2010 NORTHEASTERN NATURALIST 17(3):493–504
Trends in Piping Plover Reproduction at Jones Beach State
Park, NY, 1995–2007
Annie F. McIntyre1, Julie A. Heath2,*, and Joseph Jannsen3
Abstract - We evaluated thirteen years (1995–2007) of Charadrius melodus (Piping
Plover) nesting activity at Jones Beach State Park (JBSP) to examine temporal
trends in the number of pairs, productivity, total young produced per year, and nest
initiation. The number of plover pairs has decreased over time at JBSP, while productivity
(and total number of young produced per year) has increased. Productivity
has increased more at JBSP compared to other areas in their range. In addition, nest
initiation was significantly later in the nesting season in recent years. Increased
productivity indicates that management efforts to increase this population parameter
may have had a positive effect on nesting plovers. The decreased numbers of
nesting pairs, however, suggests that management techniques to increase numbers
of breeding pairs, such as nesting-habitat restoration, may also be necessary to
increase local population size. In the absence of suitable nesting habitat for recruitment
of locally fledged birds, JBSP may be functioning as a “source” for other
regional breeding populations.
Development of beaches along the Atlantic Coast has been a contributing
factor in the decline of Charadrius melodus Ord. (Piping Plover)
populations (USFWS 1996, Wilcox 1959). Piping Plovers breed on open,
sparsely vegetated beaches between the primary dune and the high tide line.
Increased human and predator activity in these nesting areas have negative
impacts on breeding Piping Plovers. In addition, large-scale engineering
projects, such as jetty construction and beach nourishment, have changed the
frequency and magnitude of natural processes that drive coastal dynamics
and create or maintain Piping Plover nesting and foraging habitat (Dean and
Dalrymple 2002, Loegering and Fraser 1995, USFWS 1996).
The United States Fish and Wildlife Service (USFWS) revised recovery
plan for the Atlantic Coast Piping Plover population sets a productivity
goal of 1.5 young per pair per year (USFWS 1996). Since this target is infrequently
met on unmanaged Atlantic Coast beaches, several management
efforts have been widely employed to either decrease the human-plover
conflict on public beaches or to decrease Piping Plover losses to predators
(Hecht and Melvin 2009a). Exclosures that prevent predators from consuming
eggs have led to increases in hatching success (Larson et al. 2002,
1Theodore Roosevelt Nature Center, Jones Beach State Park, PO Box 1000, Wantagh,
NY 11793. 2Department of Biological Sciences, Boise State University, Boise, ID
83725. 3The Nature Conservancy, Uplands Farm Sanctuary, 250 Lawrence Hill Road,
Cold Spring Harbor, NY 11724. *Corresponding author - email@example.com.
494 Northeastern Naturalist Vol. 17, No. 3
Melvin et al. 1992, Rimmer and Deblinger 1990). Similarly, symbolic fencing
that prevents people from trampling eggs or disturbing incubating birds
may also increase hatching success (Ikuta and Blumstein 2003). Predator
removal is used occasionally to reduce predation rates of chicks and adults
(Reynolds and Tapper 1996). Finally, restoration or creation of new habitat
may attract breeding Piping Plovers to areas of low human use and minimize
conservation and recreation conflicts (Cohen et al. 2009; McIntyre
and Heath, in press).
Evaluation of Piping Plover population recovery requires a large-scale
analysis of monitoring data from multiple sites and land managers. The
USFWS Atlantic Coast Piping Plover recovery coordinator and state coordinators
organize monitoring efforts, and a recent analysis suggested that Piping
Plovers populations are increasing (Hecht and Melvin 2009b). On a smaller
scale, local population trends may be useful for understanding the effectiveness
of site-specific management strategies, especially within an adaptive
management context. Site managers may use local population trend analysis
to make decisions about management actions or restoration projects.
Our objectives were to quantify temporal trends in the reproductive ecology
of Piping Plovers from 1995 to 2007 at Jones Beach State Park (JBSP).
We assessed changes in the number of breeding pairs, productivity, total
number of young produced per year, and nest-initiation dates. We also compared
Piping Plover reproductive trends at Jones Beach to populations along
the Atlantic coast, and examined the effects of management practices on
Piping Plover reproduction at JBSP. We predicted that management would,
over time, increase the number of Piping Plover pairs and productivity. We
also hypothesized that plover population trends at JBSP would be similar to
other areas along the Atlantic Coast using similar management practices.
Jones Beach State Park is located on the western end of Jones Island, a
barrier island off the south shore of Long Island, NY. The Park, officially
opened in 1929, encompasses approximately 10 km of un-vegetated shoreline
that varies in width between 50–130 m. A large beach nourishment project
added 4.6 million m3 of sand to low-lying Jones Island to allow for the
construction of permanent park facilities (Blakelock 1953). Early managers
planted Ammophila breviligulata Fern. (American Beachgrass), to help stabilize
deposited sand. Jones Island is now densely vegetated by native and
introduced shrubs. In 1959, a 1600-m jetty extending southward from the
western tip of Jones Island was built to stabilize the entrance to Jones Inlet
and facilitate navigation. The jetty catches sand moved by long shore currents,
which flow westward along the south shore of the island. Since jetty
construction, sand has accreted east of the jetty and increased the breadth
of the island. The initial beach nourishment project, followed by extensive
vegetation plantings and sand accretion subsequent to jetty construction, has
2010 A.F. McIntyre, J.A. Heath, and J. Jannsen 495
all but eliminated storm overwash, allowing the dune ecosystem to flourish.
Currently, Jones Island is covered by an extensive dune field that extends
north from the primary dune at the sandy ocean beachfront. The areas farther
from the shore support Morella pensylvanica Mirbel. (Northern Bayberry),
Rosa rugosa Thunb. (Beach Rose), Prunus maritima Marsh. (Beach Plum),
and a few Populus tremuloides Michx. (Quaking Aspen).
In 1951, Piping Plovers were reported nesting on the western end of
Jones Island, then referred to as “Short Beach”, with an estimated population
of at least 75 pairs (Bull 1964). Since that time, the barrier islands
along the south shore of Long Island, NY have been extensively modified
and have become popular tourist destinations during the spring and summer
months when shorebirds are nesting. Monitoring and management
activities started on Jones Beach State Park (JBSP) in 1983. The National
Audubon Society’s Scully Sanctuary oversaw activities from 1983–1986,
and then The Nature Conservancy (TNC) conducted monitoring and management
activities from 1987–2002. In 1995, other groups such as the
Citizen’s Campaign for the Environment (CCE) and New York State Parks
(NYSP) began to participate in plover monitoring and management activities
that were coordinated among the three groups by TNC. Since 2003,
NYSP has been solely responsible for management and monitoring at Jones
Beach and a year-round, on-site biologist was stationed at the Jones Beach
Nature Center from 2000–2007.
Piping Plover reproduction
To monitor Piping Plover reproduction, 2–3 crews of 2 people systematically
searched for nesting plover pairs between mid-April and mid-August
1995–2007. All of the shoreline (≈10 km) was monitored every year. Annual
total pair counts were the sum of all the pairs found nesting along JBSP
each season. Once nests were found, observers monitored plover incubation
4–5 days a week by observing the birds from a distance of 30–100 m
away from the nest. Observers verified egg presence and counted eggs by
direct approach to the nest. The first day that a complete clutch (four eggs;
Cairns 1982) was present was recorded as the incubation initiation date. If
a nest was found with four eggs, then we back-dated from the hatch date to
estimate incubation initiation. If a nest failed during incubation, a second
nesting attempt was common, especially if failure occurred early in the
season. In such cases, adults typically remained in the area, and after 5–7
days, they began laying eggs in a second nest that was within 10 m of the
failed nest. Often at least one plover could be identified by plumage as being
from the failed nest. For each nesting attempt, stewards recorded nest
location by GPS, clutch size, incubation start date, exclosure date, hatching
and fledging date, and egg and chick fate. Plover chicks leave the nest
bowl shortly after hatching, but young were monitored regularly enough to
connect specific broods to specific nests based on chick ages. Annual productivity
was calculated as the total number of chicks fledged divided by
the total number of pairs per year.
496 Northeastern Naturalist Vol. 17, No. 3
Symbolic fencing and predator exclosures were placed around most
plover nests. The symbolic fence was constructed from mason twine strung
between metal fence posts, marked with bright flagging so it was clearly visible
to people. It posed no true physical barrier, and as such it was considered
“symbolic”. The fencing was installed prior to the breeding season (before
1 April) and was placed around all areas known to have had Piping Plover
nests in the past. This procedure typically resulted in a buffer of 3–200 m
between plover nests and areas frequented by beachgoers.
Predator exclosures were constructed from 1.5-m-tall and 10-m-long galvanized
turkey wire with a 5.1-cm by 10.2-cm mesh that was formed into a
3-m-diameter circle. The exclosures were dug 30 cm into the sand to prevent
predators, primarily mammals, from digging underneath. Fruit netting was
attached across the top to deter avian predators. Exclosures were installed
around most nests, though the need for exclosures was evaluated on an individual
nest basis. Nests were not exclosed if previous exclosures in that area
had higher predator activity and subsequent failure. Potential egg predators
included Vulpes vulpes L. (Red Fox), Procyon lotor L. (Raccoon), Corvus
brachyrhynchos Brehm. (American Crow), Corvus ossifragus Wilson (Fish
Crow), Haematopus palliatus Temminck (American Oystercatcher), Larus
argentatus Pontoppidan (Herring Gull), Larus marinus L. (Great Blackbacked
Gull), feral Felis catus L. (Domestic Cat), and Ocypode quadrata
Fabricius (Atlantic Ghost Crab). Once plover eggs hatched, we located pairs
and chicks daily, until the chicks were 25 days old, at which point they were
considered fledged (Elliot-Smith and Haig 2004).
We examined temporal trends in number of pairs, productivity, and total
number of young produced per year using linear, quadratic, and interceptonly
(null) models. We evaluated model fit by comparing each model’s
Akaike's information criterion adjusted for small sample sizes (AICc) and
selected the model with the lowest AICc (Anderson 2008). To examine shifts
in nesting chronology among years, we evaluated trends in the timing of the
first, average, and last incubation dates of first nests for each year. Renesting
attempts were excluded from this analysis. To compare plover breeding
trends on Jones Beach to other nesting areas, we used ANCOVAs to evaluate
differences in slopes among regressions of productivity and pair numbers
between 1995 and 2007 at JBSP, all nesting areas in the New York and New
Jersey recovery unit (NY/NJ), and all United States Atlantic Coast (USAC)
nesting areas (USFWS 2004, 2007).
Between 1995 and 2007, the total number of plover pairs has decreased
in a quadratic pattern (βyear
2 = 0.24, βyear = -980.38; Fig. 1). The number of
pairs ranged from 13–30, with the highest total number of pairs nesting in
1995. There was a drop in number of pairs from 1995 to 1999, and the lowest
2010 A.F. McIntyre, J.A. Heath, and J. Jannsen 497
number of pairs nested in 2002, a 57% decrease from 1995. Since 2002, the
population fluctuated around a median of 18 pairs. The total number of nests
exhibited a similar trend from 1995 to 2007 (Table 1).
Pair productivity ranged from 0.46 in 1997 to 1.72 in 2005. The
model that best described productivity from 1995 to 2007 had a positive
linear trend (β = 0.08 ± 0.02, 95% CI = 0.05–0.12; Table 2). The target
Figure 1. Temporal changes in the total number of nesting Piping Plover pairs at
Jones Beach State Park from 1995–2007.
Table 1. Total numbers of Piping Plover nest attempts, eggs, young, and productivity for nesting
birds at Jones Beach State Park, NY from 1995–2007.
Year Nest attempts Eggs Young Productivity
1995 43 152 24 0.80
1996 40 123 24 0.86
1997 40 138 12 0.48
1998 33 118 12 0.52
1999 18 66 12 0.80
2000 23 78 16 0.89
2001 16 53 20 1.33
2002 15 50 20 1.54
2003 24 79 28 1.56
2004 29 105 32 1.60
2005 23 73 31 1.72
2006 21 76 21 1.17
2007 23 88 24 1.33
498 Northeastern Naturalist Vol. 17, No. 3
productivity of 1.5 set by the USFWS Revised Recovery Plan was achieved
in 2002–2005 (Table 1). As productivity increased, the number of second
nesting attempts decreased (rs = -0.79, P = 0.001). Although pair number
has decreased, the total number of young fledged per year has increased
slightly during the study period (β = 0.86 ± 0.43, 95% CI = 0.03–1.70;
Table 2. Evaluation of trend patterns to explain annual changes in total number of breeding
pairs, productivity, and number of young for Piping Plovers breeding at Jones Beach State Park
from 1995–2007. Number of Parameters (K), Akaike information criterion adjusted for small
sample size (AICc), difference in AICc (Δ AICc), and model weights (Wi) are shown.
Model K AICc Δ AICc Wi
Total # of pairs
Quadratic 3 65.40 0.00 0.995
Linear 2 76.23 10.84 0.004
Intercept-only 1 80.77 15.37 0.000
Linear 2 7.27 0.00 0.776
Quadratic 3 9.86 2.59 0.213
Intercept-only 1 15.83 8.56 0.010
Total # of young
Linear 2 87.60 0.00 0.497
Intercept-only 1 88.32 0.72 0.347
Quadratic 3 89.92 2.33 0.155
Figure 2. Annual trends in first, mean, and last incubation-initiation dates for first
nests of Piping Plovers nesting at JBSP between 1995–2007.
2010 A.F. McIntyre, J.A. Heath, and J. Jannsen 499
Table 2). There was no significant relationship between the number of pairs
and productivity (rs = -0.52, P = 0.07), suggesting that productivity was
not density-dependent. The average incubation-initiation date was significantly
later as the years of the study progressed (β = 0.95 ± 0.26, 95% CI =
0.43–1.46, P = 0.0003; Fig. 2).
Trends in productivity were significantly different among all three plover
breeding areas (JBSP, NY/NJ, and the US Atlantic Coast), with greater
increases in productivity at JSBP compared with other areas (F2,33 = 10.30,
P = 0.0003; Fig. 3). Trends in pair numbers also differed between JBSP and
NY/NJ and US sites, with decreases at JBSP and slight increases in other
areas (F2,33 = 20.09, P = 0.0001; Fig. 4).
Over the past thirteen years, the Piping Plover productivity at JBSP
has increased while number of nesting pairs has decreased. Because there
has been an increase in reproductive success, one might predict that the
JBSP breeding population would increase, particularly since breedingsite
fidelity has been documented in other studies (Haig and Oring 1988,
Wiens and Cuthbert 1988, Wilcox 1959). Plover population declines despite
increased reproduction suggest that emigration or mortality of either
Figure 3. Piping Plover productivity at JBSP, New York /New Jersey (NY/NJ), and
the US Atlantic Coast (USAC) differed significantly among nesting areas. JBSP had
a higher rate of increase in productivity compared to the either the NY/NJ recovery
unit or the entire USAC population.
500 Northeastern Naturalist Vol. 17, No. 3
juveniles or adults affects the JBSP plover breeding population. Pauliny et
al. (2008) evaluated the effects of nest exclosures and found similar patterns
of increased nest survival, with no concomitant changes in breeding
population. In addition, Cohen et al. (2006) found that adults breeding on
eastern Long Island Beaches had high return rates but juveniles had very
low return rates.
Piping Plover pairs produced at JBSP may be attracted to favorable
conditions in other areas. The decrease in nesting pairs at Jones Beach
roughly coincides with a dramatic jump in pairs at West Hampton Dunes
(WHD), which is another barrier island along the southern shore of Long
Island (Cohen et al. 2009). The West Hampton Dunes pair numbers increased
from 0 in 1992 to 39 in 2000 after a winter storm resulted in a
large overwash fan in the bay on the north side of the island (Houghton
2005). Cohen et al. (2009) concluded that this increase could only be the
result of immigration (not local recruitment alone). Birds produced at
JBSP may have been attracted to this site. Unfortunately, this explanation
is impossible to assess without having information on movements from
Figure 4. Change in the number of Piping Plover pairs at JBSP compared to New
York /New Jersey (NY/NJ) (USFWS 2004, 2007) and the United States Atlantic
Coast (USAC) (USFWS 2004, 2007). JBSP had a significant decline in pair numbers
while NY/NJ and USAC populations experienced stable or slightly increasing pair
numbers. Number of pairs for each area was converted to the natural log + 1 to allow
for comparison across areas with markedly different abundances.
2010 A.F. McIntyre, J.A. Heath, and J. Jannsen 501
Emigration to other areas may be driven by loss of nesting or foraging
areas. It is possible that there is less nesting habitat in current years
compared to pervious years as processes such as shoreward migration of
dunes may result in less open beach area. Future management of JBSP may
benefit from annual habitat assessment. A Piping Plover foraging ecology
study conducted at Jones Beach in 1992–1993 indicated that Piping Plovers
nesting on the western portion of the beach had access to ephemeral pools
(10–35,000 m2), and these pairs had higher chick peck rates and survival
rates than broods that foraged on the ocean beach (Elias et al. 2000). By
1999, these ephemeral pools were overgrown with vegetation and no longer
provided suitable plover foraging habitat. The loss of these quality foraging
areas may have contributed to pair decline. In other years, plover pairs
have quickly responded to ephemeral pools by nesting near (less than 50 m) storm
overwash and rainfall pools, even in areas of high pedestrian traffic (A.F.
McIntyre, pers. observ.). This observation supports the idea that foraging
habitat is an important factor in nest-site selection and that the benefits of
good foraging habitat may outweigh the costs of human disturbance. Conservation
projects aimed at restoring brood foraging and nesting habitat could
have a positive influence on the local population at JBSP. Restoration efforts
that create both nesting and foraging habitat and maintain connectivity between
these two habitat types would be most effective, particularly if these
were in areas where human disturbance could be managed.
In recent years, plovers have had more successful first nesting attempts
and, because of this success, they made fewer second and third nests. Most
likely, management efforts that decreased egg depredation and human disturbance
had a positive effect on plover productivity. This result is consistent
with other studies that showed the effect of disturbance on productivity. For
example, a study in Nova Scotia found that human disturbance caused significantly lower fledging success (Flemming et al. 1988). In a Rhode Island
study, plover chicks that spent less time responding to human disturbance
had higher survival rates (Goldin and Regosin 1998).
In recent years, Piping Plovers initiated nesting at later times during the
nesting season. The reason for this shift remains unclear. Perhaps the portion
of the population that nested early has moved to other breeding areas
and only late breeders chose to nest at JBSP. Cairns (1982) noted that high
pair densities speed up territory establishment and courtship. It is possible
that the decrease in pair numbers, and therefore lower pair concentrations,
may have an effect on nest initiation. A comparison of JBSP nest timing with
other breeding sites would identify whether the shift is national, regional, or
isolated to JBSP, and perhaps provide insight into contributing factors. More
study is needed to identify the underlying causes for later incubation.
Management recommendations that have been suggested in the USFWS
revised recovery plan for Atlantic Coast Piping Plovers have been
regularly implemented at JBSP, resulting in positive productivity trends
that outpace trends for other sites within the NY/NJ recovery unit, as well
502 Northeastern Naturalist Vol. 17, No. 3
as trends for the larger Atlantic Coast population. Although Atlantic Coast
Piping Plover Recovery is coordinated at the regional scale of recovery
units, and evaluated for the whole population during 5-year status reviews,
implementation of on-the-ground management is the administrative
responsibility of a relatively small number of land managers and conservation
partners, whose authorities are often limited by property lines (Hecht
and Melvin 2009a). Endangered Species Act (ESA) compliance for many
of these partners is evaluated in several contexts (review of beach management
plans, special event permits, and other ESA consultations) based on
locally collected monitoring data, such as those reported here, that are often
restricted to a single property. This multi-year examination of patterns
in reproduction at JBSP illustrates the effectiveness of regular implementation
of management practices targeted to increase plover productivity by
one set of conservation partners at one property used by a small fraction of
the total Atlantic Coast population, but representing a significant fraction
of the Piping Plovers nesting within the NY-NJ recovery unit. The degree
to which effective management at the scale of a single property can contribute
to population recovery at larger scales is difficult to assess without
information regarding immigration and emigration from other properties,
which would require large-scale mark-recapture studies. Regardless, high
productivity documented at JBSP did not translate into local population
increases, perhaps due to the absence of suitable nesting habitat to accommodate
more pairs. Given the high productivity that is possible at this site,
nesting habitat restoration (particularly if it were combined with foraging
habitat restoration) could potentially contribute to both local and regional
We are thankful for the help of plover stewards. This work was supported
by New York State Department of Environmental Conservation and Hofstra
University. R.L. Burke, P.J. Doherty, C.A. Lott, and J.D. Williams made helpful
suggestions on the manuscript. Comments from P. Paton and two anonymous reviewers
improved the manuscript.
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