Surviving in the city: higher apparent survival for urban birds

but worse condition on noisy territories

JENNIFER N. PHILLIPS ,1,2,  KATHERINE E. GENTRY ,3,4 DAVID A. LUTHER,4,5 AND ELIZABETH P. DERRYBERRY6
1
Department of Ecology and Evolutionary Biology, Tulane University, 400 Boggs, New Orleans, Louisiana 70118 USA
2
Department of Biological Sciences, California Polytechnic State University, 1 Grand Avenue, San Luis Obispo, California 93407 USA
3
Department of Biological Sciences, Purdue University, West Lafayette, Indiana 47906 USA
4
Biology Department, George Mason University, Fairfax, Virginia 22030 USA
5
Smithsonian Mason School of Conservation, 1500 Remount Road, Front Royal, Virginia 22630 USA
6
Department of Ecology & Evolutionary Biology, University Tennessee Knoxville, Knoxville, Tennessee 37996-1610 USA

Citation: Phillips, J. N., K. E. Gentry, D. A. Luther, and E. P. Derryberry. 2018. Surviving in the city: higher apparent
survival for urban birds but worse condition on noisy territories. Ecosphere 9(9):e02440. 10.1002/ecs2.2440

Abstract. Anthropogenic landscapes and soundscapes impose strong selective pressures on a number of
species, which can manifest in changes in vocalizations, foraging strategies, predator vigilance, and repro-
ductive success. However, few studies have examined survival rates, a major component of fitness, across
urban landscapes and soundscapes. White-crowned sparrows (Zonotrichia leucophrys) persist in both urban
and rural landscapes and change their behavior in response to the soundscape. We color-banded adult
white-crowned sparrows and collected noise levels on their territories in the urban San Francisco Bay Area
and adjacent rural Point Reyes National Seashore in California. We collected mark-encounter data on terri-
torial males from 2014 to 2017. Using the Program MARK, we tested the effects of habitat (urban/rural)
and territory noise level on annual survival rates and body condition. We predicted that survival and body
condition would be lower in urban habitats and decrease with increasing background noise level on terri-
tories. We found that survival estimates vary according to year, and males in urban landscapes have higher
survival. Noise levels best predict body condition, such that soundscape negatively correlates with male
body condition. Taken together, the urban landscape and soundscape shape the survival and health of
birds in and near cities.

Key words: body condition; MARK; soundscape; survival; urban ecology; white-crowned sparrow.

Received 28 March 2018; revised 10 August 2018; accepted 20 August 2018. Corresponding Editor: Paige Warren.
Copyright: © 2018 The Authors. This is an open access article under the terms of the Creative Commons Attribution
License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.
  E-mail: jnphilli@calpoly.edu

INTRODUCTION around tall structures such as buildings with

Environmental conditions affect the relative fit-
ness of populations, which is measured by repro-
duction and survival. On an evolutionary
timescale, urban habitats are relatively new and
create novel selective pressures on animal popula-
tions that could affect their fitness (Swaddle et al.
2015). For example, in cities, some animals use
new types of ornamental plants for food and nest
sites (Corlett 2005), interact and compete with
introduced species and predators that thrive in
cities (Koenig 2003, Shochat et al. 2010), navigate
many reflective surfaces (Jung and Kalko 2010),
and are affected physiologically by environmental
pollutants where body condition is decreased in
urban areas (Bailly et al. 2017). Human activities
also pose novel soundscapes or background
noise, and many animals modify communication
behaviors and predator vigilance in anthro-
pogenic noise (Luther et al. 2015, Ware et al. 2015,
Kleist et al. 2016). These adjustments to noise
may ultimately affect relative fitness. Therefore,
the evolutionarily recent urban landscape and
soundscape may have various costs and benefits,

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and researchers are just beginning to understand
how urban areas affect relative fitness.
Anthropogenic environments are thought to
have negative effects on wildlife by decreasing
survival (Aronson et al. 2014). Domestic preda-
tors, such as cats, are often abundant, which can
have profound effects on wildlife (Loss et al.
2013). Parks, yards, and agricultural fields are
often treated with pesticides (Moore 1967, Kohler
and Triebskorn 2013). The fragmentation of suit-
able habitat makes it difficult for juveniles to dis-
perse (Fischer and Lindenmayer 2007), and
roadways can cause a high occurrence of mortality
across taxa (Fahrig and Rytwinski 2009). The land-
scape is not the only thing that changes with
anthropogenic activity—soundscapes also change.
Urban soundscapes are dominated by the
noise produced by traffic, generators, and the
day-to-day hum of other human-built machines.
Correspondingly, animals, such as birds, increase
the pitch (Slabbekoorn 2013, Derryberry et al.
2016) and change the tempo of their vocaliza-
tions (Luther and Derryberry 2012, Slabbekoorn
2013), probably in order to increase signal detec-
tion for receivers in noise. Nestlings raised in
noisy conditions are exposed to increased stres-
sors and have decreased body condition, which
may have cascading effects into future reproduc-
tive success (Crino et al. 2013, Salmo n et al.
2016, Raap et al. 2017). Animals may also have
to change their territorial behaviors in noisy
urban conditions, such as approaching more clo-
sely to be able to hear an acoustic signal, which
could lead to aggressive territorial interactions
and potentially affect body condition, reproduc-
tion, and survival (Phillips and Derryberry 2018).
Although the detriments of anthropogenic
activity may be numerous, anthropogenic envi-
ronments may also have some benefits to wild-
life. Supplemental feeding, such as backyard bird
feeders, can increase the value of otherwise pat-
chy habitat (Marzluff and Rodewald 2008).
Although predator abundance can be higher in
cities, predator diversity is often lower which
might help explain the increased survival of
some species in cities (Moller 2005, Shochat et al.
2006). Humans also often introduce nest boxes
which can increase important nesting resources
and thus the ability of species to reproduce in
urban habitats (Vincze et al. 2017). Great tits
(Parus major) are one such cavity nester that
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PHILLIPS ET AL.
seems to benefit from urban habitats (Horak and
Lebreton 1998). However, there are some costs to
these activities. For example, supplemental feed-
ing may favor introduced species (Galbraith
et al. 2015), increase disease transmission (Robb
et al. 2008), or increase predator abundance at
feeding locations (Malpass et al. 2017). Thus, the
true survival trade-offs of an urban life for many
wild species remain unknown.
A handful of studies have examined differ-
ences in survival between urban and rural popu-
lations (Horak and Lebreton 1998, Ryder et al.
2010, Stracey and Robinson 2012, Evans et al.
2014), yet few have delved deeper into specific
pressures associated with urban life. In particular,
no study has examined whether both anthro-
pogenic habitat (landscapes) and anthropogenic
noise (soundscapes) together influence adult ani-
mal survival and body condition (Shannon et al.
2016). Here, we test whether anthropogenic land-
scapes and soundscapes affect annual survival in
Nuttall’s white-crowned sparrows (Z. l. nuttalli).
Because Nuttall’s white-crowned sparrow is a
year-round resident with high site fidelity (Bap-
tista 1975, Dewolfe et al. 1989), and occurs in
both urban and rural areas, it is an ideal subject
to test for a relationship between survival, land-
scape, and soundscape. We focus only on males,
as females disperse from the natal area, but males
tend to hold a territory close to their natal area
for their lifetime (Petrinovich and Patterson 1982).
First, we predict that urban areas will have lower
apparent survival and body condition, because of
higher mortality due to a number of potential fac-
tors, such as environmental pollution, increased
predator abundance, and higher vehicular colli-
sion rates. Alternatively, urban areas may
increase survivorship and body condition, poten-
tially due to an increase in anthropogenic year-
round food resources (i.e., bird feeders and trash).
Second, we predict that males holding territories
with relatively high noise levels will have lower
survival and body condition, potentially due to
an inability to detect approaching predators and
increased vigilance and/or stress levels.
METHODS
Study locations and survey methods
We captured 258 adult male white-crowned
sparrows during the breeding seasons across
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 PHILLIPS ET AL.

2014, 2015, and 2016 in two regions—urban San
Francisco, California (N = 162), and rural Point
Reyes, California (N = 96; Fig. 1). Point Reyes
National Seashore consists of a mix of grassland,
native scrub, and Douglas fir forest (Pseudotsuga
menziesii) with some agricultural structures scat-
tered throughout the landscape (Fig. 1, upper left
portion of map). San Francisco is the fourth lar-
gest and the most densely populated city in Cali-
fornia, with little native vegetation remaining
outside of public parks (Fig. 1, lower right por-
tion of map). We considered sites within these
two regions to be urban or rural based on noise
at the regional level (Lee and MacDonald 2011,
2013) following (Gentry and Luther 2017, Fig. 1).
We sampled four rural sites and five urban sites
(Fig. 1, Table 1). We captured males using mist
nets and playback of song or alarm calls. Age
was determined by crown plumage, where sec-
ond-year Z. l. nutalli males often retain some
brown in the crown, and after second year
(ASY), birds have white and black crowns (Pyle
1997). A second-year bird or older is considered
a breeding adult. Sex was determined by cloacal
protuberance during banding and confirmed
with subsequent singing and territory defense
behaviors. We took morphological measures,
including tarsus length and mass, and calculated
scaled mass index (SCI; Peig and Green 2009).
Each male received a unique combination of four

Fig. 1. Study locations in the San Francisco Bay Area (urban) and Point Reyes National Seashore (rural) in
California, USA. Map created in ArcMap 10 using ESRI World Imagery. (Sources: ESRI, DigitalGlobe, GeoEye,
Earthstar Geographics, CNES/Airbus DS, USDA, USGS, AeroGRID, IGN, and the GIS User Community).

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 PHILLIPS ET AL.

Table 1. Banding data and noise levels for each urban and rural location by year.

Year Location No. banded No. males Avg. noise (LAeq) Noise range (LAeq)

Urban
2014 Golden Gate 10 7 59.3 59.3
2014 Lobos Dunes 24 19 48.4 43.7–52.0
2014 Inspiration Point 23 20 45.3 42.1–49.6
2014 Lake Merced–Fort Funston 41 29 54.4 47.0–65.7
2015 Lobos Dunes 11 6 48.3 43.6–5
2015 Land’s End 8 5 50 46.1–58.5
2015 Lake Merced–Fort Funston 1 0 58.5 45.5–66.8
2016 Golden Gate 32 30 56.1 45.0–66.3
2016 Lobos Dunes 13 10 56.4 46.1–60.9
2016 Lake Merced–Fort Funston 27 21 55.1 45.9–65.1
2016 Richmond 17 15 51.3 45.7–57.9
All Urban 207 162 53.0 42.1–65.7
Rural
2014 Commonweal 30 16 39.8 35.1–45.4
2014 Abbott’s Lagoon 22 8 40.1 38.0–44.1
2014 Schooner Bay 14 8 44.7 42.2–47.0
2015 Abbott’s Lagoon 8 8 40.9 40.0–41.8
2016 Commonweal 28 21 47.8 41.3–57.2
2016 Abbott’s Lagoon 31 21 42.9 35.9–50.3
2016 Limantour 20 14 47.4 42.3–53.4
All Rural 153 96 43.4 35.1–57.2
Total 360 258
Notes: No. banded includes total banded (males, females, and juveniles), while No. males indicates only the number of
males. Of the 87 rural and 154 urban non-transient males banded, 42.5% and 62.3% were resighted across years, respectively.
Average noise is the mean noise for that site in a given year, and noise range shows the lowest and highest noise levels in a
given site for that year.

colors, including a silver Fish and Wildlife alu-
minum band (Permit 23900). After banding, sites
were revisited in the same year to determine
whether males were holding territories on or
near the site of banding or whether they were
transient. If males were never seen after banding,
they were considered transient and were
excluded from analyses. Territories were re-
surveyed opportunistically. Each neighboring
territory was also searched, and when new
unbanded males were observed, they were pre-
sumed to have taken over the territory.
Noise level data
To estimate territory background noise, we
used a calibrated sound-level meter (SLM; Lar-
son Davis model 831, firmware version 2.206
with the settings of A-frequency weighting, fast
time weighting, linear averaging, and 0.0 dB
gain) that calculated 1 s LAeq, or A-weighted
equivalent continuous level value, based on the
sound pressure levels (SPL) sampled 10 times a
second. For each territory, a four-minute (240 s)
A-weighted equivalent continuous level value
(LAeq) was calculated from the 1–s LAeq values,
where the meter was pointed one minute in each
cardinal direction (Brumm 2004). LAeq is a mea-
surement of equivalent continuous sound that
contains the same sound energy as a noise that
varies over time. The A-weighting filter is an
international standard that approximates the
human hearing range, which is similar to avian
hearing (Dooling and Popper 2007). All reported
sound pressure levels are in absolute units of
LAeq dB re: 20 lPa (8–20 kHz). These territory
noise levels were recorded opportunistically at
the time of banding or during subsequent
resighting surveys or before playback experi-
ments. All noise readings were taken in the
absence of wind noise (Beaufort Wind Scale of 3
or less) and away from singing birds within
30 m (Francis et al. 2012). Because noise levels
were sampled opportunistically, not all males
had an individual territory noise level value. For
these males (N = 59), we instead used a calcu-
lated site average noise level value.

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 PHILLIPS ET AL.

MARK analysis and statistics
We analyzed the mark-encounter (resight) data
in Program MARK (White and Burnham 1999) to
investigate the effects of noise and habitat on
apparent annual survival (Ф), and we considered
the effect of habitat on resighting probability (p).
Hereafter, we use “survival” to describe appar-
ent annual survival. We used a Cormack-Jolly-
Seber data structure to fit a fully grouped and
time-dependent global model and then tested for
goodness of fit using the median c-hat approach.
The median c-hat generated from the goodness-
of-fit test confirmed model fit (^c = 1.51,
SE = 0.06), so we constructed a set of 15 candi-
date models based on the global model structure
(Table 2). The influence of habitat, territory noise
level, and temporal period (year) was considered
when developing the candidate model set. Can-
didate models with constant and time-dependent
survival parameters were included, as well as
both interactive and additive effects. Resighting
effort was consistent across years, so we did not
include candidate models with a time-dependent
resighting parameter.
An information-theoretic approach and multi-
model inference was used to account for model
selection uncertainty and to obtain robust parame-
ter estimates. Candidate models were ranked using
the Quasi-likelihood Akaike’s information criterion
(QAIC) based on the variance inflation factor
(^c = 1.51; White and Burnham 1999). Full-model
averaging was used to estimate relative variable
importance (RVI; Burnham and Anderson 2002).
Apparent survival and resighting estimates were
model-averaged across all candidate models, and
results were interpreted based on the weighted
average annual estimate, unconditional standard
errors (SEs), and 95% confidence intervals.
Body condition
We estimated whether body condition was
influenced by landscape (habitat type) or sound-
scape (noise level; N = 249). Males that lacked
morphological data or site ambient noise levels
were dropped from the analysis. To reliably esti-
mate body condition, we calculated a scaled
mass index following Peig and Green (2009).
All statistical analyses were done in R
(R Core Team 2016). The qqp function in pack-
ages car and MASS was used to confirm our data
were normally distributed (Venables and Ripley
2002, Fox and Weisberg 2011). Akaike’s informa-
tion criterion adjusted for small sample sizes
(AICc) was used to examine the fit of linear
mixed models by maximum-likelihood using
nlme (Pinheiro et al. 2013), where we tested all
combinations of habitat and territory noise level
as fixed effects and site as a random effect. Linear
mixed effects model residuals were examined
visually in QQ-plots for validation.
Models were ranked according to the strength
of support of each model, and models with delta

Table 2. QAIC candidate models.

Candidate model DQAIC QAICc Model weight Likelihood K QDeviance

{Ф(time) p(.)} 294.39 0 0.46 1 4 286.27

{Ф(time) p(habitat)} 296.45 2.06 0.16 0.36 5 286.27
{Ф(habitat 9 time) p(.)} 296.56 2.17 0.15 0.34 7 282.23
{Ф(habitat 9 time + LAeq),p(.)} 298 3.61 0.07 0.16 8 281.57
{Ф(habitat 9 time) p(habitat)} 298.66 4.27 0.05 0.12 8 282.23
{Ф(habitat 9 time + LAeq)p(habitat)} 300.11 5.72 0.03 0.06 9 281.574
{Ф(habitat) p(.)} 300.56 6.17 0.02 0.05 3 294.49
(NULL MODEL){Ф(.) p(.)} 301.13 6.74 0.02 0.03 2 297.09
{Ф(LAeq)p(.)} 302.27 7.89 0.009 0.02 3 296.21
{Ф(habitat + LAeq)p(.)} 302.60 8.22 0.008 0.02 4 294.49
{Ф(.) p(habitat)} 303 8.61 0.006 0.01 3 296.93
{Ф(habitat 9 LAeq)p(.)} 304.06 9.67 0.003 0.007 5 293.88
{Ф(LAeq)p(habitat)} 304.32 9.93 0.003 0.007 4 296.21
{Ф(habitat + LAeq)p(habitat)} 304.67 10.28 0.003 0.006 5 294.49
{Ф(habitat 9 LAeq)p(habitat)} 306.13 11.74 0.001 0.003 6 293.88
Notes: Model parameters are denoted as Ф() for survival and p() for resighting, and “.” denotes a constant. LAeq is the noise
measurement parameter. The top model is time, with the second model including time + habitat.

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 PHILLIPS ET AL.

AICc < 4 were considered to be equally well followed by the interactive effect of time and
supported. For the models with delta AICc < 4, habitat (RVI = 0.31). The noise level LAeq param-
we calculated model-averaged parameter esti- eter and its interactive effect with habitat were
mates with unconditional standard errors (SEs) the least important predictor variables of sur-
and confidence intervals using the AICcmodavg vival (RVI = 0.21, RVI = 0.05, respectively),
R package (Mazerolle 2016). We report the rela- which suggests that there is not a strong effect of
tive variable importance (RVI) of model parame- background noise level on survival in either
ters and consider a fixed effect to influence body urban or rural habitats. The resighting probabil-
condition if the 95% confidence interval of its ity was estimated on the boundary (1.00), and
parameter estimate did not include zero (Naka- thus, the standard error could not be computed
gawa and Cuthill 2007). We evaluated the signifi- properly; a 95% profile likelihood interval for
cance of models with delta AICc lower than 4 constant resighting probability in the top-ranked
using an F test with Kenward-Roger approxima- model was estimated as (0.999, 1.000).
tion in comparison with a null model through
the KRmodcomp function in the pbkrtest pack- Soundscape has a greater effect on body
age (Halekoh and Højsgaard 2014). We used the condition than landscape
doBy package (Højsgaard and Halekoh 2016) to The top model for body condition included
compute least squares means for habitat using only noise level as a fixed effect with an evidence
adjusted degrees of freedom. ratio of 45.2 over the null model (Table 4) and is
highly significant (F1, 103 = 9.2, P = 0.003). The
RESULTS next models within 4 AICc were Habitat and
then Noise 9 Habitat (Table 4). Model averaging
Landscape but not soundscape affects survival of the most supported models suggests that body
The top-ranked model (Фtime, p.) had an AICc condition significantly decreases as noise levels
weight of 46%, with an evidence ratio of 2.88 rel- increase (Fig. 3), and noise is more important
ative to the next ranked model (Фtime, phabitat; than habitat for scaled mass index (noise:
Table 2). Of the survival parameters, time was by b  SE = 0.07  0.02, 95% CI = 0.1, 0.02,
far the most important (RVI = 0.93), which indi- RVI = 0.98, LMS estimate = 31.4  0.27; habitat:
cates survival estimates vary according to year. b  SE = 0.01  0.09, 95% CI = 1.4, 0.3,
For each year, the urban survival rate was esti- RVI = 0.02, LMS  SE = 31.38  0.42).
mated to be consistently higher than the rural
survival rate (see Table 3 for urban and rural sur- DISCUSSION
vival estimates per year; Fig. 2). The survival
parameter habitat ranked next highest in terms Despite many anthropogenic stressors in cities,
of importance (RVI = 0.35) and was closely we found that urban male birds have a higher
survival rate than rural male birds. Territory
noise level did not have a strong effect on sur-
Table 3. Model-averaged apparent survival (Ф) and
recapture (p) estimates, unconditional standard error
vival relative to time or habitat type. However,
(SE), and the 95% confidence interval for urban and
the noise did have an important effect on body
rural male white-crowned sparrows for each study
condition, such that males holding territories
with higher noise levels had lower body condi-
year time interval from 2014 to 2017 (from MARK).
tion. Together, our results show that anthro-
Parameter Estimate SE Lower Upper pogenic landscapes affect apparent survival and
Urban: Ф soundscapes affect our measured index of body
Time 1 0.68 0.07 0.52 0.80 condition, which may lead to long-term evolu-
Time 2 0.74 0.10 0.52 0.88 tionary consequences for populations adapting
Time 3 0.47 0.06 0.36 0.59 to city life.
Rural: Ф Although it is well known that many species
Time 1 0.59 0.13 0.34 0.80 cannot persist in cities, there is mounting evi-
Time 2 0.71 0.12 0.44 0.88
dence that some have higher survival in urban
Time 3 0.46 0.06 0.34 0.58
than in rural habitats (Chamberlain et al. 2009,

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 PHILLIPS ET AL.

Fig. 2. Mean and SE of the number of years resighted for male white-crowned sparrows banded in (a) 2014
(Rural N = 32, Urban N = 75) with 4 yr of resight surveys; (b) 2015 (Rural N = 8, Urban N = 11) with 3 yr of
resight surveys; and (c) 2016 (Rural N = 56, Urban N = 76) with 2 yr of resight surveys. Overall, urban males
tend to live longer than rural males, although 2015 has a slight opposite trend when less birds were marked
compared to other years.

Table 4. AICc model ranking for body condition (N = 249).

Model K AICc DAICc AICcWt Wi Evidence ratio Likelihood

Noise 4 1001.90 0.00 0.64 0.64 45.7 496.87

Habitat 5 1003.90 2.10 0.22 0.87 15.7 496.87
Noise 9 Habitat 6 1005.50 3.60 0.10 0.97 7.1 496.59
Null 3 1009.50 7.60 0.014 0.99 1 501.71
Noise + Habitat 4 1009.80 7.90 0.012 1.00 0.9 500.83
Note: Noise is the top model, followed by Habitat, then Noise 9 Habitat.

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Fig. 3. Scaled mass index is a measure of body con-
dition (y-axis), and LAeq is a measure of background
territory noise (x-axis). As noise increases, body
condition decreases in both urban and rural birds
(N = 239).
Marzluff 2017). A number of non-exclusive
reasons could explain higher urban survival (re-
viewed in Marzluff 2017), but one important rea-
son in the white-crowned sparrow system might
be year-round food resources in urban habitats
compared to fluctuating food resources in rural
habitats. Seasonal changes and environmental
events, such as drought, affect rural areas more
than urban areas (Diffenbaugh et al. 2015, Mann
and Gleick 2015). Notably, there was a significant
drought in the region during the years of the
study. Therefore, it is likely that irrigated urban
managed lands provide more food and less
vegetation die-off (i.e., more cover) than rural
non-irrigated areas. In times of limited resources
such as drought, birds may be putting more effort
into finding food resources, rather than increas-
ing signaling opportunities provided by quiet
areas, which may play a more important part in
overall fitness. These landscape-level factors may
play an important role in why habitat is more
important than noise in our survival models.
Similarly, disproportionate supplemental feed-
ing in cities can offset seasonal variation and has
the potential to increase adult survival, allowing
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PHILLIPS ET AL.
less fit individuals to breed and reproduce, lead-
ing to differences in survival between urban and
rural populations (Evans et al. 2014, Marzluff
et al. 2016). Supplemental feeding was observed
in at least two of our five urban sites (Lake
Merced, Land’s End) on multiple occasions, and
the remaining urban sites have backyard bird
feeders and trashcans nearby that may also pro-
vide food sources. However, even if food sources
are abundant in cities, anthropogenic food
sources may be less nutritious, potentially lead-
ing to lower body condition (Meillere et al.
2015).
Our data are the first to support reduced body
condition on noisy territories. We found lower
body condition among males holding territories
in areas with higher noise levels in both urban
and rural habitats. Our results are consistent with
other studies that found reduced body size, mass,
or condition in urban conditions (Liker et al. 2008,
Meillere et al. 2015), but we suggest that noise,
rather than solely habitat, may be the driver. For
example, one study did not find nutritional stress
differences between urban and rural adult house
sparrows (Passer domesticus), but juvenile fat
scores were reduced in urban areas (Meillere et al.
2015). Similarly, food supplementation in cities
was associated with reduced body condition in
great tits (Parus major; Demeyrier et al. 2017).
These studies suggest urban habitats, food sup-
plementation, and predation pressures as mecha-
nisms to explain lower body condition in urban
areas, but did not consider background noise as a
contributing factor. Our data suggest that future
studies should also consider territory noise levels
and test the relative impact of these different
potential mechanisms on body condition.
There are several potential mechanisms that
could explain how noise levels affect body condi-
tion. Physiological studies show noise can be a
stressor for birds (e.g., fecal corticosteroid
metabolites, Blickley et al. 2012, glucocorticoids
reviewed in Potvin 2017), and stressors often
lead to lower body condition (e.g., increased cor-
ticosterone and decreased fat stores; Wingfield
et al. 1983). A recent study links exposure to
noise with decreased baseline corticosterone in
nestlings and female birds and increased acute
stressor-induced corticosterone in nestlings,
which suggests chronic stress (Kleist et al. 2018).
In addition, high levels of anthropogenic noise
8 September 2018 ❖ Volume 9(9) ❖ Article e02440

tend to result in increased vigilance and less time
foraging for food, which could also lower body
condition (Ware et al. 2015). Predator abundance
or diversity may change based on noise levels
and also could affect vigilance behavior and con-
dition (Shochat et al. 2006, Francis et al. 2012).
Experimental work is needed to test the relative
importance of these different potential mecha-
nisms linking the soundscape to body condition.
Our finding that noise levels do not affect
apparent survival of male white-crowned spar-
rows is not consistent with previous studies in
other species. One study that removed the con-
founding effects of urban–rural dynamics while
retaining increased noise levels showed that
predator–prey dynamics changed due to noise,
such that small birds that nested closer to the
noise source were less affected by predatory
birds (Francis et al. 2011). Such a finding sug-
gests increased survival in areas with higher
noise levels. Crino et al. (2013) found that
increased noise in a natural forest positively
affected chick condition, which may give birds a
survival advantage, although survival was not
measured. Thus, while noise is not the most
important factor driving our apparent survival
models, it may still play an active and complex
role in shaping how animals such as small birds
survive in a city.
We examined adult males, but other life stages
may also be sensitive to urban soundscape and
landscape conditions. Chronic noise is known to
elevate haptoglobin immune response, which can
be energetically costly (Raap et al. 2017),
although a recent study found no correlation
between decreased body size and nutritional
stress (Meillere et al. 2015). Other recent studies
show that fledglings are negatively affected by
suburbs, which act as sinks rather than sources,
due mainly to cats and crows (Balogh et al. 2011).
Although adult male survival is increased in our
urban study area, the first year of life might be
much tougher for urban than for rural birds. For
a fuller picture of how fitness and survival shape
populations in cities, future work should continue
to examine body condition and survival across
age classes, including adult females, fledglings,
and first-year birds. Together, such data would
allow a more complete picture of the evolutionary
consequences of city life.
❖ www.esajournals.org
PHILLIPS ET AL.
CONCLUSIONS
Anthropogenic landscapes and soundscapes
selected for higher survival but worse body con-
dition in individual adult male white-crowned
sparrows. Our results are the first to show that
body condition of adult birds decreases with
noise level, suggesting that exposure to higher
noise levels in the city may have fitness conse-
quences. While these consequences were not seen
in our four-year measure of apparent survival,
further research should combine survivorship of
adults with monitoring of nestlings, fledglings,
and adult females to better understand the long-
term impact of cities on populations and fitness
of animals. Understanding whether noise levels
affect different life stages separately or
cumulatively over a lifetime can elucidate overall
fitness impacts of anthropogenic activities on
populations.
ACKNOWLEDGMENTS
The authors would like to thank Kathleen Grady,
Michael Kryzwicki, Irene Koulouris, Mae Berlow, and
Leanne Norden for field assistance. Point Blue staff
Diana Humple and Mmark Dettling provided training
and housing, while Bill Merkle, Michael Chasse
(Golden Gate National Recreation Area), Ben Becker
(Point Reyes National Seashore), Doug Bell (East Bay
Regional Parks), and Lisa Wayne (San Francisco Parks
and Recreation) provided site access and permits. This
work was funded by NSF IOS—1354763 and 1354756,
Tulane Gunning Award, Tulane One-Term Dissertation
Award, Wilson Ornithological Society Grant, and an
American Ornithologist’s Union Van Tyne Award. JNP,
KEG, and DAL collected data, JNP and KEG analyzed
data, JNP wrote the manuscript with input and final
approval from KEG, DAL, and EPD. The authors
declare no conflict of interest.
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