Écoscience

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A landscape perspective of bird nest predation in a

managed boreal black spruce forest

Marylène Boulet, Marcel Darveau & Louis Bèlanger

To cite this article: Marylène Boulet, Marcel Darveau & Louis Bèlanger (2000) A landscape
perspective of bird nest predation in a managed boreal black spruce forest, Écoscience, 7:3,
281-289, DOI: 10.1080/11956860.2000.11682597

To link to this article: https://doi.org/10.1080/11956860.2000.11682597

Published online: 24 Mar 2016.

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 7 (3) : 281-289 (2000)

A landscape perspective of bird nest predation

in a managed boreal black spruce forest1
Marylène BOULET2, Marcel DARVEAU & Louis BÉLANGER, Centre de recherche en biologie forestière,
Faculté de Foresterie et de Géomatique, Université Laval, Sainte-Foy, Québec G1K 7P4, Canada.

Abstract: Several landscape level studies have reported that bird nest predation increases as forest cover decreases. These studies
have mainly been conducted in agricultural or urban regions. However, few studies have explored relationships between
forest cover and nest predation in boreal forests managed for timber harvesting. In 1997 and 1998, we evaluated bird nest
predation in a mosaic of clearcuts and forest remnants dominated by black spruce (Picea mariana [Mill.] B.S.P.) and located
north of Lake Saint-Jean, Québec. We used a 7 km × 9 km grid of sampling points to determine nest predation at four landscape
scales (local vegetation, and 250 m, 500 m, and 1000 m radii around sampling points). Artificial nests (ground and arboreal)
containing a common quail (Coturnix coturnix L.) egg and a plasticine egg were used to calculate predation pressure and to
identify nest predators. Nest predation was high over the entire study area. Dominant predators were the gray jay (Perisoreus
canadensis L.) and the red squirrel (Tamiasciurus hudsonicus Erxleben). Depredation by squirrels was influenced by local
variables in 1997 and by landscape variables in 1998. In the latter case, depredation by squirrels increased as spruce cover
increased. Depredation by gray jays was positively related to water body area and jack pine (Pinus banksiana Lamb.) cover.
Squirrels preyed more on ground nests than on arboreal nests, while gray jays preyed almost exclusively on arboreal nests.
We conclude that these predators probably impose different threats to different songbird species in boreal black spruce forests.
Our results show that, in the short term, timber harvesting did not seem to increase predation in a boreal black spruce forest.
Keywords: bird nest predation, landscape scales, Perisoreus canadensis L., Tamiasciurus hudsonicus Erxleben., boreal black
spruce forest, clearcutting.

Résumé : Plusieurs études réalisées à l’échelle du paysage ont démontré que la prédation des nids d’oiseaux augmente
lorsque le couvert forestier diminue. Ces études ont toutefois été surtout réalisées dans des régions agricoles ou urbaines. Par
contre, peu d’études ont exploré les relations entre les types de couvert forestier et la prédation dans les forêts boréales. En
1997 et 1998, nous avons évalué la prédation des nids d’oiseaux dans une mosaïque de parterres de coupe et de lambeaux de
forêt dominée par l’épinette noire (Picea mariana [Mill.] B.S.P.) au nord du Lac Saint-Jean, Québec. Nous avons établi une
grille de points d’échantillonnage de 7 km × 9 km pour déterminer le risque de prédation à quatre échelles différentes
(végétation locale et rayons de 250, 500 et 1000 m autour des points). Nous avons utilisé des nids artificiels (0 et 5 m de
hauteur) dans lesquels nous avons placé un œuf de caille des blés (Coturnix coturnix L.) et un œuf de plasticine pour faciliter
l’identification des prédateurs. La prédation des nids était importante dans l’ensemble de l’aire d’étude. Le mésangeai du Canada
(Perisoreus canadensis L.) et l’écureuil roux (Tamiasciurus hudsonicus Erxleben) constituaient les principaux prédateurs.
La prédation par l’écureuil était influencée par des variables locales en 1997 et par des variables de paysage en 1998.
Cette dernière année, la prédation par l’écureuil était positivement associée à la superficie en épinettes. La prédation par le
mésangeai du Canada était positivement corrélée à la présence de plans d’eau et au couvert en pins gris (Pinus banksiana Lamb.).
L’écureuil attaquait un peu plus les nids au sol que les nids à 5 m, alors que le mésangeai s’en prenait presque exclusivement
aux nids à 5 m. Nous concluons donc que ces prédateurs exercent probablement des pressions différentes sur les différentes
espèces de passereaux dans la pessière noire boréale. À court terme, nos résultats indiquent que la récolte forestière ne semble
pas augmenter la prédation des nids dans une pessière noire boréale.
Mots-clés : prédation de nids d’oiseaux, échelles de paysage, Perisoreus canadensis L., Tamiasciurus hudsonicus Erxleben,
pessière noire boréale, coupes forestières.

Introduction
Three types of fragmentation studies on bird nest
predation can be distinguished: (i) stand level studies; (ii)
“simple-contrast” landscape studies; and (iii) “multi-contrast”
landscape studies. Stand level studies, which focus on the
impacts of forest fragmentation within a patch of habitat,
are the most common type (Wilcove, 1985; Vander Haegen
& DeGraaf, 1996; Huhta, Jokimäki & Helle, 1998). Patch
size and edge effects are now well documented, particularly
in agricultural areas (e.g., predation increases near edges and
in small patches of forest habitats; Paton, 1994; Andrén,
1995). However, the understanding of landscape level pheno-
1Rec. 1999-10-26; acc. 2000-03-20.
2Author for correspondence. Present address: Department of Biology, LSB, McMaster
University, 1280 Main Street West, Hamilton, Ontario L8S 4K1, Canada, e-mail:
bouletm@mcmaster.ca
mena is somewhat limited with this approach, because the
spatial context of the study area is not taken into account.
Landscape level studies have emerged from the
development of new technologies, in particular geographic
information systems, global positioning systems, and satellite
imagery. Simple-contrast landscape studies usually simplify
the ecosystem by contrasting only two types of land cover:
forest cover versus open habitat cover. For example,
Robinson et al. (1995) used a landscape level approach and
reported that nest predation and parasitism by cowbirds
increased with decreasing forest cover in nine landscapes
(located in midwestern U.S.A.) that varied from 6% to 95%
forest cover within a 10 km radius around study plots. Hartley
& Hunter (1998), in a meta-analysis summarizing 13 studies
on artificial nest predation which were conducted in eastern
U.S.A., showed that daily nest predation rate decreased as
BOULET, DARVEAU & BÉLANGER: BIRD NEST PREDATION
forest cover increased at three spatial scales (5, 10, and 25 km
radius). In eastern and central North America, generalist
species like corvids, raccoons, and skunks are abundant and
often associated with high predation rates (Andrén, 1995;
Faaborg et al., 1995).
In multi-contrast landscape studies, the ecosystem is
simplified into several types of land cover (e.g., water,
coniferous forest, deciduous forest, and field). Such studies
often focus on two questions: who are the predators and
how do they use the landscape? In Europe, Andrén (1992)
reported that different species of corvids responded differ-
ently to the type of land cover. For example, the carrion
crow (Corvus corone L.) correlated positively with croplands
and fields and negatively with young and mature forests,
whereas the Eurasian jay (Garrulus glandarius L.) correlated
positively with mature forests and negatively with croplands.
Andrén (1992) also reported that the hooded crow caused
increased predation pressure near forest-farm edges and in
small fragments of forest surrounded by agricultural land.
In North America, the raccoon (Procyon lotor L.) is
often reported as an important nest predator at forest edges
(Gates & Gysel, 1978; Wilcove, 1985; Small & Hunter,
1988). Recently, Pedlar, Fahrig & Merriam (1997) studied
habitat use by this species at the microhabitat (10-m radius)
and macrohabitat (1000-m radius) scales. They found that
raccoons used microhabitats with woody vegetation features
that were associated with fencerows, den trees, and deciduous
stands, and macrohabitats with extensive agricultural edges
and wooded fragments in areas with extensive corn cover.
This preference for agricultural edges provides more evidence
for this species causing increased predation near edges. If we
consider the diversity of habitat preferences and territory
size of nest predators, relationships between landscape
composition and nest predation may vary considerably with
landscape contexts and predator communities.
The local vegetation also influences predator for-
aging. High levels of spatial heterogeneity decrease foraging
efficiency (e.g., number of bird nests found per unit of time)
by the raccoon (Bowman & Harris, 1980). Similarly, Seitz
& Zegers (1993) found that nest success was greater in
early successional stands than in adjacent deciduous or
coniferous stands. These authors concluded that dense
vegetation impeded movements of predators and decreased
nest visibility. Moreover, high levels of heterogeneity offer
greater number of potential nest sites (Martin, 1993). Forest
type also modulates nest predation, particularly if nests are
located in the preferred habitat of an abundant predator.
For example, artificial nests in coniferous forest suffered
higher rates of predation than in deciduous forest, as a result
of higher abundance of red squirrels in coniferous forest
(Bayne, Hobson & Fargey, 1997; Sieving & Willson, 1998).
In a boreal black spruce (Picea mariana [Mill.]
B.S.P.) forest, we investigated bird nest predation at four
different scales (local vegetation, and radii of 250 m, 500 m
and 1000 m). We identified important nest predators in our
study area and we determined how they responded to land-
scape composition. We predicted that nest predation would
increase as preferred habitats of forest-specialist predators
increase. As our study site was partially clearcut, we also
evaluated how clearcuts affected nest predation and predator
communities.
282
Material and methods
STUDY AREA
We conducted this study in an area of 100 km2
located north of Lake Saint-Jean, Québec (49° 09’ N ,
72° 58’ W). This area is located in the southern range of the
black spruce and moss bioclimatic domain, in the boreal
ecozone (Grondin, 1996). Mean annual temperature at the
nearest meteorological station (Normandin, 50 km farther
south) was 1.0°C and mean annual precipitation was 866 mm
(Environment Canada, 1993). Snow covered the ground
from November to end of April each year. The landscape
was undulating and several rivers and lakes were present. In
the 1950s, part of this area was clearcut and stands that had
not reached merchantable size at that time were harvested in
the 1990s (Figure 1). Recent clearcuts represented 50% of
the study area (Figure 1). Forest remnants left after timber
harvesting included riparian buffer strips (20 to 30 m wide)
along all permanent rivers and lakes, “non-riparian leave”
strips (60 m wide), and some blocks of forest (> 100 m wide).
Forest remnants often remained connected to adjacent
remnants or to intact forest, which generally consisted of
patches that had not reached merchantable size.. The area
was dominated by black spruce stands, intermixed with jack
pine (Pinus banksiana Lamb.), balsam fir (Abies balsamea
[L.] Mill.), trembling aspen (Populus tremuloides Michx.),
and white birch (Betula papyrifera Marsh; N. Bertrand,
C. Paquet & F. Potvin, unpubl. data). Clearcuts were colo-
Québec
Study area
Québec
city
Legend
Sampling point
Clearcut
Deciduous
Fir
Pine
Sapling
Spruce
Wetland
Water
3 0 3 6 kilometers
FIGURE 1. Layout of sampling points over the reclassified land cover
map of the study area, north of Lake Saint-Jean, Québec. Areas clearcutted
in 1998 (western sector) are delimited by bold lines.

nised by sheep laurel (Kalmia angustifolia L.), blueberry
(Vaccinium spp.), Labrador tea (Ledum groenlandicum
Retzius), black spruce, balsam fir, aspen and white birch.
Based on Banfield (1974), Gauthier & Aubry (1995),
and Prescott & Richard (1996), possible bird nest preda-
tors within this area included: red fox, short-tailed weasel
(Mustela erminea L.), pine marten (Martes americana Turton),
mink (Mustela vison Schreber), red squirrel (Tamiasciurus
hudsonicus Erxleben), northern flying squirrel (Glaucomys
sabrinus Shaw), eastern chipmunk (Tamias striatus L.),
small mammals (possibly recruited among deer mouse
[Peromyscus maniculatus Wagner], southern red-backed vole
[Clethrionomys gapperi Vigors], and woodland jumping
mouse [Napaeozapus insignis Miller]), gray jay, common
raven (Corvus corax L.), and American crow (Corvus
brachyrhynchos Brehm).
EXPERIMENTAL GRID
In 1997, we set up a 7 km × 9 km grid divided into
1 km × 1 km cells (Figure 1). We placed a sampling point
in each accessible cell (method shared by the Landscape
Structure Biodiversity Project [LSBP] of the Sustainable
Forest Management Network, Natural Sciences and Eng-
ineering Research Council of Canada). Our grid included
45 sampling points in 1997, of which five were clearcutted
during winter 1997-1998 and replaced with six new points
in 1998, all located within forest remnants. We also added
five sampling points that were located in highly forested
areas to cover a larger continuum of forest cover, for a total
of 50 sampling points in 1997 and 51 in 1998.
LANDSCAPE COMPOSITION DATA
The study site was mapped with a geographic
information system (ArcView), using the same stratification
as Québec’s Forest Inventory, with the minimal size of
stands fixed to 1 ha. We reclassified land cover into height
classes more relevant for birds (Table I) and generated three
landscape buffers (250 m, 500 m and 1000 m) for each
sampling point with ArcView. For each class, we then cal-
culated the total area (a variable reflecting the composition
of the ecosystem) and the mean edge contrast index (a
variable reflecting the structure of the ecosystem) within
the three buffers using FRAGSTATS (McGarigal &
Marks, 1995). We measured Pearson correlations among
our variables of structure and composition. As the mean edge
contrast of a particular class usually significantly (P < 0.05)
correlated with the total area of the same class, we excluded
the mean edge contrast from our analysis and concluded
that the ecosystem structure was highly associated with its
composition and did not allow us to distinguish between
composition and structure.
LOCAL VEGETATION
In 1998, we sampled vegetation in four 20 m × 10 m
plots, one located within the 50 m radius and the other three
located north, southeast, and southwest of the first one,
between the 50 m and 100 m radius (n = 46, see previous
paragraph). Quadrats were displaced in a counter-clockwise
direction when an obstacle was present (e.g., a road, lake, or
river). We tallied all trees (> 2 m height, by species and
ÉCOSCIENCE, VOL. 7 (3), 2000
TABLE I. Landscape composition variables and local vegetation
variables used in stepwise logistic regressions. Local vegetation
variables used are mean values of the four quadrats (20 m × 10 m)
for each sampling point, located in a black spruce forest north of
Lake Saint-Jean, Québec.
Classes Descriptions
LANDSCAPE COMPOSITION VARIABLES
Spruce Stands dominated by black spruce > 7 m high
Pine Stands dominated by jack pine > 7 m high
Fir Stands dominated by balsam fir > 7 m high
Deciduous Stands dominated by deciduous trees > 7 m high
Sapling Young stands 1.5 to 7 m high
Clearcut Very young stands (< 8 years) < 1.5 m high, bare ground,
rocky outcrops
Wetland Swamps, wet stands, alder stands, peatlands forested
or not, islands
Water Lakes, large rivers, ponds
LOCAL VEGETATION VARIABLES
Intolerant trees Mean number of trembling aspen and white birch trees
> 8 cm dbh
Fir trees Mean number of balsam fir trees > 8 cm dbh
Spruce trees Mean number of black spruce trees > 8 cm dbh
Pine trees Mean number of jack pine trees > 8 cm dbh
Debris Mean number of coarse woody debris > 8 cm dbh
and ≥ 50 cm long
Snags Mean number of snags > 8 cm dbh and ≥ 50 cm high
Alder and willow Mean cover (%) of alder and willow shrubs < 2 m high
diameter at breast height [dbh] class) and all snags (> 8 cm
dbh and ≥ 50 cm height, by species, dbh class, and decay
class) present within the four plots. We measured coarse
woody debris (> 8 cm dbh, ≥ 50 cm long) present along the
two 20-m lines and noted their decay class and species
when possible. We classified the percentage of shrub cover
(by species) and ground cover into five categories: rare
(< 1%); few occurrences (1-10%); several occurrences
(10-20%); frequent (20-50%); and very common (> 50%)
in each plot. We estimated the height of low plants and
shrubs and we measured height of the subcanopy and
canopy with a clinometer. Canopy cover was estimated by
walking along the two 20-m parallel line transects and
scoring the cover 20 times with an ocular tube. Subcanopy
cover was assessed from below.
We measured Pearson correlations among 38 vari-
ables of structure and composition from our local vegetation
data set. We excluded 22 variables that were correlated with
other variables or had a priori poor biological relevance for
predation events and summarized six other variables into
three new variables, for a total of 14 independent variables.
A principal coordinate analysis (in Q-mode and with the
Chi-square metric coefficient) was used to identify the vari-
ables that best summarized the local structure and compo-
sition of our ecosystem, using R Statistical Software
(Legendre & Legendre, 1998). Three eigenvalues were sig-
nificant according to the broken stick model (Legendre
& Legendre, 1998) and explained 7% of the variance.
Examination of the resulting graphs revealed that the
ecosystem structure was again highly associated with its
composition. According to the position of the 14 variables
in graphs and a priori biological significance for nesting
birds, we kept seven of these variables, thereafter referred to
as local vegetation variables (Table I).
283
BOULET, DARVEAU & BÉLANGER: BIRD NEST PREDATION
ARTIFICIAL NEST EXPERIMENT
Two arboreal (5-m high) and two ground nests
were placed within a radius of 25 m of each sampling point.
Arboreal nests consisted of a chicken wire matrix filled with
arboreal lichens and mosses. They were suspended from a
branch of a coniferous tree, close to the trunk. Ground nests
were hollows made by compacting mosses, lichens, or litter
by hand. A common quail egg and a painted plasticine egg
mimicking the quail egg in size, shape, and colour pattern
were put in each nest. We used a small wire to attach the
plasticine egg to the nest or to a shrub root to prevent its
removal by predators (Darveau et al., 1997). Nests were
placed out from June 5 to 17 in 1997 and from June 2 to 13
in 1998, and were inspected three times, e.g. 7-9 days, 14-
17 days, and 21 days later. This exposure covers the period
of laying and incubation of songbird species found in our
area (usually around 17 days in most species, but can extend
up to 21 in some species; Gauthier & Aubry, 1995) and
has been used in other studies (Song & Hannon, 1999).
Predation was assumed when at least one egg was pecked,
broken, bitten, or removed. We compared bite prints in the
plasticine eggs with prints taken from mammal skulls and
bird specimens to identify predators. We also established a
key (based on four years of experimentation) based on
tooth size and dental characteristics for mammals, and on
peck size and shape for birds (I. Chouinard, M. Boulet,
M. Courteau, & M. Darveau, unpubl. data). To minimize
identification errors, two observers independently identified
prints left by predators and corrected each other.
OBSERVATION OF PREDATORS
We censused red squirrels and gray jays using a
standard point count technique (Ralph, Sauer & Droege,
1995). Three 15-minute visits were made at each point in
June 1997 and 1998. We used radii grossly corresponding
to the size of the territory of predators, e.g. 50 m radius for
squirrels (territory size usually ranging from 0.24 to
0.98 ha; Steele, 1998) and unlimited radius for gray jays
(territory size ranging from 41 to 146 ha; Strickland &
Ouellet, 1993).
STATISTICAL ANALYSIS
We used a stepwise logistic regression with the
following ratio entered as the dependent variable: number
of nests preyed on at each sampling point/total number of
nests at each sampling point, e.g. 0-4 nests/4 nests (SAS
Institute, 1990; Agresti, 1996). In a first series of analyses,
we ran our model eight times, e.g. four scales (local, 250 m,
500 m and 1000 m) in two years with n = 46 sampling
points for the local scale and n = 50 in 1997 and n = 51 in
1998 for the landscape scales. In a second series including
both years, we used the year as a random variable and sub-
sampled points that were identical for both years of our
experiment, e.g., had not been modified by logging activities
(n = 40, 36, 34, and 28 sampling points for local, 250 m,
500 m, 1000 m scales, respectively). In a third series includ-
ing all scales (e.g., global model), we subsampled points for
which we had data at all scales (n = 40 in 1997 and n = 46
in 1998). We repeated these procedures three times, for:
(i) all predation events; (ii) predation by squirrels as
revealed by marks on plasticine egg; and (iii) predation by
284
gray jays. P values for entry and staying in the model were
set to 0.05 and 0.10, respectively. Alpha was also set to 0.05
in the following tests/correlations. We compared predation
events among nest heights using McNemar’s test (Agresti,
1996). G-tests were used to test for associations between
presence of predators and evidence of predation events. We
also calculated Pearson’s coefficients of correlation
between the abundance of predators and the number of eggs
that they preyed on.
Results
NEST PREDATION
Nest predation was high in both years of the study:
67% (133/200) of nests were manipulated by predators after
21 days of exposure in 1997, and 74% (150/204) in 1998.
Predation was influenced by local variables, which differed
between 1997 and 1998 (Table II). Based on the model
including both years, predation was higher in sites domi-
nated by black spruce trees and with fewer snags. Only one
landscape variable (fir cover) entered into the model at the
250 m scale, and none entered at the 500 m and 1000 m
scales. The global models for 1997 and 1998 selected dif-
ferent variables (Table II), but both landscape (1997) and
local (1998) abundance in pines correlated negatively with
nest predation. Local abundance of black spruce was still a
significant variable (global model 1997).
NEST PREDATORS
One hundred and twenty-seven plasticine eggs
were attacked in 1997 and 143 in 1998. In 1997 and 1998
respectively, 26 and 10 plasticine eggs were taken despite
their metallic attachment, 10 and five eggs remained
unidentified, and two and four eggs had marks of two dif-
ferent species of predators, for a total of 91 identifiable
marks recorded in 1997 and 132 in 1998. The most impor-
tant predators were gray jays, with 39 identifications in
1997 and 60 in 1998, and small sciurids (red squirrels,
northern flying squirrels, or eastern chipmunks), with
41 identifications in 1997 and 49 in 1998 (Figure 2). Other
predators included small mammals (voles, mice and shrews,
8-14 identifications for 1997-1998), snowshoe hare (Lepus
americanus Erxleben; 3-7 identifications for 1997-1998),
short-tailed weasel (1 in 1998), and a small falcon (1 in
1998). Depredation by squirrels differed marginally among
nest heights in 1997 (McNemar’s test, Statistic = 3.5,
P = 0.06) but not in 1998 (McMemar’s test, Statistic = 0.8,
P = 0.4; Figure 2). Depredation by gray jays was much
more significant on arboreal nests than on ground nests
(McNemar’s test, Statistic = 31.1, P = 0.001 in 1997;
Statistic = 40.3, P = 0.001 in 1998; Figure 2).
Predation attacks by squirrels were negatively
associated with local abundance of alder and willow shrubs
in the local model 1997, in the model including years 1997
and 1998, and in the global model 1997 (Table II). Two
other local variables, namely snags and fir trees, were
entered into the global model 1997. In 1998, predation by
squirrels was greater in sites with greater cover of black
spruce and clearcuts at 250 m and 500 m scales. Predation
by squirrels was also negatively related to the area of water
(1000 m scale) in 1998. Only black spruce cover (250 m
 ÉCOSCIENCE, VOL. 7 (3), 2000

TABLE II. Results of stepwise logistic regressions that entered landscape composition variables and or local vegetation variables for: (i) all
predation events; (ii) predation by squirrels; and (iii) predation by gray jays, in a black spruce forest north of Lake Saint-Jean, Québec.
+ and – indicate positive or negative relationships that were significant at P ≤ 0.05.

Scales 1997 1998 1997-1998

ALL EVENTS
Local spruce trees (+), debris (–) alder and willow (+) spruce trees (+), snags (–)
250 m fir (–)
500 m
1000 m
Global pine 500 m (–), water 1000 m (+), fir 500 m (–), deciduous 250 m (–),
spruce trees (+) pine trees (–) N/A
SQUIRRELS
Local alder and willow (–) alder and willow (–)
250 m spruce (+), clearcut (+) spruce (+)
500 m spruce (+), clearcut (+)
1000 m water (–)
Global alder and willow (–), snags (–), spruce 500 m (+), N/A
fir trees (–) spruce 250 m (+)
GRAY JAYS
Local
250 m water (+) water (+) water (+)
500 m water (+) water (+)
1000 m water (+) water (+)
Global pine 1000 m (+), alder and willow (+) water 500 m (+), spruce 500 m (–), N/A
fir trees (–)
N/A: non applicable; number of variables exceeded the number of sampling points we could use for this analysis.

1997 scale) was significant in the analysis of both years. For gray
Small falcon jay, only one landscape variable, i.e., the area of water, was
5m significant. However, the global models revealed that pine
0m cover (1000 m) and local abundance of alder and willow
Weasel
positively influenced predation by gray jays in 1997, where-
as spruce cover (500 m) and fir trees negatively influenced
Snowshoe hare predation in 1998.
Small mammal OBSERVATIONS OF PREDATORS
Nest predators were abundant in our study area.
Squirrel During point counts, we detected the presence of nest
predators in 94% (46/49) of the sampling points in 1997 and
Gray jay in 88% (45/51) in 1998 (the same sites where artificial nests
had been placed). Predators most often seen were gray jays
0 20 40 60 80 100 and red squirrels (Table III). Ravens and American crows
were rare. We did not observe any northern flying squirrels
1998 or eastern chipmunks but we know they were present in low
Small falcon numbers in our study area (Côté, 1999).
We generated graphs presenting sites where we
Weasel observed red squirrels and gray jays and where we found
plasticine eggs showing marks of these predators (Figure 3).
Snowshoe hare In general, predation events occurred at sites where squir-
rels or jays were observed, but the observation of a predator
did not necessarily imply that nests would be preyed upon.
Small mammal
We found nearly significant associations between the
observation of squirrels and predation by squirrels in 1997
Squirrel (G = 2.7, P = 0.10) and in 1998 (G = 3.1, P = 0.08). We did
not find an association between observation of gray jays and
Gray jay predation by gray jays in 1997 (G = 0.4, P = 0.60), but this
association was significant in 1998 (G = 4.8, P = 0.03). The
0 20 40 60 80 100 abundance of red squirrels was not correlated with the
Number of eggs preyed upon number of nests depredated by this predator (P > 0.05 in
FIGURE 2. Frequency distribution of predators that left marks on plas- 1997-1998). Abundance of gray jays correlated significantly
ticine eggs placed in artificial nests in relation to nest heights in a black with the number of nests depredated by this predator in
spruce forest north of Lake Saint-Jean, Québec. 1998 (r = 0.27; P = 0.05).
285
 BOULET, DARVEAU & BÉLANGER: BIRD NEST PREDATION

TABLE III. Number of predators observed during point counts predators were not precise, e.g., artificial nests differ from
(three visits of 15 minutes) in 49 sampling points (over 50) in natural nests (Whelan et al., 1994, Wilson, Brittingham &
1997 and 51 sampling points (over 51) in 1998 where artificial Goodrich, 1998). However, another explanation for such
nests were placed, in a black spruce forest north of Lake Saint-
Jean, Québec. Values are maximum number of individuals seen or
results is that predators are more or less opportunistic, e.g.,
heard in a 50 m radius for squirrels and in an unlimited radius for they find nests simply by chance, and do not rely exclu-
birds. Number of sampling points where predators were observed sively on eggs and nestlings for their survival. Eggs and
are in parenthesis. nestlings are of minor importance in the diet of squirrels
and their major food sources consist of seeds of coniferous
Species 1997 (49) 1998 (51) and deciduous trees and fungi (Rush & Reeder, 1978;
BIRDS Steele, 1998). In contrast, eggs and nestlings are assumed to
Gray jay 40 (20) 55 (30 points) represent an important component in the diet of gray jays
American crow 6 (5) 1 (1) (Ouellet, 1970; Strickland & Ouellet, 1993). Our results
Common raven 7 (7) 11 (11) agree with this assumption, since gray jays attacked as
MAMMALS
Red squirrel 71 (42) 50 (36)
many nests as squirrels (39 for jays versus 41 for squirrels
in 1997; 60 versus 49 in 1998), but were at least half as
abundant as squirrels considering the same area of detection
Discussion (18 jays versus 71 squirrels in 1997; 23 versus 50 in 1998
within a radius of 50 m). We suggest that gray jays could be
WHO PREYS ON NESTS? greater “egg-specialists” than squirrels. In a western conif-
Red squirrels were probably responsible for most erous forest, Craig (1997) suggested that corvids (Steller’s
attacks attributed to small sciurids, because they accounted jays [Cyanocitta stelleri Gmelin], common raven, American
for 90% of captures of small sciurids in a concurrent study crow, black-billed magpie [Pica pica L.]) have nest-oriented
in the same study area in spring 1997 and 1998 (Côté, search images, whereas rodents (red squirrels, chipmunks)
1999). Our tests of association of predator presence with discover nests at random. As an alternative to this hypothesis,
predation events revealed that the observation of a predator Schmidt (1999) recently suggested that avian predators
did not imply that nests of birds would be attacked and vice (sight searchers) have higher foraging efficiencies than
versa. One could argue that the methods used to sample mam-malian predators (olfaction searchers). Temporal vari-
ations in distribution and abundance of alternative food
25 Squirrel 1997 can also modify the response of predators toward nest
1998 abundance (Schmidt & Whelan, 1999).
20 Gray jays and red squirrels directed their attacks
toward different nest heights. Gray jays preyed more on
Number of points

arboreal nests and red squirrels showed a slight preference

15 for ground nests in 1997 in this study, as well as in another
study conducted within the same area in 1997 and 1998
10 (Boulet, 1999). Gray jays are known to visually detect nests
by scanning surroundings from perch to perch (Strickland &
Ouellet, 1995), whereas squirrels probably detect nests
5 olfactively. Arboreal and ground-nesters may therefore have
developed different adaptations to reduce depredation by
0 these predators. Ouellet (1970) reported two cases where
songbirds (Lincoln’s sparrows [Melospiza lincolnii
25 Gray jay Audubon], fox sparrows [Passerella iliaca Merrem]) had
successfully chased away an adult gray jay and seven cases
20 where songbirds (American robin [Turdus migratorius L.],
Number of points

Swainson’s thrush, hermit thrush [Catharus guttatus

Pallas], magnolia warbler [Dendroica magnolia Wilson],
15
white-crowned sparrow [Zonotrichia leucophrys Forster]
and white-troated sparrow [Zonotrichia albicollis Gmelin])
10 had been unable to defend their nest against single or groups
of gray jays. Increased nest concealment and aggressive
5 defence might be a good strategy to reduce vulnerability of
nests. Most songbird species in our study area are probably
unable to defend their nests against red squirrels, as these
0 mammals are able to kill adult songbirds (Sullivan, 1991;
no-ni o-i no-i o-ni
Steele, 1998). Sieving & Willson (1998) suggested that
Observation versus identification small nest size reduces nest discovery and thus may help
FIGURE 3. Number of sampling points where gray jays and squirrels small-bodied ground-nesters like the dark-eyed junco (Junco
were observed and/or marks identified on plasticine eggs placed in artifi- hyemalis L.) and winter wren (Troglodytes troglodytes L.),
cial nests. Sampling points included are those where we sampled predators
with both methods (n = 49 in 1997 and n = 51 in 1998). No-ni refers to cases
that nest successfully in coniferous understory. In our area,
where predators were not observed and not identified; o-i, observed and iden- hermit thrushes tended to nest on the ground in moss clumps
tified; no-i, not observed but identified; o-ni, observed but not identified. surrounded by dense sheep laurels (M. Boulet, pers. observ.).
286

CAN WE PREDICT PREDATION?
Local vegetation was not a strong predictor of pre-
dation events when we considered predation events by all
predators. Four vegetation variables were retained in the
landscape models (numbers of black spruce trees, snags,
coarse woody debris, and alder and willow shrubs) and two
were retained in the global models (numbers of black spruce
trees and jack pine trees). None of these variables were con-
sistent between years. In their study conducted in a boreal
mixed-wood forest in Alberta, Song & Hannon (1999)
reported that vegetation features were not good predictors
of predation and that the relationships they found were
inconsistent between years. These authors emphasized the
importance of long-term studies that can separate true
effects from stochastic effects. We believe that because
predation was caused by several species of predators that
have different foraging strategies and habitat preferences,
nest predation was difficult to predict locally. The same argu-
ment may also account for the fact that land cover poorly
explained predation when we did not discriminate predators.
Global models suggest that local abundance of spruce trees
increased predation, whereas abundance of pines at local
and landscape (500 m) scales decreased predation.
CAN WE PREDICT ATTACKS BY SPECIFIC PREDATORS?
Here again, local vegetation variables were not
consistent predictors of predation events caused by squirrels
or jays. Squirrels responded mostly to local vegetation
variables in 1997, but to landscape variables in 1998. We
conclude that animals may respond to both local and land-
scape scales, but their degree of “specialization” toward one
or the other scale could vary considerably. This degree of
specialization may vary according to disturbances occurring
in their habitat (e.g., clearcuts). Studies considering both
scales are rare and focus more on habitat selection as a
whole than on foraging habitat (but note that the latter is
often included in the previous).
In 1998, habitat features at the landscape level
better explained nest predation by squirrels than did local
vegetation features. We found an association between
spruce cover at the 250 m (19.6 ha) and 500 m (78.5 ha)
scales and predation events by squirrels. Squirrels there-
fore responded to landscape variables at a scale exceeding
their territory area (territory size: 0.24 to 0.98 ha). This
association between spruce and squirrel is not surprising
considering that preferred habitat of red squirrel is conifer-
ous forest (Rush & Reeder, 1978; Steele, 1998). In a boreal
mixed-wood forest, Song & Hannon (1999) found that nest
predation (by different predators including red squirrel)
tended to increase as the amount of conifer cover increased
(450 m radius), but this relationship was only significant in
one of two years. In western U.S.A., Tewksbury, Hejl &
Martin (1998) observed that bird nest predation and abun-
dance of red squirrel were greater in landscapes dominated
by coniferous forests than in landscapes dominated by
agricultural lands where the remaining forest cover was
deciduous trees.
We also found positive associations between gray
jay, water body area, and local abundance in alder and
willow, possibly because the diet of gray jay also includes
insects (Strickland & Ouellet, 1993). We hypothezise that
ÉCOSCIENCE, VOL. 7 (3), 2000
gray jay may forage along lakeshores and in moist patches
because these habitats may offer high densities of insects.
This riparian effect has been found in other studies, e.g.,
lakeshores had greater biomass of insects than roadsides for
the tree swallow (Tachycineta bicolor Vieillot; Dunn &
Hannon, 1992), while flooded habitats had higher arthropod
abundance than dry habitats for the prothonotary warbler
(Protonotaria citrea Boddaert; Petit & Petit, 1996). Predation
by gray jay may also be associated with a feature we have
not identified that is correlated with water body area and
moist patches.
In global models, predation by gray jay correlated
positively with jack pine cover (1997) and negatively with
black spruce cover (1998). We suspect that gray jay pre-
ferred foraging in jack pine stands compared to black spruce
stands because: (i) considering the shape of the tree, nest
concealment could be lower in jack pine trees; and (ii) jack
pine stands were more open (M. Darveau, unpubl. data).
DOES CLEARCUTTING AFFECT NEST PREDATORS ?
We found a positive association between clearcuts
and squirrels. Although this result is equivocal, we know
that as clearcuts increased in our study area, spruce stands
decreased. We hypothezise that squirrels concentrated in the
remaining spruce stands, which may have increased the
probability they would find nests. In Alaska, red squirrels
are known to abandon recent clearcuts and relocate into
residual forests (Wolff & Zasada, 1975). In a managed
forest of New Hampshire, King, Griffin & DeGraaf (1998)
also reported increases of red squirrel near clearcut borders.
They hypothezised that forest clearcut edges may represent
barriers to dispersal (Bider, 1968; Oxley, Fenton & Carmody,
1974). A concurrent study comparing densities of squirrel in
fragments of forest in the same study area as us revealed
that squirrel inhabit wide riparian buffer strips as well as
non-riparian leave strips, but not clearcuts (Côté, 1999). In
the case of gray jays, we do not have evidence that clearcuts
affected them, at the current level of fragmentation. Jays
were observed at most sampling points and seemed to be
active predators in all sectors of the study area. Their area
requirements are considerable (between 41 to 146 ha;
Strickland & Ouellet, 1993), but these birds can tolerate
clearcuts in their territory; they compensate for the presence
of clearcuts by increasing their home range (J. Ibarzabal,
pers. comm.).
We offer two conclusions to this study. On the one
hand, the predation pressure was high in our study area. It is
unlikely that clearcutting could lead to detrimental edge
effects resulting from generalist predators like crows, that
remained nearly absent and did not prey upon our nests. On
the other hand, we found associations between red squirrels,
clearcut cover, and spruce cover, suggesting that there
might be subtle changes in communities of predators, e.g., a
crowding effect of squirrels in residual forest. Such changes
would not be surprising, because the preferred habitat of two
predators (red squirrel and gray jay) is disappearing. We
believe that more studies on nest predation and landscape
composition are needed to help understand the fragmenta-
tion effect. Moreover, distinguishing how different types of
land cover influence predation may provide important
insights on how predators use landscapes.
287
BOULET, DARVEAU & BÉLANGER: BIRD NEST PREDATION
Acknowledgements
We are greatly indebted to D. Lavoie, N. Bergeron, C.
Fournier, I. Gilbert, and I. Chouinard for field work and to V.
Delage for GIS and FRAGSTATS work. We are grateful to N.
Bertrand, F. Potvin, C. Paquet for logistic assistance and to M.
Côté for sharing his results. We also thank A. Desrochers, E. M.
Bayne, and W. A. Montevecchi for constructive comments on ear-
lier drafts of the manuscript. C. Barrette and G. Rochette gave us
access to bird and mammal collections. Funding was provided by
the Sustainable Forest Management Network (National Sciences
and Engineering Research Council of Canada [NSERC]), the
Ministère des Ressources naturelles du Québec (via the Programme
de mise en valeur du milieu forestier - Volet 1), and the Province
of Québec Society for the Protection of Birds, in collaboration
with Faune et Parcs Québec and Produits Forestiers Donohue. M.
Boulet benefited from a NSERC postgraduate scholarship.
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