chapter 4 et al 2011.pdf · in: woodlands: ecology, management and conservation isbn...

In: Woodlands: Ecology, Management and Conservation ISBN 978-1-61122-542-6

Editor: Erwin B. Wallace ©2011 Nova Science Publishers, Inc.

Chapter 4

PARTIAL HARVESTING IN OLD-GROWTH

BOREAL FORESTS AND THE PRESERVATION

OF ANIMAL DIVERSITY FROM ANTS

TO WOODLAND CARIBOU

Daniel Fortin 1,a, Christian Hébert

2,b, Jean-Philippe Légaré 3,c,

Nicolas Courbin 1, Kyle Swiston

1, James Hodson

1,

Mélanie-Louise LeBlanc 1, Christian Dussault

4,d,

David Pothier 3, Jean-Claude Ruel

3,

and Serge Couturier 4

1. Département de biologie, NSERC–Université Laval industrial research chair in

silviculture and wildlife, Laval University, Québec, Canada 2. Natural Resources Canada, Canadian Forest Service, Laurentian Forestry Centre,

Québec, Canada 3. Département des sciences du bois et de la forêt, NSERC–Université Laval industrial

research chair in silviculture and wildlife, Université Laval, Québec, Canada 4. Direction de l'expertise sur la faune et ses habitats, Ministère des Ressources naturelles

et de la Faune, Québec, Canada

ABSTRACT

Current forest management must maintain biodiversity, an objective that has led to

the rapid development of new forestry practices in recent years. However, empirical

evaluation of the impact that these practices have on biodiversity has not kept the same

pace. For example, small merchantable stems are now frequently protected during the

harvest of old-growth boreal forests in eastern Canada. This silvicultural practice

a 1045 Av. de la Médecine, pavillon Alexandre-Vachon, Québec, QC G1V 0A6, Canada.

b 1055 du P.E.P.S., P. O. Box 10380, Stn. Sainte-Foy, Québec, G1V 4C7, Canada.

c Sainte-Foy, Québec G1V 0A6.

d 880 chemin Sainte-Foy, Québec, QC G1S 4X4, Canada.

The exclusive license for this PDF is limited to personal website use only. No part of this digital document may be reproduced, stored in a retrieval system or transmitted commercially in any form or by any means. The publisher has taken reasonable care in the preparation of this digital document, but makes no expressed or implied warranty of any kind and assumes no responsibility for any errors or omissions. No liability is assumed for incidental or consequential damages in connection with or arising out of information contained herein. This digital document is sold with the clear understanding that the publisher is not engaged in rendering legal, medical or any other professional services.

Daniel Fortin, Christian Hébert, Jean-Philippe Légaré, Nicolas Courbin et al. 116

(referred to as CPPTM in Québec) ends up protecting all stems with a diameter at breast

height of 9-15 cm, and is therefore expected to maintain some of the irregular attributes

of old-growth forest structure. Whether or not this approach is sufficient to maintain local

biodiversity remains unclear. We evaluated the short-term impact of CPPTM harvesting,

mostly 2-3 years after logging, on a broad range of animal species differing largely in

size and resource requirements. More specifically, we estimated the abundance,

occurrence or local intensity of habitat use by ants, beetles, forest birds, snowshoe hare

(Lepus americanus), moose (Alces alces) and woodland caribou (Rangifer tarandus

caribou) in a boreal ecosystem. The 19,000 km2 study region is dominated by >270 year

old stands with irregular structure mostly comprised of black spruce (Picea mariana) and

balsam fir (Abies balsamea). Harvesting by CPPTM caused a 75-85% reduction in tree

basal area. This decrease was sufficient to alter animal assemblages of all taxonomic

groups. Ants and beetles associated with open areas were more abundant in CPPTM sites

than in uncut stands. Conversely, species associated with mature forests were much lower

in CPPTM than in uncut stands. Most birds associated with late-successional habitats

were also less likely to be observed in CPPTM sites than in uncut stands. Snowshoe hares

significantly decreased their use of harvested stands following CPPTM. Woodland

caribou displayed a strong avoidance for CPPTM sites, while moose did not display any

such aversion. CPPTM thus alters species assemblages, potentially reshaping trophic

interactions with consequences for wildlife conservation. For example, moose did not

avoid CPPTM cuts as much as they avoided stands that were harvested more intensively.

Wolves that generally focus their hunting on moose might become attracted to CPPTM

cuts, which could result in a higher concentration or even in a numerical response of

wolves. At a regional scale, the outcome might lead to an increase in local predation risk

for woodland caribou, a threatened species. While CPPTM could still be useful in an

ecosystem-based management context, our study shows that the protection of small

merchantable stems is often insufficient for the short-term maintenance of species

composition which characterizes old-growth boreal forests.

INTRODUCTION

Current forest harvesting practices have to sustain our long-term wood supply needs,

while also maintaining primary ecosystem attributes, including regional biodiversity (Côté

and Bouthillier 2002). Forest management strategies are increasingly developed following the

principles of ecosystem-based management by emulation of natural disturbances (Bergeron et

al. 2007, Rosenvald and Lõhmus 2007). The premise is that biodiversity and essential

ecological functions are most likely to be maintained by emulating the spatio-temporal

patterns of natural disturbances (Hunter 1999, Fenton et al. 2009) because local populations

have been shaped by the processes leading to these patterns.

The natural disturbance regime responsible for the dynamics of local ecosystems varies

widely across the boreal forest. For example, the average time between wildfire recurrences

can be less than a century in some areas, while in others, it can exceed 500 years and even

thousands of years (Bergeron et al. 2002, Bergeron et al. 2007, Bouchard et al. 2008, Fenton

et al. 2009, Johnstone et al. 2009). Forest composition and structure strongly depend on the

length of the regional fire cycle. Balsam fir (Abies balsamea) generally increases in

abundance as black spruce (Picea mariana) dominated forests get older (De Grandpré et al.

2000). Approximately 80-90 years after the last fire, the local basal area and density of trees

(>9 cm in diameter at breast height, dbh) cease to increase, and even start declining

Partial Harvesting in Old-Growth Boreal Forests and the Preservation… 117

(Bouchard et al. 2008). Under a long fire cycle regime, tree mortality due to windthrow,

insects, pathogens, and senescence lead to fine scale canopy gap dynamics creating

landscapes dominated by irregularly structured old-growth stands (Boucher et al. 2003, Pham

et al. 2004).

In response to strong regional variations in natural disturbance regimes, an ecosystem-

based approach to forest management should prescribe different silvicultural practices across

the boreal forest (Rosenvald and Lõhmus 2007). Currently, clearcutting (or closely related

harvest techniques, such as cutting with protection of advanced regeneration and soils

[CPRS], Groot et al. 2005, Courtois et al. 2008) is the most common logging practice in the

boreal forest (Bergeron et al. 2007, Rosenvald and Lõhmus 2008, Ruel et al. 2007).

Clearcutting brings stands back to an early seral-stage from which they develop with a regular

structure (Groot et al. 2005). This practice thus appears particularly relevant in regions with

relatively short fire cycles (Bergeron et al. 2007). In contrast, the emulation of natural

disturbances in regions with long fire cycles, dominated by old-growth forests, should require

the use of partial harvesting to maintain the irregular structure of forest stands (Bergeron et al.

2007, Ruel et al. 2007, Rosenvald and Lõhmus 2007, Drapeau et al. 2009).

A large number of partial harvesting techniques have emerged in recent years (Groot et

al. 2005, Ruel et al. 2007, Courtois et al. 2008, Rosenvald and Lõhmus 2008, Cimon-Morin et

al. 2010, Poulin et al. 2010). Since the 1990s, the protection of small merchantable stems has

become increasingly frequent when harvesting old-growth boreal forests in eastern Canada.

This silvicultural technique (taking the acronym ―CPPTM‖ in Québec and HARP in Ontario,

Groot et al. 2005) protects all stems with a dbh of 9-15 cm, thereby maintaining some of the

irregular structure of old-growth forests (Ruel et al. 2007). Not spending time and money

harvesting these relatively small stems can lead to financial gains because small merchantable

stems are worth less to the forest industry than larger stems (Liu et al. 2007, Ruel et al. 2007).

However, the impact of CPPTM on biodiversity has remained largely unexplored, and the

industry still does not know what role CPPTM serves in maintaining local biodiversity and

ecosystem functioning of old-growth boreal forests.

Clearcutting of mature forests generally alters local animal communities by favouring

species associated with early-successional habitats at the expense of late-successional species

(Annand and Thompson 1997, Simon et al. 2000, Vanderwel et al. 2007). In contrast, partial

harvesting can maintain species assemblages, depending on logging intensity (Vanderwel et

al. 2009). Whether the protection of small merchantable stems during CPPTM suffices to

maintain local biodiversity remains an open question.

Because it is impractical to evaluate the impacts of forest harvesting on all species,

investigators often rely on indicator species (Potvin et al. 1999, Thompson et al. 2008), which

are generally selected from a single taxonomic group. This approach may not fully inform us

of the potential contribution of a given silvicultural practice to sustainable forest

management, because the impacts of logging vary among animal groups (Venier and Pierce

2004). To acquire a comprehensive understanding of the impact of forest harvesting, it is

therefore critical to consider a diversity of taxa for adequate wildlife conservation and

management.

We investigated the short-term impact of CPPTM (mostly 2-3 years after harvesting) on

target species chosen from multiple taxa. These species were selected because they varied

broadly in movement capacity, home-range size, and habitat requirements, as well as in their

associations with different stages of forest succession. First, we considered two saproxylic ant


species: Camponotus herculeanus, which is generally more abundant in closed forests, and

Formica neorufibarbis, which is typical of open areas (Jennings et al. 1986, Francoeur 2001,

Higgins and Lindgren 2006). We also considered 10 beetle species, five of which have been

found to be associated with closed coniferous forest stands (Silvanus bidentatus (Majka

2008), Enicmus tenuicornis (Jacobs et al. 2007b, Majka et al. 2009), Oxypoda grandipennis

(Csy.) (Klimaszewski et al. 2006), Epuraea planulata (Saint-Germain et al. 2004, Janssen et

al. 2009), Rhizophagus dimidiatus (Saint-Germain et al. 2004, Jacobs et al 2007a, Janssen et

al. 2009), whereas the other five are typical of open areas (Leiodes spp. from the assimilis-

group (Baranowski 1993, Unpublished data, C. Hébert), Clypastraea fusca (Boulanger and

Sirois 2007), Xylita laevigata (Boulanger and Sirois 2007), Hylobius congener (Welty and

Houseweart 1985, Boulanger and Sirois 2007), Pseudanostirus triundulatus (Randall) (Saint-

Germain et al. 2004). For instance, the last four species are particularly abundant shortly after

fire in the boreal forest (Saint-Germain et al. 2004, Boulanger and Sirois 2007). Boreal

Chickadee (Poecile hudsonsicus), Brown Creeper (Certhia familiaris), and Ruby-crowned

Kinglet (Regulus calendula) were selected to represent closed-canopy bird species because

they are abundant on our study area (Lemaître 2009), and they should respond negatively to

low tree retention levels (Vanderwel et al. 2007, 2009, St-Laurent et al. 2008). White-throated

Sparrow (Zonotrichia albicollis) and Dark-eyed Junco (Junco hyemalis) are also locally

abundant species (Lemaître 2009), and they are expected to respond positively to the

openings of the tree canopy resulting from logging (Vanderwel et al. 2007, 2009). Finally, we

evaluated the response of snowshoe hare (Lepus americanus) and moose (Alces alces), two

species favouring rather early (10-30 years) successional forests (Pastor et al. 1999, Dussault

et al. 2005, Fisher and Wilkinson 2005, Newbury and Simon 2005), and forest-dwelling

woodland caribou (Rangifer tarandus caribou), a late-successional species (Fortin et al. 2008,

Courbin et al. 2009). The field methods used varied among taxa, from an evaluation of

changes in abundance (ant and beetle species) or probability of occurrence (bird species) of

individual species, to an assessment of habitat use and selection by mammalian herbivores

(snowshoe hare, moose and caribou).

STUDY AREA

The study took place in the eastern spruce-moss subdomain of the boreal forest, in the

Côte-Nord region of Québec, Canada (Figure 1). The 19,000 km2 study area is typical of the

Canadian Precambrian Shield, with a rolling and hilly landscape, and an altitude varying

between 300 and 1,000 m. Mean annual temperature ranges from -2.5 to 0.0 °C, and annual

precipitation ranges from 1,000 to 1,400 mm (Boucher et al. 2003). The abundant

precipitation results in a long fire cycle (average stand age: >270 years old, Bouchard et al.

2008). The study area is therefore characterized by old-growth forest stands that are irregular

in structure and composition (Boucher et al. 2003). The dominant tree species are black

spruce, balsam fir, white birch (Betula papyrifera Marsh.) and trembling aspen (Populus

tremuloides Michx).


Figure 1. Study area located in the Côte-Nord region of Québec, Canada. Harvested areas are displayed

in white.

METHODS

Estimation of Ant Abundance

In 2004 or 2005, ants were sampled in 14 stands (>4 ha) that had been harvested by

CPPTM 1 to 8 years earlier (average: 4.2 ± 2.1 years). Ants were captured concurrently in 70

uncut conifer stands (median age: 120 years; range: 70 to >120 years). All sites were >2 km


from one another. At each site, ants were captured in four pitfall traps buried at ground level,

10 m apart (following Janssen et al. 2009). The traps had a diameter of 10 cm, and were

screened with a wire mesh (1 × 1 cm) to avoid capture of vertebrates. Trapped insects were

collected every two weeks, from 2 June until 17 August, and preserved in a 40% ethanol

solution with a trace of acetic acid (5%), before being identified in the laboratory.

The abundance of each of the two focal ant species was contrasted between CPPTM and

uncut stands with one-sided, nonparametric Wilcoxon Two-Sample Tests, evaluating the

specific prediction that CPPTM opens the canopy sufficiently so that the closed-habitat

species, C. herculeanus, would be less abundant in CPPTM sites than in uncut conifer stands,

whereas the open-habitat species, F. neorufibarbis, would be more abundant in CPPTM sites

than in uncut stands. Because the time between CPPTM harvesting and ant sampling varied

by as much as 8 years, we used a Spearman rank correlation to test whether the abundance of

each of these ant species varied with the time since partial harvesting.

Estimation of Beetle Abundance

Beetles were sampled from 5 June to 22 August 2007 in 16 old-growth forest stands

(>120 years), and in 16 CPPTM conifer stands (average size: 19 ha; range: 12-24 ha). Eight

of these sites had been harvested in 2004 and eight in 2005. All sites were >100 m apart and

>100 m from the stand edge. For logistical reasons, these sites were grouped in four sectors

(e.g., each near a forestry camp or associated with different forestry companies). The

potential non-independence of data within a sector was considered through the use of mixed-

effects modelling (see below).

Four pitfall traps were located at each site to capture ground-dwelling beetles, together

with one flight-interception trap to capture flying beetles. The multi-directional flight-

interception trap was located at the centre of each sampling site, 0.5-1 m above ground. This

trap was built using four 15 × 40 cm panels (two made of Plexiglas and two of mosquito net)

mounted into a cross pattern, along a 10-cm diameter black ABS cylinder, with two funnels

located above and below the cylinder, and leading to collecting vials (Saint-Germain et al.

2004). As with the ants, beetles were collected every two weeks, and preserved in a solution

of ethanol, acetic acid and water, before being identified in the laboratory at the species level.

The log-transformed abundance (log [x+0.1]) of each of the 10 focal beetle species was

compared between cut and uncut stands using mixed-effects models with a Gaussian

distribution. The four sectors were considered as a random-effect term.

Estimation of the Occurrence Probability and Abundance of Bird Species

Birds were surveyed in 2006 and 2007 between 3 June and 30 June at 12 point count

stations (Verner, 1985) located in old-growth forests, and at 12 stations located in conifer

stands harvested by CPPTM (average size: 19 ha; range: 12-24 ha) in 2004 or in 2005.

Stations were aggregated into three sectors, a design controlled for by the use of mixed-

effects models. Point count stations were >150 m from one another and >100 m from the

stand edges.


Point count stations were visited by four observers three times each year, between 05:00

and 10:00, in the absence of wind or heavy rain. To reduce the potential risk of observer bias,

the four observers surveyed different sites during the three yearly visits. All birds seen or

heard within a 50-m radius of the point count stations were recorded during 10 minutes.

Observers trained together to become consistent in their evaluation of bird distances.

A species was coded as 1 for a given point count station if it was recorded during any of

the visits. Otherwise, the species was coded as 0 for that station. We used this approach

because of low bird count at individual stations: only 0-2 individuals were observed per

station, except for White-throated Sparrow which was represented by up to 4 individuals per

station. We used mixed-effects logistic regression for each of the five focal bird species to

evaluate whether it was more likely to be observed at CPPTM sites or in the uncut stands. The

three sectors were considered as a random-effect term to allow for potential inter-sector

variations in the probability of occurrence of the different species. We also carried out mixed-

effects Poisson regression for the White-throated Sparrow, the most abundant bird species, to

test whether its abundance was higher in CPPTM sites than in uncut stands.

Evaluation of Habitat Use by Snowshoe Hare

Habitat use by snowshoe hare was assessed in four uncut conifer stands paired with four

CPPTM sites harvested in 2004 or 2005. In 2007, we evaluated browse history at all sites by

identifying years in which white birch stems (an abundant and preferred browse species,

Newbury and Simon 2005) were browsed by snowshoe hare. This approach allowed us to

quantify the intensity of stand use by snowshoe hare prior to logging, even though field

sampling was conducted only 2 to 3 years after harvesting. During winter, this herbivore

generally clips the terminal leaders of the previous summer‘s growth, which kills the terminal

bud (Pease et al. 1979). The following spring, vertical growth will resume from a dormant

lateral bud further down the stem (Keigley and Frisina 1998). In contrast, vertical growth of

non-browsed stems resumes from the terminal bud, leaving a bud scar for each year of

uninterrupted growth. The number of years of growth following the mortality of a terminal

leader due to browsing can then be determined by counting the number of terminal bud scars

along the stem originating from a lateral bud which resumed vertical growth (Keigley and

Frisina 1998). We can thus determine the year in which each twig was clipped by counting

the bud scars backwards from the current year‘s growth to the browsed segment. Bud scars

are generally discernable for up to five years of previous growth. We were interested in the

previous 4 years of growth, which included the winter preceding the earliest cuts (2004) in the

experimental blocks.

We evaluated the browse history of 18 white birch stems distributed across six quadrats

(6 × 75 m) located in each CPPTM and each uncut site (i.e., 36 stems per pair of cut-uncut

sites). In each quadrat, we sampled the first three white birch stems that met the following

criteria: 1) stems had to be at least 5 years old (determined by counting terminal bud scars) so

that they could provide information on pre-treatment use by hare, 2) stems also had to be at

least 50 cm in height 5 years prior to the survey so that they would be above the snow during

a large part of each winter, and 3) stems had to have at least one browse mark so that they

could provide a historical record of use. For each stem, we identified the year in which twigs

were clipped by snowshoe hare starting from the winter before the harvest treatment took


place (i.e. the winter of 2003/2004 or 2004/2005) until the winter of 2006/2007 (2-3 years

after harvesting took place). We then used the presence or absence of scars in each year to

classify each stem as used or unused for each of the previous 4 years.

To assess whether snowshoe hare used sites with similar intensity before harvesting took

place, we first tested a model predicting the probability of white birch browse use by

snowshoe hare in the winter before harvesting occurred. We used mixed-effects logistic

regression with the dependent variable being whether a stem was browsed (coded as 1) or not

(coded as 0) in a given year. To account for the hierarchical structure of our sampling design

and the non-independence among the 18 stems selected within each site, we included random

effects which consider that stems were nested within habitats, and that habitats (CPPTM

versus uncut stands) were paired. To model temporal changes in the probability of birch stem

browsing by snowshoe hare, we used a second mixed-effects logistic regression for repeated

measures. We considered browsing from the winter before harvest treatments (year = 0) took

place until 2 to 3 years after logging. Habitat (CPPTM = 1, uncut = 0), year and ―habitat ×

year‖ interaction were included as fixed-effects terms. Birch stems were considered the

experimental units upon which repeated measures were taken (i.e. each stem was classified as

used or unused during each year) and we used an autoregressive (order 1) correlation

structure to account for the fact that the probability of stem use was more likely to be similar

in successive years than 2 or 3 years apart in time. We used the same random effects structure

as specified for the pre-harvest model.

Evaluation of Habitat Selection by Moose and Caribou

From mid-March 2005 to mid-March 2007, we followed the movement of 18 adult

female caribou and 10 adult female moose equipped with global positioning system collars

(Lotek Engineering Inc., Newmarket, Ontario or Telonics Inc., Mesa, Arizona) taking

locations at 4-h intervals for caribou and every hour for moose. Due to deaths or battery

exhaustion, each caribou was followed during an average of 13 months (range: <1–25

months), whereas each moose was followed for a mean of 11 months (range: <1–19 months).

We considered only 2D locations with a horizontal dilution of precision (HDOP) < 10 and 3D

locations with a HDOP < 15, resulting in a precision of approximately 25 m (Dussault et al.

2001).

The landscape was characterized from Landsat Thematic Mapper images taken in 2000

and having a 25-m resolution. Satellite images were classified using the land cover types:

mature conifer forest, open area, lake, conifer forest with lichen and lichen-heath community,

and mixed/deciduous forest. In addition, we georeferenced recent clearcuts (<5 years),

regenerating clearcuts (5–10 years), CPPTM, and roads based on information provided by the

local forestry companies. We further delineated stands that had been harvested while

retaining tall, but not yet merchantable stems (10 cm in dbh). This harvest practice takes the

acronym ―CPHRS‖ in Québec (Groot et al. 2005, Courtois et al. 2008).

We evaluated habitat selection using resource selection functions (RSF, Manly et al.

2002). One RSF was developed for each ungulate species based on a comparison between

landscape characteristics at the observed locations and at an equal number of random

locations drawn within the 100% minimum convex polygon (MCP) of each individual‘s

annual home range. We estimated RSF parameters using mixed-effects logistic regression


with ‗individual‘ being used as a random factor (i.e., random intercept) to account for the

non-independence of the multiple observations per individual. We also added random slopes

for CPPTM and for CPHRS to allow for the response to these silvicultural practices to vary

among individuals. Multicollinearity was absent from both caribou and moose RSFs, with

condition indices being <7.5 in both cases (Belsley et al. 1980).

RESULTS

Ants

Camponotus herculeanus, a closed-habitat ant species, was 11 times more abundant in

uncut conifer forests than in CPPTM sites (One-sided Wilcoxon Two-Sample test: Statistic:

479.5, Z = -1.6, P = 0.05, Figure 2). In contrast, F. neorufibarbis, a species typical of open

habitat, was most abundant in CPPTM, with 29 times more individuals being captured in

CPPTM sites than in uncut stands (Statistic: 744.0, Z = 3.2; P < 0.001) (Figure 2).

Figure 2. Mean abundance (+ SE) of two ant species in uncut stands and in stands harvested while

protecting small merchantable stems (CPPTM), according to their preference for closed- or open-

canopy habitats.

The time elapsed since logging (range: 1-8 years) did not influence the abundance of either

species (rs < 0.39 and P > 0.16 for any species).


Figure 3. Mean abundance (+ SE, on a logarithmic scale) of 10 beetles in uncut stands and in stands

harvested while protecting small merchantable stems (CPPTM), according to their preference for

closed- or open-canopy habitats.

Figure 4. Predicted probability (± 95% CI) of paper birch stem use by snowshoe hare in forest stands

harvested while protecting small merchantable stems (CPPTM) paired with uncut stands, from the

winter before the harvest took place (Year = 0) up until three years following logging. Square symbols

represent uncut forest stands, circles represent harvested stands, grey arrows indicate the summer when

harvesting took place in treated stands.


Beetles

All five beetle species typical of closed habitats were more abundant in uncut conifer

forests than in CPPTM (F1, 27 > 5.70 and P < 0.02 for any species). R. dimidiatus and E.

planulata were the most abundant of the 10 focal species (Figure 3). Although the abundance

of these two closed-canopy species was >3.5 higher in uncut forests than in CPPTM sites,

they remained among the most abundant beetles in the CPPTM sites. In fact, among the open-

canopy species, P. triundulatus was the only species more abundant in CPPTM than the two

closed-canopy species. All five species typical of open habitats were more abundant in

CPPTM than in uncut stands (F1, 27 > 12.50 and P < 0.002 for each species). For example,

Leiodes sp. was only found in CPPTM, whereas X. laevigata was 34 times more abundant in

CPPTM than in uncut stands (Figure 3).

Forest Birds

The Boreal Chickadee and the Ruby-crowned Kinglet, two closed-canopy species, were

more likely to be observed in uncut forests than in CPPTM sites (Table 1). However, we did

not detect a difference in the probability of occurrence of the Brown Creeper, also a closed-

canopy species, between uncut and harvested stands. The White-throated Sparrow and Dark-

eyed Junco had a higher probability of occurrence in CPPTM than in uncut forests (Table 1).

For example, we only observed 2 White-throated Sparrows at two point count stations in

uncut conifer stands (i.e., 2/12 stations in uncut stands), while we recorded the species at all

the CPPTM stations (i.e., 12/12 stations), totalling 26 individuals observed in the harvested

stands. In fact, White-throated Sparrows were significantly more abundant in CPPTM sites

than in uncut stands (Mixed-effects Poisson regression: F1,20 = 485.7, P < 0.001).

Table 1. Coefficients () from a mixed-effects logistic regression testing for a difference

between the probability of occurrence of closed- and open-habitat bird species

in CPPTM sites versus in uncutconifer stands in the Côte-Nord region of Québec,

Canada.Negative s imply a lower probability of occurrence in CPPTM cuts than

in uncut conifer stands

Species ± SE F1,18 P

Closed-canopy species

Boreal Chickadee -2.40 ± 1.01 5.66 0.03

Brown Creeper -1.30 ± 1.09 1.42 0.25

Ruby-crowned Kinglet -3.09 ± 1.08 8.18 0.01

Open-canopy species

White-throated Sparrow 21.23 ± 0.76 785.97 <0.0001

Dark-eyed Junco 2.73 ± 1.05 6.75 0.02

Snowshoe Hare

The winter before stands were harvested, birch stems in stands that were eventually

harvested had a similar probability of being browsed by snowshoe hare as those in stands that


remained uncut (Habitat effect: F1, 136 = 0.92, P = 0.34, Figure4). Following logging,

temporal patterns of browse use varied between CPPTM and uncut stands (habitat × time: F1,

494 = 21.36, P < 0.0001). Snowshoe hares did not change their use of birch stems over time in

uncut stands. In contrast, the probability of stem use was more than three times lower 3 years

after harvesting than before logging (Figure 4).

Moose and Woodland Caribou

Radio-collared moose and woodland caribou had access to 627 CPPTM patches, which

covered a total of 0.33% of the 19,000 km2 area (100% MCP) occupied by the 28 individuals.

We collected 37,447 locations from the 18 radio-collared caribou (range: 17 to 2,128 per

caribou per year) and 71,039 locations among the 10 radio-collared moose (range: 115 to

8,473 per moose per year). These locations were not distributed randomly with respect to

habitat features (Table 2). We observed some similarities in habitat selection by caribou and

moose, but also major differences. Relative to mature conifer forests, both species avoided

open areas, lakes, open forest stands, roads, clearcuts, and stands harvested by CPHRS (Table

2). Both species also selected areas comprised of conifer forest with lichen and lichen-heath

plant community. On the other hand, caribou avoided mixed and deciduous forest stands,

whereas moose strongly selected these areas. Moreover, caribou displayed a strong aversion

for CPPTM sites, while moose did not (Table 2).

Table 2. Coefficients () from mixed-effects resource selection functions developed

from 37,447 observations on 18 radio-collared female woodland caribou

and from 71,039 observations on 10 radio-collared female moose

in the Côte-Nord region of Québec, Canada

Covariate† Woodland caribou Moose

β SE P β SE P

Open area -0.57 0.13 <0.0001 -0.61 0.19 0.001

Lake -2.03 0.19 <0.0001 -2.25 0.29 <0.0001

Conifer forest with lichen and

lichen-heath community

0.36 0.07 <0.0001 0.36 0.06 <0.0001

Open canopy -0.57 0.13 <0.0001 -0.61 0.19 0.001

Mixed/deciduous forest -0.64 0.11 <0.0001 1.29 0.15 <0.0001

Road -2.19 0.23 <0.0001 -1.95 0.41 <0.0001

Recent clearcut -1.95 0.32 <0.0001 -0.72 0.25 0.004

Regenerating clearcut -3.06 0.48 <0.0001 -0.65 0.14 <0.0001

CPPTM -3.24 0.71 0.0004 -1.36 1.02 0.22

CPHRS -3.38 0.48 <0.0001 -2.36 1.04 0.05 †Mature conifer forest was used as the reference category.


DISCUSSION

We evaluated the impact of CPPTM, a partial harvesting technique, up to 8 years post-

logging on 20 animal species belonging to multiple taxa and differing in both body size and

resource requirements. By focusing on species expected to be associated with closed- or

open-canopy areas, our approach assessed whether preserving small merchantable stems

maintained sufficient structure to sustain a suite of local animal communities. We found that

the null hypothesis was not supported: CPPTM supported early-successional species and were

not used as much by mid- and late-successional species as uncut stands. Overall, the structural

and compositional changes induced by this harvest technique altered species assemblages of

ants, beetles, forest birds, and mammals.

Although the single-grip harvester does not collect stems 15 cm in dbh during CPPTM,

the impacts of this silvicultural technique on wildlife habitat remains significant, at least in

the short-term. Approximately 80% of tree basal area is harvested in stands using CPPTM

(Ruel et al. 2007). The average tree diameter and the range of tree diameters are also greatly

reduced (Cimon-Morin et al. 2010). CPPTM has been shown to increase the proportion of

basal area of balsam fir in the canopy, from an initial value of 49% to 69% after logging

(Cimon-Morin et al. 2010). Furthermore, the density of arboreal lichens is nearly five times

lower, the biomass of terrestrial lichen >50 times lower (Courtois et al. 2008), the canopy

cover 75% lower (Hodson et al. 2010), and the lateral cover 28% lower (Courtois et al. 2008)

in CPPTM sites than in uncut stands. The amount of coarse woody debris is similar on

CPPTM sites to that on uncut stands (Légaré 2010), but the snag abundance is reduced

(Cimon-Morin et al. 2010, Légaré 2010).

The consistency in the response to CPPTM by a variety of species occupying very

distinct ecological niches indicated that, at least over the short term, this harvesting method

can have a profound impact on ecosystem functioning. For example, ants can be viewed as

ecological engineers due, in part, to their strong impact on soil characteristics (Jones et al.

1997). They also represent important components of forest food webs, being predators of

other insects (Way and Khoo 1992), and prey for woodpeckers (Torgersen and Bull 1995)

and black bears (Noyce et al. 1997). By building their nests in coarse woody debris and

stumps, the species studied here (Camponotus herculeanus and Formica neorufibarbis)

accelerate wood decomposition. In western Canada, for example, 49% of coarse woody

debris (including stumps) was reported to host ant colonies when located in harvested sites,

whereas they occurred on only 3% of uncut stands (Higgins and Lindgren 2006). Also,

logging could increase competitive interaction among ant species, potentially leading to

competitive exclusion (Kidd and Longair 1997). This might happen for C. herculeanus,

which is known to be non-aggressive despite its large size (Francoeur 1997).

Janssen et al. (2009) reported that E. planulata and R. dimidiatus were the most common

beetle species in the study area. We also found these species to be the most abundant beetles

in uncut forests. However, their abundance was >3.5 times lower in CPPTM sites than in

uncut stands. These saproxylic species are closely associated with recent snags (Jacobs et al.

2007a), which are five times less abundant in CPPTM sites than in uncut stands (Légaré

2010). The abundance of E. planulata and R. dimidiatus thus seemed to follow changes in the

availability of this resource. Nevertheless, these two species remained among the most

abundant beetles in the CPPTM sites suggesting that, at least on a short term, dead tree


recruitment still allows these species to persist after logging. The other closed-canopy beetles

were even more severely affected by CPPTM. These species might be more sensitive to an

increase in the fragmentation of old-growth forests. On the other hand, C. fusca, P.

triundulatus, H. congener, and X. laevigata, four saproxylic beetles, and Leiodes sp., a group

of epigaeic non-saproxylic beetles, benefited from forest harvesting. Our results indicated that

these species are favoured by canopy opening, which may lead to warmer temperatures and a

higher availability of flowering plants that can be consumed by adults. By feeding on bark of

young seedlings, H. congener may interfere in the establishment of regeneration (Welty and

Houseweart 1985). Species of the Leiodes assimilis-group are usually found in arid

environments (Baranowski 1993, Unpublished data, C. Hébert). Their abundance in CPPTM

is indicative of the important changes in abiotic conditions after harvesting. Overall, the

observed changes in species abundance could have a variety of implications, ranging from

nutrient cycling to interspecific interactions.

Many bird species are known to respond strongly to anthropogenic disturbances (Imbeau

et al. 2001, Vanderwel et al. 2007, Ghilain and Bélisle 2008, St-Laurent et al. 2008). These

responses can include changes in species assemblages, population declines (Wyshynski and

Nudds 2009), and changes in morphology as an adaptation to environmental change

(Desrochers 2010). We found that most of our focal bird species were influenced by CPPTM,

with only the Brown Creeper having a similar probability of occurrence in both CPPTM sites

and uncut forests. This result is surprising because previous studies have generally shown this

species to be negatively affected by intensive harvesting (Vanderwel et al. 2009, Poulin et al.

2010, but see also Venier and Pearce 2007 for an association of Brown Creeper with sites in a

logged landscape). Although the Brown Creeper is generally most abundant in mature and

old-growth forests in summer, its cryptic plumage, and high-pitched songs and calls

contribute to a low detection rate for this species on most surveys (Hejl et al. 2002). This low

detection rate could result in an under-estimation of numbers and distribution, making

detection of a treatment effect difficult. It is therefore possible that the number of sites, or the

sampling effort per site, was insufficient to detect the expected effect (although the trend was

consistent with expectations).

Boreal Chickadee and the Ruby-crowned Kinglet were less likely to be observed in

CPPTM sites than in uncut forests. Generally, most late-successional species are expected to

respond negatively to silvicultural practices with <30% tree retention (Harrison et al. 2005,

Vanderwel et al. 2009), as was observed following CPPTM. Other species benefited from the

forest structural changes: the White-throated Sparrow and the Dark-eyed Junco were most

likely to be found in CPPTM. This matched our expectations, as both species avoid dense

spruce forest, and have been shown to be neutral or positively affected by clearcutting and

forest fragmentation (Peters and Burleigh 1951, Schmiegelow et al. 1999, Steventon et al.

1998).

Finally, mammals were also influenced by the CPPTM practice. Snowshoe hare

gradually reduced their use of CPPTM sites for at least 3 years after logging, while they

maintained their use of uncut sites. The impacts of CPPTM were similar to those observed for

clearcut stands with protection of regeneration, which were found to be unsuitable for

snowshoe hares at least up to 4 years after logging (Ferron et al. 1998). Snowshoe hare is a

mid-successional herbivore that is generally most abundant in 20-30 year old stands because

they offer an advantageous combination of protective cover and browse availability

(Thompson et al. 1989, Koehler 1990, Newbury and Simon 2005). CPPTM seems to mainly


influence fitness through its effect on cover availability (Hodson et al. 2010). Given the

strong trophic influence of snowshoe hare on both predators and vegetation (Boutin et al.

1995), the reduced use of areas harvested by CPPTM could alter local predator distribution

and vegetation dynamics.

Woodland caribou and moose reacted differently to partial harvesting. Courtois et al.

(2008) evaluated the characteristics of CPPTM sites used by forest-dwelling caribou, but as

they pointed out, their relocation interval was insufficient to allow for an evaluation of

whether caribou benefit from CPPTM relative to more intensive logging practices. Our

intensive radio-tracking of individuals revealed that caribou avoided CPPTM sites just as

much as CPHRS and recent clearcut sites (e.g., Smith et al. 2000, Courtois et al. 2008, Fortin

et al. 2008, Courbin et al. 2009). Such high-intensity harvesting did not maintain habitat

quality for woodland caribou, as expected by Vanderwel et al. (2009). In contrast, CPPTM

appeared to have created short-term habitat conditions more suitable for moose compared to

those resulting from other logging techniques. Indeed, moose did not avoid CPPTM sites as

much as they avoided clearcut blocks.

Our findings on the impacts of CPPTM were evaluated in stands 1-8 years post-harvest.

Species assemblages typical of pre-harvesting conditions may not reoccur before regenerating

stands regain the characteristics of mature forest habitats (Schieck and Hobson 2000). One of

the main purposes of using CPPTM in old-growth forests would be to accelerate the return to

pre-harvest conditions (Courtois et al. 2008). Because the speed of return to original

conditions depends to some extent on the height of the residual regeneration trees (Pothier et

al. 1995), CPPTM sites are expected to become available for harvest 10 years before clearcut

sites (Ministère des Ressources naturelles, de la Faune et des Parcs 2003). The potential

benefits to wildlife species of a faster forest succession rate depend, however, on the

trajectory that this succession will take.

Courtois et al. (2008) suggested that CPPTM was most likely to be applied in stands

dominated by balsam fir because they often have an irregular structure suitable for partial

harvesting (Côté et al. 2010). The CPPTM technique could therefore favour the development

of balsam fir for 20 to 30 years post-harvest (Doucet 2000), with consequences on the direct

and indirect interactions of large mammals. In some eastern Canadian boreal forests, balsam

fir can dominate the diet of moose during winter months (Crête and Courtois 1997). If

CPPTM creates better winter habitat for moose, wolves that generally focus their hunting

efforts on this species might become attracted to CPPTM sites. This could result in a higher

concentration or even in a numerical response by wolves. Higher wolf densities or numbers

could result in negative feedback effects on woodland caribou (Seip 1992, Wittmer et al.

2007, 2010) that are established in landscapes partly harvested by CPPTM. This possible

indirect effect of recent silvicultural practices needs to be further investigated because

forestry activities are rapidly progressing towards the north, into some of the last pristine

habitats of forest-dwelling woodland caribou, a species threatened across Canada (Thomas

and Gray 2002).


CONCLUSION

Ecosystem-oriented conservation requires the use of silvicultural approaches that

maintain adequate distribution and abundance of forest cover types at landscape and regional

scales (Bergeron et al. 2002, 2007). We found that the tree retention associated with CPPTM

was insufficient to preserve animal diversity typical of old-growth boreal forests, at least up

to 8 years post-harvesting. The potential advantages of CPPTM over clearcutting strongly

depend on pre-harvest stand characteristics. The value of CPPTM for maintaining wildlife

diversity could certainly be improved by carefully choosing the stands based on the post-

harvest stand characteristics expected, given that stems >15 cm in dbh will be removed and

the smaller ones protected. Moreover, perhaps a certain number of large stems (>15 cm in

dbh) could be preserved to increase structural retention to a level more suitable to closed-

canopy wildlife species (e.g., 50-70%, Vanderwel et al. 2009).

Long-term advantages of CPPTM over clearcutting remain uncertain, but offer some

potential for wildlife conservation. CPPTM could bring back adequate structure for wildlife

more rapidly than clearcutting. The sites might need to be managed, however, to avoid major

changes in tree composition and structures (cf. Thompson et al. 2003). Specifically, managers

should avoid a conversion (outside the range of natural variability) of spruce- to fir-

dominated stands because such changes could have far-reaching consequences. Such a

conversion could reduce the future value of stands because balsam fir has weaker wood

properties and a faster rate of decay than black spruce, making balsam fir often less profitable

for the industry (Jessome 1977, Whitney 1995). Ecological consequences of stand conversion

can also emerge. For example, forests dominated by balsam fir are more vulnerable to spruce

budworm outbreaks (MacKinnon and MacLean 2004). Further, balsam fir can influence the

assemblage of beetle species (Azeria et al. 2009), and can also alter wolf-moose-caribou

interactions, and hence alter the functional characteristics of the ecosystem. There is field

evidence, however, indicating that a high abundance of balsam fir seedlings does not

necessarily result in a fir-dominated stand (Dubois et al. 2006, Doucet 2000). Additional

information is therefore needed on vegetation succession in stands harvested by CPPTM.

ACKNOWLEDGMENTS

This work was supported by the NSERC-Laval University industrial research chair in

silviculture and wildlife and its partners. We thank A. Labonne and Dr A. Francoeur for ant

identification, and G. Pelletier for help in coleopteran identification. We also thank L. Breton,

P. Dubois, B. Rochette, and D. Guay for helping with the capture of moose and caribou. We

would also like to thank the many field assistants who helped in data collection, as well as our

industrial partners for the logistical support. Finally, we are grateful to I. Thompson and J.-P.

Tremblay for their constructive comments on this paper.


REFERENCES

Annand, E. M. and Thompson III, F. R. (1997). Forest bird response to regeneration practices

in central hardwood forests. Journal of Wildlife Management, 61, 159–171.

Azeria, E. T., Fortin, D., Hébert, C., Peres-Neto, P., Pothier, D. and Ruel, J.-C. (2009). Using

null model analysis of species co-occurrences to deconstruct biodiversity patterns and

select indicator species. Diversity and Distributions, 15, 958-971.

Baranowski, R. (1993). Revision of the genus Leiodes Latreille of North and Central America

(Coleoptera: Leiodidae). Entomological Scandinavia Supplement, 42, 1-149.

Belsley, D. A., Kuh, E. and Welsch, R.E. (1980). Regression diagnostics. New York, NY:

John Wiley and Sons.

Bergeron, Y., Drapeau, P., Gauthier, S. and Lecomte, N. (2007). Using knowledge of natural

disturbances to support sustainable forest management in the northern Clay Belt. Forestry

Chronicle, 83, 326-337.

Bergeron, Y., Leduc, A., Harvey, B. D. and Gauthier, S. (2002). Natural fire regime: A guide

for sustainable management of the Canadian boreal forest. Silva Fennica, 36, 81–95.

Bouchard, M., Pothier, D. and Gauthier, S. (2008). Fire return intervals and tree species

succession in the North Shore region of Eastern Quebec. Canadian Journal of Forest

Research, 38, 1621–1633.

Boucher, D., De Grandpré, L. and Gauthier, S. (2003). Development of a stand structure

classification systems and comparison of two lichen-spruce woodlands in Quebec.

Forestry Chronicle, 79, 318–328.

Boulanger, Y. and Sirois, L. (2007). Postfire succession of saproxylic arthropods, with

emphasis on Coleoptera, in the North boreal forest of Quebec. Environmental

Entomology, 36, 128-141.

Boutin, S., Krebs, C. J., Boonstra, R., Dale, M. R. T., Hannon, S. J., Martin, K., Sinclair, A.

R. E., Smith, J. N. M., Turkington, R., Blower, M., Byrom, A., Doyle, F. I., Doyle, C.,

Hik, D., Hofer, L., Hubbs, A., Karels, T., Murray, D. L., Nams, V., O'Donoghue, M.,

Rohner, C. and Schweiger, S. (1995). Population changes of the vertebrate community

during a snowshoe hare cycle in Canada's boreal forest. Oikos 74: 69-80.

Cimon-Morin, J., Ruel, J.-C. and Darveau, M. (2010). Short term effects of alternative

silvicultural treatments on stand attributes in irregular balsam fir-black spruce stands.

Forest Ecology and Management, 260, 907-914.

Côté, G., Bouchard, M., Pothier, D. and Gauthier, S. (2010). Linking stand attributes to

cartographic information for ecosystem management purposes in the boreal forest of

eastern Québec. Forestry chronicle, 86, 511-519.

Côté, M. A. and Bouthillier, L. (2002). Assessing the effect of public involvement processes

in forest management in Quebec. Forest Policy and Economics, 4, 213-225.

Courbin, N., Fortin, D., Dussault, C. and Courtois, R. (2009). Habitat management for

woodland caribou: the protection of forest blocks influences wolf-caribou interactions.

Landscape Ecology, 24, 1375-1388.

Courtois, R., Gingras, A., Fortin, D., Sebbane, A., Rochette, B. and Breton, L. (2008).

Demographic and behavioural response of woodland caribou to forest harvesting.

Canadian Journal of Forest Research, 38, 2837-2849.


Crête, M. and Courtois, R. (1997). Limiting factors might obscure population regulation of

moose (Cervidae: Alces alces) in unproductive boreal forests. Journal of Zoology, 242,

765–781.

De Grandpré, L., Morissette, J. and Gauthier, S. (2000). Long-term post-fire changes in the

northeastern boreal forest of Quebec. Journal Vegetation Science, 11, 791-800.

Desrochers, A. (2010). Morphological response of songbirds to 100 years of landscape

change in North America. Ecology, 91, 1577-1582.

Doucet, R. (2000). L‘envahissement des parterres de coupe par le sapin est-il inévitable?

L‟Aubelle, 132, 11–13.

Drapeau, P., Nappi, A., Imbeau, L. and Saint-Germain, M. (2009). Standing deadwood for

keystone bird species in the eastern boreal forest: Managing for snag dynamics. Forestry

Chronicle, 85, 227-234.

Dubois, J., Ruel, J.-C., Élie, J.-G. andArchambault, L. (2006). Dynamique et estimation du

rendement des strates de retour après coupe totale dans la sapinière à bouleau jaune.

Forestry Chronicle, 82, 675-689.

Dussault, C., Courtois, R., Ouellet, J.-P. and Huot, J. (2001). Influence of satellite geometry

and differential correction on GPS location accuracy. Wildlife Society Bulletin, 29, 171–

179.

Dussault, C., Ouellet, J.-P., Courtois, R., Huot, J., Breton, L. and Jolicoeur, H. (2005).

Linking moose habitat selection to limiting factors. Ecography, 28, 619-628.

Fenton, N. J., Simard, M. and Bergeron, Y. (2009). Emulating natural disturbances: the role

of silviculture in creating even-aged and complex structures in the black spruce boreal

forest of eastern North America. Journal Forestry Research, 14, 258–267.

Ferron, J., Potvin, F. and Dussault, C. (1998). Short-term effects of logging on snowshoe

hares in the boreal forest. Canadian Journal of Forest Research, 28, 1335-1343.

Fisher J. T. and Wilkinson, L. (2005). The response of mammals to forest fire and timber

harvest in the North American boreal forest. Mammal Review, 35, 51-81.

Fortin, D., Courtois, R., Etcheverry, P., Dussault, C. and Gingras, A. (2008). Winter selection

of landscapes by woodland caribou: behavioural response to geographical gradients in

habitat attributes. Journal of Applied Ecology, 45, 1392-1400.

Francoeur, A. (1997). Ants (Hymenoptera: Formicidae) of the Yukon. In: H. V. Danks, and J.

A. Downes (Eds.), Insects of the Yukon (pp. 901-910). Ottawa, ON: Biological Survey of

Canada (Terrestrial Arthropods).

Francoeur, A. (2001). Les fourmis de la forêt boréale (Hymenoptera, Formicidae). Le

Naturaliste Canadien, 125, 108-114.

Ghilain, A. and Bélisle, M. (2008). Breeding success of tree swallows along a gradient of

agricultural intensification. Ecological Applications, 18, 1140-1154.

Groot, A., Lussier, J.-M., Mitchell, A. K. and MacIsaac, D. A. (2005). A silvicultural systems

perspective on changing Canadian forestry practices. Forestry Chronicle, 81, 50–55.

Harrison, R. B, Schmiegelow, F. K. A. and Naidoo, R. (2005). Stand-level response of

breeding forest songbirds to multiple levels of partial-cut harvest in four boreal forest

types. Canadian Journal of Forest Research, 35, 1553–1567.

Hejl, S. J., Newlon, K. R., Mcfadzen, M. E., Young, J. S. and Ghalambor, C. K. (2002).

Brown Creeper (Certhia americana) [online]. 2002 [cited 2010-07-10]. Available from:

http://bna.birds.

http://bna.birds.cornell.edu/bna/species/669


Higgins R.J. and Lindgren B. S. 2006. The fine scale physical attributes of coarse woody

debris and effects of surrounding stand structure on its utilization by ants (Hymenoptera:

Formicidae) in British Columbia, Canada. In S. J. Grove and J. L. Hanula (Eds.), Insect

biodiversity and dead wood: Proceedings of a symposium for the 22nd International

Congress of Entomology. Gen. Tech. Rep. SRS-93. Asheville, NC: U.S. Department of

Agriculture, Forest Service, Southern Research Station.

Hodson, J., Fortin, D., Le Blanc, M.-L. and Bélanger, L. (2010). An appraisal of the fitness

consequences of forest disturbance for wildlife using habitat selection theory. Oecologia,

164, 73-86.

Hunter, M. L. Jr. (1999). Maintaining biodiversity in forest ecosystems. Cambridge, UK:

Cambridge University Press.

Imbeau, L., Monkkonen, M. and Desrochers, A. (2001). Long-term effects of forestry on birds

of the eastern Canadian boreal forests: A comparison with Fennoscandia. Conservation

Biology, 15, 1151-1162.

Jacobs, J. M., Spence, J. R. and Langor, D. W. (2007a). Influence of boreal forest succession

and dead wood qualities on saproxylic beetles. Agricultural and Forest Entomology, 9, 3-

16.

Jacobs, J. M., Spence, J. R. and Langor, D. W. (2007b). Variable retention harvest of white

spruce stands and saproxylic beetle assemblages. Canadian Journal of Forest Research,

1631-1642.

Janssen, P., Fortin, D. and Hébert, C. (2009). Beetle diversity in a matrix of old-growth boreal

forest: Influence of habitat heterogeneity at multiple scales. Ecography, 32, 423-432.

Jennings, D. T., Houseweart, M. W. and Francoeur, A. (1986). Ants (Hymenoptera:

formicidae) associated with strip-clearcut and dense spruce-fir forest of Maine. Canadian

Entomologist, 118, 43-50.

Jessome, A. P. (1977). Résistance et propriétés connexes des bois indigènes au Canada.

Laboratoire des produits Forestiers de l'Est. Rapport Technique de Foresterie 21. Ottawa,

ON: Pêches et environnement Canada.

Johnstone, J. F., Hollingsworth, T. N., Chapin III, F. S. and Mack, M. C. (2009). Changes in

boreal fire regime break the legacy lock on successional trajectories in the Alaskan boreal

forest. Global Change Biology, 15, 1281-1295.

Jones, C. G., Lawton, J. H. and Shachak, M. (1997). Positive and negative effects of

organisms as physical ecosystem engineers. Ecology, 78, 1946-1957.

Keigley, R. B. and Frisina, M. R. (1998). Browse evaluation by analysis of growth form.

Helena, MT: Montana Fish, Wildlife, and Parks.

Kidd, M. G. and Longair, R. W. (1997). Abundance and diversity of ant (Hymenoptera:

Formicidae) assemblages in regenerating forests of northern Saskatchewan. Canadian

Field-Naturalist, 111, 635-637.

Klimaszewski, J., Pelletier, G., Germain, C., Work, T. and Hébert, C. (2006). Review of

Oxypoda species in Canada and Alaska (Coleoptera, Staphylinidae, Aleocharinae):

systematics, bionomics and distribution. The Canadian Entomologist, 138, 737-852.

Koehler, G. M. (1990). Population and habitat characteristics of lynx and snowshoe hares in

North Central Washington. Canadian Journal of Zoology, 68, 845-851.

Légaré, J.-P. (2010). Traitements sylvicoles alternatifs en forêt boréale irrégulière sur la Côte-

Nord: la réponse des communautés de coléoptères. MSc thesis. Québec, Qc: Département

des sciences du bois et de la nature, Université Laval.


Lemaître, J. (2009). Déterminants de la diversité des oiseaux et des micromammifères en

forêt boréale naturelle et aménagée. PhD thesis. Québec, Qc: Département de biologie,

Université Laval.

Liu, C., Ruel, J.-C. and Zhang, S. Y. (2007). Immediate impacts of partial cutting strategies

on stand characteristics and value. Forestry Ecology Management, 250, 148-155.

MacKinnon, W. E. and MacLean, D. A. (2004). Effects of surrounding forest and site

conditions on growth reduction of balsam fir and spruce caused by spruce budworm

defoliation. Canadian Journal of Forest Research, 34, 2351–2362.

Majka, C. J. (2008). The flat bark beetles (Coleoptera, Silvanidae, Cucujidae,

Laemophloeidae) of Atlantic Canada. ZooKeys, 2, 221-238.

Majka, C. J., Langor, D. and Rücker, W. H. (2009). Latridiidae (Coleoptera) of Atlantic

Canada: new records, keys to identification, new synonyms, distribution, and

zoogeography. The Canadian Entomologist, 141, 317-370.

Manly B. F. J., McDonald L. L., Thomas D. A., McDonald T. L. and Erickson W. E. (2002).

Resource selection by animals, statistical design and analysis for field studies (2nd

edition). Dordrecht, The Netherlands: Kluwer Academic Publishers.

Ministère des Ressources naturelles de la Faune et des Parcs. (2003). Manuel d‘aménagement

forestier du Québec (4th

edition). Charlesbourg, Qc: Ministère des Ressources naturelles

de la Faune et des Parcs.

Newbury, T. L. and Simon N. P. P. (2005). The effects of clearcutting on snowshoe hare

(Lepus americanus) relative abundance in central Labrador. Forest Ecology and

Management 210:131-142.

Noyce, K. V., Kannowski, P. B., Riggs, M. R. (1997). Black bears as ant-eaters: seasonal

associations between bear myrmecophagy and ant ecology in north-central Minnesota.

Canadian Journal of Zoology, 75, 1671-1686.

Pastor, J., Cohen, Y. and Moen, R. (1999). Generation of Spatial Patterns in Boreal Forest

Landscapes. Ecosystems, 2, 439-450.

Pease, J. L., Vowles, R. H. and Keith, L. B. (1979). Interaction of snowshoe hares and woody

vegetation. Journal of Wildlife Management, 43, 43-60.

Peters, H. S. and Burleigh, T. D. (1951). The birds of Newfoundland. St.John‘s, NL:

Department of Natural Resources.

Pham A. T., De Grandpré, L., Gauthier, S. and Bergeron, Y. (2004). Gap dynamics and

replacement patterns in gaps of the northeastern boreal forest of Quebec. Canadian

Journal of Forest Research, 34, 353–364

Pothier, D., Doucet, R. and Boily, J. (1995). The effect of advance regeneration height on

future yield of black spruce stands. Canadian Journal of Forest Research. 25, 536-544.

Potvin, F., Courtois, R., and Bélanger, L. (1999). Short-term response of wildlife to clear-

cutting in Quebec boreal forest: Multiscale effects and management implications.

Canadian Journal of Forest Research, 29, 1120-1127.

Poulin, J.-F., Villard, M.-A. and Haché, S. (2010). Short-term demographic response of an

old forest specialist to experimental selection harvesting. Ecoscience, 17, 20-27.

Rosenvald, R. and Lõhmus, A. (2007). Breeding birds in hemiboreal clear-cuts: tree-retention

effects in relation to site type. Forestry, 80, 503–516.

Rosenvald, R. and Lõhmus, A. (2008). For what, when, and where is green-tree retention

better than clear-cutting? A review of the biodiversity aspects. Forest Ecology and

Management, 255, 1–15.


Ruel, J.-C., Roy, V., Lussier, J.-M., Pothier, D. Meek, P. and Fortin, D. (2007). Mise au point

d‘une sylviculture adaptée à la forêt boréale irrégulière. Forestry Chronicle, 83, 367-374.

Saint-Germain, M., Drapeau, P. and Hébert, C. (2004). Comparison of Coleoptera

assemblages from a recently burned and unburned black spruce forests of northeastern

North America. Biological Conservation, 118, 583–592.

Schieck, J. and Hobson, K. A. (2000). Bird communities associated with live residual tree

patches within cut blocks and burned habitat in mixedwood boreal forests. Canadian

Journal of Forest Research, 30, 1281–1295.

Schmiegelow, F. K. A. and Hannon, S. J. (1999). Forest-level effects of management on

boreal songbirds: the Calling Lake Fragmentation Studies. In J. A. Rochelle, L. A.

Lehmann and J. Wisniewski (Eds.), Forest fragmentation, wildlife and management

implications (pp. 201-221). Boston, MA: Brill Press.

Seip, D.R. (1992). Factors limiting woodland caribou populations and their interrelationships

with wolves and moose in southeastern British Columbia. Canadian Journal of Zoology,

70, 1494–1503.

Simon, N. P. P., Schwab, F. E. and Diamond, A. W. (2000). Patterns of breeding bird

abundance in relation to logging in western Labrador. Canadian Journal of Forest

Research, 30, 257–263.

Smith, K. G., Ficht, E. J., Hobson, D., Sorensen, T. C. and Hervieux, D. (2000). Winter

distribution of woodland caribou in relation to clear-cut logging in west-central Alberta.

Canadian Journal of Zoology, 78, 1433-1440.

Steventon, J. D., MacKenzie, K. L. and Mahon, T. E. (1998). Response of small mammals

and birds to partial cutting and clearcutting in northwest British Columbia. Forestry

Chronicle, 74, 703-713.

St-Laurent, M.-H., Ferron, J., Haché, S. and Gagnon, R. (2008). Planning timber harvest of

residual forest stands without compromising bird and small mammal communities in

boreal landscapes. Forest Ecology and Management, 254, 261-275.

Thomas, D. C. and Gray, D. R. (2002). Update COSEWIC status report on the woodland

caribou Rangifer tarandus caribou in Canada: COSEWIC Assessment and Update Status

Report on the Woodland Caribou Rangifer tarandus caribou in Canada. Ottawa, ON.

Committee on the Status of Endangered Wildlife in Canada.

Thompson, I. D., Baker, J. A., Jastrebski, C., Dacosta, J., Fryxell, J. and Corbett, D. (2008).

Effects of post-harvest silviculture on use of boreal forest stands by amphibians and

marten in Ontario. Forestry chronicle, 84, 741-747.

Thompson, I. D., Baker, J. A. and Ter-Mikaelian, M. (2003). A review of the long-term

effects of post-harvest silviculture on vertebrate wildlife, and predictive models, with an

emphasis on boreal forests in Ontario, Canada. Forest Ecology and Management, 177,

441–469.

Thompson I. D., Davidson I. J., Odonnell S. and Brazeau F. (1989). Use of track transects to

measure the relative occurrence of some boreal mammals in uncut forest and regeneration

stands. Canadian Journal of Zoology, 67, 1816-1823.

Torgersen T. R. and Bull E. V. (1995). Down logs as habitat for forest dwelling ants – the

primary prey of Pileated Woodpeckers in northeastern Oregon. Northwest Science, 69,

294-303.


Vanderwel, M. C., Malcolm, J. R. and Mills, S. C. (2007). A meta-analysis of bird responses

to uniform partial harvesting across North America. Conservation Biology, 21, 1230–

1240.

Vanderwel, M. C., Mills, S. C. and Malcolm, J. R. (2009). Effects of partial harvesting on

vertebrate species associated with late-successional forests in Ontario‘s boreal region.

Forestry chronicle, 85, 91-104.

Venier, L. A. and Pearce, J. L. (2004). Birds as indicators of sustainable forest management.

Forestry Chronicle, 80, 61-66.

Venier, L.A. and J.L. Pearce. (2007). Boreal forest landbirds in relation to forest composition,

structure, and landscape: implications for forest management. Canadian Journal of

Forest Research 37: 1214–1226.

Verner, J. (1985). Assessment of counting techniques. Current Ornithology, 2, 247-302.

Way, M.J. and Khoo, K.C. (1992). Role of ants in pest management. Annual Review of

Entomology, 37, 479-503.

Welty, C. and Houseweart, M. W. (1985). Site influences on Hylobius congener (Coleoptera:

Curculionidae), a seedling debarking weevil of conifer plantations in Maine.

Environmental Entomology, 14, 826-833.

Whitney, R. D. (1995). Root-rotting fungi in white spruce, black spruce, and balsam fir in

northern Ontario. Canadian Journal of Forest Research, 25, 1209-1230.

Wittmer, H. U., Ahrens, R. N. M. and McLellan, B. N. (2010). Viability of mountain caribou

in British Columbia, Canada: Effects of habitat change and population density. Biological

Conservation, 143, 86-93.

Wittmer, H. U., McLellan, B. N., Serrouya, R. and Apps, C. D. (2007). Changes in landscape

composition influence the decline of a threatened woodland caribou population. Journal

of Animal Ecology, 76, 568-579.

Wyshynski, S. A. and Nudds, T. D. (2009). Pattern and process in forest bird communities on

boreal landscapes originating from wildfire and timber harvest. Forestry chronicle, 85,

218-226.

Reviewed by Ian D. Thompson (Canadian Forest Service) and Jean-Pierre Tremblay

(Université Laval)

chapter 4 et al 2011.pdf · in: woodlands: ecology, management and conservation isbn...

Documents