 2005

C1 C2 C3 C4 C5 Frequency Landscape (%)

C1 0.37 0.11 0.04 0.19 0.30 27 20.8

C2 0.10 0.28 0.28 0.31 0.03 29 22.3

C3 – 0.50 0.50 – – 2 01.5

C4 – 0.04 0.50 0.31 0.15 26 20.0

C5 0.09 0.04 0.17 0.35 0.35 46 35.4

Frequency 17 14 32 38 29 130

Landscape 13.0 10.8 24.6 29.3 22.3 100

(%)

 a C1: Picea spp.–Abies; C2: Picea spp.-dominated; C3: deciduous–conifers; C4: Betula spp.–Abies;
C5: Abies-dominated.
 “No change” transition types are in bold.

For the 1925 period, the adjusted R2 coefficients indicate that the two selected explanatory
variables (fires prior to 1925 [FIRE < 1925] and moderate drainage [MOD_DR];
Appendix S4) explain 23.0% of the variation of taxa distribution in the RDA model
(Fig. 4a). We interpret the first axis (21.2% of the variation) as the influence of fire on
vegetation. The second axis, which only explains 3.0% of the variation, is likely associated
with a drainage gradient. No groups or outliers seem to drive the model, since the circles
are quite evenly distributed across the multidimensional space of the RDA. The distribution
of taxa and community types in the multidimensional space agrees with the life history of
the various taxa: the presence of the late successional coniferous taxa Abies is negatively
correlated with fire; the relative abundance of early successional deciduous taxa
(Populus spp. and Betula spp.) follows the opposite trend and is positively correlated with
fire (Fig. 4a).

Figure 4

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RDA ordination triplot illustrating the correlation of natural disturbances, logging history and
physiographical variables on taxa relative abundance in 1925 (a) and 2005 (b) and the observed
difference (2005–1925) (c). The colour of each circle corresponds to forest community type,
following the legend of Fig. 3. The distributions of taxa are depicted with the solid red lines,
whereas the explanatory variables are depicted with the arrows. The length of each vector
indicates its importance to the ordination. The angles between vectors reflect their linear
correlation. Direction of each vector indicates its correlation with each of the axes. See
Appendix S4 for acronyms.

For the 2005 period, the four explanatory variables selected for the RDA (Appendix S4)
explain 44.0% (with the adjusted R2) of the variation of the distribution of the taxa
(Fig. 4b). The relative basal area of Abies and Picea spp. are negatively correlated with old
(<1925) and recent fires (>1925) and to some extent with poor drainage
conditions. Populus spp. and Pinus spp. are strongly correlated with fire,
whereas Betula spp., Thuja and the ‘other spp.’ group are correlated with SBOs and
negatively correlated with poor drainage. Early and modern logging variables were not
selected in the model (Fig. 4b).

The explanatory variables (n = 4; Appendix S4) explain a relatively small fraction (13.5%)
of the variation related to compositional changes of taxa from 1925 to 2005 (Fig. 4c). The
triplot shows that the relative basal area increase of Populus spp. is strongly correlated with
fires prior to 1925, and even more with more recent fires. Modern logging also shows a
weak positive correlation with Populus spp. In contrast, the decrease of late successional
coniferous taxa (Abies and Picea spp.) is positively correlated with modern logging and
recent fires. The increase of Betula spp. is positively correlated with SBOs. The increase of
the ‘other spp.’ group is positively correlated with SBOs, fires >1925 and modern logging.
The explanatory variables associated with natural disturbances explain 96.0% of the total
unique variation, while modern logging explains only 4.0% (Appendix S5).

Discussion
The combination of historical and contemporary data is an effective way to unravel long-
term effects of multiple disturbances on forest landscapes (Thompson et al. 2013). The
present-day forest composition cannot be fully understood without considering processes
acting over previous decades, centuries or more (Foster et al. 2003). By linking disturbance
history maps with historical surveys, our study isolates the relative impacts of logging and
natural disturbances on forest composition over the last century.

The preindustrial forest landscape composition changed substantially over the 20th century
in response to disturbances. In 1925, the preindustrial forest matrix was strongly dominated
by late successional coniferous taxa (Abies and Picea spp.). Early successional deciduous
taxa, associated with burned areas, were a minor component of the landscape. From 1925 to
2005, the relative abundance of early successional deciduous taxa (Populus spp.,
Betula spp. and ‘other spp’. [now dominated by red maple]) increased sharply at the
expense of late successional coniferous taxa. These observations concur with several
studies conducted in the eastern Canadian boreal forest (Jackson et al. 2000; Pinto
et al. 2008; Boucher et al. 2009; Laquerre et al. 2009; Dupuis et al. 2011; Danneyrolles
et al. 2016) and in similar ecosystems in the USA Great Lakes region (Michigan,
Wisconsin, Minnesota; Friedman & Reich 2005; Schulte et al. 2007; Rhemtulla
et al. 2009).

However, in contrast to many studies (see Jackson et al. 2000; Pinto et al. 2008; Laquerre
et al. 2009) that assumed that logging was the main or only cause of these compositional
changes, our comparative analysis of several disturbance agents strongly suggests that
natural disturbances were more important drivers of these changes than logging. Prior to
human influence, fires and SBOs shaped the composition of the preindustrial forest
landscape. Throughout the Holocene, fire return interval fluctuations controlled the
proportion of early successional deciduous species at the landscape scale (Bergeron 2000;
Cyr et al. 2009), while cyclical (30–40 yr), large-scale SBOs of varying severity
constrained the dominance of Abies and Picea spp. (Blais 1983). In our study
area, Betula spp. (B. papyrifera) and Populus spp. are both fire-adapted taxa. They produce
large numbers of wind-dispersed seed and either sprout (Betula spp.) or sucker
(Populus spp.) vigorously after fire (Bergeron 2000). In addition to fire, successive SBOs
during the 20th century (in the 1920s, 1950s and 1980s; Blais 1965, 1983) have affected the
study area and caused various levels of mortality in coniferous stands, especially those
dominated by Abies, the most vulnerable species to the insect (Blais 1983). The various-
sized openings created by the death of coniferous trees then allowed recruitment of B.
papyrifera, a species well adapted to fill gaps (Bergeron 2000; Bouchard et al. 2007). The
‘other spp.’ group, now dominated in the modern inventory by the ‘super-generalist’ red
maple (Abrams 1998), also spread throughout the study area, profiting from multiple
disturbances and a more favourable climate in the second half of the 20th century (Zhang
et al. 2000). A sharp increase in red maple at its northern range limit has been observed
elsewhere in eastern Canada (Duchesne & Ouimet 2008; Dupuis et al. 2011) and across the
eastern USA (Schulte et al. 2007; Nowacki & Abrams 2015).

While logging history has modified the age structure of the studied landscape (Boucher &
Grondin 2012; Boucher et al. 2014) and of most managed boreal forest regions (Cyr
et al. 2009; Hellberg et al. 2009), it has had a relatively minor impact on landscape
composition in the study region. During the early logging period (<1965), non-mechanized
clear-cutting was generally conducted during winter in pure conifer-dominated stands. This
harvesting technique protected soils and allowed the retention of pre-established coniferous
seedlings, and therefore, allowed the rapid re-occupation of shade-tolerant conifers
(mostly Abies) in post-harvest stands (Hatcher 1960).
In contrast, sites that were logged during the modern period (>1965) show a slight positive
relationship with Populus spp. basal area. This situation may be explained by several
factors associated with the harvesting technique used during the modern period. Clear-
cutting was practiced year-round, and the heavy machinery often (1) exposed mineral soils
that constitute good seed beds for deciduous species, (2) fostered Populus spp. suckering,
and (3) destroyed a fraction of conifer seedlings (Harvey & Bergeron 1989). Another
decisive factor is that a large fraction of stands harvested after 1965 were of post-fire origin
(fires 1820–1925; see Appendix S1) and contained a proportion of Populus spp. that could
invade logged sites by suckering. Finally, 20th century clear-cut logging mainly targeted
coniferous trees. This practice has reduced the relative dominance of late successional
coniferous taxa to the profit of early successional deciduous taxa.

Modern logging history was a less important driver of the expansion of early successional
deciduous species than fire. Fires, alone or followed by logging, are more likely to have
increased the abundance of early successional deciduous taxa at the regional scale. A
similar outcome can be observed in the USA Great Lakes region, where Populus spp.
and Betula spp. became dominant after slash fires during the post-settlement era, and have
since been maintained by 20th century clear-cutting practices (Graham et al. 1963; Schulte
et al. 2007). Despite the minor role of logging on compositional changes, the influence of
human activities on forest composition remains determinant. Indeed, regional fire history
reconstructions show that many burned areas result from escaped settlement fires (Boucher
& Grondin 2012; Boucher et al. 2014). This greater influence of fire, associated with land-
clearing activities over the 19th and early 20th centuries, has been observed throughout
eastern North America (Whitney 1994; Dupuis et al. 2011) and explains the present-day
forest composition in these regions.

Yet, future disturbances regimes will most likely differ from the past, and predicting the
trajectory of forest composition remains difficult. Considering that fire activity is expected
to remain low relative to the settlement period, logging could progressively become a more
important driver of tree species composition during the 21st century. We also expect that
red maple will continue its increase in dominance and northward expansion (Duchesne &
Ouimet 2008; Nowacki & Abrams 2015) in a context of rising temperatures and higher
disturbance rates than in the 20th century (Wang et al. 2015).
Conclusion
Combining historical and contemporary data is an effective way to unravel the long-term
effects of multiple disturbances on forest landscapes (Thompson et al. 2013). Present-day
forest composition cannot be fully understood without considering processes that acted
over previous decades, centuries or more (Foster et al. 2003). By linking disturbance
history maps with historical surveys, our study isolates the relative impacts of logging and
natural disturbances on forest composition over the last century. Using detailed forest
inventories made prior and after 20th century logging, we were able to assess that fires and
insect outbreaks have been the most influential drivers of forest composition compared to
logging, which had a minor effect. In the current context of ecosystem-based forest
management (Lindenmayer et al. 2006), land managers should continue to elaborate
management targets and consider mitigation measures in order to keep forest landscape
composition within its natural range of variability, in order to preserve biodiversity and
ecosystem resilience (Landres et al. 1999). However, global change may move the entire
ecosystem outside its range of variability if environmental changes are too important and
mitigation measures prove inefficient (Gauthier et al. 2015). In this context, in order to
properly manage forest landscapes, more prospective research is required to model
interactions between climate, natural disturbances and forest management on tree species
composition.

Acknowledgements
This study was financed by Quebec's Ministère des Forêts, de la Faune et des Parcs (project
no. 142332085). We would like to thank V. Poirier for geomatics and M.-H. Brière for
integrating manuscript archives into a GIS. We also thank P. Jasinski and D. Tousignant for
English revision. The staff of Archives nationales du Québec and Resolute Forest Products
Inc. also provided valuable help with the archives.