Biological Conservation 126 (2005) 363–370

www.elsevier.com/locate/biocon

Edge eVects on nesting success of cavity-nesting

birds in fragmented forests
a,¤ b
Wen-Hong Deng , Wei Gao
a
Ministry of Education Key Laboratory for Biodiversity Sciences and Ecological Engineering,
College of Life Sciences, Beijing Normal University, Beijing, 100875, China
b
College of Life Sciences, Northeast Normal University, Changchun, 130024, China

Received 11 February 2005

Available online 21 July 2005

Abstract

Forest fragmentation leads to the creation of isolated forest patches and habitat edges with subsequent impact on forest-interior
bird species. Although the eVects of fragmentation and edge on avian nesting success are well documented for open cup-nesting
species in eastern deciduous forests in North America, it is unclear whether these eVects are common for all birds nesting in pre-
dominantly forested landscapes. In particular, edge eVects on nesting success of cavity-nesting birds are poorly understood. Using
natural cavity nests, we examined nesting success of four species of cavity-nesting birds (two nonexcavators and two excavators),
the yellow-rumped Xycatcher (Ficedula zanthopygia), the great tit (Parus major), the great spotted woodpecker (Picoides major),
and the grey-faced woodpecker (Picus canus) in relation to forest edges in Zuojia Nature Reserve, Jilin province, northeastern
China. Our primary objective was to assess whether distance to the edge of agricultural lands was related to nesting success for cav-
ity-nesting birds in fragmented forests. A total of 439 natural cavity nests of the four species were located and monitored during
four breeding seasons. Probability of nest success was inXuenced by distance to forest edge for nonexcavators, but not for excava-
tors. The rate of nesting success of the two nonexcavators, yellow-rumped Xycatcher and the great tit, increased with distance from
the edges. For all cavity nests, nesting success was 0.43 at 0–100 m, 0.56 at 101–200 m, 0.68 at 201–300 m, 0.61 at 301–400 m, 0.77 at
401–500 m from the edges. Nesting success ranged from 0.57 for the yellow-rumped Xycatcher to 0.89 for the Grey-faced Wood-
pecker. Failed nests were often occupied by nest-site competitors (accounting for 68%). However, predation only accounted for
20% of all nest failures. Our results suggest that negative edge eVects do exist for some cavity-nesting birds, especially for nonexca-
vator species.
 2005 Elsevier Ltd. All rights reserved.

Keywords: Cavity-nesting birds; Nesting success; Edge eVects; Nest-site competition; Forest fragments

1. Introduction
Forest fragmentation aVects the distribution and abun-
dance of organisms by reducing the amount and
proximity of remnant patches of suitable habitat and
increasing the amount of edges (Andrén, 1992; Boulinier
*
Corresponding author. Tel.: +86 10 58805121; fax: +86 10
58807721.
E-mail address: dengwh@bnu.edu.cn (W.-H. Deng).
et al., 1998; Ford et al., 2001; Maina and Jackson, 2003).
Edges may aVect the organisms by causing changes in
the biotic and abiotic conditions some distance into the
forest, such as increased amounts of sunlight, high wind
speeds, and larger Xuctuations in temperature and
humidity (Saunders et al., 1991; Murica, 1995). Avian
species may respond to one or a combination of these
changes in the landscape as a result of diVerent biologi-
cal mechanisms (Robinson et al., 1995; Donovan et al.,
1997). Species that require forest interior may avoid

0006-3207/$ - see front matter  2005 Elsevier Ltd. All rights reserved.
doi:10.1016/j.biocon.2005.06.013
364 W.-H. Deng, W. Gao / Biological Conservation 126 (2005) 363–370
edges due to altered microclimate, vegetation structure,
or high density of predators or brood parasites
(Yahner and Scott, 1988; Malcolm, 1994; Marini et al.,
1995; Pasitschniak-Arts et al., 1998; Stephens et al.,
2003). On the other hand, although predators and brood
parasites may be more abundant or active in the adja-
cent habitat and edge, some bird species were concen-
trated near to edge areas. This paradox of high bird
abundance and richness but low nesting success near
edges was termed an ecological trap by Gates and Gysel
(1978).
Many studies have tested the occurrence of edge
eVects on nesting success of forest birds and concluded
that high predation and brood parasitism rates near
edges were main causes of nest failures (Hannon and
Cotterill, 1998; Manolis et al., 2002). However, an edge
eVect has not been found in all studies (Angelstam, 1986;
Santos and Telleria, 1992; Hanski et al., 1996; Morrison
and Bolger, 2002). Early studies suggested that abrupt or
permanent edges generally were thought to be associated
with higher rates of predation and parasitism than grad-
ual or regenerating edges (Suarez et al., 1997; Deng et al.,
2003), but many contradictory results exist, particularly
in forest landscapes (Yahner and Scott, 1988; Rodewald,
2002). These conXicting results may be explained in part
by the mediation of edge eVects by life history traits of
organisms (e.g., open- vs. cavity-nesting bids) and land-
scape characteristics.
Most early studies were completed on open cup-nest-
ing birds. However, nest predation and brood parasitism
are unlikely to provide a general explanation for the
edge eVects on bird species that nest in more protected
sites such as cavities (Matthysen and Adriaensen, 1998).
Nesting success of cavity-nesting birds near edges may
be inXuenced by other factors such as changes in compe-
tition for nest sites (Deng, 2001). Very few studies have
assessed nesting success of cavity-nesting birds near
edges in fragmented forests (Nour et al., 1993; Matthy-
sen and Adriaensen, 1998). Many studies of eVects on
nesting success have used artiWcial nests (Wilcove, 1985;
Picman and Schriml, 1994; Hartley and Hunter, 1998;
Githiru et al., 2005). Such studies can identify general
patterns of variation in nest predation intensity (Haskell,
1995; Sloan et al., 1998). However, predation and brood
parasitism rates on artiWcial nests may be poorly or
inconsistently correlated with predation and parasitism
rates on real nests (Ortega et al., 1998; Moore and
Robinson, 2004).
Here, we examine edge eVects on nesting success for
natural nests of four cavity-nesting bird species (two
excavators and two nonexcavators) in a forested land-
scape in which edges were created by farmland reclama-
tion. We also used data from these four species to
determine whether life history characteristics (excava-
tors vs. nonexcavators) aVect the relationship between
nest success and distances to edge.
2. Materials and methods
2.1. Study site
Our study site, approximately 200 km2 in size, was
located in Zuojia Nature Reserve and included the
Tumengling Mountains and Zhujia Mountains ranging
from the eastern Changbai Mountains to the western
plain (126°1⬘127°2⬘N, 44°6⬘45°9⬘E). The climate is east
monsoon, characterized by hot, dry summers and cold,
snowy winters. Mean monthly temperatures ranged
from ¡20.5 °C in January to 23.6 °C in August. Vegeta-
tion within the study area was quite diverse, although
the forest type was secondary forest. The age of the for-
ests is 40 years. The seven tree species mainly present on
the study area were Mongolian oaks (Quercus mongo-
lica), dahurian birchs (Betula davurica), Manchurian lin-
den (Tilia mandschurica), Japanese elm (Ulmus japonica),
Scotch pine (Pinus sylvestris), Korean larches (Pinus
koraiensis) and masson pines (Pinus massoniana) (Deng
et al., 2003). In the study area, Dahurian rose (Rosa
dahurica), Korean rose (Rosa d oreana), willowleaf spi-
raea (Spiraea salicifolia), ural falsespiraea (Sorbaria
sorbifolia), and sakhalin honeysuckle (Lonicera maxi-
mowiczii) dominated the shrub layer (Deng, 2001).
We established six study plots (mean size D11.2 ha,
range 7.6–32.8 ha) to evaluate edge eVects on nesting
success of cavity-nesting birds. The six study plots were
chosen because they had (1) similar vegetation chara-
cteristics (tree age, tree species, composition); (2) similar
management histories; and (3) similar ectones (embed-
ded farmland). The plot terrain was relatively Xat and
elevation ranged from 325 to 388 m above sea level. We
measured the distance from each nest to the forest edge
by pacing, with a known meter, or by measuring the dis-
tance on a gridded map of each plot.
2.2. Nest monitoring
Four to Wve people searched for nests daily in each
study plot from mid-April to the end of July during
1998–2001. Nests were found by observing birds during
nest building, food carrying and by searching tree holes.
Workers distributed their search eVort as evenly as pos-
sible throughout the plots. The location of each nest was
marked with a Xag that was usually placed at least 10 m
from the nest. Upon Wnding a nest, we recorded the stage
of nesting, location of the nest, and the species of the
bird. Nests were revisited every three or four days and
every other day near anticipated Xedging date to moni-
tor the fate of the nests. We checked the cavities using a
device consisting of mirrors and a torch. When checking
nests, we strived to avoid damaging vegetation and mak-
ing trails near nests. Nests that Xedged at least one
young were considered successful. Observation of Xedg-
ing, Xedglings near nests, and parents feeding Xedglings
 W.-H. Deng, W. Gao / Biological Conservation 126 (2005) 363–370
were taken as evidence of a successful nest. Nesting
attempts were considered failed if (1) there was clear evi-
dence of predation (broken egg, scattered feathers); (2)
the cavity was empty before Xedging was possible; (3)
chicks were never seen; (4) the cavity was taken over by
other species. If those cues were not available to deter-
mine success or failure, the outcome was considered
uncertain. Uncertain nest fate was rare in this study. We
also noted when nests were abandoned before the Wrst
egg was laid, although we did not consider these early
abandonments in our analyses. Daily survival rates were
estimated following MayWeld (1961, 1975).
2.3. Statistical analyses
Nesting success rates were estimated using the May-
Weld method (MayWeld, 1961, 1975). For nest fates con-
sidered uncertain, exposure period was terminated on
the last day that nests were known to be active. For nests
known to be successful or failed, exposure period was
terminated on the midpoint between the last day the nest
was observed as active and the Wrst day the nest was
observed as inactive. Because nest success rates often
vary between nesting stages, nest success was calculated
separately for the incubation and nestling periods.
Donovan et al. (1995) suggested if partial nest loss were
common, MayWeld estimates of nest success might not
be appropriate for calculating productivity. Partial nest
loss in this study was rare (<3%). Comparisons of May-
Weld nest success values between plots and years were
done using the computer program Contrast, with a
2
analysis and multiple comparisons (Sauer and Williams,
1989).
We grouped nests and compared nest success in two
zones (0–200 m vs. 201–500 m) because these contained
roughly equal numbers of nests for all species. Deng
(2001) suggested that a threshold edge eVect on nesting
success appears to exist for several bird species at
approximately 200 m in the study area. We used contin-
gency tables to test the hypothesis that nest success was
diVerent in the two zones. While we recognize the impor-
tance of species-speciWc nest success as a unit of analysis,
for some analyses, and to increase sample sizes, we
pooled species according to bird life history traits (i.e.,
nonexcavator or excavator). For comparisons of nest
success at diVerent distances from the edge, we ranked
the six plots in order of plot-level nest success for each
species. We then compared patterns of plot-level nest
success for nonexcavators and excavators. Also, we
tested for species-speciWc year and plot eVects using
logistic regression models. For species that showed sig-
niWcant diVerences among nest success in the two zones,
we then grouped nests into smaller zones and tested mul-
tiple hypotheses related to eVects at 100 m intervals (0–
100, 101–200, 201–300, 301–400, 401–500 m) from forest
edge. Because the number of nests further than 500 m
365
from the forest edges was small (<20 nests), we only used
the nests within 500 m from forest edges in the analysis
during this step. We used logistic regression to examine
the inXuence of distance to forest edge on nest fate. This
technique does not require that data from the indepen-
dent variable be normally distributed (Flaspohler et al.,
2001). For multi-species calculations of nest success, we
used an average weighted by exposure days to estimate
nest success for pooled groups of birds. For comparisons
of nest success at distances from the edge, we used
 D 0.05. Data were given as means §1 SE.
All logistic models were calculated with forward,
backward, stepwise, and score methods to achieve the
best model. All methods yielded similar results. To eval-
uate whether the relationship between nest success and
distance to forest edge was independent of bird species,
log-linear models were Wtted to three-way contingency
tables for all species. For all logistic regression analyses,
we used simple nest fate (success D 1 or failure D 0;
unoccupied D 1, occupied D 0), either from a Wrst or a
repeated nest. In all logistic regression models, we used
an information-theoretic approach and calculated
Akaike’s Information Criterion (AIC) based on log-like-
lihood values to identify the best models (Burnham and
Anderson, 1998). The model with the smallest AIC is the
best approximating model for the data. All statistical
analyses were conducted using SAS (1990).
3. Results
3.1. Nest success by distance to forest edge
A total of 439 cavity nests of four species were located
and monitored during 1998–2001 (Table 1). Of this total,
422 nests were entered into our analysis of edge eVects.
Nests abandoned before egg laying (n D 12) or of uncer-
tain outcome (n D 5) were excluded from analyses. Nest
success rates of each cavity-nesting bird did not vary
between years (
2 D 6.13, df D 3, P D 0.11) and plots
(
2 D 9.08, df D 5, P D 0.91). However, nest success rates
among species within years were signiWcantly diVerent.
Nest success varied among individual species, with the
lowest value for yellow-rumped Xycatcher and the high-
est for grey-faced woodpecker (Table 2). On all six plots,
excavator species had higher nest success than non-exca-
vator species (Table 2). For four years of pooled data,
nest success was lower in the 0–200 m zone than in the
201–500 m zone from the forest edge for the nonexcava-
tors (
2 D 6.94, df D 1, P < 0.01), but not for excavators
(
2 D 2.89, df D 1, P D 0.09). Logistic regression models
including a distance to forest edge variable provided a
better Wt to nest fate data for the yellow-rumped
Xycatcher and the great tit (Table 3), but not for the
great spotted woodpecker and the grey-faced wood-
pecker. The best models for yellow-rumped Xycatcher
366 W.-H. Deng, W. Gao / Biological Conservation 126 (2005) 363–370

Table 1
Nest fates of cavity-nesting birds in Zuojia Nature Reserve, Northeastern China, 1998–2001
Species and year Total nests Total nests Number (percentage) of nest failures, by cause
in analysis failed
Occupationa Predation Abandonmentb Uncertainc
Yellow-rumped Xycatcher
1998 19 9 7 (78) 2 (22)
1999 27 12 6 (50) 2 (17) 3 (25) 1(8)
2000 26 10 7 (70) 3 (30)
2001 23 7 5 (71) 2 (25)
All years 95 38 25 (66) 7 (16) 5 (13) 1 (5)
Great tit
1998 18 8 5 (62) 2 (25) 1 (13)
1999 21 9 7 (78) 2 (22)
2000 20 5 3 (60) 1 (20) 1 (20)
2001 25 10 5 (50) 2 (20) 1 (10) 2 (20)
All years 84 32 20 (63) 5 (16) 5 (16) 2 (5)
Great spotted woodpecker
1998 26 2 2 (100)
1999 27 6 4 (67) 2 (33)
2000 31 4 3 (75) 1 (25)
2001 34 5 5 (100)
All years 118 17 12 (70) 4(24) 1 (6)
Grey-faced woodpecker
1998 28 2 2 (100)
1999 30 3 2 (67) 1 (33)
2000 35 5 3 (60) 1(20) 1 (20)
2001 32 4 3 (75) 1(25)
All years 125 14 10 (72) 1 (7) 1 (7) 2 (14)
a
Nests were re-occupied by another species after laying.
b
Abandonment prior to laying.
c
If cues were not available to determine success or failure.

Table 2
Nest success estimates of cavity-nesting birds in Zuojia Nature Reserve, northeastern China, 1998–2001
Species Nest success § SEa
1998 1999 2000 2001 All yearsb
Yellow-rumped Xycatcher 0.67 § 0.08 (19)c 0.64 § 0.06 (27) 0.71 § 0.11 (26) 0.66 § 0.04 (23) 0.67 § 0.09 (95)
Great tit 0.72 § 0.12 (18) 0.69 § 0.09 (21) 0.79 § 0.06 (20) 0.66 § 0.12 (25) 0.73 § 0.10 (84)
Great spotted woodpecker 0.86 § 0.03 (26) 0.83 § 0.05 (27) 0.69 § 0.17 (31) 0.87 § 0.04 (34) 0.84 § 0.04 (118)
Grey-faced woodpecker 0.91 § 0.02 (28) 0.88 § 0.06 (30) 0.84 § 0.09 (35) 0.90 § 0.11 (32) 0.89 § 0.10 (125)
a
MayWeld (1961, 1975) nest success values calculated from exposure.
b

2 comparison of diVerences species for all years nest success (P > 0.05).
c
Number of nests.

included just the single predictor log distance to forest
edge, but for great tits, log distance and year were the
predictors that produced the best-Wtting model (Table 3).
The probability of failure for yellow-rumped Xycatcher
nests decreased with increasing distance to the edge in a
roughly linear function (Fig. 1), but this trend was not
apparent for other cavity-nesting species.
3.2. Causes of nest failure
Potential predators on cavity-nesting birds (eggs, nes-
tlings, adults) that were observed on the study plots dur-
ing four years were Eurasian sparrow hawk (Accipiter
nicus), besra sparrow hawk (Accipiter virgatus), long-
eared owl (Asio otus), Siberian weasel (Mustela aibirica),
Siberian chipmunk (Tamias sibiricus), and beauty snake
(Elaphe taeniura). The white-cheeked starling (Sturnus
cineraceus) was the most important nest-site competitor
for the four cavity-nesting bird species.
A total 101 failure nests occurred in our analyses.
Failed nests were often a results of nest-site competitors
(competitors which accounted for 68% of such losses).
Occupation rate by nest-site competitors was higher
(66%, n D 46) during the egg period than during the nest-
ling period (34%, n D 22). Seventy percent of nests lost to
competitors were lost to white-cheeked starlings. In con-
trast, nest predation only accounted for 20%. The four
species also suVered from nest site occupation by the
white-backed woodpecker (Picoides leucotos) and some-
times by Siberian chipmunk. Also, the occupation rate
 W.-H. Deng, W. Gao / Biological Conservation 126 (2005) 363–370 367

Table 3 1 yellow-rumped flycatcher

Logit transformation models of the nest fate for the yellow-rumped
Xycatcher and great tit during four breeding seasons in northeastern

Nest success
China 0.8

Models Number of Log-likelihood AIC

parameters 0.6
Yellow-rumped Xycatcher 21 19 14 16
25
FDC 1 ¡81.32 164.64
0.4
FDC+D 2 ¡78.26 160.52
F D C + LD 2 ¡77.49 158.98 1 great tit
F D C + LD + Y 3 ¡77.28 160.56
F D C + LD + P 8 ¡76.11 168.22

Nest success
F D C + LD + P + Y 9 ¡73.37 164.74 0.8

Great tit
FDC 1 ¡65.43 132.86 0.6
FDC+D 2 ¡64.26 132.52 20 12
23 16 13
F D C + LD 2 ¡64.11 132.22
0.4
F D C + LD + Y 3 ¡63.05 132.10
F D C + LD + P 8 ¡61.57 139.14 great spotted woodpecker
F D C + LD + P + Y 9 ¡59.44 136.88
F indicates nest fate; C indicates constant; D indicates distance to for- 1
est edge; LD indicates log10 (D); Y indicates studying years; P indicates
Nest success
plots; AIC indicates Akaike’s Information Criterion. F D C + LD is 0.8
the best Wt for the yellow-rumped Xycatcher; F D C + LD + Y is the
best Wt for the great tit. 0.6 31 26 22 19 20

0.4
by the white-cheeked starlings was signiWcantly greater
for the yellow-rumped Xycatcher in the 0–200 m zone grey-faced woodpecker
than the 201–500 m (
2 D 5.16, df D 1, P D 0.02). Nest pre- 1
dation did not follow this same trend (
2 D 1.18, df D 1,
P D 0.29) when comparing between the two zones.
Nest success

0.8
Lastly, we did not Wnd nest parasitism in this study.
0.6
29 39 12 23
22
4. Discussion
0.4
0-100 101-200 201-300 301-400 401-500
4.1. Edge eVects on cavity-nesting birds Distance to edge (m)

We found some evidence for an edge eVect on nesting
success of cavity-nesting species near anthropogenic
edges, especially for nonexcavator species. Our results
concur with Wndings from several recent studies on
open-cup nesting species (Flaspohler et al., 2001; Mano-
lis et al., 2002). Our results are also consistent with Wnd-
ings from several studies of cavity-nesting birds. Denny
and Summers (1996) found that cavity birds were
aVected by edge-related factors. Huhta and Jokimaki
(2001) found pairing success and nesting success of two
hole-nesting passerines, the pied Xycatcher (Ficedula
hypoleuca) and the cavity-nesting redstart (Phoenicurus
phoenicurus), were aVected by proximity to edge. How-
ever, Matthysen and Adriaensen (1998) reported that
reproductive success of the great tit was not aVected by
forest fragmentation in western Europe. Most early
studies of edge-related nest success of avian species sug-
gested that high nest predation and parasitism rates were
main causes of high nest failure near edges (Gates and
Gysel, 1978; Gibbs, 1991; Eriksson et al., 2001; Manolis
Fig. 1. MayWeld nest success for the four cavity-nesting species in rela-
tion to distance to the forest edge in Zuojia Nature Reserve, North-
eastern China, 1998–2001. The number of nests used to estimate nest
success is given inside each histogram bar.
et al., 2002). However, in our study, the main cause of
nest failure near edges was not nest predation or parasit-
ism, but rather, competition for nest sites among cavity
nesters. We also found that the rate of nesting success of
yellow-rumped Xycatcher and the great tit increased
with distance from the edges, but the great spotted
woodpecker and the grey-faced woodpecker did not fol-
low the same trend. This indicates local scale habitat per-
turbations can have species-speciWc eVects on nesting
success of cavity-nesting birds.
Several ideas have been proposed to explain the
absence of edge-related nest predation in forested land-
scapes. Angelstam (1986) suggested that edge-related
nest predation rates could be explained by diVerences in
productivity of the adjacent habitats. This explanation
368 W.-H. Deng, W. Gao / Biological Conservation 126 (2005) 363–370
indicated that an edge eVect on nest predation is most
likely to occur where there is a steep gradient in primary
productivity across the edge and is least likely to occur
where this gradient is less pronounced (Lahti, 2001).
Lack of predation may also occur if there is no increase
in prey density along edges or if predators are habitat
specialists (i.e., they do not move between habitats and
cross edges) (Rodewald, 2002). Morrison and Bolger
(2002) suggested that the lack of edge eVect on nest pre-
dation rate of Rufous-crowned Sparrow (Aimophila ruW-
ceps) appeared to be due to a lack of edge sensitivity of
snakes.
Excavator species (the great spotted woodpecker and
the grey-faced woodpecker) had signiWcantly higher nest
success than nonexcavator species (the yellow-rumped
Xycatcher and the great tit). Nonexcavator species are
generally smaller than excavator species. This smaller
size may reduce ability to compete for high quality nests.
Nonexcavator species may have had relatively low nest
success in part because they tend to nest in old cavities
and nest closer to the ground (Sonerud, 1985; Li and
Martin, 1991).
4.2. Nest-site competitors
Numerous studies of avian reproductive success have
concluded that the destruction of eggs and nestlings by
predators is the most signiWcant inXuence on avian nest
success (Askins, 1995). But in our results, nest predation
was not the main causes of nest failures and accounted
for only 20% of all nest failures. Most of the failed nests
(68%) were attributable to nest-site competitors. Indeed,
nest takeovers by the white-cheeked starlings were com-
mon both in edge and interior habitats. A potential cost
of cavity-nesting birds is increased competition for nest
sites, especially for nonexcavator species. Raphael and
White (1984) found a signiWcant correlation between
cavity density and nonexcavator species density in the
coniferous forests of the Sierra Nevada, suggesting den-
sities were limited by nest site availability. In our study
area, because the forest age was only about 40 years, lack
of natural tree cavities probably was the main reason for
drastic competition among the species, although we did
not investigate cavity resources throughout the study
area.
Some researchers found that some cavity-nesting
birds prefer edges and open understories in forested
landscapes (Belles-Isles and Picman, 1986; Finch, 1989).
Dobkin et al. (1995) explained that birds that use multi-
ple habitat types might nest on edges to reduce commut-
ing time between resources. Cavity nesters may select
nest sites with open understories to increase their ability
to detect approaching predators (Finch, 1989). We did
not Wnd the same phenomenon during our study period.
The number of cavity nests was roughly equal between
edge and interior habitat.
Brush (1983) observed Xycatchers, starlings, and
other species competing for nest cavities, but the star-
lings had no eVect on other species because abundant
alternate nest sites were available. However, the
potential for interference competition seems greatest
in habitats where nest sites are limited, as was the case
in our studies. Weitzel (1988) reported that starlings
were serious competitors for certain native birds in
Nevada. Kerpez and Smith (1990) found evidence for
competition and a negative relationship between star-
lings and gila woodpeckers (Melanerpes uropygialis)
in Arizona. Ingold (1998) suggested that the European
Starling (Sturnus vulgaris) was having a signiWciantly
adverse eVect on the reproductive success of northern
Xickers (Colaptes auratus). Despite these clear exam-
ples of starling having detrimental eVects on many
cavity-nesting species, Koenig (2003) did not Wnd that
starlings had had a severe impact on populations of
native cavity-nesting birds in a study of 27 species in
America.
Starlings may nest in forest, but they do not feed in
forests (Feare, 1984). They commute to open habitats
from their nest sites where they forage, just as does the
brown-headed cowbird (Molothrus ater) of North Amer-
ica. It seems likely that starlings prefer to nest near edges
because it reduces their travel costs from their breeding
areas to their feeding areas. Like the parasitic cowbird
phenomenon, this is a clear example of a situation in
which the matrix surrounding a fragment directly aVects
birds nesting within the fragment. The disturbed land-
scape supports a nest “pirate” that depends on the
matrix for food, but nest in forests.
Our results suggest that local land-use practices or
natural events can have a profound inXuence on the
nesting success of birds that depend upon disturbed hab-
itats for nesting. Our results also suggest that forest
management strategies should consider not only stands
in forest planning, but also the edges.
Acknowledgments
We thank Yan-Hui Li, Yang Liu, Zheng-Xin Sun,
Ren-Kai Song, Gui-Quan Xiang, Hai-Tao Wang, Jing-
Run Cheng, Nan Li, Quan Zhao, and Long Sun for
their involvement in the collection of Weld data. We
thank the teachers and students who are in Depart-
ment of Wildlife, Special Plant and Animal College
that they found and monitored the success of the nests.
Members of the ecology team, Beijing Normal Univer-
sity, were extremely patient and helpful in providing
assistance with data analysis. We thank fund of
National Nature Science of China to support this
research (No. 30470300). We also thank anonymous
reviewers for their helpful and excellent remarks on
this paper.
 W.-H. Deng, W. Gao / Biological Conservation 126 (2005) 363–370
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