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Trends in Piping Plover Reproduction at Jones Beach State Park, NY, 1995–2007
Annie F. McIntyre, Julie A. Heath, and Joseph Jannsen

Northeastern Naturalist, Volume 17, Issue 3 (2010): 493–504

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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. Introduction 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 - julieheath@boisestate.edu. 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. Methods Study site 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). Data analyses 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). Results 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 Productivity 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). Discussion 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 marked birds. 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 population increases. Acknowledgments 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. Literature Cited Anderson, D.R. 2008. Model Based Inference in the Life Sciences. Springer, New York, NY. 182 pp. Blakelock, C. 1953. History of Jones Beach State Park. Long Island Forum 53:23– 28. Bull, J. 1964. Birds of the New York Area. Harper and Row Publishers, Inc., New York, NY. 540 pp. Cairns, W. 1982. Biology and behavior of breeding Piping Plovers. Wilson Bulletin 94:531–545. Cohen J.B., J.D. 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