Journal of Wildlife Management 74(1):81–93; 2010; DOI: 10.

2193/2008-442

Management and Conservation Article

Habitat and Landscape Effects on Brood

Parasitism, Nest Survival, and Fledgling
Production in Swainson’s Warblers
THOMAS J. BENSON,1,2 Department of Biological Sciences and Environmental Sciences Program, Arkansas State University, P.O. Box 599, Jonesboro,
AR 72467, USA
NICHOLAS M. ANICH, Department of Biological Sciences, Arkansas State University, P.O. Box 599, Jonesboro, AR 72467, USA
JEREMY D. BROWN, Department of Biological Sciences, Arkansas State University, P.O. Box 599, Jonesboro, AR 72467, USA
JAMES C. BEDNARZ, Department of Biological Sciences and Environmental Sciences Program, Arkansas State University, P.O. Box 599, Jonesboro, AR
72467, USA

ABSTRACT Numerous factors, including nest predation and brood parasitism, may limit populations of neotropical migratory birds.
However, nest predation and brood parasitism are not constant, and temporal, biological, habitat, and landscape factors can affect the likelihood
of these events. Understanding these patterns is important for species of conservation concern for which managers seek to provide quality
habitat. One such species, the Swainson’s warbler (Limnothlypis swainsonii), is a neotropical migrant that breeds primarily in bottomland
hardwood forests of the southeastern United States. Little is known of factors influencing reproductive success of this rare, yet locally abundant,
species. From 2004 through 2007, we examined factors influencing reproductive success of Swainson’s warblers at 2 sites in eastern Arkansas,
USA, St. Francis National Forest and White River National Wildlife Refuge. We used 2-stage modeling to assess the relationship between 1)
temporal and biological, and 2) habitat and landscape factors and brood parasitism, nest survival, and fledgling production. Brood parasitism
M
was greater in this population (36%) than reported elsewhere ( 10%), but decreased throughout the breeding season. Nest survival was
comparable to or lower than in other populations of this species and increased throughout the breeding season. The probability of brood
parasitism was greater near forest edges. Although nests of Swainson’s warblers were often associated with giant cane (Arundinaria gigantea),
nest survival had a weak negative association with cane density. For nests that were successful, the best predictor of number of Swainson’s
warblers fledged was brood-parasitism status: nonparasitized nests fledged 2.75 young, whereas parasitized nests fledged 0.60 Swainson’s
warblers. Our findings suggest that managing and restoring relatively high-elevation bottomland forests that are located far from agricultural
edges should increase Swainson’s warbler productivity.

KEY WORDS Arundinaria gigantea, brood parasitism, brown-headed cowbird, fledgling production, habitat, landscape,
Limnothlypis swainsonii, Molothrus ater, nest survival, Swainson’s warbler.

Numerous factors limit bird populations by affecting
reproduction or survival, including availability of food or
other resources, predation, and brood parasitism (Newton
1998). Although knowledge of habitat use is important for
avian conservation, perhaps more crucial is an understanding
of factors affecting demography. Nest predation often
results in reproductive failure (Ricklefs 1969, Martin
1992). Resource limitations and brood parasitism may also
affect the number and condition of young fledged (Martin
1987, Dearborn et al. 1998, Trine et al. 1998). Habitat,
landscape, and temporal factors affect the abundance and
habitat use of the predators, brood parasites, and resources
that ultimately drive reproductive success of avian popula-
tions (Thompson et al. 2002).
In addition to providing resources, habitat may influence
productivity by affecting the probability of a nest’s discovery
by a predator or brood parasite. Birds may reduce the
probability that a nest will be discovered by increasing nest
concealment (total foliage hypothesis; Martin 1992, 1993)
or by placing nests in areas with large numbers of potential
nest substrates, thereby decreasing the search efficiency of
predators (potential prey site hypothesis; Martin 1992,
1
E-mail: tjbenson@gmail.edu
2
Present address: Illinois Natural History Survey, 1816 S Oak Street,
Champaign, IL 61820, USA
1993). Additionally, the number of young that are fledged
from successful nests may be influenced by food abundance,
microclimate, and habitat characteristics that promote
successful foraging (Pärt 2001, Lambrechts et al. 2004,
Lloyd and Martin 2004).
The landscape context of a nest also has an influence on
productivity. Forest songbird nests located in fragmented
and agriculture-dominated landscapes often have greater
predation and brood parasitism rates than nests located in
more forested landscapes (Donovan et al. 1995, Robinson et
al. 1995, Rodewald and Yahner 2001). By altering
microhabitat conditions, fragmentation may also alter the
availability of food resources and influence the number of
young fledged (Burke and Nol 1998). Proximity and density
of edges may be associated with predator abundance
(Chalfoun et al. 2002a, b), nest predation, and brood
parasitism (Suarez et al. 1997, Batáry and Báldi 2004), and
the edge type may influence the severity of these effects.
Temporal variation may also affect productivity. For
example, numbers of predators or nest parasites or their
behaviors may change during the breeding season (Lowther
1993, Payne and Payne 1998). Similarly, nest stage, brood
size, and the presence of parasitic young in a nest may affect
parental behavior and, thereby, the conspicuousness of the
nest, thus influencing the nest’s susceptibility to discovery by
predators (Dearborn et al. 1998, Martin et al. 2000b,

Benson et al. N Swainson’s Warbler Reproductive Success 81

Tewksbury et al. 2002). Because birds should select sites that
maximize productivity, understanding the effects of tempo-
ral, biological, habitat, and landscape factors on reproductive
success is important for clarifying the adaptive nature of
habitat selection. This knowledge is particularly important
for species of conservation concern, for which managers seek
to improve habitat conditions.
One such species of concern, the Swainson’s warbler
(Limnothlypis swainsonii), is a relatively rare, medium-sized
wood warbler that may be locally abundant in appropriate
habitat throughout the southeastern United States (Brown
and Dickson 1994). Although sometimes found in mid–
seral-stage upland pine plantations and montane thickets
(Meanley 1971, Bassett-Touchell and Stouffer 2006,
Lanham and Miller 2006), Swainson’s warblers typically
breed in mesic bottomland hardwood forests with dense
understory vegetation. A secretive and poorly understood
species, Swainson’s warbler is among the species of greatest
conservation concern both regionally (Hunter et al. 1993,
Twedt et al. 1999) and nationally (Rich et al. 2004).
Historical accounts of Swainson’s warblers noted that
giant cane (Arundinaria gigantea), a bamboo species native
to the southeastern United States, was characteristic of
occupied areas (Brewster 1885, Meanley 1971), but
subsequent studies have shown Swainson’s warblers are
routinely found in noncane habitats (Graves 2001, Bednarz
et al. 2005). Even so, Swainson’s warblers appear to favor
cane habitats (Wright 2002, Anich 2008, Benson et al.
2009, Brown et al. 2009). Understanding the quality of
alternative habitats used by Swainson’s warblers is important
for effective conservation and management of bottomland
hardwood forests, especially relatively high-elevation forests
where cane was formerly abundant (Noss et al. 1995, Platt
and Brantley 1997, Twedt and Loesch 1999).
We studied reproductive success in Swainson’s warblers
from 2004 to 2007 as related to 1) brood parasitism, 2) nest
survival, and 3) production of fledglings. Specifically, we
examined the effect of temporal, biological, habitat, and
landscape factors on reproductive success. We developed
several competing hypotheses to explain variation in
Swainson’s warbler reproductive success (Table 1) that were
based on previous research on habitat use and nest-site
selection of this species (Benson et al. 2009) and on habitat
and landscape factors that influenced reproductive success of
other species.
STUDY AREA
We studied Swainson’s warbler reproductive performance at
2 locations in eastern Arkansas, USA: St. Francis National
Forest (SFNF) and White River National Wildlife Refuge
(WRNWR). Although in close geographic proximity, these
locations differed in vegetation structure and composition.
The .8,500 ha of upland and bottomland forest of SFNF,
in Lee and Phillips counties, were within an area bordered
by the Mississippi and St. Francis rivers to the east and
extending west to Crowley’s Ridge, a relatively narrow
upland ridge within the expansive Mississippi alluvial
floodplain. Areas used by Swainson’s warblers on SFNF
82
were dominated by elm (Ulmus spp.), box elder (Acer
negundo), sweetgum (Liquidambar styraciflua), maple (Acer
spp.), oak (Quercus spp.), hickory (Carya spp.), and tulip tree
(Liriodendron tulipifera). Dominant understory vegetation
included greenbrier (Smilax spp.), poison ivy (Toxicodendron
radicans), Virginia creeper (Parthenocissus quinquefolia),
spicebush (Lindera benzoin), and paw paw (Asimina triloba),
whereas giant cane was sparsely distributed.
The 62,000 ha of WRNWR, located in Arkansas, Desha,
Phillips, and Monroe counties, were among the largest
remaining continuous tracts of bottomland hardwood forest
in the Mississippi Alluvial Valley (Twedt and Loesch 1999).
Swainson’s warblers used areas on WRNWR that were
dominated by sugarberry (Celtis laevigata), sweetgum, box
elder, elm, oak, American sycamore (Platanus occidentalis),
and hickory. Dominant understory vegetation in our study
area included greenbrier, Virginia creeper, peppervine
(Ampelopsis arborea), grape (Vitis spp.), spicebush, box elder,
and relatively dense cane thickets.
METHODS
Nest Searching
After first locating territorial males based on previous
research (Brown et al. 2009) and passive and song-playback
surveys, we searched for Swainson’s warbler nests at the 2
study areas from late April to early August in 2004 through
2007. We located nests both systematically, by first
attempting to estimate the extent of each uniquely marked
male’s territory and then searching these territories with 1–6
observers, and opportunistically while engaging in other
research activities. While searching, we looked for nests
both based on appearance and placement of nest structures
(Brown and Dickson 1994) and by observing the behavior of
individual Swainson’s warblers. We recorded Global Posi-
tioning System coordinates of located nests and returned at
1–4-day intervals to determine nest status and fate. At each
visit, we determined the nest’s parasitism status (i.e., no. of
cowbird eggs or young), total clutch or brood size, and nest
stage (a combined laying and incubation stage or nestling
stage). In 2006 and 2007, we used time-lapse video to
identify predators at a subset of nests at WRNWR; these
camera systems did not appear to influence nest survival
(Benson et al. 2010).
Habitat Features
After termination of nesting, we collected habitat data at
nests and a paired site, located a random distance and
direction between 25 m and 75 m from each nest, using a
modified Breeding Biology Research and Monitoring
Database protocol (Martin et al. 1997). We chose this
distance to eliminate overlap between nest and random
samples yet constrain random locations within the home
range of the nesting birds. The mean home range at our study
areas was 9.4 ha (range 5 1.6–30.8 ha; Anich et al. 2009).
At each pair of nest location points, we collected data at
the nest-site scale (i.e., within a 5-m radius) and at the nest-
patch scale (i.e., within an 11.3-m radius). At the nest-site
scale, we estimated percent ground cover of forbs and leaf
The Journal of Wildlife Management N 74(1)
Table 1. A priori candidate models used to examine differences in brood parasitism, nest survival, and fledgling production of Swainson’s warbler nests as a
function of temporal and biological, habitat, and landscape factors at St. Francis National Forest and White River National Wildlife Refuge (location),
Arkansas, USA, 2005 through 2007. We fit habitat and landscape models while including variables from the best-fit temporal–biological model. Variables
included in the habitat and landscape models are described in the Methods.
Temporal–biological models Habitat models Landscape models
Day of yr Cane stems Any edge
Yr Cane stems + litter vola Forest-edge density within 250 m
Yr + day of yr Cane stems 3 locationb Forest-edge density within 250 m 3 locationb
Yr 3 day of yr Cane differencec Forest-edge density within 1,000 m
Parasitismd Cane differencec 3 locationb Forest-edge density within 1,000 m 3 locationb
Parasitism + day of yra Canopy covera Location
Parasitism + yra Canopy cover + litter vola Location + parasitisma
Brood sizeb Concealmente Location 3 yra
Multiple parasitismb Density differencec % forest within 1,000 m
Nest stageb,f Density differencec 3 locationb Road edge
Nest stage 3 parasitismb Forb covera Road edge 3 locationb
Nest stage 3 brood sizeb Litter vola Road or trail edge
Nest stage 3 day of yrb Mature forestg Road or trail edge 3 locationb
Nest stage 3 day of yr + yrb Nest-patch density
Nest stage + yrb Nest-patch density + cane stems
Nest-patch density + canopy cover
Nest-patch density + litter vola
Nest-patch density + understory ht
Nest-patch density 3 locationb
Nest-patch heterogeneityh
Nest-site densityi
Noncane stems
Noncane stems 3 locationb
Noncane differencec,i
Noncane differencec 3 locationb
Stem typej
Total stems
a
Model used only to evaluate fledgling production.
b
Model used only to evaluate nest survival.
c
Difference between values for nest sites or patches and paired random sites or patches.
d
Model used only to evaluate nest survival and fledgling production.
e
Model includes nest-patch density, sub-canopy cover, and sub-canopy ht.
f
Nest stage refers to either the combined laying and incubation period, or the nestling period.
g
Model includes medium and large trees, total canopy cover, and overstory ht.
h
Model includes horizontal CV, vertical CV, horizontal difference.
i
Model used only to evaluate brood parasitism and nest survival.
j
Model includes cane-, shrub-, and vine-stem density.

litter in each of 4 quadrants of the 5-m-radius circle. At a
distance of 5 m in the 4 cardinal directions, we counted the
number of cane, vine, and shrub stems within a 1-m2
quadrat at a height of 0.3 m, and recorded the leaf-litter
depth to the nearest millimeter using a ruler. Using our
estimates of leaf-litter cover and depth, we calculated litter
volume as plot area 3 percent leaf-litter cover 3 litter depth
in meters. From the center of each plot, we estimated the
angular canopy cover (Nuttle 1997) of forest in the 4
cardinal directions using a spherical densiometer, and
visually estimated sub-canopy cover. Using a clinometer,
we measured height of overstory canopy and sub-canopy
within the nest patch. Also at the nest-patch scale, we
counted the number of medium (23–38-cm dbh) and large
trees (.38-cm dbh). We estimated understory density at
both the nest-site and nest-patch scales. We did this by
placing a 2.5-m-tall cover board in the center of each plot,
and recording the percent of the board that was obscured by
vegetation in 5 different height classes: 0–0.5 m, 0.5–1.0 m,
1.0–1.5 m, 1.5–2.0 m, and 2.0–2.5 m (Nudds 1977). We
estimated these 5 values from both 5 m and 11.3 m from
the board in the 4 cardinal directions, and we calculated the
mean density based on 20 readings at each distance. Because
heterogeneity in vegetation structure may be an important
factor for Swainson’s warblers, we computed the coefficient
of variation of understory density readings at each point for
the 4 cardinal directions averaged over all height intervals
(horizontal CV) and the 5 vertical height intervals averaged
over all cardinal directions (vertical CV). As another
measure of heterogeneity at nests, we calculated the
difference in mean density (i.e., averaged over all ht intervals
and cardinal directions) between the nest-patch and nest-
site scale.
Horizontal difference and coefficient of variation estimat-
ed patchiness only within 11.3 m. To evaluate patchiness at
a larger, territory-level scale, we used the difference between
nest-patch values of understory density and height and
corresponding values from the paired random patch to
create 2 new variables: density difference and understory
height difference. We calculated these same differences for
cane-stem and noncane-stem densities (cane difference and
noncane difference) and these variables differentiate isolated
from relatively continuous patches of cane and noncane
habitat. Positive values for these differences represented nest

Benson et al. N Swainson’s Warbler Reproductive Success 83

sites that have a relatively dense, tall, cane-dominated, or
noncane-dominated understory relative to random locations.
Landscape Context
We obtained digital land-cover data for summer 2004 and
color infrared aerial photographs taken between 1999 and
2004 (Arkansas State Land Information Board 2002, Center
for Advanced Spatial Technologies 2005). We classified data
into 14 categories (low-intensity urban, high-intensity urban,
barren land, water, woody–herbaceous, forest and woodland,
soybeans, rice, cotton, sorghum–corn, other cropland, bare
soil–seedbed–fallow, warm-season pasture–forages, and cool-
season pasture–forages). Using these aerial photographs, and
shape-files for secondary roads, all-terrain vehicle (ATV)
trails, and streams acquired from agency personnel (R. Hines,
United States Fish and Wildlife Service, and T. Hocut,
United States Forest Service, unpublished data), we digitized
agricultural edges, roads, and streams and ditches within and
surrounding our study areas. We confirmed the accuracy of
spatial data based on ground-truthing at both study areas and
modified misclassified data to account for discrepancies.
Using these digital spatial data, we estimated the distance
of each nest from 4 different edge types: 1) agriculture,
defined as the interface between forest and row-crop, hay, or
pasture; 2) primary roads, which were well-maintained, had
open canopies, and were open to vehicular traffic under most
circumstances; 3) secondary roads that are not well-
maintained, had partially open canopies, and were not
regularly open to vehicular traffic and trails that were open
to horse or ATV traffic; and 4) streams, including bayous
and perennial ditches. We determined percent forest cover
and forest-edge density (the interface between forest and
another land-cover type in m/ha) within 250 m of each nest
(a 19.6-ha area that likely encompassed the home range of
most Swainson’s warblers) and within a 1,000-m radius
(314 ha) that assessed the nest’s landscape context. We also
generated estimates of landscape heterogeneity within these
2 radii using the Shannon diversity index based on the
proportion of 14 different land-cover types in each radius as
2Spi lnpi, where pi is the relative proportion of the ith land-
cover type (Magurran 1988).
Data Analyses
To determine which variables influenced brood parasitism,
nest survival, and fledgling production, we generated a set of
a priori models for each analysis based on factors we
hypothesized as important based on previous research and
personal observations (Table 1). We first evaluated correla-
tions among these variables and did not include correlated
variables (|r| . 0.70) in the same model. Because of
redundancy among landscape metrics, we removed agricul-
tural edge distance, stream distance, percent forest cover at
the 250-m scale, and landscape heterogeneity at both the
250-m and 1,000-m scales, all of which were highly
correlated with percent forest cover within 1,000 m. We
ranked these candidate models using Akaike’s Information
Criterion adjusted for small sample sizes (AICc) and
computed model weights (Burnham and Anderson 2002).
84
We evaluated 2 types of models: 1) temporal and
biological, and 2) habitat and landscape (Table 1). Because
the temporal and biological factors likely reflect important
factors that ultimately affect the influence of habitat or
landscape features, we first evaluated these factors and used
the variables from the best-fit temporal or biological model
based on AICc in subsequent landscape and habitat models.
We considered models with DAICc
M
2 to be competing
models and generated model-averaged parameter estimates
and 95% confidence intervals based on this set (Burnham
and Anderson 2002).
We examined our models for brood parasitism using
logistic regression (SAS PROC LOGISTIC; Allison 1999).
We assessed model fit for the global model using a Hosmer
and Lemeshow goodness-of-fit test, and assessed evidence
of overdispersion using the ratio of the Pearson chi-squared
statistic to the degrees of freedom (Allison 1999).
We investigated the nest survival models using the
logistic-exposure method (Shaffer 2004). To account for
potential nonindependence of nest fates due to proximity,
we first classified nests into blocks based on proximity and
examined logistic-exposure models with a random block
effect using SAS PROC NLMIXED (Shaffer 2004, Littell
et al. 2006, Rotella et al. 2007, Benson 2008). Inclusion of
block effects did not improve models, so we proceeded with
evaluating candidate models using logistic-exposure models
in SAS PROC GENMOD (Shaffer 2004). We confirmed
overall model fit using a likelihood ratio test between the
global and intercept-only models (x2 5 53.31, P , 0.005),
and used the ratio of the chi-squared statistic to degrees of
M
freedom to evaluate evidence of overdispersion (ĉ 1.2).
We generated model-averaged values of predicted daily
survival rates and 95% confidence intervals for variables of
interest while holding all other variables at their mean values
(Shaffer and Thompson 2007).
For nests that successfully fledged young, we evaluated
temporal, biological, habitat, and landscape models that may
affect this production. Although nests that fledged no
L
Swainson’s warblers but did fledge 1 cowbirds are not
considered successful, we included these nests for estimating
fledgling production as fledging zero Swainson’s warbler
young. Because the number of fledged young represents
count data, we evaluated fledgling production using Poisson
regression (SAS PROC GENMOD; Allison 1999). To
assess evidence of overdispersion, we used the ratio of the
chi-squared statistic to the degrees of freedom (Allison
1999). Limited evidence of overdispersion (ĉ
M
1.2) and
approximate equality of the mean (2.1) and variance (1.8)
for this variable confirmed that the Poisson distribution was
suitable for this analysis.
RESULTS
From 2004 to 2007, we found 269 nests, of which 135 were
active (i.e., containing viable eggs or young). Of these 135
nests, 48 (36%) were parasitized by brown-headed cowbirds,
of which 14 (10%) contained multiple cowbird eggs. Mean
clutch size of Swainson’s warbler eggs was 3.43 6 0.07 (SE)
for nonparasitized nests (n 5 68 incubation-stage nests),
The Journal of Wildlife Management N 74(1)
Table 2. Results of the best-fitting logistic regression models predicting
brown-headed cowbird parasitism of Swainson’s warbler nests (n 5 129) as
a function of temporal factors and habitat and landscape factors at St.
Francis National Forest and White River National Wildlife Refuge,
Arkansas, USA, 2005 through 2007. We fit habitat and landscape models
while including variables from the best-fit temporal model.
Modela Kb AICcc DAICcd wie
Temporal models
Day of yr 2 168.19 0 0.69
Constant parasitism 1 171.25 3.06 0.15
Habitat and landscape models
Forest-edge density (250 m) +
day of yr 3 163.92 0 0.38
% forest (1,000 m) + day of yr 3 166.75 2.83 0.09
Nest-patch density + Nest-patch
ht + day of yr 4 167.13 3.21 0.08
Day of yr 2 168.19 4.27 0.05
a
For a list of all candidate models see Table 1.
b
No. of parameters.
c
Akaike’s Information Criterion for small sample sizes (Burnham and
Anderson 2002); AICc 5 22 log L + 2K + 2K(K + 1) / (n 2 K 2 1).
d
DAICc 5 AICci 2 minAICc; we calculated DAICc values separately for
each model set. Figure 1. Predicted probability (with 95% CL) of brown-headed cowbird
e (BHCO) parasitism of Swainson’s warbler as a function of day of year and
wi 5 exp[2(DAICci / 2)] / S exp[2(DAICci / 2)]. We calculated wi
values separately for each model set. forest-edge density within 250 m of a nest site at St. Francis National
Forest and White River National Wildlife Refuge, Arkansas, USA, 2005
through 2007.
2.57 6 0.17 for singly parasitized nests (n 5 30), and 1.45
6 0.28 for multiply parasitized nests (n 5 11). Of the
parasitized nests, 34 contained one, 11 contained 2, 2 0.958) and there was limited evidence of overdispersion of data
M
contained 3, and one contained 4 cowbird eggs (x ¯ 5 1.38 6 (ĉ 1.2).
0.10 [SE]). Of 52 nests fledging either Swainson’s warbler
Nest Survival
or cowbird young (x ¯ 5 2.13 6 0.19 [SE] Swainson’s
The best-fitting temporal or biological model for nest
warblers/nest), 43 fledged Swainson’s warblers (6 with
survival incorporated the interactive effect of nest stage and
cowbirds; ¯x 5 2.58 6 0.14 [SE]), and 9 fledged only
day of year (Table 3). The sum of Akaike weights for
cowbirds. The presumed cause of most nest failures was
models incorporating an effect of nest stage was 1.00, and
predation (n 5 72, 78%), followed by nests that fledged only the value for day of year was 0.66, indicating strong support
cowbirds (n 5 9, 10%), abandonment (n 5 6, 7%), or failed for both of these effects. The estimated daily survival rate
due to weather (n 5 5, 5%). We had complete habitat data was greater for the combined laying and incubation period
for 129 of the 135 active nests and we included only these in than for the nestling period. Nest survival increased as the
subsequent analyses: 35 in 2005, 47 in 2006, and 47 in 2007 breeding season progressed (Fig. 2). Two habitat or
(Appendix A). landscape variables improved the fit of the nest stage 3
Brood Parasitism day of year model, and there were 6 other competing models
M
The only strongly supported temporal model for brown- with DAICc 2. In addition to the nest stage and day-of-
headed cowbird parasitism was a negative effect of day of year year effects, daily survival rate decreased with increasing
(Table 2; Fig. 1). Three landscape or habitat models fit better density of cane stems at a nest site (Fig. 3). Although the
than the day of year model, although only one with DAICc
M model-averaged 95% confidence intervals for all habitat and
2 (Table 2). The top 3 models accounted for 55% of the landscape parameters included zero, this is partly a
Akaike weight and the landscape variables from these models consequence of model selection uncertainty (averaging
indicated that the probability of cowbird parasitism was greater across all competing models, by treating the absence of a
with increased forest-edge density (Fig. 1) and decreased variable as a parameter estimate of zero, causes model-
percent forest cover. The third-ranked model incorporated a averaged parameter estimates to tend toward zero; Burnham
weak positive effect of nest-patch density and a positive effect and Anderson 2002). Confidence intervals from some
of understory height. The best-fitting model was (p 5 individual models did not include or only slightly overlapped
probability of cowbird parasitism; SE in parentheses): zero (Benson 2008). Based on these individual models and

 the summed weights, there was limited support for a few
loge p 1{p ~2:495ð1:553Þz0:024ð0:010Þ|Forest variables, including the overall effect of cane density, and of
edge density{0:022ð0:010Þ|Day of year noncane patchiness at SFNF (Table 4) although no habitat
or landscape variables were particularly strong predictors of
Concordance (a metric of prediction accuracy) for the top nest survival. The overall model-averaged daily survival rate
model was 68%; the models fit the data well (x2 5 2.57, P 5 was 0.950 (95% CI 5 0.937, 0.960; n 5 129; 1,678

Benson et al. N Swainson’s Warbler Reproductive Success 85

Table 3. Results from the best-fitting logistic-exposure models predicting
daily survival rate of Swainson’s warbler nests (n 5 129) as a function of
temporal and biological factors, and habitat and landscape factors at St.
Francis National Forest and White River National Wildlife Refuge,
Arkansas, USA, 2005 through 2007. We fit habitat and landscape models
while including variables from the best-fit temporal–biological model.
Modela Kb AICcc DAICcd wie
Temporal and biological models
Nest stage 3 day of yr 3 519.75 0 0.40
Nest stage 3 day of yr + yr 5 520.64 0.89 0.26
Constant survival 1 532.68 12.93 ,0.01
Habitat and landscape models
Cane stems + nest stage 3 day
of yr 4 518.97 0 0.09
Noncane difference 3 location
+ nest stage 3 day of yr 5 519.13 0.16 0.08
Nest stage 3 day of yr 3 519.75 0.78 0.06
Road distance + nest stage 3
day of yr 4 520.00 1.03 0.05
Noncane stems + nest stage 3
day of yr 4 520.40 1.43 0.04
Stem type + nest stage 3 day
of yr 6 520.52 1.54 0.04
Cane stems 3 location + nest
stage 3 day of yr 5 520.52 1.55 0.04
Any edge distance + nest stage
3 day of yr 4 520.79 1.82 0.04
Nest-patch density + cane stems
+ nest stage 3 day of yr 5 520.93 1.95 0.03
a
For a list of all candidate models see Table 1.
b
No. of parameters.
c
Akaike’s Information Criterion for small sample sizes (Burnham and
Anderson 2002); AICc 5 22 log L + 2K + 2K(K + 1) / (n 2 K 2 1).
d
DAICc 5 AICci 2 minAICc; we calculated DAICc values separately for
each model set.
e
wi 5 exp[2 (DAICci / 2)] / S exp[2 (DAICci / 2)]. We calculated wi
values separately for each model set.
Figure 2. Model-averaged predicted daily survival rate (with 95% CL) of
exposure-days). The model-averaged daily survival rate for Swainson’s warbler nests as a function of day of year for (A) the combined
the egg stage was 0.967 (95% CI 5 0.954, 0.976), and was laying and incubation stages, and (B) the nestling stage at St. Francis
National Forest and White River National Wildlife Refuge, Arkansas,
0.927 (95% CI 5 0.901, 0.946) for the nestling stage (while USA, 2005 through 2007. We generated estimates while holding all other
holding all other variables at their mean values and day of yr variables from competing models at their mean values.
at 160 and 175, respectively).

Fledgling Production
In the analysis of fledgling production, the best-fitting
Poisson regression model incorporated an effect of brown-
headed cowbird parasitism and models with this variable
accounted for 100% of the Akaike weight (Table 5).
Although a few models that incorporated habitat or
landscape factors received some support, the 95% confidence
intervals for the coefficients of these effects included zero.
Production from nests that successfully fledged Swainson’s
warblers, cowbirds, or both was drastically reduced from
2.75 (95% CI 5 2.27, 3.35) Swainson’s warbler fledglings
in nonparasitized nests to 0.60 (95% CI 5 0.30, 1.13)
Swainson’s warbler fledglings in nests parasitized by
cowbirds. No landscape or habitat variable improved upon
this parasitism model, although there were 7 other models
M We observed high (36%) rates of cowbird parasitism. This
with DAICc 2 (Table 5). Included among these
competing models was evidence for positive effects of
cane-stem density and nest-patch density, as well as a
negative effect of litter volume although the 95% confidence
intervals for these parameters included zero (Table 4).
DISCUSSION
Generally, Swainson’s warbler reproductive success at our
study areas was similar to that observed in previous Swainson’s
warbler studies in other locations. Our estimated daily nest
survival rate of 0.950 was similar to estimates from Louisiana,
USA, pine plantations and hardwood habitats (0.949 and
0.956, respectively; Henry 2004) but less than estimates from
managed bottomland hardwood forests in South Carolina,
USA (0.961 and 0.979; Peters et al. 2005, Thompson 2005),
and Missouri, USA, canebrakes (0.980; Thomas et al. 1996).
Assuming a 26-day nesting period (Thompson 2005), our nest
success estimate was 26% (95% CI 5 18–35%). The clutch
size for nonparasitized nests at our study areas, 3.43, was
similar to the values from Missouri, South Carolina, and
Louisiana (3.65, 3.10, and 3.30, respectively).
contrasts markedly with other studies, with no reported
brood parasitism in Virginia, USA (n 5 11; Meanley 1982),
or Missouri (n 5 17; Thomas et al. 1996), 3–8% in
Louisiana (n 5 138; Henry 2004), and 6–10% in South

86 The Journal of Wildlife Management N 74(1)


Figure 3. Model-averaged predicted daily survival rate (with 95% CL) of
Swainson’s warbler nests as a function of cane density for (A) the combined
laying and incubation stages, and (B) the nestling stage at St. Francis
National Forest and White River National Wildlife Refuge, Arkansas,
USA, 2005 through 2007. We generated estimates while holding all other
variables from competing models at their mean values.
Carolina (n 5 17, 79; Peters et al. 2005, Thompson 2005).
The 2.75 fledglings produced per successful nonparasitized
nest at our study areas was similar to the 3.0 fledglings per
successful nest observed in Louisiana and greater than the
2.4 and 2.1 fledglings per successful nest in South Carolina
and Missouri, respectively (Thomas et al. 1996, Henry 2004,
Thompson 2005). However, because clutch size in parasit-
ized nests was reduced to 2.27 and these nests produced only
0.60 Swainson’s warbler fledglings when successful, fledg-
ling production from all successful nests on our study areas
was 2.13 (SE 5 0.19).
Nest predation was the most important cause of
reproductive failure, but brown-headed cowbird parasitism
reduced productivity of successful nests. Both daily survival
rate and probability of cowbird parasitism displayed
temporal trends, with survival increasing and parasitism
decreasing throughout the breeding season. A nest initiated
on 5 May had a 53% probability of being parasitized by a
cowbird, a 23% probability of being multiply parasitized,
and an estimated nest success of around 17%, whereas a nest
Benson et al. N Swainson’s Warbler Reproductive Success
initiated on 25 June had a 37% chance of being parasitized
by a cowbird, a 6% chance of being multiply parasitized, and
a 37% probability of successfully producing young. Given
these temporal trends, renesting after a failure was vital to
successful reproduction and potential viability for the
Swainson’s warbler populations we studied (Benson 2008).
Swainson’s warblers routinely renest following nest failure;
often 2 and up to 3 renesting attempts have been observed in
our populations and in North Carolina (T. J. Benson,
Arkansas State University, unpublished data; N. Chartier,
North Carolina State University, personal communication).
Likewise, females that successfully produce broods early in
the season may attempt a second brood (Thompson 2005; T.
J. Benson, unpublished data; N. Chartier, personal commu-
nication). Because first broods are likely to be parasitized and
have decreased production of Swainson’s warbler young when
successful, renesting may be important to the maintenance of
populations. However, additional study with color-banded or
radiomarked females is needed to determine probabilities of
renesting and double-brooding.
The reasons for these temporal changes in predation and
parasitism risk are likely caused by the ecology of brown-
headed cowbirds and the major nest predators at these study
areas (Benson et al. 2010). Others have observed that
brown-headed cowbird breeding activity is more intense in
May and June than later in the breeding season (e.g.,
Lowther 1993, Payne and Payne 1998, Gannon 2005,
Chapa-Vargas and Robinson 2007). Moreover, predation of
nests by cowbirds and raptors, major nest predators at
WRNWR, decreased throughout the breeding season
(Benson et al. 2010). This same temporal pattern of nest
parasitism and predation was observed for indigo buntings
(Passerina cyanea) at WRNWR (Welch 2000). However,
studies of nest success in other areas have found increases in
predation as the season progressed, which may be related to
differences in nest predators in these areas (Grant et al.
2005, Lloyd and Slater 2007, Peak 2007).
In addition to temporal patterns, the increased probability
of survival during the combined egg-laying and incubation
stages compared to that during the nestling stage was likely
caused by differences in activity at nests; the high rate of
feeding by both parents during the nestling period
contrasting markedly with the relatively infrequent trips to
and from the nest during incubation (T. J. Benson,
unpublished data). Indeed, past studies of other species
have found similar differences in predation risk during these
nest stages and direct effects of increased visitation rates
during the nestling or incubation periods on predation risk
(Martin et al. 2000a, b). Even so, we did not observe an
effect of brown-headed cowbird parasitism or increased
brood size on predation, although these factors increased
activity at nests (Pappas et al. 2010). The nest stage
difference in survival suggests that the predators responsible
for most nest failures may be visually oriented. Time-lapse
camera monitoring of a subset of nests at WRNWR (45 of
91 nests in 2 yr) indicated rat snakes (Elaphe obsoleta) and
raptors were the primary predators during the nestling
87
Table 4. Model-averaged parameter estimates (b), 95% confidence limits (LCL, UCL), and variable importance values (Swi) for competing models (DAICc
M 2) of nest survival and fledgling production from Swainson’s warbler nests at St. Francis National Forest (SFNF) and White River National Wildlife
Refuge (WRNWR), Arkansas, USA, 2005 through 2007.
Nest survival Fledgling production
Parameter b LCL UCL Swi b LCL UCL Swi
Intercept 0.921 21.147 2.988 20.284 21.536 0.969
Incubation 3 day of yr 0.016 0.003 0.029
Nestling 3 day of yr 0.010 20.002 0.022
Parasitism
No brown-headed cowbird 1.550 0.862 2.237
Brown-headed cowbird 0 0 0
Cane stems (/m2) 0.010 20.027 0.046 0.16
SFNF 20.026 20.103 0.051 0.21
WRNWR 20.029 20.110 0.052 0.21
Shrub stems (/m2) 0.016 20.050 0.081 0.04
Vine stems (/m2) 20.001 20.009 0.008 0.04
Noncane stems (/m2) 0.004 20.013 0.022 0.08 20.001 20.009 0.006 0.05
Noncane difference (/m2)
SFNF 0.028 20.071 0.126 0.10
WRNWR 20.008 20.043 0.026 0.10
Nest-patch density (%) 0.001 20.001 0.002 0.12 0.001 20.003 0.005 0.17
Litter vol (m3) 20.052 20.224 0.120 0.22
Any edge distance (100 m) 0.011 20.038 0.059 0.04
Road distance (100 m) 20.006 20.032 0.019 0.05
% forest within 1,000 m 20.002 20.009 0.005 0.10

period (Benson et al. 2010). These species use visual cues to parasitism; on average, parasitized nests that fledged young
locate nests (Mullin and Cooper 1998, Dykstra et al. 2008). produced ,1 Swainson’s warbler fledgling, with singly
Despite changes in the probability of nest survival and brood parasitized nests producing 0.82 Swainson’s warbler young
parasitism, successful nests produced the same number of per nest (SE 5 0.30; n 5 11) and multiply parasitized nests
young both early and late in the breeding season. Fledgling producing zero (n 5 4 nests). Although decreased productivity
production from successful nests was most affected by brood due to egg removal and competition between cowbird and
host young has been reported in other species (Trine et al.
1998), the magnitude of this effect for Swainson’s warblers
Table 5. Results from best-fitting Poisson regression models predicting
fledgling production from successful Swainson’s warbler nests (n 5 52) as a was large. Some host species can successfully raise multiple
function of temporal and biological factors, and habitat and landscape host young even in multiply parasitized nests (Hoover 2003),
factors at St. Francis National Forest and White River National Wildlife although many smaller species have significant reductions in
Refuge, Arkansas, USA, 2005 through 2007. We fit habitat and landscape
models while including variables from the best-fit temporal/biological the number of fledglings (Payne 1998, Trine et al. 1998,
model. Lorenzana and Sealy 1999). Thus, for some species,
Modela Kb AICcc DAICcd wie
parasitism-induced reductions in fledgling numbers combined
with high parasitism rates may have significant population-
Temporal and biological models
level effects (Ward and Smith 2000).
Parasitism 2 28.57 0 0.57
Constant 1 55.74 27.17 ,0.01 We expected nest success to increase, and brood parasitism
Habitat and landscape models
to decrease, with increased nest concealment and increased
Parasitism 2 28.57 0 0.12 availability of potential alternative nest substrates. However,
% forest (1,000 m) + parasitism 3 28.82 0.25 0.10 we found little evidence of concealment benefits at our study
Litter vol + parasitism 3 29.10 0.53 0.09 areas. Additionally, density of cane stems, the preferred nest
Cane stems + parasitism 3 29.50 0.93 0.07
substrate in both locations (Benson et al. 2009), was
Cane stems + litter vol +
parasitism 4 30.00 1.43 0.06 associated with slightly decreased nest survival, although
Nest-patch density + parasitism 3 30.08 1.50 0.06 density of noncane stems relative to surrounding areas was
Noncane stems + parasitism 3 30.21 1.64 0.05 associated with increased nest survival at SFNF.
Nest-patch density + litter vol
+ parasitism 4 30.54 1.97 0.04 The observed relationships with cane density and noncane
a
patchiness likely reflect the ecology and searching behavior
For a list of all candidate models see Table 1.
b
No. of parameters.
of cowbirds and major predators. Cane is the preferred nest
c
Akaike’s Information Criterion for small sample sizes (Burnham and substrate for Swainson’s warblers largely because at these 2
Anderson 2002); AICc 5 22 log L + 2K + 2K(K + 1) / (n 2 K 2 1). study areas few other substrates provide a suitable structure
d
DAICc 5 AICci 2 minAICc; we calculated DAICc values separately for (Benson et al. 2009). Indeed, cane at these locations is used
each model set.
e
wi 5 exp[2(DAICci / 2)] / S exp[2(DAICci / 2)]. We calculated wi extensively as a nest substrate by other species, including
values separately for each model set. northern cardinals (Cardinalis cardinalis), indigo buntings,

88 The Journal of Wildlife Management N 74(1)

and hooded warblers (Wilsonia citrina); the resultant
increased density of nests may lead predators, including
cowbirds, raptors, and rat snakes, to focus search efforts on
cane-dominated areas and away from isolated patches of
noncane habitat (Benson et al. 2010). Relative to the large and
expansive canebrakes historically present (Noss et al. 1995),
the isolated and fragmented nature of extant cane patches may
also contribute to our high observed predation rates.
In forests, shrub-layer nests often have the highest
predation rates (Martin 1995). Although some studies have
found effects of concealment or abundance of potential nest
substrates, including cane, on nest success (Hoover and
Brittingham 1998, Moorman et al. 2002, Chapa-Vargas and
Robinson 2006), others have found no effect of these
variables (Holway 1991, Howlett and Stutchbury 1996) or
no habitat differences between successful and unsuccessful
nests (Filliater et al. 1994, Wilson and Cooper 1998).
Because predation pressure on these shrub-layer nesting
birds is likely high, concealment may offer few benefits,
although there was weak evidence of an effect of nest-patch
density on nest survival in this study, and sub-canopy
density at WRNWR may obscure nests from aerial
predators (Benson 2008). Because many of the potential
predators at our study locations are visually oriented, the
effect of concealment may depend on parental behavior
(Remeš 2005). Similarly, we found no survival benefit for
nests surrounded by other potential nest substrates. This
contrasts with the results of Martin and Roper (1988), who
found that hermit thrush (Catharus guttatus) nests sur-
rounded by other potential nest sites had greater success.
Swainson’s warblers generally select nest sites that are well-
concealed and surrounded by other potential nest substrates
and may select areas with significant numbers of leaf clumps
that resemble nests in these surrounding substrates (Henry
2004, Lanham and Miller 2006). However, the effect of
nesting near these potential nest substrates and decoy nests
may be negated if predators search for prey based on
behavior rather than the appearance of nests, or if these
groups of potential nests are singled out by some predators.
Based on the observed nest survival relationship, the
apparent preference of Swainson’s warblers for cane-domi-
nated areas seems maladaptive. This type of ecological trap,
where preferred habitats are less productive, has been observed
in other studies and often results from human activities
(Misenhelter and Rotenberry 2000, Weldon and Haddad
2005, Robertson and Hutto 2006). The observed negative
association between Swainson’s warbler nest survival and cane
density may result from recent changes in land use. The
conversion of bottomland forests to agriculture has largely
focused on the relatively high-elevation areas that, in addition
to being suitable for cane, are also drier and more suitable for
agriculture (Noss et al. 1995, Platt and Brantley 1997, Twedt
and Loesch 1999). Thus, cane-dominated areas were
historically much more common in bottomland forests.
Indeed, in our study areas cane-dominated habitat was often
surrounded by lower elevation areas or on the periphery of
protected areas close to agricultural edges. These isolated areas
may serve as islands of habitat for understory-dependent birds
Benson et al. N Swainson’s Warbler Reproductive Success
and they may also be targeted by predators because of their
concentrated resources.
As expected, brood parasitism was associated with decreased
forest cover, increased forest-edge density, and increased
landscape heterogeneity. Brown-headed cowbirds are capable
of large movements and, although they often feed in
agricultural areas later in the day, they often venture into
forests to parasitize hosts (Thompson 1994, Goguen and
Mathews 2001). Indeed, in fragmented landscapes, cowbirds
often penetrate far into the forest interior leading to high
parasitism rates throughout the forest (Donovan et al. 1997,
Hahn and Hatfield 1995). Nests at our study areas ranged
from being immediately adjacent to an agricultural edge or
road to .3.5 km from an agricultural edge (Appendix B).
The primary nonforest land cover at our study areas was
agriculture, so forest cover, forest-edge density, and hetero-
geneity are all largely driven by proximity to an agricultural
edge. Our observed relationship between parasitism and
agriculture is consistent with results from previous studies
(Morse and Robinson 1999, Hoover et al. 2006).
Despite the relationship between parasitism and proximity
to agriculture, even nests in completely forested areas were
susceptible to parasitism; during the first 3 weeks of May, a
nest surrounded by 60–85% forest had a predicted
probability of parasitism between 63% and 84%, whereas
the probability of parasitism for nests completely surrounded
by forest was 40–48%. Cowbirds were often found near
roads even far from agricultural areas, so these high
parasitism rates were not surprising, and nests located close
to roads had high probabilities of multiple parasitism
(Benson 2008). At WRNWR, Gannon (2005) found the
probability of parasitism decreased significantly at approx-
imately 1.4 km from levees for Acadian flycatchers (Empi-
donax virescens) and at 2.2 km for prothonotary warblers
(Protonotaria citrea). The Swainson’s warbler nests we
studied were generally within 2 km of a levee (Appendix B).
Thus, the current landscape context at our study areas may
foster higher rates of nest parasitism than occurred prior to
extensive human modifications of the landscape and the
productivity of these populations may be depressed com-
pared to historical rates.
No landscape variables were good predictors of nest
survival. Nest predation was the most frequent cause of
complete failure, and the lack of consistent landscape
predictors is likely caused in part by contrasting effects of
landscape features on each important predator species. Data
from cameras monitoring nests at WRNWR (Benson et al.
2010) indicated nest predation by rat snakes increased
farther from roads. This is consistent with the weak effect
we found for roads on nest success and as reported elsewhere
(King and DeGraaf 2002, Rodewald 2002).
We found only a weak effect of edge proximity on nest
survival compared to other studies (Flaspohler et al. 2001,
Batáry and Báldi 2004). In particular, agricultural edges may
be particularly dangerous for some forest species (Suarez et
al. 1997, Batáry and Báldi 2004) and density of forest edges
in general has had a reported negative effect on nest survival
for some species (Weldon and Haddad 2005, Peak 2007).
89
However, other studies have found no effect or a varying
effect of edge proximity, depending on edge type (Kaiser
and Lindell 2007, Newell and Kostalos 2007).
Although relationships between brood parasitism and
habitat or landscape factors were reasonably strong, those
between nest survival and these factors had less support and
were more variable. However, there was evidence for an
impact of several different factors on nest survival. Even
though temporal or biological factors may have had stronger
and more consistent effects on nest survival, these factors are
not easily managed. Conversely, management focused on
habitat or landscape factors may result in small changes in
nest survival but these may markedly influence the viability
of these populations (Benson 2008).
Because frequent flooding alters the suitability of potential
habitat (Benson and Bednarz 2010), management for
Swainson’s warblers and other understory-dependent birds
of conservation concern in the Southeast should be focused on
conserving, restoring, and managing relatively high-elevation
bottomland hardwood forests (e.g., Graves 2001, 2002; Twedt
et al. 2006). We recommend management of these areas
emphasize creating and maintaining dense understory vege-
tation through silviculture (Heltzel and Leberg 2006, Twedt
and Wilson 2007, Twedt and Somershoe 2009). The
immediate effect of recommended silvicultural treatments
will likely be a decrease in density of Swainson’s warblers with
a peak in density between 10 years and 20 years postharvest
(Heltzel and Leberg 2006, Twedt and Wilson 2007, Twedt
and Somershoe 2009), so managed forests should be
maintained as a mosaic of different-aged stands to ensure
landscape-level availability of suitable habitat and care should
be taken to avoid extirpating existing isolated Swainson’s
warbler populations. For isolated populations, aggressive
management of marginal habitat may promote colonization
of unoccupied habitat whereas less-aggressive management
should be used within occupied habitat. However, additional
monitoring and demographic studies are necessary to
determine the effects of alternative management regimes.
MANAGEMENT IMPLICATIONS
To provide quality nesting habitat, managers should
maintain or enhance existing forested canebrakes where
practical, but canopy disturbances that promote the
development of dense shrub- or vine-dominated understo-
ries should benefit Swainson’s warblers and may be more
effectively implemented given the difficulty of establishing
and restoring cane. Because isolated cane patches may be
targeted by some predators, silvicultural treatments such as
thinning, group selection, group selection with thinning, or
shelterwood cuts may promote the development of uni-
formly dense understory structure that is beneficial for
Swainson’s warblers (Heltzel and Leberg 2006, Twedt and
Wilson 2007, Twedt and Somershoe 2009). Additionally,
management should be focused on higher elevation (i.e., less
flood-prone) bottomland forests that are relatively far from
agricultural edges and on restoring forest to buffer existing
habitat from the negative effects (e.g., cowbird parasitism)
of agricultural-edge proximity.
90
ACKNOWLEDGMENTS
Funding for this research was provided by the Arkansas
Game and Fish Commission (AGFC) and United States
Fish and Wildlife Service (USFWS) through a State
Wildlife Grant. Additional funds were received from a
cost-share program with USFWS and Arkansas State
University. We thank C. W. Rideout and L. M. Barnhill,
formerly of AGFC, for continued support of this research
and for help securing funding. R. E. Hines, W. C. Hunter,
and S. R. Reagan of USFWS provided invaluable assistance.
D. J. Twedt, P. B. Hamel, T. S. Risch, R. S. Grippo, G. S.
Guha, and 2 anonymous reviewers provided valuable
comments on an earlier version of this manuscript. We
thank M. C. Roa, J. M. Sardell, M. B. Albrechtsen, K. E.
Ballantyne, R. J. Baxter, E. J. Huskinson, J. A. Jackson, K.
A. Jones, C. A. McCarroll, J. L. O’Connell, B. A. Paterson,
A. M. St.-Pierre, D. M. Townsend, and W. D. Edwards for
providing valuable field assistance.
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The Journal of Wildlife Management N 74(1)
Appendix A. Mean, standard error, and range of values for habitat measurements taken at Swainson’s warbler nests at St. Francis National Forest (SFNF; n
5 38) and White River National Wildlife Refuge (WRNWR; n 5 91), Arkansas, USA, 2005 through 2007.
SFNF WRNWR
Variable ¯x SE Range ¯x SE Range
Nest site (5-m radius)
Cane stems (/m2) 3.6 0.5 0–13.0 4.1 0.3 0–12.3
Shrub stems (/m2) 1.0 0.1 0–4.3 0.6 0.2 0–15.5
Vine stems (/m2) 2.5 0.4 0–11.0 2.8 0.3 0–16.3
Noncane stems (/m2) 3.5 0.5 0–14.5 3.4 0.4 0–17.3
Total stems (/m2) 7.1 0.7 0.5–16.3 7.5 0.4 1.3–20.5
Nest-site density (%) 47.1 3.2 9.5–83.3 42.6 1.9 8.5–77.3
Forb cover (%) 8.0 1.6 0–48.8 3.4 0.5 0–23.5
Litter vola (m3) 1.4 0.2 0.1–4.0 1.9 0.1 0.3–4.4
Nest patch (11.3-m radius)
Total canopy (%) 95.3 0.4 87.2–99.5 94.6 0.3 79.4–99.2
Sub-canopy (%) 66.8 3.6 13.8–97.3 61.5 2.1 17.0–97.0
Canopy ht (m) 27.3 1.1 14.6–45.3 24.2 0.8 14.8–52.4
Sub-canopy ht (m) 10.8 0.7 4.4–21.6 10.1 0.4 4.4–26.6
Medium–large treesb 5.0 0.4 1.0–9.0 5.5 0.3 0–13.0
Understory ht (m) 2.1 0.1 0–4.0 2.1 0.1 0–3.7
Nest-patch density (%) 75.3 3.0 34.3–99.7 74.5 1.6 27.3–99.0
Horizontal CVc 27.6 3.7 0.5–90.5 30.2 2.2 1.4–113.6
Vertical CVd 16.4 1.6 0.5–37.3 17.8 1.2 2.0–66.1
Patchiness
Horizontal differencee (%) 28.2 2.3 12.6–69.1 31.9 1.6 220.9–85.9
Cane differencef (stems/m2) 3.4 0.5 20.8–13.0 1.8 0.3 27.5–7.5
Noncane differencef (stems/m2) 0.3 0.5 24.5–7.0 0.8 0.4 29.8–12.3
Density differenceg (%) 45.3 4.0 26.8–90.7 30.1 2.6 243.6–91.6
Understory ht differenceg (m) 0.4 0.1 21.6–3.4 0.3 0.1 22.0–1.7
a
Litter vol 5 plot area 3 % leaf litter cover 3 litter depth in m; units are m3/plot.
b
No. of trees/11.3-m-radius plot.
c
Coeff. of variation in density-board estimates for the 4 cardinal directions averaged over all ht intervals.
d
Coeff. of variation in density-board estimates for the 5 ht intervals averaged over all horizontal directions.
e
Calculated as difference between nest-patch and nest-site density.
f
Calculated as the difference between the nest-site and paired random site values.
g
Calculated as the difference between the nest-patch and paired random patch values.

Appendix B. Mean, standard error, and range of values for landscape measurements taken at Swainson’s warbler nests at St. Francis National Forest (SFNF;
n 5 38) and White River National Wildlife Refuge (WRNWR; n 5 91), Arkansas, USA, 2005 through 2007.
SFNF WRNWR
Variable ¯x SE Range ¯x SE Range
Edge distance (m)
Agricultural 1,905 233 6–3,866 1,158 82 23–2,531
Any 35 6 3–131 85 9 0–436
Road 67 12 3–232 293 29 3–973
Road or trail 51 8 3–165 168 17 2–656
Stream 217 32 5–568 348 35 0–1,164
250-m buffer
% forest 97 1 80–100 94 1 54–100
Forest-edge density (m/ha) 15 3 0–62 15 2 0–72
Heterogeneitya 0.12 0.03 0–0.68 0.20 0.03 0–1.09
1,000-m buffer
% forest 91 1 76–100 85 2 44–100
Forest-edge density (m/ha) 26 2 4–56 15 1 1–34
Heterogeneitya 0.37 0.03 0.03–0.84 0.47 0.04 0–1.20
a
Heterogeneity 5 2Spilnpi, where pi is the relative proportion of the ith land-cover type (Magurran 1988).

Benson et al. N Swainson’s Warbler Reproductive Success 93