Allen Press

Riparian Forest Strips as Habitat for Breeding Birds in Boreal Forest

Author(s): Marcel Darveau, Patrick Beauchesne, Louis Bélanger, Jean Huot and Pierre Larue
Source: The Journal of Wildlife Management, Vol. 59, No. 1 (Jan., 1995), pp. 67-78
Published by: Wiley on behalf of the Wildlife Society
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 J. Wildl. Manage. 59(1):1995 BIRD RESPONSETO FIRE * Wilson et al. 67

NEAL,J. C., ANDW. G. MONTAGUE. 1991. Past and
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NUTTALL,T. 1980. A journal of travels into the
Arkansas Territory during the year 1819. Univ.
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ROBBINS,C. S., J. R. SAUER,R. S. GREENBERG,AND
S. DROEGE. 1989. Population declines in North
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STAUFFER,D. F., AND L. B. BEST. 1980. Habitat
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8:1-14.
WALDROP, T. A., D. L. WHITE, AND S. M. JONES.
1992. Fire regimes for pine-grassland commu-
nities in the southeastern United States. For. Ecol.
and Manage. 47:195-210.
WEBB, J. S., AND D. K. WETHERBEE. 1960. South-
eastern breeding range of the brown-headed
cowbird. Bird-Banding 31:83-87.
WILSON, C. W. 1994. Breeding bird community
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Received 14 December 1993.
Accepted 17 August 1994.
Associate Editor: Ryan.

RIPARIANFORESTSTRIPSAS HABITATFOR
BREEDINGBIRDSIN BOREALFOREST
MARCELDARVEAU,D6partementde Biologie,Universit6Laval,Ste-Foy, PQ G1K7P4, Canada
PATRICKBEAUCHESNE,' D4partementdes Sciences du Bois et de la Foret,UniversiteLaval,Ste-Foy, PQ G1K7P4, Canada
LOUISBELANGER,D6partementdes Sciences du Bois et de la Fordt,Universit6Laval,Ste-Foy, PQ G1K 7P4, Canada
JEANHUOT,Departementde Biologie,Universit6Laval,Ste-Foy, PQ G1K 7P4, Canada
PIERRELARUE,2
D'partementdes Sciences du Bois et de la Foret,UniversiteLaval,Ste-Foy, PQ G1K7P4, Canada

Abstract: Riparian forest strips are usually protected from logging for their buffer effect on aquatic habitats.
However, their value to terrestrial wildlife is unknown. From 1989 to 1992, we compared bird abundance
and species composition in 5 experimental riparian forest strips (20-m, 40-m, 60-m, and control [>300 m
wide], intact strips, and 20-m-wide thinned strips), in boreal balsam fir (Abies balsamea) stands, for 3 years
following clear-cutting. Bird densities increased 30-70% (P < 0.05) in all strips the year after cutting and
decreased (P > 0.05) thereafter to approximately pretreatment levels. The 20- and 40-m-wide riparian strips
had highest mean bird densities, but also the fastest (P < 0.05) decreases thereafter. By the third year after
clear-cutting, forest-dwelling species were less (P = 0.01) abundant than ubiquitous species in the 20-m strips.
The golden-crowned kinglet (Regulus satrapa), Swainson's thrush (Catharus ustulatus), blackpoll warbler
(Dendroica striata), and black-throated green warbler (D. virens) became nearly absent in 20-m strips. The
removal of 33% of the trees in some 20-m strips resulted in a <20% decline of bird densities, a moderate
effect that combined with the greater effect of strip narrowness. There was evidence that 60-m-wide strips
are required for forest-dwelling birds. Bird populations may continue to decline in strips before regeneration
of adjacent clear-cuts provides suitable habitat for forest-dwelling species.
J. WILDL. MANAGE.59(1):67-78
Key words: boreal, breeding birds, buffer strip, forests, fragmentation,logging, Neotropics, Quebec, ri-
parian.

Ecologists have highlighted the ecotone prop- to conciliate conflicting land-use demands, man-
erties of riparian systems and promoted conser- agers have begun to consider riparian habitats
vation of these areas (Odum 1971). In an effort in land-use planning (Swanson and Franklin

I
Present address: Ministere de l'Environnement et
de la Faune du Qu6bec, 2360 chemin Ste-Foy, Ste-
Foy, PQ G1V 4H2, Canada.
2
Present address: Ministere de l'Environnement et
de la Faune du Qu6bec, 150, boulevard Rene L6-
vesque est, Qu6bec, PQ GIR 4Y1, Canada.

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68 BIRDSIN RIPARIANFORESTSTRIPS* Darveau et al.
1992, Naiman et al. 1993). Forest strips, which
are retained adjacent to rivers, streams, and lakes
to protect aquatic habitats (Barton et al. 1985,
Hornbeck et al. 1986, Budd et al. 1987), cor-
respond to priority areas for protection of ter-
restrial species (Cline et al. 1980, Stauffer and
Best 1980, Johnson and Brown 1990, Naiman et
al. 1993). Comparisons of riparian and upland
forests have shown that the former often have
higher mammal species richness and densities
and, moreover, that riparian areas act as pop-
ulation sources for many species (Doyle 1990).
Riparian forests often have higher bird species
richness and densities compared with upland
forests of similar composition and structure
(Stauffer and Best 1980, LaRue 1990, Gates and
Giffen 1991; but see McGarigal and McComb
1992). However, Gates and Giff en (1991) sug-
gested that certain riparian edges were favor-
able to the brown-headed cowbird (Molothrus
ater) and that they could be ecological traps for
forest-dwelling passerines.
We evaluated the effect of width and forest
stand thinning on the ability of riparian strips
to support an avifauna similar to that of large
forest tracts. Our first objective was to compare
bird species composition and densities in 3 widths
of riparian forest strips (20, 40, and 60 m) and
control (>300 m wide) intact strips, for 3 years
after clear-cutting. Our second objective was to
evaluate the effect of thinning 33% of the trees
(>10 cm diam at breast height [DBH]) in nar-
row strips (20 m wide), which is the practice in
Quebec.
We are grateful to field assistants A. Desro-
siers, P. Filiatrault, G. L6vesque, R. McNicoll,
R. Ouellet, S. St-Onge, and C. Savignac. R. J.
Hughes provided editorial assistance. A. Des-
rochers and J. P. L. Savard commented on the
manuscript. This study was funded by the Que-&
bec Ministry of Environment and Wildlife, the
forestry practices component of Forestry Can-
ada's Green Plan, the Quebec Wildlife Foun-
dation, Wildlife Habitat Canada, the Quebec
Ministry of Natural Resources, and the Daish-
owa Company, with contributions from the Ca-
nadian Wildlife Service and from the Faculty
of Forestry and Geomatics, the Department of
Biology, and the Center for Northern Studies of
Laval University.
STUDY AREA
We conducted the study in Montmorency
Forest, in the Laurentian Mountains, 60 km north
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J. Wildl. Manage. 59(1):1995
of Quebec city. Mean annual temperature was
0.3 C, and mean annual rainfall was 142 cm;
33% of this fell as snow, which covers the ground
until mid-May (n = 25 yr; Bdlanger et al. 1991).
The landscape was hilly, with elevations ranging
from 600 to 1,100 m, and had several lakes,
rivers, and streams. Rivers were generally fringed
with speckled alder (Alnus rugosa). Balsam fir
dominated mature forest riparian stands, ac-
companied by black spruce (Picea mariana),
white birch (Betula papyrifera), and white
spruce (P. glauca; Beauchesne 1991). From 1974
to 1986, the eastern spruce budworm (Choris-
toneura fumiferana) killed 15% of the trees in
the area (Bdlanger et al. 1991, Dupont et al.
1991). Clear-cut sites were colonized by com-
mon raspberry (Rubus idaeus), balsam fir, and
white birch, and locally by graminoids.
METHODS
In 1988, we delineated 25 400-m long ripar-
ian segments that (1) bordered rivers 5-15 m
wide, (2) had a slope <30%, (3) included an
alder fringe 1-7 m wide, (4) held fir stands >50
years old, >15 m high, and had <25% dead
trees (Beauchesne 1991). Our experimental de-
sign consisted of 5 randomized blocks with 4
types of treated strips and a control strip. Be-
cause of operational constraints such as steep
slopes or the absence of roads or bridges to access
sites, we placed 4 of 5 control strips on 1 side
of rivers and treated strips on the other (Beau-
chesne 1991). We pooled the remaining seg-
ments and distributed treated strips at random
within the pool. Later, we performed a cluster
analysis on site, physiognomic, and floristic char-
acteristics of the segments to define blocks on
an ecological basis (Sokal and Rohlf 1981:349).
The 25 segments, <6 km apart from each other,
showed few ecological differences (Beauchesne
1991). We avoided bias related to harvesting
operations by applying the same logging meth-
ods to all treated strips: clear-cutting was done
by harvesting conifers and felling deciduous
trees, snags > 1 m high, and shrubs > 2 m high,
without scarifying the soil. Thinning consisted
of an even harvest of 33% of the largest trees
within strips, using light equipment and reduc-
ing the basal area of strips by 50%. We harvested
the first 2 experimental blocks in fall 1988 and
the other 3 after the 1989 nesting season. Hence,
blocks 1-2 and 3-5 were 1 year old in 1989 and
1990, respectively.
We counted birds by territory mapping (Rob-
J. Wildl. Manage. 59(1):1995 BIRDS IN RIPARIANFORESTSTRIPS* Darveau et al. 69

CFA

80m0 m
t ZZ Z
-- 411--117777771
River 11111- ---

20-m forest strip 40-m forest strip 60-m forest strip Control
\[
-- 400 m --+
H
Uncut forest
II Harvested area

Fig. 1. Schematicview of harvestedareas and forest stripsof differentwidthswithinthe 80-m-wideriparianzone in the boreal
forest, Quebec, 1989-92. The strips, randomlydistributedin the field, are orderedby size in the figureto improveclarity.The
constant forest area (CFA)is the first 20 m adjacentto the river.Riverwidthnot drawnto scale.

bins 1970) in 80- x 350-m strips centered lon-
gitudinally within the 400-m segments. This
method was the most appropriate to small and
elongated strips, especially to record the exact
position of birds in a mosaic of forest and clear-
cut habitats (Bibby et al. 1992:43, Ralph et al.
1993:35). In 1988, prior to harvesting, we sam-
pled 10 of 15 strips in blocks 1-3. In 1989, we
counted birds in the 10 strips of blocks 1-2, and
thereafter we surveyed all strips each year. We
visited strips 8 times/year in June, between 0500
and 1000, during clear weather with little or no
wind. In 1 hour, an observer, using a system-
atically located 20- x 25-m grid, mapped po-
sitions of all birds heard or seen. We rotated
order of strip visits, entrance points on grids,
and observers (4-5/yr, 10 total). Two observers
later independently delimited territories, and,
we used the averages to calculate bird species
richness and the number of territories overlap-
ping the riparian zone (absolute no. of territories
could not be obtained by sampling small and
elongated plots of 80 x 350 m = 2.8 ha). We
also calculated species richness and number of
territories for forest strips and harvested areas
separately to determine the contribution of each
to bird populations in the riparian zone (Fig. 1).
Because the latter indices were subjected to hab-
itat area effects (i.e., they did not control for
levels of sampling intensity in forest strips of
different sizes; Haila 1986), we also calculated
indices for a constant-forest-area plot corre-
sponding to the largest forest area common to
all treatments (the first 20 m adjacent to the
river; hereafter referred to as constant forest
area).
We defined 3 bird guilds on the basis of use
of wooded and clear-cut areas by species ob-
served in strips over the 3 years of study and in
concurrent studies (E. Melanqon and J. F. Rail,
Universit6 Laval, Ste-Foy, Que., unpubl. data):
(1) forest-dwelling species (i.e., those for which
- 80% of all territories were in uncut forests >50
yr old), (2) open-habitat species with a corre-
sponding of all territories in clear-cuts,
-80%
(3) ubiquitous species including all other species
such as forest-edge species and species occurring
in all successional stages (Table 1).
Using repeated measures analysis of variance
(RM-ANOVA), we compared bird count data
with analysis of estimated coefficients, which
focuses upon the response curves of each ex-
perimental plot by treating estimated coeffi-
cients of the curves as primary data (Meredith
and Stehman 1991). We tested first- and second-
degree polynomial models in all analyses, but
the latter were never significant (P > 0.05), and
we present only first-degree models in this pa-
per. Similarly, we did not consider the random
blocks' effects in RM-ANOVAs, because prelim-
inary analyses for randomized blocks yielded no
effects (P > 0.05). Because bird count data col-
lected prior to cutting were incomplete, we
omitted them from RM-ANOVAs, and they are
shown as reference on figures or compared with
control strips of subsequent years, using a Krus-
kal-Wallis test. When the treatment effect was
significant (P < 0.05), we made multiple com-
parisons among means using Steel-Dwass's test
(Day and Quinn 1989).
RESULTS
We identified 52 bird species, including 25
observed in >2 strips each year (Table 1). We
recorded a yearly mean of 15.1 ? 4.6 territories
(?SD; pooled among years) belonging to 7.5 ?
2.2 species in the 25 strips. No species was ob-
served in all strips, but 7 species had a yearly

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Table 1. Numberof territoriesof each birdspecies in the 80-m-wideriparianzone in the 3 years followingthe establishmentof 5 types of r
1989-92.

Treatment
20 m Tb 20 m 40 m
Habitatguilda
Species 1C 2 3 1 2 3 1 2 3

Forest-dwelling species
Spruce grouse (Dendragapus canadensis) d
Ruffed grouse (Bonasa umbellus) + + + + +
Black-backed woodpecker (Picoides arcticus) + + +
Northern flicker (Colaptes auratus)
Yellow-bellied flycatcher (Empidonax flaviventris) + + +
Alder flycatcher (Empidonax alnorum)
Gray jay (Perisoreus canadensis) 1 1 + +
Black-capped chickadee (Parus atricapillus) +
Boreal chickadee (Parus hudsonicus) + +
Red-breasted nuthatch (Sitta canadensis) + + +
Brown creeper (Certhia americana) + +
Winter wren (Troglodytes troglodytes) 5 3 1 2 + 1 5 2 2
Golden-crowned kinglet 5 5 2 4 6 4 8 8 9
Ruby-crowned kinglet (Regulus calendula) 1 + 1 1 2 4 1 2
Bicknell's thrush (Catharus bicknelli) 1
Swainson's thrush 4 5 1 8 5 1 9 7 4
Solitary vireo (Vireo solitarius) + 1 1 3 + +
Philadelphia vireo (Vireo philadelphicus) 1
Tennessee warbler (Vermivora peregrina) +
Cape May warbler (Dendroica tigrina) + 1 1 +
Black-throated green warbler 3 + 1 4 1 1 4 3
Blackburnian warbler (Dendroica fusca) + + 1
Bay-breasted warbler (Dendroica castanea) 5 3 1 9 4 1 11 8 1
Blackpoll warbler 11 9 6 15 11 8 19 11 4
Black-and-white warbler (Mniotilta varia)
American redstart (Setophaga ruticilla) + 3 3 1 1 1 + +
Northern waterthrush (Seiurus noveboracensis) + + + 1 + 1
Wilson's warbler (Wilsonia pusilla) 2 3 + 2 3 2 + +
Canada warbler (Wilsonia canadensis) + + +
Pine grosbeak (Pinicola enucleator)
White-winged crossbill (Loxia leucoptera) +
Pine siskin (Carduelis pinus) + + 1 1 +
Subtotal 37 32 16 46 35 21 64 42 27

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Table 1. Continued.

Treatment
20 m Tb 20 m 40 m
Habitat guilda
HaSpecies 2 3 2 1 3 1 2 3

Ubiquitous species
Tree swallow (Tachycineta bicolor) + + + 1 1
American robin (Turdus migratorius) 5 4 4 3 3 1 2 4 2
Cedar waxwing (Bombycilla cedrorum) + +
Nashville warbler (Vermivora ruficapilla) + 1 + + + 1 1
Magnolia warbler (Dendroica magnolia) 3 4 2 + 2 2 +
Yellow-rumped warbler (Dendroica coronata) 5 9 6 11 8 8 11 9 8
Chipping sparrow (Spizella passerina) + +
Fox sparrow (Passerella iliaca) +
White-throated sparrow (Zonotrichia albicollis) 8 10 7 8 8 9 10 9 8
Dark-eyed junco (Junco hyemalis) 10 9 9 9 10 6 9 7 5
Rusty blackbird (Euphagus carolinus) +
Purple finch (Carpodacus purpureus) + + 1 2 2 2 +
Subtotal 31 37 26 34 31 24 37 35 24
Open-habitat species
Killdeer (Charadriusvociferus) +
Spotted sandpiper (Actitis macularia) 1 +
American crow (Corvus brachyrhynchos)
Eastern bluebird (Sialia sialis) +
American pipit (Anthus rubescens) +
Mourning warbler (Oporornis philadelphia) + + + +
Common yellowthroat (Geothlypis trichas) + + +
Lincoln's sparrow (Melospiza lincolnii) 1 2 2 + + 2 1 + 2
Subtotal 1 3 2 2 1 2
Total 69 72 44 70 66 47 102 77 53 1

aForest-dwelling and open-habitat species had -80% of all territories in forest strips and adjacent harvested areas, respectively.
b 20 m T = 20-m-wide forest strip thinned of 33% of trees, 20 m = 20 m unthinned, 40 m = 40 m unthinned, 60 m = 60 m unthinned, and control = >300-m-wi
c The no. of years since creation of forest strips by harvesting trees in adjacent areas.
d Values are the sum of 5 replicates for each treatment each year. "Plus" signs indicate the presence of birds observed only once (nonterritory holders).

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72 BIRDSIN RIPARIANFORESTSTRIPS* Darveau et al. J. Wildl. Manage. 59(1):1995

30 20
Forest-dwelling
Riparianzone
16 c
I61 I

11o j111 l 2
10 2 I
6i

0 45
o
)
0o I
12

2 Ubiquitous
-I I
4I" I!IIi
6
S11
I! I

zO
2.5

Constantforest area Open-habitat

2-
0 1 2 3

Yearsaftercutting 1.5

Fig.2 Totalnumberof birdterritoriesinthe 80-m-wideriparian

zone and the constant-forest-areaplots (first20-m adjacent
to the river)in the 3 years followingthe establishmentof 5
types of riparianforest strips in borealforest, Quebec, 1989-
92. Meanvalues and ranges (n = 5 replicates)are shown for
the 5 striptypes: T = 20-m-widestripthinnedof 33%of trees,
I 2
2 = 20 m unthinned,4 = 40 m unthinned,6 = 60 m unthinned,
and C = >300-m-widecontrolstrip.The asteriskshows mean * I2,Ic
I 24Ic iIc T1
values for 10 strips in the year before cutting. 0 1 2 3

mean number of pairs > 1/strip: blackpoll war-
bler (2.3 pairs/strip), yellow-rumped warbler
(1.9), golden-crowned kinglet (1.8), dark-eyed
junco (1.4), white-throated sparrow (1.4), bay-
breasted warbler (1.2), and Swainson's thrush
(1.2). These species represented 74% of all pairs
observed.
Compared with the 10 strips sampled prior
to cutting, species richness of control strips did
not differ in any year (P > 0.18). However, the
number of pairs increased by 50% in control
strips in the first year after harvesting, remained
greater (P < 0.05) than preharvest levels for 2
years, and returned to preharvest levels in the
third year (P > 0.05).
Yearsaftercutting
Fig. 3. Numberof territoriesof forest-dwelling,ubiquitous,
and open-habitatbirdspecies in the 80-m-wideriparianzone
in the 3 years followingthe establishmentof 5 types of riparian
forest strips in borealforest, Quebec, 1989-92. Mean values
and ranges (n = 5 replicates)are shown for the 5 striptypes:
T = 20-m-widestrip thinnedof 33% of trees, 2 = 20 m un-
thinned,4 = 40 m unthinned,6 = 60 m unthinned,and C =
>300-m-wide controlstrip. The asterisk shows mean values
for 10 strips in the year before cutting.
Treatments did not affect species richness in
the riparian zone (Table 2). However, total
numbers of territories and species composition
differed among treatments in the riparian zone
(Table 2 and Fig. 2). Forest-dwelling species

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Table2. Estimatedmeans and standarderrors(SE) for the treatmentresponse averaged over time (z) and the linearcoefficientof the resp
types of riparianforest strips in borealforest, Qu6bec, 1989-92.

S(z) Lin
Variable 20Ta 20 40 60 Co SEb pc 20T 20 40

Species richness (no. of species/strip)

Riparian zoned 13.9 12.3 13.9 12.9 14.3 0.91 0.51 -1.7 -2.2e -2.2e
Forest-dwelling speciesf 7.3 7.1 8.3 7.9 9.9 0.57 0.03 -1.9e -1.8e -2.0e
Ubiquitous species 5.3 4.5 5.1 4.5 4.5 0.38 0.33 -0.4 -0.8e -0.5
Forest stripd 11.3 10.2 11.7 11.5 14.3 0.83 0.03 -2.6e -3.0O -3.1e
Harvested aread 7.0 5.5 6.1 3.8 0.56 0.007 0.3 -1.2 -0.7
Constant-forest-area plotg 10.9 10.1 8.6 6.4 8.7 0.69 0.002 -2.4e -2.9e -2.3e
Forst-dwelling species 6.8 6.7 6.1 4.7 5.9 0.45 0.03 -1.9e -1.8e -1.2
Ubiquitous species 4.1 3.3 2.5 1.7 2.7 0.36 0.002 -0.6e -1.1e -1.0
Bird density (no. of territories/strip)
Riparian zone 12.9 12.9 15.5 17.1 17.2 1.03 0.01 -3.2e -3.4e -5.2e
Forest-dwelling species 5.8 6.9 9.0 11.2 13.2 0.80 0.001 -2.3e -2.6e -3.9e
Ubiquitous species 6.6 5.9 6.3 5.6 4.0 0.54 0.02 -1.0e -1.0M -1.4e
Forest strip 8.0 9.7 11.5 15.1 17.2 0.84 0.001 -2.2e -2.5e -4.5e
Harvested area 4.9 3.2 4.0 1.9 0.71 0.06 -1.0 -0.9 -0.7
Constant forest area 8.0 9.5 7.3 5.4 6.0 0.62 0.001 -2.2e -2.3e -2.9e
Forest dwelling species 5.5 6.7 6.5 4.7 4.9 0.58 0.08 -2.2e -2.5e -2.5e
Ubiquitous species 2.5 2.7 0.9 0.7 1.1 0.49 0.02 0.0 0.2 -0.4
a 20T = 20-m-wide strip thinned of 33% of trees, 20 = 20 m unthinned, 40 = 40 m unthinned, 60 = 60 m unthinned, and Co = control, >300-m-wide strip.
b SEs of estimated means from the repeated measures
analysis of variance (RM-ANOVA) of 3 years' data.
c Probability of significance of the main effect (z) and of the linear time x treatment interaction (B) from the RM-ANOVA. n = 5 replicates of 5 treatments.
d The riparian zone was an 80-m-wide area adjacent to the river and included a forest strip and a harvested area of different widths depending on treatments.
e The estimated B-value is different from zero (P > 0.05).
f Forest-dwelling and open-habitat species had of all territories in forest strips and adjacent harvested areas, respectively.
g Constant-forest-area plot controlling for habitat-80%
area effect (the first 20 m adjacent to the river).

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74 BIRDSIN RIPARIAN STRIPS* Darveau et al.
FOREST J. Wildl. Manage.59(1):1995

15
Forest-dwelling
-10
- By the third year, forest-dwelling species were
O Ubiquitous
7.5
0 were negatively affected (P < 0.05; Table 3).
5-
2.5 I2 1 tent increasing trends from control to 20-m strips.
i
1 111 I[
1 41.
0 1 2 3
0?I
Yearsaftercutting
Fig.4. Numberof territoriesof forest-dwelling and ubiquitous green
birdspecies in the constant-forest-areaplots (first20 m ad-
jacent to the river)in the 3 years followingthe establishment est for 20-m strips (5.0 ? 0.6) and lowest for
of 5 types of riparianforest strips in borealforest, Quebec,
1989-92. Meanvaluesand ranges(n 5 replicates)are shown 60-m strips (1.8 + 0.6; P = 0.004). There were
for the 5 striptypes: T = 20-m-wide-stripthinnedof 33% of
trees, 2 = 20 m unthinned, 4 = 40 m unthinned, 6 = 60 m
unthinned,and C = >300-m-widecontrolstrip.The asterisk
shows mean values for 10 strips in the year beforecutting.
richness decreased from 9.9 in control strips to
7.1 in 20-m strips, whereas the number of ter-
ritories varied from 13.2 in control strips to 5.8
in 20-m-thinned strips (P < 0.001; Fig. 3). Con-
versely, ubiquitous species increased in number
of territories from 4.0 in control strips to 6.6 in
20-m-thinned strips (P = 0.02). Thus, forest-
dwelling species, which held, on a 3-year av-
erage, 13 of 17 territories in control strips, de-
clined to 6 of 13 in 20-m-thinned strips (Table
2). Open-habitat species accounted for ?1 ter-
ritory in all strips (Fig. 3).
Comparisons for constant-forest-area plots
suggested an opposite trend as species richness
(P = 0.002) and number of territories (P = 0.001;
Fig. 2) decreased from 20-m-thinned to control
strips. Forest-dwelling species held, on a 3-year
average, 5 of 6 of the territories in constant-
forest-area plots of the control compared with
7 of 10 in the 20-m-thinned strips.
We found an overall decreasing trend in bird
populations over 3 years (Fig. 2; P < 0.05, Table
2). In the constant-forest-area plots, there were
declines in 20- and 40-m-wide strips (P < 0.05),
but not in 60-m and control strips. Declines in
20- and 40-m strips reflected a decrease in the
number of territories of forest-dwelling birds (P
< 0.05), whereas those of ubiquitous species
remained stable (P > 0.05; Fig. 4).
less abundant than ubiquitous species in 20-m
strips (P = 0.01; Fig. 3). The golden-crowned
kinglet, blackpoll warbler, and Swainson's thrush
The black-throated green warbler was absent
from 20-m-thinned strips. Conversely, the Lin-
coln's sparrow, white-throated sparrow, Amer-
ican robin, and dark-eyed junco showed consis-
On 240 occasions in 3 years, observers noted
birds crossing rivers during counts (? = 3.2
cross/8 hr). Six species accounted for 55% of all
observations: the blackpoll warbler (17%), yel-
low-rumped warbler (11%), bay-breasted war-
bler (8%), winter wren (7%), black-throated
warbler (6%), and Swainson's thrush (6%).
Mean number of crossings per 8 hours was high-
no temporal trends in crossing data (P = 0.56).
DISCUSSION
PopulationTrends
Avian populations increased in treated con-
trol strips the year after cutting and decreased
the 2 years thereafter. The 1990-91 trends from
Breeding Bird Survey data from central Quebec
and Ontario do not suggest a continental or re-
gional phenomenon: 4 of the 7 most abundant
species in our stands increased over this period
whereas 2 decreased (data missing for the sev-
enth species; Erskine et al. 1992). However, pop-
ulation increases observed in our study area may
have been influenced by disturbances at the lo-
cal scale. Within 4 km of our strips, 21% of 4,100
ha of coniferous forest >50 years old were har-
vested in 1988-89 (compiled from Que. Minist.
For. data). More locally, all strips, except control
strip 8, were located within 100 m of clear-cuts

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J. Wildl. Manage. 59(1):1995 BIRDS IN RIPARIANFORESTSTRIPS * Darveau et al. 75

Table3. Birddensities(no.of territories/strip)

inthe80-m-wide zoneinthethirdyearfollowing
riparian of 5
theestablishment
typesof riparian
foreststripsinborealforest,Qu6bec,1989-92.Onlyspeciescountedin >4 stripsareincluded.
Speciesare
orderedfromthemostassociatedwithcontrolsto themostassociatedwithnarrowstrips.

: no. of territoriesa

Species 20 m Tb 20 m 40 m 60 m Control PC Steel-Dwassd

Golden-crownedkinglet 0.4 0.8 1.8 2.8 3.0 0.001 20 m T, 20 m < 60 m, Control

Blackpollwarbler 1.2 1.6 0.8 1.8 2.6 0.007 20 m T, 40 m < Control
Swainson'sthrush 0.2 0.2 0.8 1.0 1.2 0.04
Black-throatedgreen warbler 0.0 0.2 0.6 0.4 1.0 0.10
Ruby-crownedkinglet 0.4 0.4 0.4 0.6 1.2 0.44
Yellow-rumpedwarbler 1.2 1.6 1.6 2.0 2.2 0.24
Winter wren 0.2 0.2 0.4 1.0 0.4 0.23
American redstart 0.6 0.2 0.0 0.2 0.6 0.31
Dark-eyed junco 1.8 1.2 1.0 1.4 1.0 0.47
Americanrobin 0.8 0.4 0.2 0.0 0.6 0.08
White-throatedsparrow 1.4 1.8 1.6 1.0 0.6 0.19
Lincoln'ssparrow 0.4 0.4 0.4 0.2 0.0 0.65
a f of 5
replicates.
= 20 m unthinned, 40 m = 40 m unthinned, 60 m = 60 m unthinned, and control
b 20 m T = 20-m-wide strip thinned of 33% of trees, 20 m
= >300-m-wide strip.
c Probability level for 1-way Kruskal-Wallis test.
d Multiple comparisons among means (Steel-Dwass test). A "<" sign separates treatments that are different (P < 0.05).

>20 ha. Studies of banded migratory passerines
have shown that individuals, which are philo-
patric (within a distance <40-100 m), may move
53 km when nests were unsuccessful in the
previous year (Blancher and Robertson 1985,
Bollinger and Gavin 1989). Birds whose nesting
habitat had been destroyed may have settled in
the remnant wooded areas nearby without nec-
essarily nesting successfully. These birds may
nevertheless have contributed to a short-term
increase in population densities (Bierregaard and
Lovejoy 1988). Such crowding is not likely to
persist for >2 years after cutting because of
short life-spans of passerines (assuming a 50%
annu. mortality rate; Newton 1989) or move-
ment to other sites.
DifferencesAmongTreatments
Averaged over 3 years, the 20-m-wide forest
strips had the highest indices of bird species
richness and number of pairs. However, they
also had the fastest population declines, and in
the third year, they did not have higher densities
than did other forest strips (Fig. 3). Crowding
of birds in remnant wooded areas may have
been greater in 20-m-wide forest strips, where
more birds (from nearer areas) were forced to
settle in smaller refuges. Similarly, the more
rapid decline in bird populations in narrower
forest strips over the first 3 years after cutting
may be, at least in part, explained by the greater
initial inflation of populations. Since clear-cut-
ting, windfall has killed an average 23% of trees
in forest strips compared with 8% in control
strips (Darveau et al. 1994). Dieback, which of-
ten affects trees located near newly created edg-
es (Perkins et al. 1992), also may have affected
a greater proportion of trees in narrow strips.
Time lags on bird responses to both treatments
and windfall also may complicate results. Such
a phenomenon, which may stem from philo-
patry by breeding individuals, has been docu-
mented for experimental (Wiens et al. 1986) and
natural (Knopf and Sedgwick 1987) distur-
bances.
In addition to a more rapid decline of bird
populations in narrow forest strips, other factors
contribute to reduce the value of narrow strips.
In spite of higher densities, the 20-m-wide strips,
with areas equivalent to 50 and 33% of the 40-
and 60-m-wide strips, respectively, nevertheless
supported fewer species and a lower number of
pairs. Moreover, observed high frequencies of
river crossings suggest that birds spent less time
within narrower strips compared with wider
ones. This may have biased our population in-
dices, leading to an overestimation of species
richness and densities in narrower forest strips.
Some species, such as the golden-crowned king-
let, blackpoll warbler, Swainson's thrush, and
black-throated green warbler, became nearly
absent in narrow strips. We speculate that at
least some pairs of those species frequently ob-
served crossing rivers were likely to have ter-
ritories extending on both sides of the river. Such
fragmented territories could be of lower value

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76 BIRDSIN RIPARIANFORESTSTRIPS* Darveau et al.
for these species because of increased risk of
predation and traveling cost when feeding young
(Zach and Falls 1979).
The removal of 33% of the trees in some 20-m-
wide forest strips, although reducing total basal
area by 50%, did not result in large differences
compared with unthinned strips: species rich-
ness remained at least as high and the number
of pairs declined by <20%. Because our exper-
iment combined both effects of width and thin-
ning, moderate effects of thinning may have
been masked by the greater effects of strip width.
However, studies specifically addressing effects
of thinning in hardwoods (Webb et al. 1977,
Freedman et al. 1981, DellaSala and Rabe 1987)
and coniferous forests (Virkkala 1990, Virkkala
and Liehu 1990) have shown that, even at in-
tensities >50%, thinning does not result in de-
clines of bird species richness and density of the
same amplitude as reducing strip width. Nev-
ertheless, some species are negatively affected
and experience lower nesting success or de-
creased population densities following thinning
(Virkkala 1990).
Edge habitats are recognized as poor nesting
sites for forest-dwelling passerines because of a
higher incidence of brood parasitism and egg
predation (Brittingham and Temple 1983, Wil-
cove 1985, Andren and Angelstam 1988, Yahner
and Scott 1988, Askins et al. 1990). Therefore,
it might be predicted that birds nesting in 20-
60-m-wide strips would experience reduced
nesting success compared with those nesting in
large strips such as our control. However, the
absence of the brown-headed cowbird in our
plots and an observed trend of a lower incidence
of artificial nest predation in the 20-40-m-wide
strips compared with the 60-m and control strips
(Darveau, unpubl. data) suggest these factors
are not of major importance in our area (Rud-
nicky and Hunter 1993).
MANAGEMENT
IMPLICATIONS
In the 3 years following clear-cutting, forest
birds used all remnant forest strips. However,
bird numbers, particularly those of forest dwell-
ing species, increased initially then declined
sharply. Because bird populations may not have
reached their lowest densities in strips by the
third year, longer monitoring will be necessary
to assess the value of forest strips as refuges for
forest-dwelling species. We conclude that 20-
m-wide strips, despite their short-term use by
forest-dwelling species, are more favorable to
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J. Wildl. Manage. 59(1):1995
ubiquitous species than to forest-dwelling ones.
Because our experimental forest strips were sep-
arated from large stands by only a narrow bar-
rier consisting of 15-m-wide river fringed with
alder, we consider that this was a best-case sce-
nario for forest strips and that more isolated,
20-m-wide forest strips would probably shelter
fewer forest-dwelling birds (van Dorp and Op-
dam 1987). The narrow strips may be qualified
as "line corridors" (i.e., narrow bands essentially
dominated by edge species; Forman and God-
ron 1986:131). Conversely, there was evidence
that 60-m-wide strips can support avifauna sim-
ilar to that of a large forest tract. They corre-
spond to "strip corridors" (i.e., wider bands with
a central environment that contains an abun-
dance of interior species; Forman and Godron
1986:142).
From a local perspective, clear-cutting a large
proportion of a wooded area triggers a succes-
sion of events in remnant forest patches. Two
early stages of this succession are a crowding of
birds in the year following clear-cut and a de-
cline in subsequent years. A simple comparison
of bird populations in the year prior to cutting
with the year after would have been misleading
and could have negative effects if management
practices were based on these results alone.
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NONGAMEBIRDUSE OF HABITATIN CENTRALAPPALACHIAN

RIPARIANFORESTS
NORMANL. MURRAY,'Departmentof Fisheriesand WildlifeScience, VirginiaPolytechnicInstituteand State University,
Blacksburg,VA24061-0321, USA
DEANF. STAUFFER,Departmentof Fisheriesand WildlifeScience, VirginiaPolytechnicInstituteand State University,Blacks-
burg,VA24061-0321, USA

Abstract: Most studies that have shown higher bird species richnessand abundancein riparianzones have
been conducted where there is contrast between vegetation in riparian and upland habitats. In central
Appalachianhardwoodforests, substantialriparianhabitat is potentiallysubjectto habitatalterations.How-
ever, little informationis available on the importanceof riparianhabitatsto songbirdswhere the riparian-
upland vegetation gradient is relatively uniform. We sampled relative bird abundanceand habitat charac-
teristics along a gradient from a stream to adjacent upland habitat at 16 sites in southwest Virginia. We
evaluated total relative bird abundance, species richness, and relative abundance of 28 bird species to
determine whether riparianhabitatsinfluencedbird assemblages.Total bird relative abundanceand species
richnessshowed no (P > 0.10) riparianinfluence;however, Acadian flycatchers(Empidonaxvirescens) and
Louisianawaterthrushes(Seiurusmotacilla) were most associated(P < 0.001) with streams.Americanrobins
(Turdus migratorius) showed a weaker but positive association(P = 0.068) with streams. Eastern wood-
pewees (Contopus virens) (P = 0.01), black-and-whitewarblers(Mniotilta varia) (P < 0.001), pine warblers
(Dendroica pinus) (P < 0.001), worm-eatingwarblers(Helmintheros vermivorus)(P < 0.001), and scarlet
tanagers(Piranga olivacea) (P = 0.041) were negatively associatedwith streams.We used cluster analysisto
group the 28 bird species into 5 assemblages on the basis of their distributionsat 69 sampling stations
distributedamong the 16 sites. Species were classifiedas belonging to riparian(n = 2 species), upland forest
(n = 6 species), mesic forest (n = 3 species), xeric upland forest (n = 6 species), and mature hardwoods
generalist(n = 11 species) assemblages.Managementor protectionof riparianzones often is justifiedon the
basis of higher bird species richness and abundance found in these sites. Our data did not support this.
However, Acadian flycatchersand Louisianawaterthrusheswere dependent on riparianhabitats,and such
habitatsshould be protected for these species.
J. WILDL. MANAGE.59(1):78-88

Key words: forests, habitat, neotropicalmigrants,nongame birds, riparian,Virginia.

Riparian areas have been considered unique
habitats, supporting greater wildlife species di-
versity, densities, and productivity than do
neighboring habitats (Brown et al. 1978). This
occurs not only because riparian zones are eco-
' Present address:OklahomaDepartmentof Wild-
life Conservation,P.O. Box 53465, Oklahoma City,
OK 73152-3465, USA.
tones between aquatic and upland habitats but
also because they often have unique attributes
(Hubbard 1977, Anderson 1980) resulting from
the presence of water. Bull (1978) noted that
soils in riparian zones supported greater plant
biomass, more plant diversity, and enabled fast-
er plant growth than soils in more xeric upland.
Many studies have shown that bird species
richness and abundance were greater in riparian
habitats than in adjacent habitats (Smith 1977,

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