Marie Cook

Taxonomy
 Kingdom: Animalia
 Phylum: Vertebrata
 Class: Aves
 Order: Scolopacidae
 Family: Charadriiformes
 Genus: Calidris
 Species: canutus
 Subspecies: islandica, rufa, roselaari,
rogersi, piersmai, and canutus
Genus Calidris
 Commonly referred to as “waders” or “peeps”
 Small to medium-sized shorebirds with long
wings and relatively short bills
 Found locally in large mixed flocks along
coastal estuaries, feeding via plucking, probing,
and plowing
 Arctic breeders that undergo long-hop
migrations
 Members include Sandpipers, Dunlins, and
Stints
 Calidris canutus: Brief Description
 Largest member of Calidris with stocky body
 Comparatively short bill, short legs
 About the size of an
American Robin
 Dull grey non-breeding
and bright red breeding
plumages
 Breed monogamously by
season in high arctic
climates and winter in
tropical or subtropical
climates
 Make one of the longest
annual migrations,
utilizing stopovers between jumps
 Six subspecies recognized, varying geographically, and morphologically
in terms of plumage, and wing and bill lengths.
 Global population ≈ 1,050,000 birds
Calidris canutus Subspecies
Subspecies (cont’d)

North American subspecies C. c. rufa shows

lowest estimated population size.
Differences Among Subspecies
 Divergence
 Last common ancestor can be traced to about 20,000
years before present.
 This was about the time of the end of the last Ice Age.
 As ice retreated, populations dispersed throughout new
suitable habitats.
 North American subspecies, rufa and roselaari diverged
about 1,200 years ago.
 Differences in Morphology
 Wing and bill length
 Brightness and patterns of plumage
Migration Strategy
 Fly thousands of miles without rest = long hop
migrant
 Travel in species-specific flocks due to differences in
speed, altitude, and flock configuration preferences
among shorebirds
 Fly in wind tunnels to decrease energy costs of flight.
 Decrease in digestive organ size for increased space for
fat reserves
 Increase flight muscle mass to endure long-distance
migration
 Departure times, ranges, routes, and stopovers vary
among subspecies
 Non-breeding juveniles may not migrate to Arctic.
 Metabolism During Migration
 Products of lipid and protein metabolism are associated with
flight.
 Highest in birds just arriving at
stopover
 Intermediate in refueling birds
 Lowest in inactive birds
 Energy required during long-distance
migratory flight is largely acquired from
fat.
 Supplemented by protein metabolism
 Found to be mostly proportional to
intensity of activity
 Provides compounds for citric acid cycle
 Explains why so many birds arrive at stopovers with largely
decreased flight muscles
 Organ reduction serves as evidence for protein use.
 Tissue and Organ Plasticity
 Brain seems to be only homeostatic organ
 Large changes seen mainly in digestive organs
 Negative relationship suggested between fat storage and
reduction of size in kidneys, liver, and intestine
 Organs aren’t used during migration
 Lessens maintenance cost
 Lowers basal metabolic rate
 Releases protein to be metabolized
 Allows protection from breakdown of more crucial organs
 Gizzard sizes are also reduced.
 HSC eggs are easily digested, allowing maintenance of
reduced gizzard during refueling.
 Lean muscle mass increases while staging
Reasons for Northward Migration
 Reduced Competition and Predation
 The harsh environment of the high arctic
is conducive to less organisms, including
parasites.
 Cryptic plumage and widely dispersed
large nest territories make knots harder
for predators to find.
 Increased Resources
 During the short arctic summer, the sun
shines all or most of the day long,
creating a superabundance of plants and
insects.
 This nutrition is crucial for courtship,
reproduction, and nest success.
 Knots time their migration on the
availability of resources.
Predation at High Latitudes
 Nest predation negatively
proportional to latitude
 Reduced predation
compensates for risks that
increase with latitude:
 Migration
 High mortality
 Poor weather conditions
 At right, percent decrease of
predation on artificial nests
with latitude (McKinnon
2010).
C. c. rufa  Population decline
 1980s = 100,000-150,000 knots
 2006 = 17,200 knots
 Four wintering ranges
 Mainly Florida, but also Georgia, and
South Carolina
 Texas
 Northern Brazil
 Tierra del Fuego (TDF)
 Staging sites
 Northward
 Argentina
 Venezuela
 Delaware Bay
 Southward
 James and Hudson Bays
 Massachusetts, Connecticut, Rhode Island
 Brazil
 Breeding range
 Central Canadian Arctic from
Southampton Island to Victoria Island
Distribution of C. c. rufa
 Breeding Range
 Arrive in Arctic tundra in June to claim nesting
sites, depart in mid-July to early August
 Live territorially on slopes and cliffs that are within
easy access to wetlands
 Territory area 1.5 km2
 Show high site fidelity
 Thrive on grasses and
their seeds until insects
emerge upon snowmelt
 Predators include Arctic
Fox, Snowy Owls, and a
variety of Jaegers, which
prey upon eggs and chicks.
 Nest success ≈ 75% (conservative)
 Reproduction
 Reach sexual maturity around
age 2
 Males usually arrive first to claim
nesting territory.
 Males build up to 5 scrapes out of
vegetation
 Males perform displays including
song flights during courtship.
 Female is fertilized via cloacal kiss
from male.
 One brood laid per year
 Clutch size = 3 or 4 eggs
 Incubation by males and females for
around 22 days; females first to leave
 Chicks born precocial; fledge after 18
days
Limiting Factors of Recruitment
 Density-dependent:
 Food, foraging success
 Suitable nest sites
 Predation
 At right, % juveniles vs.
index of wintering
population numbers
 A = before 1969
 B = 1969-1995
(Boyd & Piersma 2001)
 Density-independent:
 Weather conditions in
breeding grounds
 Stochastic events
Predation of Chicks
 Largely by Arctic fox
 Affected by predatory/prey
cycles of Arctic fox and
lemmings
 Lemmings on 3-4 year cycle
 When lemming population is
low, chicks are alternate prey.
 Predation of lemmings releases
pressure on knot population.
 Eggs and chicks predated increase
predator population.
Prevention of Nest Predation
 Adult plumage, nests, and egg coloration are cryptic.
 Males protect territory with ground and aerial displays.
 Shift in chemistry of preen wax
 Monoester waxes are easily detected by olfactory hunters.
 Maintained in ranges without
mammalian predators
 Upon breeding, composition
switches to diester.
 Less concentrated
 Harder to detect olfactorily
 Shift is energetically costly
 At right (Reneerkens, et al. 2005),
detection success of sniffing dog
 Solid line = monoester wax
 Dashed line = diester wax
 Coastal Range: Wintering and Staging
 Arrive in wintering sites in October and depart in late January and early
February
 Live gregariously in coastal wetlands
 Found in mixed flocks of shorebirds like dowitchers, plovers,
turnstones, and other sandpipers.
 Forage by pecking and probing soft substrate for marine invertebrates
like bivalves, crustaceans, and small snails
 Predators and competitors include Peregrine Falcons, harriers, Great
Black-Backed Gulls, and other birds of prey.
 Foraging
 Probe upper 3cm of moist
sandy substrate
 Herbst corpuscles
 Pressure sensory organs in
bill
 Allow detection of buried
prey and stones
 Uses water molecules in
pores of sand to sense
changes in pressure around
object
 Electron microscope images
at right; bottom image x100
Foraging (Cont’d)
 Efficiency affected by:
 Tides – foraging area only
available at low tide
 Prey density – positive
relationship
 Predation – negative
relationship
 Disturbance – presence of
humans, dogs, gulls, etc.
 Ideal
 Timed with prey availability
 Increased energy storage
 Non-free
 Travel from foraging to roosting
sites is energetically costly
 Seem to choose foraging sites
based on closeness to safe
roosting sites
 At left, observations match
most closely to predicted model
for ideal, non-free foragers
Delaware Bay Stopover
 Second largest stopover in the
world, utilized by 425,000 to
1,000,000 migratory shorebirds
 Last stopover before over 9,000
mile migration to reach
breeding grounds
 Knots begin to arrive in early
May and depart for breeding
grounds in late May and early
June.
 Shorebirds depend on horseshoe
crab eggs to refuel.
 Delaware Bay is a prime spawning
area for Horseshoe Crabs.
 Knots need to about double body
mass from 90-120g – 180-220g to
ensure survival and breeding success.
 Refueling success depends on arrival
time and prey availability.
Horseshoe Crabs (Limulus polyphemus)
 Harvesting increased in the 1990s by almost 10 times
 Eel and conch bait
 LAL medical testing to detect bacterial endotoxins
 Responsible for 90% decrease in egg availability
 From 1992 to 1997, the reported harvest of crabs increased by 20
times from about 100,000 to more than 2 million.
 By 2004, harvest still exceeded production.
 Recovery will be slow since HSCs don’t mature until about age 10.
 Importance of eggs to knots
 Easily digested when digestive
organs are reduced
 High in fat and protein
 Stopover at Delaware Bay is
synchronized to HSC spawning.
 Overharvesting has significantly
contributed to C. c. rufa decline.
Importance of Delaware Bay
 Utilized by 80% of North American Knot population
 No suitable stopovers between DE Bay and breeding
grounds in Arctic
 When eggs were overabundant, late-arriving knots could
still reach a healthy departure weight.
 In recent years, shortage of eggs slows weight gain in late
birds, reducing survival.
 Declines in wintering numbers from staging populations
suggest high mortality during migration due to poor
body condition upon departure from DE Bay.
 Poor health in part limits reproductive success.
 Foraging on Delaware Bay

Knots show a Type II functional response to prey density.

Cross-species Comparison
 Hudsonian Godwits (right)
 Share wintering grounds with
C. c. rufa in TDF
 Do not stop at Delaware Bay
 Show stable population
demographics

 Semipalmated Sandpipers,
Sanderlings (left)
 Use Delaware Bay as a crucial
stopover
 Show population declines
 Implications of importance of
Delaware Bay to shorebirds who
stage there during migration.
Life Table: 1995-1998 survival rates

(Baker, et al., 2004)

Life Table: 5% decline in 1995-1998 rates

(Baker, et al., 2004)

Life Table: 1998-2001 survival rates

(Baker, et al., 2004)

 Demography Models

Best Case: 1995-1998

Adult Survival = 85%
Juvenile Survival = ½ adult
Nest success = 75% (3/4 eggs)

(Baker, et al., 2004)

Compromised Adult Survival

Adult Survival = 80%

Juvenile Survival = ½ adult
Nest success = 75% (3/4 eggs)
Demography Models (cont’d)

Pending Extinction: 1999-2000

Adult Survival = 60%

Juvenile Survival = ½ adult
Nest success = 75% (3/4 eggs)
(Baker, et al., 2004)
 Comparison of Demography Models (Baker et al. 2004)

95% Confidence
95% Confidence
Interval
Interval

Annual adult Annual adult

survival = 85% survival = 56%
 Threats to Red Knot Population
 Before 1955, hunting accounted for majority of predation.
 Habitat Destruction
 Dredging
 Coastal Development
 Oil pollution
 Shortage of Resources
 Harvesting of HSCs in DE Bay
 Harsh arctic winters yield food late
 Disturbance
 Research efforts
 Beach use
 Dogs
 Climate Change
 Short term: advantageous because of lengthened Arctic summer
 Long term: disadvantageous because of decrease in tundra habitat needed for
breeding.
 Small population size
 Since large numbers, and sometimes entire populations, winter and stage together,
they are especially vulnerable to stochastic events.
 Decreased genetic variability can lead to the accumulation of detrimental traits.
 Conservation
 Status
 NJ – Listed as threatened in 1999
 US – Not listed
 Canada – Listed as endangered in 2007
 Western Hemisphere Shorebird Reserve Network
 Protect habitat in Argentina, Brazil,
Peru, Suriname, Mexico, U.S., and Canada.
 Manage about 81 million hectares of habitat
 Management of HSCs in DE Bay
 2008 harvest moratoriums
 Female crabs in Delaware
 All crabs in New Jersey
 All harvesting is prohibited during spawning (May 1-June 7)
 Shorebird Steward Program
 Volunteers educate the public about how their use of the beach could disturb
important shorebird habitats.
 Beginning in 2003, beaches in DE Bay used by migratory bird populations have
been closed to the pubic during staging.
 Since C. c. rufa is threatened in NJ, conservation officers can cite beachgoers for
disturbance and trespassing.

Literature Cited
2006. Magnificent Shorebird Migration. Prince William Network.
http://migration.pwnet.org/pdf/Magnificent_Shorebird_Migration.pdf
 Baker, Allen J, et al. 2004. Rapid population decline in red knots: fitness consequences of decreased refueling
rates and late arrival in Delaware Bay. Royal Society. 271, 875–882
 Battley, Phil, et al. 2000. Empirical evidence for differential organ reductions during trans-oceanic bird flight.
The Royal Society. 276: 191-195.
 Boyd, Hugh, Theunis Piersma. 2001. Changing balance between survival and recruitment explains population
trends in red knots Calidris canutus islandica wintering in Britain, 1969-1995. ARDEA. 89: 301-317.
 Brown, S., et al. 2000. National Shorebird Conservation Assessment: Shorebird Conservation Status,
Conservation Units, Population Estimates, Population Targets, and Species Prioritization. Manomet Center for
Conservation Sciences. http://www.Manomet.org/USSCP/files.htm
 Buehler Deborah M., A. J. Baker. 2005. Population Divergence Times and Historical Demography in Red Knots
and Dunlins. Condor. 107: 497-513.
 Cornell Lab of Ornithology. 2009. All About Birds. Cornell University.
http://www.allaboutbirds.org/guide/Red_Knot/lifehistory
 Gilg, Olivier, Nigel Yoccoz. 2010. Explaining Bird Migration. Science. 327:276-8.
 Gillings, S, et al. 2007. Shorebird predation of horseshoe crab eggs in Delaware Bay: species contrasts and
availability constraints. Journal of Animal Ecology. 76: 503–514.
 Greenberg, Russel. 2005. Birds of Two Worlds. Johns Hopkins University Press. Baltimore. 263-5.
 Harrington, Brian . 2001. The Birds of North America. Cornell Lab of Ornithology.
http://bna.birds.cornell.edu/bna/species/563/articles/introduction
 Hernandez, Daniel. Ornithology Lecture. Spring 2010.
 Literature Cited
 Landys, Meta, et al. 2005. Metabolic profile of long-distance flight and stopover in a shorebird. The
Royal Society. 272: 295-301.
 McKinnon, L., et al. 2010. Lower Predation Rates for Migratory Birds at High Latitudes. Science.
327:326-7.
 Morrison, Guy, R.K. Ross, L. Niles. 2004. Declines in Wintering Populations of Red Knots in Southern
South America. Condor. 106: 60-70.
 Niles, Lawrence et al. 2008. Status of the Red Knot (Calidris Canutus) in the Western Hemisphere.
Studies in Avain Biology. 36:1-185.
 Niles, Lawrence et al. 2009. Effects of Horseshoe Crab Harvest in Delaware Bay on Red Knots: Are
Harvest Restrictions Working? Bioscience. 59: 153-164.
 Piersma, Theunis et al. 1998. A new pressure sensory mechanism for prey detection in birds: the use
of principles of seabed dynamics? The Royal Society. 265: 1377-1383.
 Reneerkens, Jeroen, et al. 2005. Switch to diester preen waxes may reduce avian nest predation by
mammalian predators using olfactory cues. The Journal of Experimental Biology. 208: 4199-4202.
 Sibley, David. 2001. Bird Life and Behavior. Andrew Stewart. New York. 15-50, 273-287.
 Shorebird Phenomenon. Manomet Center for Conservation Sciences.
http://www.shorebirdworld.org/template.php?g=5&c=4
 Van Gils, Jan et al. 2005. Reinterpretation of gizzard sizes of red knots world-wide emphasizes
overriding importance of prey quality at migratory stopover sites. The Royal Society. 272: 2609-2618.
 Van Gils, Jan et al. 2006. Foraging in a tidally structured environment by red knots (Calidris canutus):
Ideal, but not free. Ecology. 85: 1189-1202.