EFFECTS OF NEST HABITAT, FOOD, AND PARENTAL

BEHAVIOR ON SHOREBIRD NEST SUCCESS

Author(s): PAUL ALLEN SMITH, H GRANT GILCHRIST, JAMES N.M
SMITH
Source: The Condor, 109(1):15-31. 2007.
Published By: Cooper Ornithological Society
DOI: http://dx.doi.org/10.1650/0010-5422(2007)109[15:EONHFA]2.0.CO;2
URL: http://www.bioone.org/doi/
full/10.1650/0010-5422%282007%29109%5B15%3AEONHFA%5D2.0.CO
%3B2

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The Condor 109:15–31
# The Cooper Ornithological Society 2007

EFFECTS OF NEST HABITAT, FOOD, AND PARENTAL

BEHAVIOR ON SHOREBIRD NEST SUCCESS
PAUL ALLEN SMITH1,3, H. GRANT GILCHRIST2, AND JAMES N. M. SMITH1,4
1
Department of Zoology, University of British Columbia, Vancouver, British Columbia, V6T 1Z4, Canada
2
National Wildlife Research Centre, Carleton University, Ottawa, Ontario, K1A 0H3, Canada

Abstract. In environments such as arctic tundra, where bird densities are low and
habitats are comparatively homogeneous, suitable nest sites likely are not limited. Under
these conditions, reproductive success of birds may be determined by factors other than
the habitat characteristics of nest sites. We studied the relative influence of nest habitat,
food, nest distribution, and parental behavior on the reproductive success of tundra-
breeding shorebirds at East Bay, Southampton Island, Nunavut, Canada. From 2000 to
2002, we monitored the nests of five species: Black-bellied Plover (Pluvialis squatarola),
Semipalmated Plover (Charadrius semipalmatus), Ruddy Turnstone (Arenaria interpres),
White-rumped Sandpiper (Calidris fuscicollis), and Red Phalarope (Phalaropus fulicarius).
For each species, habitat differed between nest sites and random sites. In contrast, habitat
differed between successful and failed nest sites only for White-rumped Sandpipers.
Shorebirds did not prefer to nest in habitats where food was most abundant. Although
nest success varied among species in all years, artificial nest experiments suggested that
interspecific variation in predation rate was not related to habitat type. Instead, the
marked interspecific variation in nest success may have been related to incubation
behavior. Species taking fewer incubation recesses had higher nest success, although these
results should be viewed as preliminary. The factor with the greatest interannual influence
on nest success was fluctuating predation pressure, apparently related to the abundance of
predators and lemmings.
Key words: incubation behavior, nest distribution, nest habitat, nest predation, nest site
selection, shorebird.

Efectos del Hábitat de Nidificación, el Alimento y el Comportamiento Parental Sobre el Éxito

de Nidificación de Aves Playeras
Resumen. En ambientes como la tundra ártica, donde las densidades aves son bajas y
los hábitats son comparativamente homogéneos, los sitios de nidificación adecuados
probablemente no son limitados. Bajo esas condiciones, el éxito reproductivo de las aves
podrı́a estar determinado por factores distintos a las caracterı́sticas del hábitat en donde se
ubican los nidos. Estudiamos la influencia relativa del hábitat de nidificación, el alimento,
la distribución de los nidos y el comportamiento parental sobre el éxito reproductivo de las
aves playeras que crı́an en la tundra en East Bay, Southampton Island, Nunavut, Canadá.
Entre 2000 y 2002, monitoreamos los nidos de cinco especies: Pluvialis squatarola,
Charadrius semipalmatus, Arenaria interpres, Calidris fuscicollis y Phalaropus fulicarius.
Para cada especie, el hábitat difirió entre los sitios de nidificación y sitios ubicados al azar.
En contraste, el hábitat difirió entre nidos exitosos y no exitosos sólo en C. fuscicollis. Las
aves playeras no prefirieron anidar en hábitats en donde el alimento era más abundante.
Aunque el éxito de los nidos varió entre las especies en todos los años, experimentos
realizados con nidos artificiales sugirieron que la variación interespecı́fica en la tasa de
depredación no estuvo relacionada con el tipo de hábitat. En cambio, la marcada
variación interespecı́fica en el éxito de nidificación podrı́a haber estado relacionada con el
comportamiento de incubación. Aunque los resultados deben considerarse preliminares,
las especies que tomaron menos recesos de incubación tuvieron un éxito de nidificación
mayor. El factor con la mayor influencia interanual sobre el éxito de nidificación fue la
presión de depredación fluctuante, la cual aparentemente se relacionó con la abundancia
de depredadores y roedores.

Manuscript received 16 May 2006; accepted 24 October 2006.

3
Present address: National Wildlife Research Centre, Carleton University, Ottawa, Ontario, K1A 0H3,
Canada. E-mail: Paulallen.smith@ec.gc.ca
4
Deceased.

[15]
16 PAUL A. SMITH ET AL.
INTRODUCTION
Nest site selection can influence avian com-
munity structure and composition (Martin
1988a, 1988b), as well as adult survival and
reproductive performance (Burger 1985, Mar-
tin 1992). Many studies have demonstrated
that nest sites are selected actively, such that
microhabitat at nest sites differs from that at
random sites (Colwell and Oring 1990, Clark
and Nudds 1991). However, far fewer studies
have clearly demonstrated that such nest
microhabitat preferences are adaptive (Martin
1998, Clark and Shutler 1999, Jones 2001);
i.e., that individuals nesting in preferred
microhabitats experience higher reproductive
success.
Reproductive success may be affected when
nest habitats differ in food availability (Martin
1987), nest microclimate (Walsberg 1985), or
predation risk specific to a habitat type (Martin
1995). Variation in food, for example, is known
to affect both fecundity and reproductive
success (Lack 1968, Martin 1987). In shore-
birds, where clutch size is fixed and opportuni-
ties for second broods are rare (Arnold 1999),
variation in energy intake could influence
reproductive success by affecting parental body
condition (Carey 1980) or nest attentiveness
(Hegyi and Sasvári 1998).
Nest sites may also be selected to provide
a favorable microclimate (Horvath 1964, Wals-
berg 1981, 1985, Gloutney and Clark 1997,
Wiebe and Martin 1998). For shorebirds,
selecting a nest site sheltered from the wind
could lower convective heat loss of the in-
cubating adult, or the eggs when left unattend-
ed (Reid et al. 2002). Incubation is energetically
costly (Carey 1980, Vleck 1981, Williams 1996),
and the thermal environment of a nest can
influence both the rate and duration of in-
cubation recesses (Haftorn 1988, Reid et al.
1999, Cresswell et al. 2003).
Predation is the primary cause of nest failure
for most birds (Ricklefs 1969, Martin 1992),
and selection should therefore favor nesting
strategies that reduce predation risk. In the
montane forests of Arizona, nest sites in
nonpreferred microhabitats suffered increased
predation (Martin 1998). Indeed, habitat-spe-
cific risk of predation may explain much of the
variation in the life histories of passerines
(Martin 1995). However, these selective forces
occur when good nest sites are limited and some
individuals must select sites in nonpreferred
microhabitats (Martin 1988a).
In contrast to forests, birds breeding in
arctic tundra nest in a landscape of rock,
gravel, and low vegetation (often ,5 cm high).
In a landscape lacking structural complexity,
suitable nest sites may not be limited, and
a high overlap of nest site characteristics might
be expected between successful and failed
nests, or among species. In these habitats,
antipredator or cryptic behaviors of adults at
the nest, rather than the characteristics of the
nest site itself, may have an overriding effect
on nest success (Cresswell 1997, Ghalambor
and Martin 2002).
Shorebirds exhibit a variety of antipredator
behaviors, including distraction displays and
aggressive nest defense (Gochfeld 1984). While
aggressive behaviors should influence the risk
of predation directly, risk might also be
affected by activity near the nest, such as
incubation recesses or nest visitation (Lyon
and Montgomerie 1987, Wiebe and Martin
1997, Martin et al. 2000, Cresswell et al. 2003).
As incubation and defense strategies are
related to mating systems in shorebirds, and
often differ among sympatrically nesting spe-
cies, the influence of these behavioral factors
on nest success can be evaluated with in-
terspecific comparisons (Larsen 1991, Larsen
et al. 1996).
We compared the relative influence of nest-
ing habitat and behavioral factors on nest
success, to test the hypothesis that variation in
nest site characteristics plays a lesser role in
determining the fate of shorebird nests in
tundra systems. We tested for patterns of nest
preference by contrasting the nest sites of five
shorebird species with random sites. We then
quantified differences between the microhabitat
of successful and unsuccessful nests for each
species to test for adaptive nest site choice. We
placed artificial nests across our study area to
compare the frequency of predation across
habitat types. Relative invertebrate abundance
was measured to determine if shorebirds
preferentially nested in habitats with the most
abundant prey. Finally, we used site-specific
and literature values to test for a relationship
between the frequency of incubation recesses
and risk of predation, as a guide for future
studies.
 FACTORS AFFECTING SHOREBIRD NEST SUCCESS 17

TABLE 1. Features of the habitats of East Bay, Southampton Island, Nunavut, Canada.

Habitat type Distinguishing features

Intertidal zone Intertidal or within splash range of fall storms
Dead or dormant moss (organic crust) occurs
Bare substrate dominant, living moss and graminoids sparse and patchy
Moss carpet Pond edges in coastal areas
Living moss covers substrate
Sparse to moderate abundance of grasses and sedges
Numerous herbs, but patchy and sparse
Scrub willow Drier areas in central and northern portions of plot (0.5–1 km inland)
Salix spp. abundant
Herbs, grasses, sedges, and lichens common
Substrate of bare soil and small rocks
Dry heath Drier areas .1 km inland
Ericaceous shrubs dominant; dense cover of mountain avens (Dryas integrifolia)
Willows and lichens abundant
Herbs moderate in richness and abundance
Substrate variable: soil, rock, and gravel
Relief varies from flat to extremely hummocked
Sedge meadow Moist areas and pond edges inland
Moss covers substrate, few rocks present
Sedges and grasses tall (.50 mm) and dense
Herbs abundant and diverse
Relief varies from flat to highly hummocked
Gravel ridge Bare gravel dominant
Flora sparse and depauperate
Visibly raised from surrounding areas
Colonized sparsely by mountain avens at low edges

METHODS pioides), sedges (Carex aquatilis and C. sub-

STUDY AREA
Fieldwork was conducted from late May to
early August, 2000–2002, in the East Bay
Migratory Bird Sanctuary, Southampton Is-
land, Nunavut, Canada (63u599N, 81u409W).
Our study plot was 4 km long, and extended
from the coast to 3 km inland. Habitats within
the plot varied along an elevational gradient.
Low-lying areas within 1 km of the low tide
mark supported many brackish ponds (0.1 ha–
5 ha, ,1 m deep). These saline areas supported
few plants and sand and rock substrates were
largely exposed. Farther inland from the coast
(.1 km), 1–3 m high raised gravel beaches
occurred, and were separated by lower-lying
areas with many freshwater ponds similar in
size and depth to the brackish coastal ponds. In
all areas, ponds dried as the nesting season
progressed; large ponds decreased in size and
shallow ponds (,30 cm) dried out completely.
Vegetation in low-lying inland areas, between
the raised beaches, was dominated by mosses
(Campylium stellatum and Scorpidium scor-
spathacea), and grasses (Arctagrostis latifolia).
Drier areas were dominated by dwarf shrubs
(Dryas integrifolia, Salix arctica, and S. reticu-
lata; Fontaine and Mallory, in press). In all
habitats, vegetation was generally #5 cm in
height.
HABITAT CLASSIFICATION
We recognized six habitat types based on
appearance, salinity, vegetation, and elevation
(Table 1). These habitats were distinct, and
a detailed summary of their ground cover
characteristics appears in (Smith 2003). The
study plot was divided into a coastal and an
inland portion, with intertidal areas, moss
carpets, and scrub willow comprising the
coastal habitats, and sedge meadows, gravel
ridges, and dry heaths comprising the inland
types.
SHOREBIRDS AND THEIR PREDATORS AT
EAST BAY
Over 50 bird species have been recorded in the
East Bay Migratory Bird Sanctuary (Abraham
18 PAUL A. SMITH ET AL.
and Ankney 1986). Five shorebird species
commonly nest in the study area: Black-bellied
Plover (Pluvialis squatarola), Semipalmated
Plover (Charadrius semipalmatus), Ruddy Turn-
stone (Arenaria interpres), White-rumped Sand-
piper (Calidris fuscicollis), and Red Phalarope
(Phalaropus fulicarius), and the reproductive
ecology of these five species is varied.
The Black-bellied Plover is territorial and
monogamous, with both parents incubating and
caring for chicks (Paulson 1995). They are highly
vigilant and defend their nests aggressively from
predators through aerial attack and distraction
displays (Drury 1961). They are visual foragers
and peck invertebrates off the substrate, espe-
cially dipterans, beetles, and spiders (Paulson
1995). Semipalmated Plovers are monogamous,
territorial, and both parents incubate (Nol and
Blanken 1999). They are highly vigilant, and
both parents take part in distraction displays or
scolding (Sullivan Blanken and Nol 1998). They
feed primarily on dipteran larvae and adults
(Baker 1977). The Ruddy Turnstone is monog-
amous, territorial, and has a biparental mating
system. Turnstones are highly vigilant and
aggressively pursue predators. They feed pri-
marily on dipterans, which they pick from the
substrate or find by overturning stones (Mac-
Donald and Parmelee 1962). White-rumped
Sandpipers are polygynous and territorial; only
females provide parental care (Parmelee 1992).
They feign injury when predators approach and
anecdotal information suggests that they select
well-concealed nest sites (Parmelee et al. 1968).
They feed primarily by probing in moss for
larvae (especially tipulids), but may also take
spiders, beetles, and adult tipulids from the
substrate (Parmelee 1992). The Red Phalarope
exhibits facultative polyandry. Only males in-
cubate. They show no territoriality, distraction
displays, or aggression toward predators (Tracy
et al. 2002). Red Phalaropes feed on a variety of
aquatic and terrestrial invertebrate prey on the
breeding grounds, especially the larvae and
adults of chironomids and tipulids (Tracy et al.
2002).
The species considered in this study exhibit
a variety of nest attendance patterns; species in
which incubation is shared between the sexes
(biparental species) typically take fewer in-
cubation recesses than species in which one
adult incubates (uniparental species). Incuba-
tion behavior is poorly documented for the
Black-bellied Plover, but Mayfield (1973:83)
reports bouts ‘‘as long as 3 hr.’’ Male Semi-
palmated Plovers at Churchill, Manitoba,
Canada, have longer incubation shift lengths,
but a mean for both sexes is 201 min (Sullivan
Blanken and Nol 1998). Incubation is shared in
the Ruddy Turnstone, but the male’s contribu-
tion varies geographically and seasonally (Net-
tleship 2000). At our study site, incubating
turnstones sat for approximately 12 hr each,
leaving the nest 0–6 times during that period,
apparently in response to approaching preda-
tors (Perkins 2004). Assuming an average of
three such departures, incubation bouts last
roughly 4 hr. White-rumped Sandpipers on the
Melville Peninsula, in arctic Canada, had
a mean bout length of 47 min (Cartar and
Montgomerie 1985). Red Phalaropes had
a mean incubation bout length of 46 min at
this site (Smith et al. 2007), and 47 min on the
Taimyr Peninsula, Siberia (Tulp and Schekker-
man 2006).
Though mating systems, foraging ecology,
and breeding behavior differ among these
species, they share many basic reproductive
traits. All five species lay a maximum of four
eggs per clutch, and replacement clutches in the
event of predation are uncommon. All nest in
simple scrapes on the ground and incubate for
roughly three weeks (19–26 days). Finally, due
to the constraints of the short arctic summer, all
breed at approximately the same time, with
peak laying in mid June.
Potential nest predators are abundant at East
Bay. We regularly observed Parasitic Jaegers
(Stercorarius parasiticus), arctic foxes (Alopex
lagopus), and Herring Gulls (Larus argentatus).
Peregrine Falcons (Falco peregrinus), Long-tailed
Jaegers (Stercorarius longicaudus), Glaucous
Gulls (Larus hyperboreus), Sandhill Cranes (Grus
canadensis), and Common Ravens (Corvus
corax) were also seen. The number of these
predators observed daily (sightings per 8 hr
person-day) was used as an index of relative
predator abundance between years; such indices
are considered to reflect changes in vertebrate
population size with reasonable accuracy (Ho-
chachka et al. 2000). As the influence of
generalist predators on avian nesting success
may depend on the abundance of alternative prey
(Summers 1986, Summers and Underhill 1987,
Bêty et al. 2002), daily observations of lemmings
(Dicrostonyx torquatus) were also noted.
 FACTORS AFFECTING SHOREBIRD NEST SUCCESS
NEST SEARCHING AND MONITORING
2
In 2000 and 2001, we searched 7 km of tundra
for nests, 4.5 km2 in inland habitats and 2.5 km2
in coastal areas. In 2002, this area was expanded
to 12 km2, 8.7 km2 of inland and 3.3 km2 of
coastal habitats. To avoid bias, search effort was
allocated evenly across the study area. Search
effort was greatest at the onset of the nesting
period, from mid to late June.
Nests were found through behavioral obser-
vation, flushing birds while walking, and by two
people dragging a 30 m length of 5 mm di-
ameter rope. Nests were marked with a wooden
tongue depressor placed 10–20 m from the nest
at a random bearing (Reynolds 1985), and nest
locations were recorded (63 m) with a Global
Positioning System (GPS). Eggs were checked
at regular intervals (,7 days) and floated to
estimate developmental stage (Liebezeit et al.
2007). As clutches approached the estimated
date of hatching, nests were checked every 1–
2 days.
We used the Mayfield method to estimate
hatching success and daily mortality rates
(Mayfield 1961). Nests that hatched at least
one chick were considered successful. Most
hatching events were observed directly, but
small eggshell fragments in the nest lining were
also accepted as evidence of hatching (Mabee
1997). Both abandoned and depredated nests
were considered to have failed. Mayfield
exposure days were terminated on the last
active date for nests of unknown fate, and
halfway between the last active and first
inactive date for nests of known fate found
empty (Manolis et al. 2000). Standard errors
were calculated following Johnson (1979).
NEST SITE DECSCRIPTIONS
Nest sites of shorebirds were described at two
spatial scales. First, we quantified ground cover
in a 1 m2 circle centered on the nest (nest site).
Second, in a 75 m2 circle surrounding the nest
(nest patch), we recorded the percent cover of
each of the six habitat types. We also measured
distance (61 m) to the nearest water and to the
nearest dried pond edge (which would have
been inundated at nest initiation) with a hand-
held rangefinder. To measure concealment, we
used three 12 cm diameter white plastic disks
marked with a 1 cm black grid. Two disks were
fastened at right angles and placed atop a third,
to provide an identical silhouette from four
19
lateral directions and from overhead. This
apparatus was placed in nests, and we estimated
the proportion of markings obscured to the
nearest 5%. Estimates of lateral concealment
were made from north, south, east, and west, at
a distance of 5 m and a height of 40 cm (the
approximate height of an arctic fox). Overhead
concealment was estimated from human eye
level, directly above the nest. The height of the
rocks or vegetation directly surrounding the
nest (i.e., contacting the bottom disk) was
measured to the nearest 61 cm at the north,
south, east, and west edge. All nest habitat and
concealment data were collected in late July.
Nest coordinates (63 m) were plotted using
ArcView 3.2 (ESRI 1992). Distances between
nesting neighbors were calculated with the
‘‘Nearest Feature’’ extension of ArcView (Jen-
ness 2002), and spatial patterning for each
species was measured with the Clark and Evans
test with Donnelly’s modification (Clark and
Evans 1954, Donnelly 1978). The significance
of deviations from random patterns was de-
termined with Z-tests (Krebs 1989). For each
nest, we estimated the density of neighbors as
the mean distance to the five nearest shorebird
neighbors.
RANDOM SITE DESCRIPTIONS
In each year, we divided the study area into
a 50 m 3 100 m grid. Eighty grid intersections
were selected at random. At each selected
location, we tossed a stick backwards over our
heads and used the point of the stick as the
random site. At these sites, habitat data were
collected as for nest sites, at 1 m2 and 75 m2
scales. Sites falling in ponds were not included
in analyses.
ARTIFICIAL NEST EXPERIMENTS
We conducted an artificial nest experiment in
2002 to assess relative predation pressure in the
habitats of East Bay. ‘‘Clutches’’ of two
Japanese Quail (Coturnix japonica) eggs were
distributed in a stratified random design. As
artificial nest locations were selected for anoth-
er study, sample sizes varied among habitats.
Eggs were laid out from 8 to 10 July 2002 and
were checked every third day. After three
checks, missing or damaged eggs were replaced
(fresh eggs laid out 17–19 July) and the
experiment was repeated using the same artifi-
cial nest sites.
20 PAUL A. SMITH ET AL.
Quail eggs resemble small shorebird eggs
(e.g., White-rumped Sandpiper, Red Phalarope)
in coloration and size (ca. 32 mm 3 22 mm).
The depressions used as nest sites were similar
to the simple scrapes used by shorebirds. A
colored nail was hidden under the eggs to
facilitate finding the nest if eggs were taken by
predators. Eggs were sterilized in 50uC water
for 20 min, to avoid exposing wild birds to
agricultural pathogens. Nests with at least one
egg damaged or removed were considered
depredated.
INVERTEBRATE ABUNDANCE
AND DISTRIBUTION
To investigate spatial and seasonal patterns of
food availability for shorebirds, we sampled
terrestrial invertebrates throughout the nesting
period in 2000–2001. We placed twenty plastic
cups (diameter 11 cm, depth 8 cm) in random
locations in systematically chosen patches of
each of the six habitat categories, for a total of
120 traps. In 2000, all traps were deployed in
the western half of the study plot. In 2001, traps
were divided evenly between sites used in 2000
and new sites in the eastern half of the study
plot. An equal number of black and white
colored traps were used in each habitat, and the
distance between traps within a habitat patch
was at least 4 m. Traps were placed flush with
the substrate and filled to a level of 1 cm with
propylene glycol. Trap contents were filtered
through a reusable coffee filter for collection.
We emptied traps three times in 2000 (8 July, 16
July, and 24 July) and three times in 2001 (7
July, 15 July, and 23 July), spanning the period
from mid incubation to chick hatching.
Invertebrates were identified to family (with
the exception of mites, Acari, and springtails,
Collembola), and the number of individuals
was recorded for each trap. For each habitat,
we generated a mean volume of invertebrates
per trap by recording the amount of water
displaced by each sample.
STATISTICAL ANALYSES
We used discriminant function analysis to test
for habitat differences between: (1) nest sites of
different bird species, (2) nest sites and random
sites, and (3) successful and failed nests. An
initial principal components analysis revealed
that multicollinearity in both the 1 m2 and
75 m2 nest habitat datasets was weak. There-
fore, we used the original percent cover and
concealment variables (arcsine transformed) in
the discriminant function analyses. Variables
were entered in a stepwise fashion using the
Wilk’s Lambda method. Structure coefficients
were obtained from discriminant analyses to
determine correlations of variables with the
discriminant functions. For classification, prior
probabilities were set proportional to initial
sample size. Tests of significance were based on
chance-corrected classifications (Titus et al.
1984). These analyses were performed with
SPSS 10.0.7 (SPSS 2002). We compared surviv-
al rates of real and artificial nests using
program CONTRAST (Hines and Sauer
1989). Unless otherwise noted, P-values are
two-tailed and means are displayed 6 SE.
RESULTS
SHOREBIRD NEST SITES
We found and monitored 189 shorebird nests
from 2000 to 2002 (Table 2). The number of
nests found varied among years and sample
sizes differ in some analyses because of missing
information. No year effect was found in
habitat measurements, so data were pooled
across years where appropriate.
Four of the five common species exhibited
clear preferences for one of the six habitat
types, while the Red Phalarope used both sedge
meadow and scrub willow habitats equally
often (Table 2). Nests of the study species were
readily distinguished by discriminant function
analysis, indicating that preferred habitat dif-
fered among species. Classification based on the
discriminant functions was 72% correct (a 48%
improvement on chance; ZK 5 15.3, P ,
0.001). The first function grouped nests based
on the amount of moss and herbs at the nest site
(1 m2), or the amount of intertidal habitat in
the nest patch (75 m2), and accounted for 48%
of the variability in nest habitats. The second
function used habitat type (gravel ridge and dry
heath versus intertidal) to discriminate between
the otherwise similar (sparsely vegetated, low
cover) nest sites of Black-bellied Plover, Semi-
palmated Plover, and Ruddy Turnstone. Means
of nest site characteristics for each species are
given in the Appendix.
Concealment of nests of the Black-bellied
Plover, Semipalmated Plover, and Ruddy
Turnstone was similar to that of random sites
 FACTORS AFFECTING SHOREBIRD NEST SUCCESS
TABLE 2. The percentage of shorebird nests located in each of the six habitat categories at East Bay,
Nunavut, Canada, 2000–2002.
Gravel
Species n ridge
Black-bellied Plover (Pluvialis squatarola) 21
Semipalmated Plover 24
(Charadrius semipalmatus)
Ruddy Turnstone (Arenaria interpres) 63
White-rumped Sandpiper 24
(Calidris fuscicollis)
Red Phalarope (Phalaropus fulicarius) 57
(Fig. 1). In contrast, White-rumped Sandpipers
and Red Phalaropes had higher lateral and
overhead concealment than random sites or
than the three other species. The average height
of vegetation or rocks surrounding nests of the
five species showed a similar pattern, with taller
FIGURE 1. Lateral (A) and overhead (B) conceal-
ment of shorebird nest sites and random sites at East
Bay, Nunavut, Canada. Means are displayed 6 SE,
and sample sizes are displayed above the bars. BBPL
5 Black-bellied Plover, SEPL 5 Semipalmated
Plover, RUTU 5 Ruddy Turnstone, WRSA 5
White-rumped Sandpiper, and REPH 5 Red Phal-
arope.
21
Dry Sedge Scrub Moss
heath meadow willow carpet Intertidal
12 62 2 24 0 0
0 0 0 6 2 92
0 2 2 59 17 21
0 17 69 15 0 0
0 14 29 30 12 15
cover at the nests of White-rumped Sandpipers
and Red Phalaropes (Appendix).
Species varied not only in habitat and
concealment, but also in patterns of nest
dispersion (Table 3). For Black-bellied Plovers,
a sufficient number of nests for analysis were
available in 2002 only. These nests tended
toward a uniform distribution. The nests of
both Semipalmated Plovers and Ruddy Turn-
stones were clumped in coastal habitats. Sam-
ples of White-rumped Sandpiper nests were
small, and no significant patterns of dispersion
were found. The nests of Red Phalaropes were
significantly clumped in 2000 and 2001, but not
in 2002.
The nest sites of all species were readily
distinguished from random sites in discriminant
function analyses (Table 4). The accuracy of all
classifications exceeded 80% (.30% improve-
ment on chance). Black-bellied Plovers had
more cryptobiotic crust, lichen, and exposed
rock at the nest site, and less scrub willow
habitat in the nest patch than random sites.
Semipalmated Plovers nested in sites with more
cryptobiotic crust than random sites and
selected intertidal and moss carpet habitats.
Ruddy Turnstones avoided dry heaths and
gravel ridges and selected sites with more
exposed rock and cryptobiotic crust than
random sites. White-rumped Sandpipers select-
ed nests with more willow and lateral conceal-
ment than random sites, and preferred to nest in
sedge meadow habitats. Red Phalaropes select-
ed nest sites with higher concealment and more
herbs than random sites.
While there were clear patterns of nonran-
dom nest placement for all species, there were
few differences between successful and failed
nests with regard to the habitat variables
measured. Discrimination was weak for Semi-
22 PAUL A. SMITH ET AL.

TABLE 3. Indices of aggregation for the distribution of shorebird nests at East Bay, Nunavut, Canada.
Significant index values ,1 indicate clumping, while significant index values .1 indicate overdispersion.
Samples of fewer than 10 nests were not analyzed (Donnelly 1978), and are not presented.

Nearest conspecific
Year Species n neighbor (m 6 SE) Index of aggregation
2000 Ruddy Turnstone 8 333 6 75 0.60*
Red Phalarope 20 243 6 43 0.75*
2001 Semipalmated Plover 14 159 6 71 0.40***
Ruddy Turnstone 19 181 6 24 0.54***
Red Phalarope 16 211 6 44 0.57**
2002 Black-bellied Plover 17 494 6 65 1.05
Semipalmated Plover 10 372 6 88 0.58*
Ruddy Turnstone 38 142 6 11 0.47***
White-rumped Sandpiper 10 478 6 79 0.75
Red Phalarope 19 479 6 120 1.08
* P , 0.05, ** P , 0.01, *** P , 0.001.

palmated Plovers, Ruddy Turnstones, and Red random sites (80% correct classification, an
Phalaropes (60%–85% correct classification, 18% improvement on chance, P 5 0.1).
a 6%–15% improvement on chance, all P . Similarly, successful White-rumped Sandpiper
0.05). Successful Black-bellied Plover nests nests had more dirt and moss and less sedge
tended to be in sites with less lichen, although meadow habitat than failed sites, though they
they preferred nests with more lichen than preferentially selected nest sites in sedge mead-
TABLE 4. Habitat variables, at two spatial scales, that discriminated between nest sites and random sites for
shorebirds at East Bay, Nunavut, Canada. Separate analyses were conducted for each species, and all
discriminations were highly significant. All habitat variables measured are displayed but standardized
canonical discriminant function coefficients are shown only for variables contributing significantly to
discrimination. BBPL 5 Black-bellied Plover, SEPL 5 Semipalmated Plover, RUTU 5 Ruddy Turnstone,
WRSA 5 White-rumped Sandpiper, and REPH 5 Red Phalarope. The percentage of correct classification,
and the improvement over chance that this represents, are also displayed (Titus et al. 1984).

Species
BBPL SEPL RUTU WRSA REPH
2
1m
Rock 0.61 0.80
Dirt
Cryptogamic crust 0.97 0.56 0.69
Lichen 0.75
Moss
Willow 0.32 0.65
Avens
Sedge and grass 20.58 20.39
Herb 20.43 0.39 0.40
75 m2
Sedge meadow 0.67
Dry heath 20.57
Gravel ridge 20.64 20.29
Scrub willow 20.66
Moss carpet 0.41
Intertidal 1.12 20.38
Lateral concealment 20.51 0.44 0.68
Overhead concealment 0.58
% correct classification 98 90 87 83 82
% improvement on chancea 48 40 37 33 32
a
All chance-corrected P-values , 0.001.
 FACTORS AFFECTING SHOREBIRD NEST SUCCESS
FIGURE 2. The number of sightings per 8-hr
person-day of arctic foxes (Alopex lagopus), lem-
mings (Dicrostonyx torquatus), and jaegers (Stercor-
arius parasiticus and S. longicaudus), May–August,
2000–2002, at East Bay, Nunavut, Canada. Note that
jaegers were the most abundant predator in all years
and are represented on the right y-axis with
a different scale.
ow habitats. The discrimination between the
habitat of successful and failed nests was
significant only for White-rumped Sandpipers
(100% correct classification, a 39% improve-
ment on chance, P , 0.001). However, with
only five successful nests, these results should
be interpreted with caution.
PREDATORS AND LEMMINGS
The frequency of predator and lemming sight-
ings differed substantially among years (Fig. 2).
Lemmings were encountered frequently in 2000
and 2001, but none were seen in 43 days of
observation in 2002. Fox encounters were rare
in 2000 and more common in 2001 and 2002.
Jaegers were present in all years, but were more
than three times as abundant in 2002 than in
2000 and 2001. In all years, Parasitic Jaegers
were .4 times more abundant than Long-tailed
Jaegers. Although 20–24 pairs of Herring Gulls
bred in the study area in all years, an intensive
study indicated that they do not feed heavily on
shorebird eggs (K. Allard, University of New
Brunswick, unpubl. data). We thus assumed
that Herring Gull predation on shorebird nests
was infrequent and did not vary substantially
among years.
FACTORS AFFECTING HATCHING SUCCESS
Nest success varied markedly among years and
species (Fig. 3). Estimates of daily mortality
were generally high in 2002, when lemmings
were scarce and predators abundant, but Semi-
23
FIGURE 3. Mayfield estimates of the daily mor-
tality rate (6 SE) for shorebird nests at East Bay,
Nunavut, Canada, 2000–2002. The number of nests
monitored in each year is displayed above the bars.
BBPL 5 Black-bellied Plover, SEPL 5 Semipalmat-
ed Plover, RUTU 5 Ruddy Turnstone, WRSA 5
White-rumped Sandpiper, and REPH 5 Red
Phalarope.
palmated Plovers had similar success in 2001
and 2002 (no nests were found in 2000), and
White-rumped Sandpipers had high mortality
in both 2002 and 2000.
Interspecific patterns of habitat use were
poorly related to hatching success. The strong
variation in success among species was un-
related to variation in predation pressure across
habitat types (Fig. 4). Red Phalarope nests were
found in all habitats but gravel ridges (Table 2).
White-rumped Sandpipers nested in dry heath
and scrub willow habitats, but estimates of nest
mortality for these habitats were not higher
than for other habitats when the nests of all
species were considered. Only White-rumped
Sandpipers and Red Phalaropes nested in sedge
meadows. This accounts for the higher mortal-
ity observed in this habitat type.
Density differences around real nests are
unlikely to have caused the interspecific differ-
ences in nest success. While the mean distance
to the five nearest nesting neighbors (any
shorebird species) differed among species
(F4,184 5 12.9, P , 0.001; Table 5), densities
across the study plot were low (7.4 nests per
km2). The mean distance from a focal nest to
the nearest shorebird nest was .116 m for all
species. Within species, neither the distance to
the nearest neighbor nor the mean distance to
the five nearest neighbors differed between
successful and depredated nests (all P . 0.1).
24 PAUL A. SMITH ET AL.
FIGURE 4. Mayfield estimates of daily mortality
rate (6 SE) for all shorebird nests found in each of
the six habitat types at East Bay, Nunavut, Canada,
2000–2002. When all shorebird nests found in a given
habitat were combined, indices of predation pressure
were similar in all habitats except sedge meadows.
Sample sizes of nests are given above the bars.
Overall, habitat preference was unrelated to
nest success (Fig. 5). While Semipalmated
Plovers, White-rumped Sandpipers, and Red
Phalaropes tended toward higher nest success in
preferred habitats, Black-bellied Plovers and
Ruddy Turnstones showed the opposite trend.
Over all years, daily mortality was lower for
Black-bellied Plovers, Semipalmated Plovers,
and Ruddy Turnstones (species with biparental
care and infrequent nest recesses) than for
White-rumped Sandpipers and Red Phalaropes
(species with uniparental care and frequent nest
recesses). The correlation between the mean
length of incubation bouts and the risk of
TABLE 5. Mean distances to the nearest, or five
nearest, shorebird neighbors for each species at East
Bay, Nunavut, Canada. Both metrics represent
distances to nests of any shorebirds, not only
conspecifics.
Nearest Five nearest
Species (mean [m] 6 SE) (mean [m] 6 SE)
Black-bellied 126 6 18 238 6 22
Plover
Semipalmated 117 6 14 231 6 16
Plover
Ruddy Turnstone 116 6 9 207 6 11
White-rumped 153 6 21 342 6 31
Sandpiper
Red Phalarope 256 6 54 447 6 57
FIGURE 5. Mayfield estimates of survival over the
incubation period (6 SE) for shorebird nests placed
in preferred versus nonpreferred habitats at East Bay,
Nunavut, Canada, 2000–2002. There was no consis-
tent pattern of higher hatching success in preferred
habitats. Preferred habitat is the habitat type where
nests were most commonly placed (see Table 2); Red
Phalaropes nested preferentially in two habitat types.
Sample sizes of nests are displayed above the bars.
BBPL 5 Black-bellied Plover, SEPL 5 Semipalmat-
ed Plover, RUTU 5 Ruddy Turnstone, WRSA 5
White-rumped Sandpiper, and REPH 5 Red
Phalarope.
predation was significant (daily mortality vs.
bout length in minutes, Pearson r 5 20.90, P ,
0.05). However, with only five species and
behavioral data of varying quality, this result
must be treated as preliminary.
ARTIFICIAL NEST SUCCESS
Although White-rumped Sandpipers and Red
Phalaropes had the lowest nest success, rates of
loss of artificial nests did not suggest that
predation pressure was higher in the habitats
that they preferred (i.e., sedge meadow and
scrub willow; Fig. 6). Sedge meadow habitats
had comparatively low rates of artificial nest
lost in both the early to mid incubation period
and the late to post incubation period. With
both periods and all habitats combined, artifi-
cial nests had a daily mortality rate lower than
that of real nests in the year of the experiment
(DMRartificial nests 5 0.04 6 0.01, DMRreal nests
2
(2002) 5 0.06 6 0.01; x 1 5 3.8, P 5 0.05).
INVERTEBRATE ABUNDANCE
As shorebirds may benefit from nesting near
food resources, patterns of insect abundance
could affect nest distribution and success. The
relative abundance of invertebrates varied
 FACTORS AFFECTING SHOREBIRD NEST SUCCESS
FIGURE 6. The proportion of artificial nests taken
by predators in each of the six habitat types at East
Bay. Nests contained two Japanese Quail (Coturnix
japonica) eggs, and were deployed from 8–10 July
until 17–19 July, 2002 (Round 1), and at the same
sites from 17–19 July until 26–28 July, 2002 (Round
2). The numbers of nests placed in each habitat are
displayed above the bars.
among habitats (Fig. 7). The highest volume of
invertebrates was captured in dry heath habitats,
and the lowest in gravel ridge and intertidal
habitats. Trap contents in 2000 were dominated
(in volume) by dipterans (Chironomidae, Mus-
cidae, and Scatophagidae), carabid beetles (es-
pecially Pterostichus caribou), and spiders (espe-
cially Erigone, Alopecosa, and Pardosa spp.). In
2001, many adult fritillary butterflies (Boloria
spp.) were captured in dry heath and sedge
meadow habitats (30% of volume in dry heath
and 27% in sedge meadow). Few Boloria were
captured in 2000. In both years, adult tipulids
were most common in gravel ridge, sedge
meadow, and dry heath habitats (ml per trap
per period in 2000, 2001: gravel ridge 5 0.04,
0.04; sedge meadow 5 0.03, 0.01; dry heath 5
0.02, 0.01). Other dipterans were most abundant
in scrub willow, intertidal, and moss carpet
habitats (ml per trap per period in 2000, 2001:
scrub willow 5 0.52, 0.33; intertidal 5 0.35, 0.38;
moss carpet 5 0.29, 0.59). Beetles and spiders
were most abundant in dry heaths (ml per trap
per period in 2000, 2001: Carabidae 5 0.32, 0.37;
Araneae 5 0.50, 0.31).
Shorebirds did not preferentially use the dry
heath habitats where invertebrate prey were
most abundant. Also, invertebrates were abun-
25
dant in moss carpet habitats, though few
shorebirds nested there.
DISCUSSION
As more than 40% of shorebird clutches at East
Bay were lost to predators in each year of our
study, natural selection should strongly favor
nesting preferences and behaviors that minimize
predation. Where birds are not limited in their
choice of nest site, behavioral factors such as
nest defense or incubation recesses may have
more impact on nest success (Cresswell 1997,
Ghalambor and Martin 2002). We found strong
evidence for nest habitat preferences, but the
variation in nest success that we observed
appeared to be influenced primarily by behav-
ior and the abundance of predators.
NEST MICROHABITAT
In this study, habitat features of nest sites
differed from random sites for all five species.
Discrimination of nest site habitat was also
significant among species. Our results therefore
suggest that habitat played a significant role in
nest site selection, however differences in nest
habitat within species were not related to
differences in nest success. There was high
overlap in the habitat of successful and un-
successful nests, with significant discrimination
only for White-rumped Sandpipers.
Interspecific patterns of nest success were
also poorly related to nest habitat. The overall
rates of predation in the species’ preferred
habitats did not explain the interspecific pat-
terns of nest success, and predation on artificial
nests was comparatively low in the habitats
preferred by White-rumped Sandpipers and
Red Phalaropes, the species suffering the high-
est rates of predation.
DISTRIBUTION OF NESTS
Patterns of nest distribution could affect re-
productive success through a variety of me-
chanisms, such as density-dependent predation
(Tinbergen et al. 1967, Lack 1968) or group
defense (Larsen et al. 1996). If predators
intensify their searches in areas where they
have encountered success, risk of predation
may be positively correlated with nest density.
In this study, distance to nearest neighbors (a
measure of relative density) was not related to
nest success within or among species. Thresh-
olds for density dependence may be lower for
26 PAUL A. SMITH ET AL.
FIGURE 7. The mean volume (ml) of invertebrates
per pitfall trap captured in each of the six habitats
types at East Bay, Nunavut, Canada. Traps were
emptied three times in 2000: 8 July, 16 July, and 24
July, and three times in 2001: 7 July, 15 July, and 23
July. The mean volume per trap per collection period
(6 SE) is presented separately for 2000 and 2001.
the tundra ecosystem; however, studies in
subarctic and temperate areas have not docu-
mented density-dependent predation at the
densities found in our study (Göransson et al.
1975, Sugden and Beyersbergen 1986, Schieck
and Hannon 1993, Larivière and Messier 1998).
Predation may be negatively correlated with
nesting density for birds that respond aggres-
sively to predators and collaborate in group
defense. Alternatively, incubating adults could
benefit from the vigilance and early warning
provided by neighbors nesting nearby (Nuech-
terlein 1981). Black-bellied Plovers and Ruddy
Turnstones exhibit aggressive nest defense
(Paulson 1995, Nettleship 2000; PAS, pers.
obs.), and could benefit from group mobbing.
Semipalmated Plovers are highly vigilant, en-
gage in distraction displays (Nol and Blanken
1999), and could benefit from early warning.
Although all three species could benefit from
nesting at high densities, neither the distance to
the nearest conspecific nor the mean distance to
the five nearest neighbors differed between
successful and failed nests for these species.
NEST CONCEALMENT
Cover around the nest may conceal incubating
adults and reduce the dispersion of scent
(Bergerud and Gratson 1988), or it may create
a favorable microclimate (Walsberg 1985,
Wiebe and Martin 1998). Concealment, how-
ever, may also be costly when predators
threaten incubating birds, or when the success
of antipredator behavior depends on early
detection of predators (Götmark et al. 1995).
The nests of White-rumped Sandpipers and
Red Phalaropes had higher lateral and over-
head concealment than those of Black-bellied
Plovers, Semipalmated Plovers, and Ruddy
Turnstones. White-rumped Sandpipers and
Red Phalaropes flush at close distances (Par-
melee et al. 1968, Mayfield 1979, Ridley 1980)
and may select nest sites with high concealment
to escape detection from predators. In contrast,
Black-bellied Plovers, Semipalmated Plovers,
and Ruddy Turnstones all flush at distances
.20 m before engaging in antipredator behav-
ior (Paulson 1995, Nol and Blanken 1999,
Nettleship 2000), and these species may select
exposed sites to facilitate early detection of
predators. However, these latter three species
selected nest sites with concealment similar to
that of random sites, so concealment may be an
unimportant characteristic for their nest site
choice.
ACTIVITY NEAR THE NEST
Nest concealment may aid birds in escaping
detection by predators, but behavior can also
play a role (Cresswell 1997, Martin et al. 2000,
Ghalambor and Martin 2002). Activity near the
nest may divulge its location to predators; the
rate of nest visitation is positively related to the
risk of predation in tundra-nesting Snow
Buntings (Lyon and Montgomerie 1987), and
the survival of Semipalmated Sandpiper
clutches is higher when unattended (Ashkenazie
and Safriel 1979, Safriel 1980).
A marked reduction in the number of trips on
and off the nest may account for the higher nest
success seen in bi- versus uniparental species.
The importance of activity around the nest in
determining success is corroborated by the
observation that artificial nests had lower daily
mortality than real nests. However, the in-
cubation behavior of arctic-breeding shorebirds
is strongly influenced by factors including
weather, breeding stage, and latitude (Norton
1972, Cartar and Montgomerie 1985, Schamel
and Tracy 1987), and our use of means and
literature values is a severe limitation. Future
 FACTORS AFFECTING SHOREBIRD NEST SUCCESS
studies should examine the relationship between
incubation behavior and nest success with site-
specific behavioral data, at inter- and intraspe-
cific levels, to test the hypothesis that frequent
incubation recesses increase the risk of shore-
bird nest predation.
ANTIPREDATOR BEHAVIOR
The antipredator behaviors of shorebirds are
risky (Brunton 1986) and have evolved because
of their success at dissuading or distracting
predators (Gochfeld 1984). Because the risk of
injury to the defender must be balanced against
the reward of successful nest defense (Larsen et
al. 1996), interspecific differences in nest success
could be related to level of defense behavior.
Both aggressive species (Black-bellied Plover
and Ruddy Turnstone) had high nest success in
all years, though they suffered increased pre-
dation in 2002 when predators were abundant
and lemmings scarce. However, the highest
shorebird nest success was found in Semi-
palmated Plovers, which engage in vigorous
distraction displays (both sexes), but do not
attack predators (Nol and Blanken 1999). On
the other hand, White-rumped Sandpipers also
exhibit distraction behaviors (females only;
Parmelee 1992), but they had the lowest nesting
success of any shorebird at East Bay. Thus,
differences in the level of nest defense do not
explain our observed interspecific patterns of
nest success.
THE ROLE OF PREDATOR-PREY CYCLES
Habitat and behavior may have contributed to
nest site selection and reproductive success, but
the single greatest influence on nest success
across all species was fluctuating annual pre-
dation pressure. In tundra systems, predation of
the nests of shorebirds and waterfowl is extreme
about every fourth year when lemmings are
scarce and predators of rodents turn to the eggs
of birds (Summers 1986, Martin and Baird
1988, Bêty et al. 2002). Populations of lem-
mings at East Bay probably cycle with a period
of three to five years (Krebs 1964, Hanski and
Korpimäki 1995). Lemmings were scarce in
2002 compared to 2000–2001, and we assume
that the dramatically increased nest mortality in
2002 reflected increased predation by arctic
foxes and jaegers that year.
Many shorebirds can defend their nests from
avian predators, but none are consistently
27
successful at deterring mammalian predators
such as the arctic fox (Larsen et al. 1996). As
nest loss in years of low lemming abundance is
catastrophic for many species (Summers and
Underhill 1987, Underhill et al. 1993), there
may simply be no adaptations in nest site
preference or parental behavior that reduce fox
predation in years following a lemming decline.
FOOD AVAILABILITY AND
REPRODUCTIVE SUCCESS
Birds may select nest sites that are close to food
sources, yet food is often overlooked in studies
of nest site selection. In shorebirds, the avail-
ability of invertebrate prey could influence nest
success by affecting foraging efficiency or the
body condition of incubating adults (Carey
1980). However, the relationship between prey
availability and foraging efficiency depends on
the distance between foraging locations and the
nest. The Black-bellied Plover, Semipalmated
Plover, and Ruddy Turnstone feed within their
territories, but may also use communal feeding
areas or separate feeding territories (Paulson
1995, Nol and Blanken 1999, Nettleship 2000).
We generally observed female White-rumped
Sandpipers and male Red Phalaropes feeding
within 100 m of the nest, but the former has
been observed foraging .300 m from the nest
(Cartar and Montgomerie 1985).
The abundance of suitable shorebird prey
captured in pitfall traps varied significantly
among habitats, but habitats with the greatest
abundance of suitable prey were not preferred
by nesting shorebirds. However, our collections
do not represent the full suite of prey items used
by shorebirds. Larvae are important for prob-
ing species such as the White-rumped Sandpip-
er and Red Phalarope, and the extent to which
habitat use of larvae is related to that of winged
adults is unclear. Also, flying insects, like the
fritillary butterflies captured in 2001, may be
attracted to pitfall traps from a distance. With
these significant caveats in mind, we found no
influence of food limitation on nest site
selection of shorebirds at East Bay.
ADAPTIVE NEST HABITAT PREFERENCES?
Choice of nest microhabitat by birds is widely
assumed to be adaptive, yet this assumption has
rarely been tested (Clark and Shutler 1999).
Several studies have addressed this issue, and
found that reproductive success suffers when
28 PAUL A. SMITH ET AL.
nests are placed in nonpreferred habitats
(Martin 1988a, Martin 1998). However, this
may apply only in systems where nest sites are
limiting. Our study did not find nest sites to be
limited, nor did shorebirds consistently show
higher hatching success when nesting in pre-
ferred habitats. There were few habitat differ-
ences between successful and unsuccessful nest
sites in the habitat variables measured. This
does not imply that habitat is unimportant in
determining nest fate; merely that variation in
the habitat of occupied nest sites does not drive
variation in predation. Where shorebirds are
not limited in their choice of nest site, the
behavior of incubating adults may have a stron-
ger influence on reproductive success by affect-
ing the detectability of the nest by predators.
ACKNOWLEDGMENTS
We are indebted to David Larson, Roger Pickavance,
and Krista Bowers, of Memorial University of
Newfoundland, for identifying the invertebrate sam-
ples. We are grateful to those who contributed to
fieldwork, especially Rachel Bryant, Eric Davies,
Anna Hargreaves, Deborah Perkins, Carolyn Saun-
ders, Iain Stenhouse, and Karen Truman. We thank
Erica Nol for assistance with study design and
analysis, and Will Cresswell for his helpful comments
on the manuscript. Funding for this study was
provided by the Canadian Wildlife Service–Prairie
and Northern Region, the Polar Continental Shelf
Project, the Natural Sciences and Engineering Re-
search Council, and the Northern Scientific Training
Program. All experimental procedures were approved
by the Animal Care Committees of the University of
British Columbia and Environment Canada.
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 FACTORS AFFECTING SHOREBIRD NEST SUCCESS 31

APPENDIX. Mean values (6 SE) for habitat characteristics of shorebird nests at East Bay, Nunavut,
Canada, 2000–2002. BBPL 5 Black-bellied Plover, SEPL 5 Semipalmated Plover, RUTU 5 Ruddy
Turnstone, WRSA 5 White-rumped Sandpiper, and REPH 5 Red Phalarope.

Species
BBPL SEPL RUTU WRSA REPH
(n 5 21) (n 5 24) (n 5 63) (n 5 24) (n 5 57)
% cover at 1 m2
Water 0 6 0 0 6 0 0 6 0 0 6 0 1 6 1
Rock 29 6 5 50 6 5 37 6 2 3 6 1 26 6 3
Cryptogamic crust 23 6 4 32 6 5 29 6 2 14 6 2 18 6 2
Lichen 9 6 2 2 6 1 4 6 1 3 6 1 2 6 0
Moss 3 6 2 6 6 2 14 6 1 45 6 4 24 6 3
Dirt 9 6 2 10 6 3 8 6 1 9 6 3 8 6 2
Avens 25 6 4 0 6 0 2 6 1 9 6 3 6 6 2
Willow 3 6 1 0 6 0 6 6 1 13 6 2 7 6 1
Sedge and grass 1 6 0 5 6 2 2 6 0 15 6 2 9 6 2
Miscellaneous herbs 0 6 0 1 6 1 2 6 0 0 6 0 5 6 2
% cover at 75 m2
Gravel ridge 10 6 7 0 6 0 0 6 0 0 6 0 0 6 0
Dry heath 60 6 10 0 6 0 2 6 1 13 6 6 11 6 4
Scrub willow 19 6 8 6 6 20 45 6 5 14 6 5 23 6 5
Sedge meadow 5 6 3 0 6 0 3 6 2 70 6 7 25 6 5
Moss carpet 3 6 2 8 6 17 25 6 3 3 6 2 14 6 3
Intertidal 1 6 1 81 6 6 18 6 4 0 6 0 9 6 3
Rock 0 6 0 0 6 0 1 6 1 0 6 0 3 6 2
Water 0 6 0 0 6 0 2 6 1 0 6 0 5 6 2
Dried pond 0 6 0 4 6 7 3 6 1 1 6 1 11 6 3
Nearest water (m) 39 6 5 19 6 12 12 6 1 47 6 6 22 6 3
Nearest dried pond (m) 20 6 2 7 6 11 7 6 1 20 6 4 9 6 2
Overhead concealment 0 6 0 0 6 1 1 6 0 10 6 2 10 6 1
Avg. lateral concealment (%) 6 6 1 6 6 6 8 6 1 14 6 2 23 6 2
Avg. height of vegetation (cm) 16 6 2 12 6 8 18 6 1 54 6 4 45 6 3