A Historical Estimate of Apparent Survival of American
Oystercatcher (Haematopus palliatus
) in Virginia
Author(s): Erica Nol , Sean P. Murphy and Michael D. CadmanSource: Waterbirds, 35(4):631-635. 2012.
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A Historical Estimate of Apparent Survival of American Oystercatcher
(Haematopus palliatus) in Virginia
ERICA NOL1,*, SEAN P. MURPHY2 AND MICHAEL D. CADMAN3 1Biology Department, Trent University, Peterborough, ON, K9J 7B8, Canada 2U.S. Geological Survey, Forest and Rangeland Ecosystem Science Center, Corvallis, OR, 97331, USA 3Environment Canada, Canadian Wildlife Service, 867 Lakeshore Road, Burlington, ON L7R 5A6, Canada *Corresponding author; E-mail: Abstract.—Using mark-recapture models, apparent survival was estimated from older banding and re-sighting
data (1978-1983) of American Oystercatchers (Haematopus palliatus) nesting on beaches and in salt marshes of coastal Virginia, USA. Oystercatchers nesting in salt marshes exhibited higher apparent survival (0.94 ± 0.03) than birds nesting on beaches (0.81 ± 0.06), a difference due to variation in mortality, permanent emigration, or both. Nesting on exposed barrier beaches may subject adults and young to higher risk of predation. These early estimates of adult survival for a species that is heavily monitored along the Atlantic and Gulf Coasts can be used to (1) develop demographic models to determine population stability, (2) compare with estimates of adult survival from popula-tions that have reached carrying capacity, and (3) compare with estimates of survival from other oystercatcher populations and species. Received 24 February 2012, accepted 17 July 2012. Key words.—American Oystercatcher, apparent survival, Haematopus palliatus, mark-recapture, shorebirds, Virginia.
The American Oystercatcher (Haemato- et al. 2005). In many states numbers of birds pus palliatus), a Species of High Concern in nesting on beaches has declined, possibly the eastern United States (Brown et al. 2001), due to human disturbance (McGowan and has an estimated Atlantic Coast population of Simons 2006; Sabine et al. 2008), while num-10,971 ± 298 individuals (Brown et al. 2005). bers of birds nesting in salt marshes and shell Survival estimates of banded individuals are rakes have increased, comprising the major-critical to the development of demographic ity of nesting pairs in eastern North America models (Hitchcock and Gratto-Trevor 1997) (Lauro and Burger 1989; Wilke et al. 2005; that can be used to determine trajectories of lo- Virzi 2010). Herein, we present a survival es- cal or regional populations. Survival estimates timate of the American Oystercatcher from of marked individuals in the past are rare but a Virginia study population that was derived comparisons across historical timeframes can from a banded breeding population from also help to inform and assess management ac- 1978-1983 nesting on coastal beaches and salt tions and future risks, especially in long-lived marshes in Virginia. organisms (De La Mare and Kerry 1994; Beiss-inger and Westphal 1998).
small in numbers relative to other North From 1978 to 1983, field work was conducted from American shorebirds, has re-colonized from March through July in salt marshes and beaches to the near extirpation (Mawhinney et al. 1999), and south of Chincoteague, VA, (37° 50’ N, 75° 35’ W; Nol increased over the last 50 years in the north- et al. 1984). From 1978 to 1981, breeding adult oyster-catchers were captured during the incubation period ern part of its range, reaching over 800 birds using drop traps placed over the nest (Mills and Ryder in the states of Massachusetts and New York 1979). Birds were banded with a federal aluminum band (Melvin 2007; New York State Department of and a unique combination of 2-3 spiral color bands on Environmental Conservation). During this the tarsometatarsus. Observations of color-banded oys-recovery period, populations appear to be tercatchers were conducted over the breeding seasons of 1979 to 1983. Sex of each bird was determined by size declining in the core, Mid-Atlantic breeding and weight with females both larger and heavier (Nol areas including Virginia and South Carolina et al. 1984). The study area and survey effort remained (Davis et al. 2001; Sanders et al. 2008; Wilke constant over the study period. We used Cormack-Jolly-Seber (CJS, Cormack 1964; Jolly 1965; Seber 1965) models using Program Mark 6.0 (White and Burnham 1999) to estimate apparent surviv- We uniquely marked 58 (31 in 1978; twelve al (␸) and encounter probability (p) from live encounter in 1979; four in 1980; eleven in 1981) nesting data. This open population model assumes: 1) capture has no effect on survival or encounter probability; 2) every oystercatcher has an equal chance of survival; 3) nesting females, 13 beach-nesting females, color bands are not lost; 4) capture periods are instanta- 17 marsh-nesting males, twelve beach-nesting neous relative to the intervals between them; 5) fates are independent, and 6) emigration is permanent (White unique live encounters from 1979-1983. Us- and Burnham 1999). Observations during this study sup-port these assumptions. To reduce the risk of violating assumption 4, capture periods were held constant (15 April-15 June) over the study period. Apparent survival is live encounter data (Ȑ = 1.57), adjusted AICc the product of true survival and site fidelity and, as a re- (QAIC ) by dividing the observed deviance sult, is negatively biased. We included time-dependence of the global model by the mean expected (t) and sex (sex) in our models of ␸ and p because both parameters are known to vary between sex and across deviance, and proceeded with model selec- years in several shorebird species (Sandercock 2003). Because habitat is also known to impact survival (Van de our analysis included apparent survival as a Pol et al. 2006), nesting habitat (habitat) was grouped into function of nesting habitat and detection that We assessed the fit of competing models using an information-theoretic approach (Burnham and Ander- the only other candidate model within two 6 son 2002). Selection of the best-fit model was done using QAIC units, included an effect of habitat on corrected quasi-Akaike’s Information Criterion adjusted apparent survival and retained a time- and for small sample size (QAIC , Lebreton et al. 1992). We considered all models less than two QAIC units from This model differed by only a single parame- the model that minimized QAIC . We ranked models by 6QAIC and included normalized Akaike weights (w ). ter, so does not improve model fit (Burnham We conducted a goodness-of-fit (GOF) test for the global and Anderson 2002; Arnold 2010). Apparent strap procedure (White and Burnham 1999). The boot- nesting on Virginia beaches was 0.81 (95% strap GOF test compares the observed deviance to 1,000 CI: 0.67-0.90), whereas adults nesting in salt randomly-generated replications, detects overdispersion in the data, and estimates a variation inflation factor, Ȑ, marshes exhibited higher apparent survival which corrects the data (Cooch and White 2009).
The Web of ScienceTM and unpublished sources are probability (p) was high but varied annually used to extract other estimates of adult survival for oys- Table 1. Cormack-Jolly-Seber candidate models we used to estimate apparent survival () and recapture probabil-
ity (p
) for American Oystercatchers in Virginia, USA, 1978-1983. Quasi-Akaike’s Information Criterion for small
sample sizes (QAIC ), differences in QAIC values (
6QAIC ), normalized model weight (w ), model likelihood,
number of parameters (K), and deviance (Dev) are provided.
aModel factors included: habitat = nesting in barrier beach or marsh habitat, t = annual variation, c = constant, and sex = male or female.
Table 2. Estimates of apparent survival for Ameri-
tat effects on survival, breeding-site fidelity can Oystercatchers nesting on beaches (
) and salt
or a combination of these two life-history marshes (
), encounter probabilities (p), standard
error (SE), lower 95% confidence limit (LCL), and up-
per 95% confidence limit (UCL) in Virginia from 1978-
1983 under the best fit model (
, p ).
ing habitats, then the discrepancy between apparent survival rates is a function of site fidelity varying by habitat. Using breeding and nonbreeding encounters of oystercatch- ers in Massachusetts, Murphy (2010) disen- tangled site fidelity from survival and pro- posed that local population swings are likely the result of birds emigrating from the study population. Congeneric oystercatchers ex- hibited higher levels of site fidelity to territo-ries that successfully fledged young (Safriel et al. 1984; Ens et al. 1992; Harris and Wan- terized as having high survival (Table 3) less 1997; Hazlitt and Butler 2001). Higher and breeding-site fidelity (Tomkins 1954; apparent survival of Virginia salt-marsh Hockey 1996). Thus, apparent survival of nesting oystercatchers may indicate higher adult oystercatchers is a credible estimate, reproductive output relative to those nest-and similar to that reported for American ing on beaches, as beach-nesting American Oystercatchers in Massachusetts (Murphy Oystercatchers experience both high mam-2010). The Virginia population was at a malian predation and human disturbance in much lower breeding density than the Mas- other parts of their range (McGowan et al. sachusetts population (Lauro et al. 1992; 2005; Sabine et al. 2008). Birds on beaches Murphy 2010), so density, at least, with this may have moved into nearby salt marshes limited sample, does not appear to impact (Wasilco 2008) or northward to growing adult survival. Our final model did not in- clude the variable sex, a result that contrasts Alternatively, equal levels of nest-site fidel- with those for the similar Eurasian Oyster- ity suggest a difference in survival between catcher (H. ostralegus, Durell 2007). By con- habitats with beach-nesting oystercatchers trast, our final model suggested that adult experiencing direct mortality. At the time survival was 13% greater for birds nesting of the study, one inlet between two barrier in salt marsh habitat than for those nesting islands used for nesting was closing by move-on coastal beaches. As apparent survival es- ment of sand, reducing saltwater flow to timates are the product of true survival and oystercatcher feeding areas. Reductions in the probability that an individual returns to food supply could have contributed to lower the breeding site (i.e. breeding site-fidelity), apparent survival of beach-nesting birds. Ad-these differences may be attributed to habi- ditionally, Red Foxes (Vulpes vulpes) denned Table 3. Estimated adult survival (SE) of oystercatcher species (Haematopodidae).
1True survival; all others are apparent survival. and hunted at beach study sites (Nol, pers. Burnham, K. P. and D. R. Anderson. 2002. Model Se-obs.). Emigration in a Massachusetts study lection and Inference: A Practical Information-the-oretic Approach, 2nd edition. Springer-Verlag, New population is estimated to be between 0-11% (Murphy 2010). Thus, it is likely that the dif- Cooch E. G. and G. C. White. 2009. Program MARK: ference in survival between habitats in Vir- A Gentle Introduction. 9th edition. http://www.phi- ginia is due to both emigration and direct Cormack, R. M. 1964. Estimates of survival from the sighting of marked animals. Biometrika 51: 429-438.
Davis, M. B., T. R. Simons, M. J. Groom, J. L. Weaver standing the role of site fidelity on adult and J. R. Cordes. 2001. The breeding status of the survival. Studies estimating inter-annual American Oystercatcher on the East Coast of North movements across the U.S. population from America and breeding success in North Carolina. Massachusetts to Florida are currently un- De La Mare, W. K. and K. R. Kerry. 1994. Population dynamics of the Wandering Albatross (Diomedea exu- Group et al. 2012). Movement data linking lans) on Macquarie Island and the effects of mortal- ity from longline fishing. Polar Biology 14: 231-241.
also allow study of carry-over effects (e.g. Durell, S. E. A. le V. dit. 2007. Differential survival in Duriez et al. 2012) and can be incorporated adult Eurasian Oystercatchers Haematopus ostralegus. Journal of Avian Biology 38: 530-535.
into future, more complex, mark-recapture Duriez, O., B. J. Ens, R. Choquet, R. Pradel, M. Klaassen. models (e.g. multi-state models, White et al. 2012. Comparing the seasonal survival of resident and migratory oystercatchers: Carry-over effects of habitat quality and weather conditions. Oikos 121: Ens, B. J., M. Kersten, A. Brenninkmeijer and J. B. Hulscher. 1992. Territory quality, parental effort, A. Baker assisted MDC and EN in capturing Amer- and reproductive success of oystercatchers (Haema- ican Oystercatchers. We are indebted to him for his topus ostregalus). Journal of Animal Ecology 61: 703- expertise. M. Davis and T. Simons discovered the po- tential of these data for survival analyses. We thank Harris, M. P. and S. Wanless. 1997. The effect of remov- C. Risley, the late E. Risley and the Marine Science ing large numbers of gulls Larus spp. on an island Consortium for providing accommodation. Any use of population of oystercatchers Haematopus ostralegus: trade, firm or product names is for descriptive purpos- Implications for management. Biological Conserva- es only and does not imply endorsement by the U.S. Hazlitt, S. L. and R. W. Butler. 2001. Site fidelity and re- productive success of Black Oystercatchers in British Hitchcock, C. L. and C. Gratto-Trevor. 1997. Diagnosing American Oystercatcher Working Group, E. Nol and a local population decline with a stage-structured R. C. Humphrey. 2012. American Oystercatcher population model. Ecology 78: 522-534.
(Haematopus palliatus), The Birds of North America Hockey, P. A. R. 1996. Haematopus ostralegus in perspec- Online (A. Poole, Ed.). Ithaca: Cornell Lab of Orni- tive: comparisons with other oystercatchers. Pages thology; Retrieved from the Birds of North America: 251-288 in The Oystercatcher: From Individuals to
Populations (J. D. Goss-Custard, Ed.). Oxford Uni- Arnold, T. W. 2010. Uninformative parameters and model selection using Akaike’s information criteri- Jolly, G. M. 1965. Explicit estimates from capture–re- on. Journal of Wildlife Management 74: 1175-1178.
capture data with both death and immigration – sto- Beissinger, S. R. and M. I. Westphal. 1998. On the use of chastic model. Biometrika 52: 225-247.
demographic models of population viability in en- Lauro, B., and J. Burger. 1989. Nest-site selection of dangered species management. Journal of Wildlife American Oystercatchers (Haematopus palliatus) in Brown, S. C., C. Hickey, B. Harrington and R. Gill. 2001. Lauro, B., E. Nol and M. Vicari. 1992. Nesting density The U.S. Shorebird Conservation Plan, 2nd edi- and communal breeding in American Oystercatch- tion. Manomet Center for Conservation Sciences, Lebreton, J.–D., K. P. Burnham, J. Clobert and D. R. Brown, S. C., S. Schulte, B. Harrington, B. Winn, J. Bart Anderson. 1992. Modeling survival and testing bio- and M. Howe. 2005. Population size and winter dis- logical hypotheses using marked animals: A unified tribution of eastern American Oystercatchers. Jour- approach with case studies. Ecological Monographs nal of Wildlife Management 69: 1538-1545.
Mawhinney, K., B. Allen and B. Benedict. 1999. Status schi) on farmland in Canterbury, New Zealand. Not- of the American Oystercatcher, Haematopus pallia- tus, on the Atlantic Coast. Northeastern Naturalist Sandercock, B. K. 2003. Estimation of survival rates for wader populations: A review of mark-recapture McGowan, C. P., T. R. Simons, W. Golder and J. Cordes. methods. Wader Study Group Bulletin 100: 163-174.
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