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often forage outside of territories, and broods leave
territories after hatching. Curlews quickly disperse
after they arrive on the breeding grounds to establish
territories in suitable habitat. Returning breeders
establish territories with less agonistic interaction than
first-time breeders do. After pairing, primarily the males
defend territory boundaries only against conspecifics.
One study (Nebraska) reported that some breeders
defended a feeding territory in addition to a nest
territory (Bicak 1977). Territory size at Humboldt Bay,
California averaged 3.0 ha ± 2.1 SD (range = 1.3–7.5, n
= 8) in summer, with overlap averaging 28.5 percent ±
29.7 SD (range = 1.3–88.1, n = 12; Mathis 2000).
Away from breeding areas, territoriality has
been reported for intertidal habitat; territoriality was
not observed in wet pastures (Dugger and Dugger
2002). Not all individuals establish territories during
migration or on winter areas. Territories at Humboldt
Bay, California, which are occupied during winter and
summer, averaged 2.4 ha ± 1.6 SD (range = 1.3–4.2, n =
3), similar in size to summer territories (Mathis 2000).
Dispersal
Juvenile curlews may move extensively after
hatching. One brood that was relocated 6 days after
banding was 6.5 km from the nest site (Sadler and
Maher 1976). Curlew chicks fledge at 38 to 45 days
after hatching (King 1978, Allen 1980, Redmond and
Jenni 1986) and depart the breeding grounds relatively
early (most from mid-June to mid-August) and in small
flocks, with no real evidence of fall staging (Campbell
et al. 1990). During late summer and migration, flocks
of 10 to 50 birds are common (range = 100–500) (Allen
1980, Renaud 1980, Pampush 1981, Campbell et al.
1990, Roy 1996). Small migratory flocks in Utah during
fall contain one or two adults and two to four juveniles,
suggesting that family groups sometimes migrate
together (Paton and Dalton 1994).
There is little information on natal philopatry, but
no birds marked as chicks were ever seen in subsequent
years on breeding areas as yearlings (Redmond and
Jenni 1986). Compared to females, male curlews are
more likely to return and breed (first time) near the
natal nest site (Redmond and Jenni 1982). Return rates
of breeding adults (based on resighting data) over three
years in Idaho were 89 percent ± 0.10 SD, 64 percent ±
0.10 SD, and 0.84 percent ± 0.16 SD.
Source/sink, demographically linked
populations
There is no evidence of source-sink dynamics
in this species. Because there has been only one long-
term study of a marked population (Redmond and Jenni
1986), and few recoveries of banded individuals, there
is no information on the possible linkage of populations
or metapopulation dynamics.
Factors limiting population growth
Curlew deaths have been reported from predators,
disease, and contaminants. There are no mortalities
reported due to exposure, but the impact of climate
on prey abundance and availability may influence
population growth. Decreased food availability—either
on breeding areas or along migration routes to the
south—is presumed to be responsible for interannual
variation in clutch size, and may limit population
growth (Redmond and Jenni 1986).
With the exception of habitat loss, predation
on eggs and chicks is probably the single greatest
factor limiting population growth. Gopher snakes
(Pituophis spp.) and a variety of mammalian and avian
predators are known to depredate nests. In Idaho,
28.6 percent of the nests were destroyed by canids
(coyote [Canis latrans], red fox [Vulpes vulpes], feral
dog [C. familiaris]), badgers (Taxidea taxis), or other
undetermined mammals; 6.7 percent were destroyed by
birds (primarily black-billed magpie [Pica hudsonia]);
4.2 percent of nests were abandoned due to disturbance
by livestock (n = 119; Redmond and Jenni 1986). In
Oregon, nest predators (including badger, coyote, and
various corvids) destroyed 10 to 16 percent of 101 nests
(Pampush and Anthony 1993). Other potential nest
predators include feral cat (Felis catus), striped skunk
(Mephitis mephitis), raccoon (Procyon lotor), and
long-tailed weasel (Mustela frenata). Livestock destroy
curlew nests by trampling (Dugger and Dugger 2002).
Predation of adult long-billed curlews has not
been confirmed, but prairie falcons (Falco mexicanus)
have been observed making unsuccessful attempts on
curlews, and raptors took two radio-marked chicks in
Idaho one week after they fledged (Redmond and Jenni
1986). During one year, almost all chick deaths were
attributed to raptors (a long-tailed weasel ate one chick)
(Jenni et al. 1981).
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25
Life cycle graph and model development
The studies of Dugger and Dugger (2002)
provided the basis for formulating a life cycle graph for
long-billed curlew that comprised two stages (censused
at the fledgling stage and “adults”). The scanty data on
survival suggested highest survival of yearlings (20 of
30 males returning) and lower survival of older birds
(5 of 12 returning). We further assumed considerably
lower survival in the first year, a value for which we
solved by assuming λ (population growth rate) was
1.003. This “missing element” method (McDonald and
Caswell 1993) is justified by the fact that, over the long
term, λ must be near 1 or the species will go extinct or
grow unreasonably large. In addition, we assumed that
first-year reproduction was lower than that of “adult”
birds (Table 1). From the resulting life cycle graphs
(Figure 8), we produced a matrix population analysis
with a post-breeding census for a birth-pulse population
with a one-year census interval (McDonald and Caswell
1993, Caswell 2001). The models had two kinds of input
terms: P
i
describing survival rates and m
i
describing
number of female fledglings per female (Table 1).
Figure 9a and Figure 9b show the numeric values
for the matrix corresponding to the life cycle graph
of Figure 8. The model assumes female demographic
dominance so that, for example, fertilities are given as
female offspring per female; thus, the fledgling number
used was half the total annual production of fledglings,
assuming a 1:1 sex ratio. Note also that the fertility terms
(F
ij
) in the top row of the matrix include both a term for
fledgling production (
m
i
) and a term for the survival of
the mother (
P
i
) from the census (just after the breeding
season) to the next birth pulse almost a year later. The
population growth rate, λ, was 1.003, based on the
estimated vital rates used for the matrix. Although this
suggests a stationary population, the value was used as
an assumption for deriving a vital rate, and it should not
be interpreted as an indication of the general well-being
of the population. Other parts of the analysis provide a
better guide for assessment.
Sensitivity analysis
A useful indication of the state of the population
comes from the sensitivity and elasticity analyses.
Table 1. Parameter values for the component terms (P
i
and m
i
) that make up the vital rates in the projection matrix for
long-billed curlew.
Parameter
Numeric value
Interpretation
m
1
1.4
Number of female fledglings produced
by a first-year female
m
a
1.9
Number of female fledglings produced by an “adult” female
P
21
0.28
First-year
survival rate
P
32
0.67
Second-year survival rate
P
a
0.42
Survival rate of “older adults”
P
21
m
1
1
2
3
P
21
P
32
P
a
P
a
m
a
P
32ma
Figure 8. Life cycle graph for long-billed curlew. The numbered circles (“nodes”) represent the three stages (first-
year birds, second-year birds and “older adults”). The arrows (“arcs”) connecting the nodes represent the vital rates
– transitions between age-classes such as survival (P
ji
) or fertility (F
ij
) (the arcs pointing back toward the first node).