Biology and Conservation of American on the Outer Banks of North Carolina - 20 years of collaborative focal species research and management Ted Simons USGS Cooperative Fish and Wildlife Research Unit Department of Applied Ecology, NC State University Outline

• Overview of American Biology • AMOY Research on the Outer Banks – Changes in Distribution and Abundance – Factors affecting reproductive success • History of Working Group Oystercatchers of the Americas Haematopus palliatus Taxonomic Status

• Generally recognized as separate species from Haematopus ostralegus • Five subspecies are currently recognized • The subspecies galapagensis, endemic • Regularly hybridizes with to the Galapagos H. bachmani, and hybrids islands, may be a with H. ater are known separate species H. palliatus subspecies

• palliatus - Atlantic and coasts , , South America to southern , West Indies, and Pacific Central America

• durnfordi - (and ?)

• frazari - W.

• galapagensis - Galapagos Is.

• pitanay - W. South America to Chiloe Island, palliatus subspecies Habitats: Diverse, but Strictly Coastal Feeding • Food: almost exclusively shellfish and other marine that inhabit intertidal areas. • Foraging microhabitat: restricted to intertidal sand or mud flats, or reefs or shoals; less commonly, rocky shorelines Breeding Biology

• Strong mate and nest site fidelity • In North America, the nesting season begins in late March- early May • Oystercatchers lay 2-4 eggs in a shallow scrape • Incubation takes 27 days, during which time the eggs are vulnerable to nest predators, flooding, and exposure • Precocial chicks, 35-45 days to develop flight capability • Unlike most other shorebirds young fed by adults up to 6 weeks after fledging

• Globally: Least Concern (IUCN 2007) • US and Canada: Species of Greatest Concern • El Salvador and Mexico: Endangered • Guatemala: Near Threatened • Brazil: Considered Vulnerable in São Paulo state Population estimates: palliatus Distribution Season Individuals

Coasts E & S Non-breeding 11,000 USA Mexico Breeding 500 Caribbean Breeding 550 Non-breeding 550 Central Breeding 1751-2450 America Non-breeding 275-600 N South Breeding 200 America Brazil Breeding 6500 palliatus total - 21,326

Clay et al. in Ens, B.J. and L.G. Underhill. 2014. Conservation Status of Oystercatchers around the World. International Studies Number 20. US SHOREBIRD CONSERVATION PLAN SHOREBIRDS OF CONSERVATION CONCERN – 2016 LIST

ESA-listed

• Black-necked Stilt (knudseni) • Snowy (nivosus [Pacific Coast]) • Piping Plover • Eskimo Curlew • Red Knot (rufa)

Greatest Concern

• American Oystercatcher (North America breeding palliatus) • Snowy Plover (nivosus [Interior/Gulf Coast] and tenuirostris) • Wilson’s Plover (North America breeding wilsonia) • Mountain Plover • Bristle -thighed Curlew • Bar-tai led Godwit (baueri) • Red Knot (roselaari) Oystercatcher Research in North Carolina

• Research began on the Outer Banks in 1997 • Collaborative effort with the NC Wildlife Resources Commission, NPS, USFWS, NC Audubon • Studies of breeding biology now include data from over ~3,000 nesting attempts (~325 breeding pairs) Focal Species Conservation • Widespread along Atlantic and Gulf Coasts • Sensitive to a variety of factors affecting coastal resources – Habitat loss, coastal development – Pressure from human recreation – Pollution – Non-native predators • Large, charismatic, easily identified • Long-lived, amenable to long term mark-resight studies Research Methods

• Demographic Studies – Oystercatchers are easily marked, allowing us to identify individuals from year to year and track survival and movement – Each banded Oystercatcher has a unique code engraved on a plastic band that can be read with binoculars or a spotting scope Capture Methods

• Adult oystercatchers are captured with a decoy and noose carpet or a “Whoosh” net • Juveniles are captured just before fledging

AMOY noose trapping.mov Telemetry • Small chicks tagged with 2g glue-on transmitters • Fledglings and adults are tracked with larger transmitters glued to a leg band • Argos and GPS tags now feasible.

Reproductive Success

• The breeding season is divided into two distinct periods: nesting and chick rearing • Oystercatcher chicks are mobile shortly after hatch and do not remain in the nest • Nest survival is variable among sites and years, but is generally low; ~25% of all nests survive to hatch • 0.39 (SE 0.02) chicks produced per breeding pair (n = 2,985 nesting attempts, 1995 – 2014) Sources of Nest Failure During Incubation

Avian Ghost 5% Human 3% 3% Abandonment 6%

Mammal 54% Storm/Tides 29%

N = 478

Source of failure could not be determined for 52% of failed nests (N = 518) Sources of Nest Failure – Cape Hatteras Chick Survival

• Oystercatcher chicks are camouflaged and highly mobile, which makes them difficult to monitor

• We used radio tags to track chicks and identify sources of mortality Chicks Fledged per Breeding Pair

1.4 Cape Hatteras National Seashore

Cape Lookout National Seashore 1.2

1

0.8

Productivity 0.6

0.4

0.2

0 1999 2000 2001 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 Year Sources of Chick Mortality

Human 16% Environmental 30%

Predators 54% N = 37 Source of mortality can not be determined for ~ 50% of chick mortalities

1 0.9 0.8 0.7 0.6 0.5 0.4 0.3 0.2 Cumulative survival probability survival Cumulative 0.1 Schulte, S.A. and T.R. Simons. 2015. Factors affecting the reproductive success of American Oystercatchers 0 (Haematopus palliatus) on the Outer Banks of North 1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45 Carolina. Marine Ornithology 43: 37-47. Days observed Vehicles and Chick Survival

• Humans have direct and indirect effects on chick survival • Oystercatcher chicks are highly mobile • Small chicks use truck ruts and beach debris for hiding • Fledglings collide with vehicles at night • 13 chicks killed by vehicles on Hatteras and Lookout from 2003- 2007 • Cape Lookout closed beach sections with oystercatcher broods after two chicks were killed in 2005 • No mortality from vehicles documented on CALO after the policy was implemented Chick Survival and Vehicle Closures

0.6 N=146

0.5

0.4 N=120

0.3

0.2 Survival probability 0.1 0.267 0.476

0 Partial beach Full Beach

Closure type Daily survival rates (DSR) for American Oystercatcher broods on beaches with and without off-road vehicles.

1

0.99

0.98

0.97

0.96

DSR 0.95

0.94

0.93 No vehicles present 0.92 Vehicles present

0.91 0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30 32 34 36 38 Age (days) Vehicles, Chick Behavior, and Habitat Use

FullFull beach beach closures closures Partial beach closures

Below tide line Open beach Dunes 22% 26% 39% Chick habitat use

Open beach 39% Dunes 74% FullFull beach beach closuresclosures Partial beach closures Resting Resting 24% 31%

Out of sight 43% Chick behavior Foraging Out of sight 4% 56%

Foraging Moving 10% 16% Getting fed Moving Getting fed 1% 15% 0% Chick Survival - Access to Foraging Sites

1

0.99

0.98

0.97

0.96

DSR 0.95 0.94

0.93 Forage access 0.92 No forage access 0.91 0 3 6 9 12 15 18 21 24 27 30 33 36 39 Age (days) Hurricane Effects

• Hurricane Isabel made landfall on the Outer Banks in 2003 • Overwash and sand movement improved habitat and reduced predator populations • Large improvement in reproductive success in areas affected by the storm

0.9 0.8 Hurricane Isabel 0.7 0.6 0.5 0.4 0.3 0.2

Chicks fledged/breeding pair 0.1 0 1999 2000 2001 2002 2003 2004 2005 2006 2007 Hurricane Isabel Overwash Fans

1998 2004 Demographic Model Insights • Hurricanes can have strong positive effects on nest survival • “Bonanza” years may be very important for population stability • Oystercatchers are long-lived , so a single strong cohort can have long term effects on the population • Periodic depression of nest predator populations may be an effective management tool for inducing bonanza years

Schulte and Simons. 2016. Waterbirds. Predator Management

• Gathered baseline data on the ecology of raccoons South Core Banks in 2007 and 2008 – Population size – Home range size – Movement patterns – Population parameters • Age distribution • Survival • Recruitment • Evaluated the response of raccoon, shorebird, and turtle populations to an experimental removal of raccoons in 2008 - 2009

Parsons, A. W., T. R. Simons, A. F. O’Connell, Jr., and M. K. Stoskopf. 2012. Demographics, diet, movements, and survival of an isolated, unmanaged raccoon Procyon lotor (Procyonidae, Carnivora) population on the Outer Banks of North Carolina. Mammalia 77: 21–30. Camera Trapping Results • Total of 1,299 camera “captures” over 5 trapping periods from May 2007 to June 2008 • Total of 532 recaptures • Average population estimate based on photo- recaptures over five camera-trapping periods = 228 (SE 20) • Density ~ 10 raccoons per square mile

Parsons et al. 2015. Journal of Wildlife Management. Chicks fledged per breeding pair Chicks Fledged per Breeding Breeding per Fledged Chicks in December 2008 and April 2009 April and 2008 December in euthanized Core Banks South on and trapped were raccoons 149 Pair Effects of Human Disturbance

Objectives: • To assess if military overflights, vehicles, and recreational activities affect the behavior, reproductive success, and survival rates of breeding American Oystercatchers on Cape Lookout National Seashore in 2011 and 2012

Methods: • Baseline audio monitoring of ground noise level across the entire Park • 24- hour continuous audio and video monitoring of American Oystercatcher nests (48,000 hours) • Heart- rate monitoring of incubating Borneman, T.E., E.T. Rose, and T.R. Simons. 2014. Minimal changes in the heart rate of incubating American Oystercatchers individuals (12,000 hours) (Haematopus palliatus) in response to human activity. The Condor 116: 493-503.

Behavior of American Oystercatchers Before and During Human Activities

* * * * Heart Rate of American Oystercatchers Before and During Human Activities

* Limitations of Observational Studies - Lack of true replication, randomization, and controls

Table 1. Logistic exposure model exploring factors potentially affecting the daily survival rate of American Oystercatcher nests at North Core Banks, Cape Lookout National Seashore, USA, in 2010 and 2011 (n = 48). All factors evaluated were included in this model.

Explanatory variable Estimate (se) Z P-value Odds ratioa 95% CI for (eEstimate) odds ratio Intercept 0.800 (2.627) 0.304 0.761 Average daily aircraft events 0.013 (0.020) 0.660 0.509 Average daily vehicle events - 0.063 (0.021) - 2.941 0.003* 0.939 0.902, 0.982 Vehicle closureb Nest-only None 14.810 (929.137) 0.016 0.987 Drive-through - 1.373 (0.875) - 1.570 0.116 Ramp 0.104 (0.847) 0.123 0.902 Average daily Oystercatcher departures - 0.345 (0.575) - 0.601 0.548 Average daily nest attendance 0.043 (0.055) 0.777 0.437 Habitatc 0.038 (0.023) 1.653 0.098 Within dunes - 0.236 (0.740) - 0.319 0.750

Borneman, T.E., E.T. Rose, and T.R. Simons. 2016. Off-road vehicles affect nesting behavior and reproductive success of American Oystercatchers. Ibis 158: 261-78. Behavioral and physiological responses of nesting American Oystercatchers to off-road vehicles on North Carolina’s National Seashores

Experimental Study 2014-2015 Vehicles on Cape Hatteras ORV Drive-through Corridor Experiment

When vehicles are permitted to drive past American Oystercatcher nests 1)Does their incubation behavior change? a)Do they spend less time on their nests? b)Do they leave their nests more often? c)Do they spend less time resting? 2)Does their physiology (heart rate) change? Driving treatment

08:00 08:30 09:00 Control

08:00 08:30 09:00 Measuring incubation behavior

Rate of Leaving Head Tucking Incubation Time Measuring incubation behavior

Rate of Head Tucking Leaving Incubation Time Measuring incubation behavior

Rate of Leaving Head Tucking Incubation Time Measuring incubation behavior

Rate of Head Tucking Leaving Incubation Time Measuring heart rate

Experimental nests: 2014 Experimental nests: 2015 Does driving influence incubation behavior?

Do oystercatchers spend less time on their nests? Does driving influence incubation behavior?

Do oystercatchers spend less time on their nests? Does driving influence incubation behavior?

Do oystercatchers spend more time out of a head tucked position? Does driving influence incubation behavior?

Do oystercatchers spend more time out of a head tucked position? Does driving influence incubation behavior?

Do oystercatchers spend less time with their heads tucked? Does driving influence incubation behavior?

Do oystercatchers leave their nests more often? Does driving influence incubation behavior?

Do oystercatchers leave their nests more often? Does driving influence heart rate?

2015 Results Does driving influence heart rate?

2014 Results Does driving influence heart rate?

2014 Results Do these differences matter?

Incubating American Oystercatchers… • Stayed away from their nests for more time • Left their nests more frequently • Experienced reduced heart rates

Responses may diminish closer to hatching, and depend on time of day

Differences in heart rate, although statistically significant, are small, and there is considerable variation among individual birds

Do these observed differences influence demography? Vehicles on Cape Hatteras Implications for Management • Breeding population at CAHA has declined nearly 50% in past 15 years • Multiple factors influence the distribution, abundance, fecundity, and survival of AMOY on the Outer Banks • Predation • Storms • Human Disturbance • Birds are resilient and behavior is flexible, but they have limited tolerance of excess mortality of nests, chicks, and adults • Population level goals can help clarify management trade-offs The American Oystercatcher Working Group

• Research on American Oystercatchers in Virginia was pioneered by E. Nol in the 1980’s and summarized in her 1994 BNA account with R. Humphrey

• New research began on the Outer Banks of North Carolina in 1995. Additional research projects were initiated in Georgia and South Carolina in 2000.

• Interested biologists met at the Waterbird Society Meeting in 2001 and agreed to create an informal working group

• Research initiated in Virginia in 2002, and in MA and NJ in 2004. Today 30+ institutions and organizations participate

• Working Group meetings have been held annually since 2001 at study sites between Maine and Texas

• 2016 National Conservation Award from the USFWS NFWF Focal Species Initiative

• In 2008 working group members developed a 10 year business plan to promote the conservation of American Oystercatchers through a National Fish and Wildlife Foundation focal species initiative

• The resulting program provided $5 million in direct and $4.2 million in matching funds to support dozens of research and management projects conducted by working group members along the Atlantic and Gulf Coasts Working Group Affiliations

• Barataria-Terrebonne National Estuary Program • Clemson University • College of William and Mary • City University of New York • Delaware Division of Fish and Wildlife • Game and Freshwater Fish Commission • Georgia Department of Natural Resources • Gulf Coast Observatory • Louisiana Audubon • Manomet Center for Conservation Sciences • Maryland Department of Natural Resources • Massachusetts Audubon • Massachusetts Division of Fish and Wildlife • National Audubon Society • National Park Service • New Jersey Audubon • New Jersey Division of Fish and Wildlife • New York City Audubon • New York City Parks and Recreation • North Carolina Audubon • North Carolina State University • North Carolina Wildlife Resources Commission • Rutgers University • South Carolina DNR • Texas State University • Texas A&M • The Nature Conservancy • Trent University • University of Georgia • University of Houston • U.S. Fish and Wildlife Service • U.S. Geological Survey • Virginia Department of Game and Inland Fisheries • Wildlife Conservation Society Working Group Web Page http://amoywg.org

• A Working Group Web page created in 2002 provides: – A summary of Working Group objectives, activities, and contact information – Access to the Working Group list serve with >150 members – Banding protocols, banding summaries by state, banded bird reporting form – Descriptions of field methods – Data summaries, reports and publications AMOY BNA Account Revision

Section Authors:

Introduction: Erica Nol , Trent University Characteristics: Tracy Borneman, NCSU Coop.Unit, Distribution: Janell Brush, Florida Fish and Wildlife Conservation Commission Systematics: Theodore R. Simons, USGS NCSU Coop. Unit Migration: Shiloh Schulte, Manomet Center for Conservation Science Habitat: Jessica Stocking, NCSU Coop. Unit Food Habits: Jon Altman, National Park Service Sounds: Pam Denmon, U.S. Fish and Wildlife Service Behavior: Alex Wilke, The Nature Conservancy Breeding: Tom Virzi, Rutgers University Demography and Populations: Conor McGowan, Alabama Coop. Unit Conservation: Brad Winn, Manomet Center for Conservation Science Appearance: Sara Schweitzer, North Carolina Wildlife Resources Commission Measurements: Kim Peters, Massachusetts Audubon Future Research: Stephen Brown, Manomet Center for Conservation Science American Oystercatcher Working Group: Ruth Boettcher, Virginia Dept. of Game References: Sue Heath, Gulf Coast Bird Observatory

Other Contributors:

Robert C. Humphrey (RH), Brunswick Maine. Sean Murphy, USGS Forest and Rangeland Ecosystem Science Center Felicia Sanders, South Carolina Department of Natural Resources

American Oystercatcher Working Group. 2012. American Oystercatcher (Haematopus palliatus), The Birds of North America Online (A. Poole, Ed.). Ithaca: Cornell Lab of Ornithology. Mark-resight Banded as a chick, July 2013 Studies • ~4,000 birds marked with dual alpha- numeric bands color-coded by state, >41,000 resights Resighted, October 2013

• Winter re-sights of American Oystercatchers banded during the breeding season (map)

• Northern breeders appear to make longer movements

• Pairs show strong mate and nest site fidelity

• Winter movements of nearly ~4,000 km documented (Cape Lookout, NC to Laguna de T’erminos, Campeche, Mexico, Oct. 2013), 2015 Pacific Coast of Nicaragua and Honduras

• Generally strong connections between natal sites and nest sites for first-time nesters but some long dispersals documented

• Age at first breeding 3-5 years Online Mark-resight Database http://ancperch.org/amoy/index.html

• Launched in 2012 with NFWF funding to Audubon NC • >41,000 Records for ~4,000 individual AMOY • Reports from Maine to Mexico, 758 unique observers, 81 Working Group members • Secure, fully searchable • Includes >1,300 photos and Google Earth mapping capability • Accessible to the public • Process of archiving records older than 3 years at the BBL underway US Population - Winter Roost Surveys

• Aerial surveys from NJ to TX Winter 2003 and NY to TX Winter 2013 • Stratified sampling, 239 blocks • High tide +/- 2 hours survey window • Detection rates calculated from digital photographs and ground truth surveys – Detection rate 0.73 for flocks <50 birds – Detection rate 1.0 for flocks > 50 birds

Year Estimate S.E. 2003 10,971 298 2013 11,284 313

Brown et al. 2005. Journal of Wildlife Management 69: 1538-1545. Banding and Productivity Monitoring

# Nests # Birds Genetic Winter State # pairs monitored Source banded data surveys annually Maine 6 S. Schulte 20141 New Hampshire 0 Massachusetts 200 494 50-200 Yes No Melvin 2012 Rhode Island 27 3 No S. Schulte 20111 Connecticut 55 0 No S. Schulte 20141 New York 76 35 Yes S. Elbin 20152 New Jersey 400 339 75-150 No Yes T. Pover 20152 Delaware 15 24 Yes M. Bailey 20141 Maryland 108 2 Yes Traut et al. 2006 Virginia 730 267 100-400 No Yes Wilke et al. 20072 North Carolina 325 321 50-100 No Yes S. Schweitzer 20152 South Carolina 400 324 20-40 Yes Yes Sanders et al. 20082 Georgia 120 167 10-50 Yes Yes T. Keyes 20152 Florida 272 103 120-280 No Yes J. Brush 20152 Alabama 15 0 Yes Zdravkovic et al. 2006 Mississippi 25 0 Yes Zdravkovic et al. 2006 Louisiana 120 43 Yes E. Johnson 20152 Texas 170 295 50-90 No Yes S. Heath 20152 Total 3064 3970 Range-wide Perspectives

Winter roost surveys Breeding season surveys Linking to ~11,000 birds ~3,064 pairs = 6,128 birds With Demographic modeling of range-wide fecundity and survival estimates

Fecundity (F) Juvenile Second year Subadult survival Subadult Adult survival NC (1995-2013) survival survival w/o transition transition to adult (Sa) N=2,985 nests (Sj) (Ss1) (Ss2) (Ts2a)

0.20 0.70 0.92 0.77 0.15 0.92

5,815 Future Directions

• Strengthen linkages to Hemispheric conservation efforts • Refine range-wide monitoring methods to reduce survey bias associated with variations in sampling effort and detection probability • Expand citizen science and outreach • Measure range-wide changes in distribution, abundance, and productivity • Relate these metapopulation dynamics to changes in habitat use and availability Thanks!

The American Oystercatcher Working is comprised of over 30 institutions and organizations including; Barataria-Terrebonne National Estuary Program, Clemson University, College of William and Mary, City University of New York, Delaware Division of Fish and Wildlife, Florida Game and Freshwater Fish Commission, Georgia Department of Natural Resources, Gulf Coast Bird Observatory, Louisiana Audubon, Manomet Center for Conservation Sciences, Maryland Department of Natural Resources, Massachusetts Audubon, Massachusetts Division of Fish and Wildlife, National Audubon Society, National Park Service, New Jersey Audubon, New Jersey Division of Fish and Wildlife, New York City Audubon, New York City Parks and Recreation, North Carolina Audubon, North Carolina State University, North Carolina Wildlife Resources Commission , Rutgers University, South Carolina DNR, Texas State University, Texas A&M , The Nature Conservancy, Trent University , University of Georgia , University of Houston, U.S. Fish and Wildlife Service, U.S. Geological Survey, Virginia Department of Game and Inland Fisheries, and the Wildlife Conservation Society. I also thank; M. Bailey, J. Brush, R. Boettcher, R. DeMay, S. Elbin, S. Felton, S. Heath, N. Hostetter, E. Johnson, T. Keyes, D. LeBlanc, C. Mostello, T. Pover, S. Melvin, F. Sanders, S. Schulte, S. Schweitzer, S. Sinkevitch, S. Stanley, and A. Wilke for their individual contributions to this presentation. Students

Melissa Davis Conor McGowan Shiloh Schulte Arielle Parsons Jessica Stocking Tracy Borneman Eli Rose Nathan Hostetter Shilo Felton