WILDLIFE ECOLOGY RESEARCH CHAIR
THIRD ANNUAL REPORT of the
WILDLIFE ECOLOGY RESEARCH CHAIR
Dr. Fred Cooke
Senior Chair, Wildlife Ecology
Simon Fraser University
MARCH 1996
I. INTRODUCTION..............................................................................................................3
II. THE CHAIR IN ACTION ...............................................................................................8
A. SPECIFIC PROJECTS ..........................................................................................................8
1. The Population Dynamics of Western Sandpiper............................................................8
i. Breeding Studies (Alaska)...........................................................................................8
ii. Migration Studies ......................................................................................................9
iii. Wintering Ground Studies.........................................................................................9
2. Winter Ecology of Dunlin............................................................................................ 10
3. Migration Physiology of Arctic-Nesting Shorebirds...................................................... 11
i. Age and sex-specific modulation of body composition during the migratory cycle ...... 11
ii. Fatty acid binding protein (FABP) and flight muscle conditioning............................. 11
iii. Lipid dynamics and fatty acid composition of western sandpipers ............................ 11
4. Reproductive physiology and ecology of birds.............................................................. 12
i. Physiological basis of individual variation in reproductive effort................................ 12
ii. Molecular genetics of marbled murrelets and other seabirds...................................... 13
iii. Hematological measures of health and condition of BC seabirds .............................. 13
5. Riske Creek Project..................................................................................................... 13
6. The Triangle Island Project.......................................................................................... 14
7. Demographic Studies of the Pacific Population of the Lesser Snow Goose.................... 15
8. Winter Population Ecology of Brant in the Fraser Delta ............................................... 16
9. Population Studies of Trumpeter Swans....................................................................... 16
10. Population Studies of Harlequin Ducks...................................................................... 17
i. Strait of Georgia, BC................................................................................................ 17
ii. Jasper National Park, ALTA.................................................................................... 17
11. Winter Studies of Grebes........................................................................................... 18
12. Population Status of Marbled Murrelets..................................................................... 18
13. Ecology of Barrow's Goldeneye ................................................................................. 19
14. Population Modelling and Demography .................................................................... 19
i. Estimation of average residency time......................................................................... 20
ii. Heterogeneity and estimator bias.............................................................................. 21
iii. Metapopulation dynamics - Pacific Flyway Snow Geese.......................................... 22
iv. Estimation of breeding propensity and age of first breeding...................................... 23
v. Philopatry, movement and LRS - spatial models and estimation ............................... 24
vi. A User's Manual for Program SURGE .................................................................... 24
vii. Systematic changes in species richness ................................................................... 25
B. OTHER INITIATIVES ........................................................................................................ 26
C. COMMUNITY INVOLVEMENT .......................................................................................... 27
D. FUNDING ....................................................................................................................... 28
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1. Research Funding Obtained (1995) ............................................................................. 28
2. Research Funding Obtained * or Applied For (1996) ................................................... 30
E. PUBLICATIONS ............................................................................................................... 32
1. Refereed Publications - 1995/96 or in press................................................................. 32
2. Other publications, book reviews, etc. ......................................................................... 35
3. Publications Submitted ............................................................................................... 35
4. Books Published in 1995............................................................................................. 36
3. Papers Presented at Conferences in 1995..................................................................... 36
F. PERSONNEL ................................................................................................................... 39
1. Board of Directors ...................................................................................................... 39
2. Research Team ........................................................................................................... 39
i. Faculty and Research Associates .............................................................................. 39
ii. Research Group ...................................................................................................... 39
3. Workshops/Meetings .................................................................................................. 40
i. Attended by Chair Representatives ........................................................................... 40
ii. To be attended by Chair Representatives.................................................................. 41
III. APPENDIX I. MISSION STATEMENT (JANUARY 1995 - F. COOKE) ............... 42
A. CONSERVATION GOALS, THE CWS PERSPECTIVE .......................................................... 42
1. Habitats...................................................................................................................... 43
2. Bird Species ............................................................................................................... 43
3. Integrated Population Approach .................................................................................. 45
i. Population Ecology .................................................................................................. 46
ii. Behavioural Ecology ............................................................................................... 46
iii. Physiological Ecology ............................................................................................ 47
B. ACADEMIC GOALS, THE SFU PERSPECTIVE ................................................................... 48
1. Population Ecology..................................................................................................... 48
2. Behavioural Ecology................................................................................................... 49
3. Physiological Ecology................................................................................................. 49
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THIRD ANNUAL REPORT - MARCH 1996
I. INTRODUCTION
The aim of the Third Annual Report is to:
i) give an overview of our third year of activities
ii) outline the progress on new and continuing projects
iii) describe the personnel involved
iv) give some indicators of our scientific and community involvement.
We are now almost two and a half years into the program of the Chair. The first year was
mainly one of planning the program, defining the objectives and establishing some of the
research projects. In the second year, we put into place most of the field research projects
defined in our program and now in our third year we are well into the generation of data
and the analysis and publication of results, as the summaries in this report will show.
There have been some changes in our personnel. Ian Jones, our sea-bird specialist, was
successful in obtaining a position as one of the Associate Chairs in our Atlantic Coast
equivalent, ACWERN, and is now a tenure track professor at Memorial University. He
will be continuing to collaborate with us on the Triangle Island project and is setting up a
similar field station off the coast of Newfoundland. We wish him all success. To replace
Ian we have been fortunate to be able to attract Doug Bertram, who brings his expertise of
seabird ecology and fisheries biology to the running of our seabird program. His primary
responsibilities will be the Triangle Island and the Marbled Murrelet projects. Doug's main
office is at the Pacific Wildlife Research Centre on Westham Island.
Joanne Harrington, our Administrative Assistant, has also gone on to bigger and better
things. She is now looking after the finances of a whole department (Political Science)
rather than just that of the Chair. Her place has been filled by Barbara Sherman, whose
experience in the Biosciences Department general office made the transition to
Administrative Assistant to the Chair an easy one.
Our research projects can be grouped into seven categories. Some have been underway
since the establishment of the Chair, others were commenced only in 1995. These
categories are:
1) The research program on Triangle Island was established in 1994 through the
cooperation of the Ecological Reserves Branch of the Ministry of Environment, Land and
Parks, and we are now starting our third field season. Data has been collected on the
demography of most of the key sea bird species on the island. The first thesis resulting
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from the establishment of the field station and the first arising as a result of the Chair's
activities was produced by Yolanda Morbey, an M.Sc. student with Ron Ydenberg,
working on the reproductive ecology of Cassin's Auklets. Hugh Knechtel has been
responsible for the night banding program and is continuing his M.Sc. thesis on Cassin's
Auklets. Doug Bertram is studying the feeding ecology of Rhinoceros Auklets, and
scientists attracted to the Island through the RNP program, Julia Parrish from the
University of Washington and Colleen Cassady St Clair from the University of Alberta, are
working with Common Murres and Tufted Puffins respectively. In addition, Laura Jones
has joined us as a new M.Sc. student studying Rhinoceros Auklets.
2) The research network on Western Sandpipers is now well established under the
coordination of Dov Lank. The work is funded mainly from NSERC operating grants and
the Latin American Program of CWS. Our projects extend from Nome Alaska, where
Brett Sandercock has completed his third field season towards his Ph.D., to as far south as
Chitre, in the Province of Herrera, Panama where Patrick O'Hara is working with
Francisco Delgado and his students. Other researchers involved in this work are Bob Elner
and Rob Butler of CWS, in the Fraser Delta, Horacio de la Cueva and Nils Warnock in
Ensenada, Baja California, Mexico and other scientists through our Research Network
Program (see below). Rob Butler and Tony Williams have recently developed a new
theory to explain the factors controlling the timing of migration and will soon be
submitting their manuscript to Nature.
3) Marbled Murrelet research continued at Desolation Sound for the second year, and
with funding provided by FRBC, a consortium of 5 forest companies and a CRD grant
from NSERC, we can confidently plan for work on this summering population to continue
for several more years which will allow us to carry out the first demographic and
reproductive physiology study of this elusive species. More than 400 birds have now been
marked and Tony Williams has recently developed a sex probe which will enable us to
accurately sex the birds that we catch. The work has been carried out in close
collaboration with Gary Kaiser and Kathy Martin of CWS and Andy Derocher of the BC
Ministry of Forests. Wendy Beauchamp has been hired as the coordinator of the project.
4) Work continues on several bird populations which winter in South Western BC. This
has been possible through funds provided by the BC Waterfowl Society, Ducks Unlimited
and the Fraser River Action Plan. Greg Robertson has learned much about the ecology of
a population of Harlequin Ducks which winters near White Rock with help from Billie
Gowans, Sean Boyd, Ian Goudie and others. Brian Carter is completing his work on the
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feeding behaviour of wintering Trumpeter Swans on Westham Island. Pippa Shepherd has
been radio tracking Dunlin and uncovering new facts about their distribution and nocturnal
behaviour. Eric Reed has unraveled the relationship between migrant and winter resident
populations of Brant geese, and has provided much useful information relevant to the
management of this species in the region. James Clowater continues his study of Western
Grebes in Saanich Inlet. Work on Barrow's Goldeneye continues, with Debbie van de
Wetering's moult study in the Yukon linking in with the breeding and wintering studies
which Sean Boyd and Saul Schneider of CWS are conducting.
5) Evan Cooch coordinates our Snow Goose project with a view to developing a
comprehensive model of the demography and interrelationships of the 3 western
populations of this species. Our collaboration with Russian scientists continues with
Barbara Ganter returning to Wrangel Island, off the north-western tip of Siberia for a
second season of goose banding. Barbara Pohl made observations on the neck collared
birds among our wintering population of Snow Geese and is currently completing a
manuscript showing the relationship of migrant birds to the population of winter residents
in the Fraser Delta.
6) Our major new initiative in 1995 was the Riske Creek Project, a comprehensive study
of an area of forests, grasslands and wetlands in the Central Interior of BC. The objective
of the study is to examine the effects of various land use practices on selected bird
populations in the region. Our first year of study, funded by the Interior Wetland Program
and Forest Renewal BC, was designed mainly to ascertain which species would be the
most important ones to concentrate on. Sean Cullen began a study of the Eared Grebe
population in the area. We use our demographic approach in conjunction with the long
term data on populations of waterbirds collected by the Waterfowl Management section of
the CWS. This project is a joint one with Kathy Martin and Sean Boyd of CWS.
7) The physiological ecology component of the Chair is now firmly established, with Tony
Williams’ lab fully equipped and with a range of analytical methods set up and validated.
Christopher Guglielmo and Oliver Egeler are investigating physiological aspects of protein
and lipid utilisation, respectively, in migrating shorebirds, and Julian Christians is working
on physiology variability in reproductive effort. Brett Vanderkist will be starting as an
MSc student in Fall 1996 to expand our work on the reproductive endocrinology and
molecular biology of marbled murrelets. In addition to these specific graduate projects, the
establishment of this analytical facility has allowed a physiological (functional) component
to be added to many of the other Chair projects, e.g. application of a novel molecular
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sexing technique, and improved indicators of body condition, now being used on Triangle
Island and in the Marbled Murrelet project.
In addition to the core funding of $330,000 provided by the CWS, NSERC and SFU, we
had external grants and contracts totaling $853,203. External funds have played an
increasing part in our ability to conduct the research projects and the graph in Figure 1
illustrates the importance of the funds in relation to our core funding.
CWS/NSERC Wildlife Ecology Chair Funding
1200000
1000000
800000
Core
Generated
600000
RNP
400000
LAP
200000
0
1993-4
1994-5
1995-6
Figure 1
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THIRD ANNUAL REPORT - MARCH 1996
We continued to be able to attract top quality scientists to our program through our
Research Network Program. This money, $75,000 annually, is provided by CWS to
expand the scope of the research to university faculty members throughout BC and
beyond, and has allowed an effective networking to be established within the mandate of
the Chair. A list of research projects and scientists funded by the Research Network
Program in 1995-1996 follows.
Senior Advisor
(Affiliation)
Research Network Funding Proposal
J.H. Christy/
P.R. Backwell
(STRI)
Feeding Behaviour and Ecology of the Western Sandpiper
in the Republic of Panama
$8,365.00
M.A. Bishop
(Copper River Delta
Inst.)
Conservation of Western Sandpipers Along the Pacific Flyway
$6,000.00
T. Williams
(SFU)
Physiological Mechanisms of Fat and Protein Metabolism in Migratory
Shorebirds`
$1,500.00
L. Dill
(SFU)
Predator-Prey Gaming Between Wintering Whimbrels
and Fiddler Crabs
$4,150.00
N. Haunerland
(SFU)
Flight Muscle Conditioning in Migratory Dunlin and
Western Sandpipers: The Role of Fatty Acid Binding Protein
$9,300.00
D. Lank
(SFU)
Winter Ecology of Dunlin in the Fraser River Delta
$7,000.00
L. Oring
(Univ. of Reno)
Overwinter Survivorship in Mexico/
Spring Migration Along the Pacific Flyway
$15,000.00
I. Jones
(SFU)
The Impact of Glaucous-winged Gull Predation and
Kleptoparasitism on Other Breeding Seabirds at Triangle Island
$9,365.00
R. Routledge
/E. Cooch
(SFU)
Local Philopatry and Home Ranges of
Harlequin Ducks over the Non-breeding Season
$7,000.00
C.D. Ankney
(Univ. of Western
Ontario)
Egg Laying Interval and Nest Site Selection in
Barrow’s Goldeneye and Bufflehead
$5,000.00
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Funding
Amount
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II. THE CHAIR IN ACTION
A. Specific Projects
At present, we have made no attempt to prioritize the projects below. They are presented
to give an account of the initiatives started to date.
1. The Population Dynamics of Western Sandpiper
The multifaceted study of Western Sandpiper gathered momentum during the year. A one
day workshop in Victoria, attended by over 30 active researchers, highlighted our state of
knowledge.
The long term objective of this study is to provide an understanding of the factors
controlling the population of this long-distance migrant which occupies several locations
during its annual cycle. Important questions relate to the factors influencing the species in
each of these locations and how migration events fit into the events which occur at the
various locations. Of conservation concern is the consequence of the removal or
deterioration of one or more of these habitats on the survival and reproduction of the
species. Events in the wintering grounds may affect migration and breeding events. We
have chosen one breeding area, several migration locations and two wintering sites.
i. Breeding Studies (Alaska)
Brett Sandercock has completed his third field season at Safety Sound, near Nome,
Alaska, where he has been studying breeding populations of Western and Semi-palmated
Sandpipers. More than 100 nests were found in each season and more than 3000 adults
and juveniles banded. Some of these birds have been relocated in Boundary Bay,
Ensenada, Mexico and at our wintering field site in Panama. Reproductive success and
survival estimates for the three years of study have been collected and factors affecting
variation in these values are being studied. Brett is currently writing several papers (see
bibliography), associated with his work.
Doug Schamel, who started as a Ph.D. student at SFU this fall, brought with him several
seasons' data on the breeding biology of western sandpipers at Cape Estenberg, on the
north shore of the Seward Peninsula, ca. 500 km north of Nome. He will be in the field
again this season, concentrating on his thesis work comparing the reproductive strategies
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of westerns and red-necked phalaropes. Doug was also awarded the 1996 SCSTKendall/Hunt Outstanding Undergraduate Science Teacher award from the Society for
College Science Teachers, for his teaching at the University of Alaska, Fairbanks..
ii. Migration Studies
We have played an active role through Nils Warnock, our Post-doctoral Fellow and with
RNP support to Mary-Ann Bishop to continue detailed radio tracking study to examine
the variability of individuals in terms of migratory patterns. Individual birds were followed
to various migratory stop-over sites. Pippa Shepherd monitored the radioed migrants
around Vancouver. This provided valuable data for testing Butler and Clark's dynamic
programming model of migratory decisions. Butler and Williams have been investigating
the importance of wind directions and speeds on migratory decisions, and Ydenberg,
Butler and Lank have been preparing a synthesis of migratory strategies in this species,
which Lank presented at the Cooper Meetings in San Diego.
Boundary Bay continues to be a focal point for our migratory studies. Mary Sewell has
now completed her stay with us and she has completed her work on the productivity and
annual cycle of invertebrates in the estuary. The work continues in the broader context of
shorebird habitats through the work of Pippa Shepherd.
iii. Wintering Ground Studies
a) Ensenada, Mexico. Three Mexican students (Alejandra Buenrostro, Yolanda Sandoval
and Guillermo Fernandez) have been investigating various aspects of the wintering
population ecology of Western Sandpipers, under the tutelage of Nils Warnock and
Horacio de la Cueva. Birds have been captured and banded for 3 seasons and analyses of
banding and resighting (recapture) data have, to date, focused on habitat use and spatial
movements of individuals and age/sex classes, and on return rates. The population is
significantly male biased, and birds of different ages use different areas. Return rates to
this location are low relative to other locations. This may reflect the lower overall numbers
of birds returning to the area in 1995/6. Sandoval and Buenrostro presented talks on their
work at the Western Sandpiper Conference in Victoria and all three students gave
presentations at the Cooper Society Meetings in San Diego.
b) Chitre, Panama. Patrick O'Hara has completed his second field season in this southerly
wintering area, working in close consultation with Dr. Francisco Delgado. A second
student, from the University of Panama, Jorge Moran will start a study of those birds
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which stay in Panama during the boreal summer. The program comprises a detailed
understanding of the relationship between the Western Sandpipers and their wintering
habitat, including food predators and competitors. An emphasis will be on variation in
demographic patterns and will be built on the long-term data set collected by Dr. F.
Delgado. Patrick has been able to detect three age classes of sandpipers and has strong
evidence of age related patterns of migration and summer residency. Yearlings appear to
lose weight in March and many may remain in Panama throughout the boreal summer.
Adults on the other hand gain weight rapidly in March prior to their spring departure to
the breeding grounds. The Panama population is not male biased and return rates are much
higher than to the Ensenada location.
Patrick is also doing a major study on the food base of the sandpipers in collaboration with
Dr. John Christy and Patricia Backwell of the Smithsonian Tropical Research Institute in
Panama.
2. Winter Ecology of Dunlin
Pippa Shepherd spent the past year collecting radio telemetry data on habitat use,
movements, home ranges, and activity budgets of four segments of the Dunlin population
(male and female adults and juveniles) with reference to environmental variables (time of
day, tide, weather, season). One interesting result is that the Dunlin regularly use the
surrounding agricultural lands for feeding at night, even though by far the most and "best"
mudflat habitat is also only available at night. We have also collected over 500 prey
samples in a variety of marine and terrestrial habitats that are used and not used by the
Dunlin, and these are presently being sorted. We have made regular Dunlin and predator
surveys throughout the winter, and have also collected data on Dunlin interactions with
predators whenever possible. Data collection is still underway, but we expect to have all
of this information integrated into interactive GIS maps before the end of 1996. This data
set can then be used to set up a sentinel species program and to make recommendations
for the conservation of wintering shorebirds in the Fraser River Delta.
In recognition of her work, Pippa Shepherd was the sole recipient of the Society of
Canadian Ornithologists prestigious Baillie Award.
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3. Migration Physiology of Arctic-Nesting Shorebirds
i. Age and sex-specific modulation of body composition during the migratory cycle
Western Sandpipers (n = 150) have now been collected from all stages of the migratory
cycle: wintering and pre-migration (Panama) and Spring and Fall migration (Fraser
estuary), for birds of both sexes and ages (by Christopher Guglielmo, Tony D. Williams
and Patrick O’Hara). Detailed body composition analysis (wet, dry and lean/lipid mass)
has been completed for 90 birds, and analysis of plasma samples is underway. Preliminary
results clearly demonstrate that widely used measures of individual ‘condition’ or ‘quality’
are misleading in determining major physiological changes, and therefore potentially
‘performance’, during migration. These measures have traditionally been an important
component of the assessment of the viability and health of bird populations and/or their
habitats. For example, the body mass and ‘condition’ of adult and juvenile female western
sandpipers is identical during Fall migration, but the bird’s mass is distributed in very
different ways in the body with juveniles having larger gut, liver and heart mass and
relatively lower fat mass. These differences might be related to patterns of age-specific
mortality which occur during migration.
ii. Fatty acid binding protein (FABP) and flight muscle conditioning
Fatty acid binding protein, an intracellular protein in muscle tissue, is thought to play an
important role in lipid utilisation during periods of intense activity, such as migration.
Muscle content of FABP could therefore represent a good indicator of a bird’s ability to
successfully complete migration. Moreover, as FABP might be increased just before, and
during migratory flight, it might indicate the migratory state of birds, i.e. whether they
have recently arrived or are about to depart. This knowledge is crucial if we are to
critically interpret other information on the condition of birds in relation at annual
variation, habitat quality, etc. Christopher Guglielmo, in collaboration with Dr. Norbert
Haunerland, has successfully isolated and purified avian FABP from western sandpiper
muscle, and is currently raising antibodies for use in development of an enzyme-linked
immunosorbent assay (ELISA).
iii. Lipid dynamics and fatty acid composition of western sandpipers
Previous studies have shown that several migratory bird species undergo seasonal
variation in the composition of fatty acids they store in depot fat, and it has been
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hypothesized that a shift to more unsaturated fatty acids occurs prior to migration to aid in
fuel mobilization for prolonged flight. Oliver Egeler, a Masters student supervised by Dr.
Tony D. Williams, is currently carrying out an analysis of western sandpiper adipose tissue
in order to determine if this phenomenon occurs in shorebirds too. Preliminary comparison
of wintering birds collected in Panama by Chris Guglielmo and spring migrants from
Boundary Bay (collected by Tony Williams and Chris Guglielmo) suggests that such a
pattern indeed exists in western sandpipers. Future work will attempt to determine
whether this shift is due to dietary events or physiological modulation in preparation for
migration. We hope to make use of our captive birds in these studies in addition to
collection and analysis of prey samples from Boundary Bay and possibly Sidney island.
The aim of this work is a) to identify patterns of lipid deposition that might influence
whether birds can migrate successfully or not, b) to determine the extent to which specific
dietary requirements may influence the quality of fat deposited, and c) to investigate
whether this influences habitat choice and, in turn, is influenced by habitat quality.
4. Reproductive physiology and ecology of birds
i. Physiological basis of individual variation in reproductive effort
Julian Christians, an M.Sc. Student under the supervision of Dr. Tony D. Williams, has
recently begun an investigation of the physiological mechanisms underlying intraspecific
variation in avian reproductive performance (specifically laying date, egg and clutch size).
The goal of such work is to determine the processes and resources which are limiting steps
in egg production. Furthermore, we seek to know why some individuals are better at
breeding than others, and thereby develop diagnostic tools to assess the “quality” of
different individuals. This research may also lead to collaboration with Oliver Egeler and
Chris Guglielmo since, in a general sense, we are all asking similar questions: how do
birds “fine tune” their physiology for the task at hand, be it migration or reproduction, and
what resources are required for such modulation? Initially, this study will be carried out
on wild starlings (Sturnus vulgaris) breeding in nest boxes at Agassiz, as these provide a
tractable model for rapid testing of techniques and hypotheses. To date, 120 nest boxes
have been set up at Agassiz, and initial tissue and blood samples have been collected from
pre-breeding males and females. We have also started to band birds in this population so
that the effects of reproductive effort on survival and future fecundity may be assessed.
Ultimately the aim is to extend this work to other free-living species: seabirds and
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shorebirds, e.g. preliminary blood samples have already been obtained for 100 breeding
western sandpipers, from Nome, Alaska, by Brett Sandercock.
ii. Molecular genetics of marbled murrelets and other seabirds
Marbled murrelets are a key species being studied by the Chair, with a focus on population
studies, habitat requirements and breeding biology. However, all of these studies are
hampered by the fact that marbled murrelets are sexually monomorphic, i.e. birds can not
be sexed using external morphometric measurements. In collaboration with Dr. Richard
Griffiths, of the Institute of Molecular Biology, Oxford University, UK, we are therefore
attempting to develop a genetic sex-specific marker for this species, which will allow birds
to be reliably sexed from very small samples of DNA (from blood or feathers). This work
is being carried out by Xiao-Hua Xue, a research assistant in Dr. Williams’ lab. If
successful we will extend this technique to other seabird species in relation to the Triangle
Island project.
iii. Hematological measures of health and condition of BC seabirds
In collaboration with Dr. Scott Newman, of the Wildlife Health Centre, University of
California, we are measuring a range of baseline hematological and serum biochemical
parameters which may provide useful indicators of immune status and condition of
seabirds in relation to environmental stressors, e.g. variability in food supply,
viral/bacterial disease, toxicity, oil spills. Preliminary plasma samples and blood smears
were obtained for Tufted Puffins, Cassin’s and Rhinoceros Auklets on Triangle Island in
1995, and this work will be extended to include additional species (e.g. Marbled
Murrelets) in 1996.
5. Riske Creek Project
This project was funded in 1995 by the Interior Wetland Program and by Forest Renewal
BC, and this funding has continued in 1996. The objectives in 1995 were to document
habitat use throughout the season of the various species of waterbirds using the Becher's
Prairie near Riske Creek in the Central Interior of BC. We built on the long term
waterfowl surveys carried out by CWS, but carried out more detailed observations
throughout the breeding season in order to understand how the birds used the habitat
during the pre-nesting, nesting, brood-rearing and molting period. The objective was to
prepare for more detailed studies of a few key species in the area. We determined that
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Lesser Scaup and Green-winged Teal would become the focus for future studies. Sean
Cullen started an MSc. project on the population of Eared Grebes on the prairie,
concentrating on the movement of birds, (both intra- and inter-seasonally) among the
ponds in the region. This work was in conduction with Sean Boyd and André Breault of
CWS, who have been studying these grebes for several years. Disturbance by cattle and
researchers formed an important practical component of the study. The research was
coordinated with Kathy Martin's work on hole nesting birds in the woodlands and forests
adjacent to some of the ponds and with Sean Boyd's detailed study of Barrow's
Goldeneye.
6. The Triangle Island Project
The CWS/SFU seabird research station survived its first winter on Triangle Island. With
the assistance of the Canadian Coast Guard, and permission from the BC Parks, the
research program was initiated in mid March and continued through September. Research
protocols established in 1994 were successfully executed on all research fronts despite the
departure of the project leader, Ian Jones. Ongoing demographic studies, a cornerstone of
the Chair's seabird program, produced 1680 captures (919 recaptures) of Cassin's Auklet,
and 538 captures (268 recaptures) of Rhinoceros Auklet largely due to the efforts of Hugh
Knechtel. Comparative data for Cassin's Auklet growth, fledging mass and age, and
productivity were also collected. Intensive research on Rhinoceros Auklet chick growth,
provisioning rate, and diet were conducted by volunteers Cecilia and Lynn BetancourtLougheed. Chair Research Network funds supported a pilot study of the effects of gull
kleptoparasitism on Tufted Puffin reproductive success by Colleen Cassady-St. Clair from
the University of Alberta. A separate but complementary study on factors affecting
attendance rates of Common Murres was initiated by Julia Parrish from the University of
Washington. The unique but small colony of BC murres had low attendance rates and the
longest chick fledging period that has been reported to date for any colony. Passerine
banding, supported by the Bailey Foundation in August and September produced 897
captures.
Through networking and cooperation, research efforts on Triangle were linked with
concurrent studies at other BC seabird colonies. Standard protocols for Rhinoceros
Auklet chick growth and diet studies were executed on 1) Seabird Rocks (West of
Barkley Sound) by Gail Davoren, an M.Sc. candidate at the University of Victoria; and
Anthony Island (southern tip of Queen Charlottes) by Todd Golumbia of Parks Canada.
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For the second consecutive year, comparative data on Cassin's Auklet nestling growth,
fledging mass and age, and productivity were collected on Frederick Island by Anne
Harfenist of the CWS. For the first time data on Cassin's Auklet nestling growth was
obtained from Limestone Island (Queen Charlottes) by Tony Gaston (CWS) and the
Laskeek Bay Conservation Society. These linkages will be fundamental to the
development of time series data for BC seabirds and will play a key role in understanding
how large scale marine phenomena affect performance on seabird colonies.
Yolanda Morby completed and successfully defended her M.Sc. thesis “Fledging
variability and the application of fledging models to the behaviour of Cassin’s Auklets
(Ptychoramphus aleutiaes) at Triangle Island, British Columbia” under the supervision of
Ron Ydenberg.
7. Demographic Studies of the Pacific Population of the Lesser Snow Goose
There are 3 populations with some gene exchange occurring among them. These are:
i) Wrangel Island breeding birds which winter in the Fraser and Skagit Deltas,
ii) Wrangel Island breeding birds which winter in California, and some of which are
transients in the Fraser and Skagit Deltas
iii) Banks Island breeding birds which also winter in California.
We are continuing our collaboration with Drs. Syroechkovsky, Litvin and Baranyuk on
the population ecology of the Wrangel Island birds. Dr. Barbara Ganter joined in with the
banding of the breeding birds for a second year despite very difficult logistics. More than
1000 birds were neck collared and for the second successive season, breeding success was
very low. We continued studying these banded birds during the winter months, building on
the data collected on the wintering Fraser-Skagit population by Dr. S. Boyd of the
Ecosystems Research Division CWS. Sean successfully defended his Ph.D. Thesis in
October 1995, under the direction of Dr. Alton Harestad. Barbara Pohl was specifically
studying the relationships between the resident wintering birds and the transient birds
which stage briefly in the Fraser Delta on their way to California.
The modeling component of this study continued at a slower rate than in 1994, while we
await the results of analyses carried out by biologists from the Prairie and Northern
Region of CWS.
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We continued the banding of birds from the breeding population on Banks Island, NWT.
This work took place again with funds from the Arctic Goose Joint Venture (AGJV), and
was coordinated by Dr. Evan Cooch. Almost 2,000 birds were banded in 1995.
Other aspects of the Lesser Snow Goose work are described below under the section on
Population Modelling and Demography.
8. Winter Population Ecology of Brant in the Fraser Delta
Work continues on the wintering population of Brant from Boundary Bay. The objectives
of the study were to determine whether there is a wintering population, to determine its
size and distribution within the Puget Sound - Strait of Georgia area, and to differentiate
wintering birds from migratory ones.
Eric Reed’s data indicate there is no major movement of migratory Brant through
Boundary Bay in the fall. Most of the birds that appear in November remain throughout
the winter. Numbers remain relatively stable from December to mid-February, when the
onset of Spring migration is apparent. Boundary Bay seems to be a very important
stopover site in spring. A maximum of 3,600 Brant were recorded there in late March,
1995. There is a lot of day to day fluctuation in numbers present in the bay but it is not
known where the birds go when not in the bay, There appears to be some interchange with
populations in Northern Washington. At least four individuals have been sighted in both
Padilla and Boundary Bay this year. There is clearly a resident population in Boundary
Bay in winter and that there is a strong philopatry from year to year (approx. 70% for
residents). Population size has not been estimated with great confidence yet, but is likely
to be between 600 and 1,000 birds. The number of birds wintering in the area seems to be
increasing. Maximal number of Brant seen in Boundary Bay in 1993/94 during the winter
was 400, while it was 450 in 1994-95, and 550 in 1995-96.
9. Population Studies of Trumpeter Swans
Populations of Trumpeter Swans wintering in BC have increased considerably in recent
years and currently threaten both agricultural crops and coastal salt marches, perhaps to
the detriment of other species of birds which use them. In the late fall of 1993 we
embarked on a study of this species, with a view to understanding the causes and
consequences of this population increase. Mr. B. Carter, an M.Sc. student is now in the
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final year of this study which concentrates on assessment of the food requirements the
winter population of swans using Westham Island.
10. Population Studies of Harlequin Ducks
In parts of their range Harlequin Ducks are becoming increasingly rare and are subject to a
wide range of environmental hazards such as oil spills, rafting and coastal development.
Little is known of the demography of the species and two studies were initiated in 1993:
i. Strait of Georgia, BC
In 1992, Ian Goudie initiated a wintering study in the Strait of Georgia as part of a
broader study of wintering populations of sea ducks. Several moulting populations were
banded for demographic studies. One of these populations, near White Rock, has formed
the basis of a detailed study by Greg Robertson, a Ph.D. student, on the demography and
winter dispersal of this species. He has provided the first detailed description of the
moulting and basic plumages of the species, and has observations on the movement
patterns of more than 100 individually marked birds. He has documented re-pairing of
previously paired birds, and shown that most pairing takes place early in the winter, much
earlier than had previously been suspected. Exchange of birds among different wintering
populations has been quantified.
ii. Jasper National Park, ALTA
The investigation of the breeding biology of Harlequin Ducks in Jasper National Park
completed its third year. The study focuses on the feeding ecology of the ducks in the
Maligne Valley, with particular reference to possible disturbance from white-water rafting.
Mr. W. Hunt will defend his M.Sc. based on the work in Summer ‘96, and the study is
being carried on by a new student.
In 1995 a second study of breeding birds was started at Banff National Park, under the
direction of Cyndi Smith of Parks Canada. This was closely integrated with our studies
and several individually marked birds from our wintering studies in the Strait of Georgia
were caught in the Park.
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11. Winter Studies of Grebes
James Clowater, an M.Sc. student at SFU, is continuing his project on food utilization and
distribution of Western Grebes in the Straits of Georgia during the winter months. This
forms part of a larger picture aimed at understanding the importance of the Coastal Waters
of BC for wintering water bird populations. James has made the interesting discovery that
Western Grebes feed little during the daylight hours but start feeding actively around dusk
and dawn. This coincides with the upward vertical nocturnal migrations of Euphausids,
which may in turn attract fish species, which feed on them and may be the prey of the
Grebes.
12. Population Status of Marbled Murrelets
The long-term collaborative research project on Marbled Murrelets continues in
Desolation Sound, B.C. Gary Kaiser (CWS); Andy Derocher (BC Forest Service); Fred
Cooke, Tony Williams, Doug Bertram, Wendy Beauchamp (Simon Fraser
University/CWS Cooperative Wildlife Research Chair) and Kathy Martin (CWS/UBC) are
coordinating studies of population demography and genetics, reproductive physiology and
marine and nesting habitat use. Two SFU graduate students are also involved: Irene
Manley, an M.Sc. student with Alton Harestad, will continue a study of nesting habitat
selection and Cecilia Betancourt-Lougheed, co-supervised by Fred Cooke and Doug
Bertram, will begin an M.Sc. project on marine habitat use and movement patterns of
murrelets.
Aerial telemetry of 37 radio-tagged birds to locate nests and track movements of
individuals was carried out in 1995. Eight Marbled Murrelet nests were discovered in the
Desolation Sound area by a crew conducting systematic searches and tree-climbing. The
information obtained on nest site characteristics represents a large increase in knowledge
of habitat use as only about 20 Marbled Murrelet nests have been found in all of B.C. to
date. All of the nests found in 1995 were in high elevation patches of old-growth forests
within 5-15 km of marine foraging areas.
Gary Kaiser plans to establish a study site in the Queen Charlotte Islands in 1996 and
begin an intensive capture and banding project. This will allow future comparisons of a
population from an area where habitat is less fragmented by forestry to a population in
Desolation Sound where logging has been more intense.
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In 1995, as part of the research program of the life history traits and population status of
this elusive species we captured 208 Marbled Murrelets and banded 189 previously
unbanded individuals. This was the second year of an intensive banding project in this
study area. A total of 398 individuals in the population have been banded since 1991 when
CWS initiated a pilot capture project. In 1995, we recaptured 9 murrelets banded in
previous years: 1 from 1991, 1 from 1993 and 7 from 1994. Site fidelity, as indicated by
inter-year recaptures or resightings, is a crucial aspect for demographic studies of this
species involving capture-mark-recapture techniques.
In conjunction with the banding, blood and feather samples were collected for DNA and
hormonal analysis to help understand population structure and breeding status of captured
individuals. These genetic and endocrinology aspects are being carried out in collaboration
with Drs. Kathy Martin (CWS/UBC), Tony Williams, Vicki Friesen (Queen's University)
and John Wingfield (University of Washington).
13. Ecology of Barrow's Goldeneye
Debbie van de Wetering continued her M.Sc. project on the moulting ecology of male
Barrow's Goldeneye. She has completed her second season of capturing moulting birds in
the Old Crow Flats and is planning a final field season in 1996. Her work has shown the
habitat requirements of the moulting birds and differences in requirements of birds of
different ages. This work correlates well with the breeding ground studies carried out by
Sean Boyd and Saul Schneider at Riske Creek and their wintering observations of this
species in the Lower Mainland. Hopefully these two studies will show further insight into
the annual patterns of behaviour of this species.
14. Population Modelling and Demography
Demography is an approach to the study of populations which is based on individuals.
Modern demographic methods are aimed at incorporating as much information about
individuals as possible, while at the same time dealing with problems of deriving models
which are tractable, both in terms of estimation of parameters, and in providing a practical
tool for population analysis.
In contrast to the time and energy focused on the theoretical implications of differences in
demographies among species or taxa, comparatively little work has been done on the
actual problems of estimating these parameters. Applied biologists and wildlife managers
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have widely adopted estimation methods, perhaps motivated by the "real world" concerns
of applied biology, but applied research is often only tangentially grounded in modern
theory of population dynamics.
Demography and population ecology are important parts of the Chair's research efforts
and we are well positioned to bridge the gap between modellers and practitioners, by
thorough investigation of the modern methods of estimation and enumeration, and by use
of these estimates in a broad practical context.
Our current demographic research reflects these aims; we have a number of projects
aimed specifically at exploring the mechanics and application of robust methods of
estimation, and others where we explore the theoretical ramifications of demographic
variation and change.
Current Projects:
i. Estimation of average residency time
This is a collaborative venture between Evan Cooch of our group, Rick Routledge of the
Statistics Department of SFU, Bill Link of the National Biological Service, and John
Smith, formerly with CWS. How long animals spend during migration stopovers is of
considerable interest, both in terms of practical management of many migratory species,
and also in terms of theoretical interest in the causes and consequence of migratory
behaviour. An hypothesis to explain the pattern and process of migration stopping is that
stopovers are adaptive behaviours to minimize mortality risk commensurate with
decreasing physiological condition. Migration flight is costly, and individuals must stop to
recoup depleted nutrient reserves necessary for continued migration. Where individuals
stop is often known but not how long they stay. In order to more fully understand the
dynamics between physiological requirements, stopping decisions, and pattern and rate of
food intake, the average residency time of individual animals needs to be estimated. For
example, birds of common physiological condition prior to stopping may have to remain
resident at the stopover site longer if food supplies are either of poor quality or patchily
distributed.
Average residency time has traditionally been estimated in terms of the difference in time
elapsed between peak arrival and peak departure dates. While this is clearly a reasonable
first approach, it suffers from the general problems of ignoring individual variation, and is
most useful when arrival and departure of individuals is highly synchronous at the
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population level. More recently, there have been attempts made to estimate average
residency time directly using radio-telemetry devices. While this clearly is the optimal
approach, it is generally restricted in terms of the numbers of individuals which can be
radio-marked, and as such, there may be problems extrapolating to the population as a
whole.
Recently, two new approaches to estimation of average residency time have been
proposed. The first involves using standard Jolly-Seber capture-mark-recapture (or
resight) models (hereafter, CMR). The analyses use recapture (or resight) data from
stopover sites to estimate the proportion of animals missed by the sampling (1-resight
rate), and estimates of local "survival rate", which in fact is primarily the probability of
remaining at the sight. If we interpret our estimates of "local survival" in this way, then we
can derive a first approximation using existing estimators for mean life span - we draw the
analogy that mean interval between birth and death in equivalent to mean interval between
arrival at a stopover site ("birth") and final departure ("death"). If the period-specific
probability of leaving (S) is constant, we can express the average lifespan (or residency
time simply as -1/ln(S). The principal advantage of this approach is that it is extremely
convenient. However, the assumption of constant departure probabilities is likely to be
incorrect in most cases. Preliminary work (Cooch & Link in prep) has attempted to extend
this approach without this (and related) limiting assumptions. We plan on extending this
work with more formal statistical modeling of average residency time in a CMR context.
Recently, Routledge et al. (In press) developed a second, different approach to the
estimation problem, by deriving an estimator based on frequencies of reobservation of
marked individuals. This approach is particularly appropriate for cases where animals are
not physically recaptured. We are currently comparing the relative advantages of the two
approaches, using a combination of computer simulations and field testing using recent
work on the population of Pacific Black Brant wintering in the Vancouver area. Some of
this work is currently funded by a grant from the Research Network Program.
ii. Heterogeneity and estimator bias
Traditional methods of capture-mark-recapture (CMR) analysis for open population
("open" = populations where animals can enter or leave the population with some nonzero probability) make a number of assumptions. One of these assumptions is that every
animal present in the population has the same probability of capture. Violation of this
assumption is known as the problem of heterogeneity.
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Since many migratory populations undoubtedly contain a variable mixture of residents and
transients, and different proportions of birds with specific recapture rates among residents,
determining the effects of heterogeneity on our estimates is of considerable importance.
Early work by Carothers (1973) and Gilbert (1973) suggested that heterogeneity in
recapture rates contributed only a small degree of bias to estimates of survival rate.
However, more recent analyses using more complex "ultrastructural models"
demonstrated that, in general, heterogeneity led to a significant negative bias in the
estimate of survival during the first interval after marking (e.g., Buckland 1982; Loery et
al. 1987). We are using computer simulations to further explore this question. Our main
intent is to provide adequate null hypotheses for expectations based on extreme forms of
heterogeneity (i.e., no transients, heterogeneity in p among residents; no heterogeneity
among residents, but some proportion of transients).
Some of this work is currently funded by a grant from the Research Network Program.
iii. Metapopulation dynamics - Pacific Flyway Snow Geese
This work is a collaborative venture between, Evan Cooch, Solange Brault of the
University of Massachusetts, John Takekawa and Michael Samuel of the US Fish and
Wildlife Service. Recent concerns about the Wrangel Island (WI) population result from
the perception that the number wintering in California has decreased. Such changes in
numbers and relative wintering distributions could reflect changes in fecundity and
survival, change in fidelity to both breeding and wintering sites, or both. A smaller
subpopulation in California could reflect at least three possible causes. First, an increasing
number of birds which would normally winter in California may be "short-stopping" in the
Fraser-Skagit area. Second, survival of the southern subpopulation may be lower. Finally,
increasing numbers of the southern subpopulation may be switching from Wrangel to
Banks Island. The WI subpopulation is a small proportion of Snow Geese wintering in
California, and geese from WI may be relocating to BI.
Population dynamics of Arctic nesting geese are primarily driven by annual productivity,
harvest management, and habitat conditions. Traditional management of Arctic nesting
geese focused on dynamics of mixed populations, primarily defined on the wintering
grounds. A more recent approach was developed that aimed at management of breeding
populations of geese. However, this latter approach may not adequately account for the
common harvest management of mixed populations on the wintering grounds. An
integrated metapopulation approach is needed to optimize management and maintain the
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integrity and sustainability of certain Arctic goose populations. This management
approach needs to consider the basic population dynamics of mixed populations, such as
population size, annual production, survival rates, sources of mortality, distribution
overlap, population exchange, and disease impacts. A metapopulation approach may
better explain significant population trends, interchange, and interaction among Wrangel
Island and Banks Island populations of LSG.
Our research involves 3 main components: (1) assessment of differential mortality between
sub-populations, (2) establish the pattern of movement (and individual mortality) using
radio-collared birds, and (3) develop stage-based projection models to assess the stability
of populations in this metapopulation in the presence of these and other factors,
particularly density-dependent feedbacks on (i) individual breeding success and survival,
and (ii) dispersal probability of individuals.
Apart from the purely demographic focus of this particular study, the application of
metapopulation models, and in particular, the use of CMR will allow more precise analysis
of the population genetic structure of this and other subdivided populations (see Nichols et
al. 1989). Traditionally, such metapopulations are modeled assuming an island model
(minimally) or a permuted stepping stone, in conjunction with one or more diffusion
approximations. Recently, Rockwell & Barrowclough (American Museum of Natural
History) have advocated using a different approach, using projection models to estimate
population structure, and from this estimated structure, deriving the appropriate
predictions. However, this approach requires good estimates of both the major transition
elements, but also of the covariances among them. CMR, particularly the multi-state
models, provides a very useful way to estimates these parameters.
iv. Estimation of breeding propensity and age of first breeding
This work is a collaboration with Roger Pradel in France; Bill Kendall of the US Fish and
Wildlife Service and Jim Nichols of the National Biological Service. Age at maturity,
defined as the age of onset of sexual activity, is a critical component of life history
variation. In general, variation in age of maturity is believed to reflect a trade-off between
increased fecundity or longevity due to delayed maturity, and increased risk of mortality
and longer generation time. The longer an organism waits to breed, the more likely it will
suffer mortality before it has actually done so. In addition, younger breeding individuals
will contribute proportionately faster to succeeding generations. On the other hand, the
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WILDLIFE ECOLOGY RESEARCH CHAIR
longer an organism waits to breed, the larger, or more experienced it might be, and thus,
relatively more fecund.
In most birds, physical maturity is reached before individuals breed. Charlesworth (1980)
has demonstrated that delayed maturation was predicted whenever reproductive efficiency
increased with age. Older birds may be more efficient foragers than younger birds, and
better able to acquire nutrients needed for egg production, or to provision their young.
Thus increased reproductive success with age generally favors delayed reproduction.
Through our collaborations several new analytical approaches make rigorous
consideration of these questions tractable for the first time in wild populations. We are
addressing several hypotheses concerning breeding propensity and recruitment in birds.
v. Philopatry, movement and LRS - spatial models and estimation
While previous work has demonstrated that philopatry to deteriorating habitats can be
maladaptive (e.g., Cooch et al. 1993), predicting the ultimate demographic and
evolutionary responses to selection favoring dispersal is potentially complex. In particular,
it is important to differentiate between genetically-based dispersal behaviors, and simple
transient behaviors adopted opportunistically by some individuals.
Using the multi-state or 'movement' models described previously, we can directly estimate
rates of transition between traditional and novel nesting and foraging areas, and relate
these transition probabilities to various factors of interest. This ability to derive precise
estimates of movement patterns will, when combined with measures of fitness, allow us to
derive estimates of spatial fitness, and, ultimately, reproductive value. In addition, once
spatially-explicit population models have been derived, standard matrix-methods for
assessing proportional changes in fitness to variation in one or more demographic
parameters provide a quantitative measure of the selection gradients on various life history
traits. This approach may be particularly relevant to several of our studies where we have
good information on components of fitness and the patterns of movement among
individuals (e.g., the harlequin duck study, and the snow goose metapopulation study - see
above).
vi. A User's Manual for Program SURGE
Program SURGE is one of the most widely used software applications available for the
analysis of CMR data. Unfortunately, the documentation supplied with the software is
rudimentary at best, and it is difficult for ecologists to learn to use the software correctly
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and efficiently without intensive and often expensive hands-on training by researchers
currently using SURGE.
Drawing on 3 years of graduate-level instruction in using SURGE, a comprehensive
hands-on textbook on the use of program SURGE has recently been completed. This 150+
page book, written in collaboration with Roger Pradel (CEFE-CNRS) and Nadav Nur
(Point Reyes Bird Observatory) is intended to provide a thorough, yet extremely userfriendly means for ecologists to learn to use SURGE in their research. Scheduled for
publication in 1996, the book should prove to be of considerable use to ecologists
working with individually marked individuals.
vii. Systematic changes in species richness
One of the questions stemming from the LPB study which has broach applicability is the
degree to which the trophic cascade in the goose-plant complex has collateral impacts on
the larger salt-marsh ecosystem. One of the ways we can address this question is to
examine systematic changes in the diversity of other species at LPB. Since 1970, daily
counts of the abundance of all bird species in the area have been recorded. Often, the total
number of species seen (or identified through other sampling procedures) is used as an
estimate of ‘species’ richness. Unfortunately, this estimate is biased, and the extent of the
underestimate is a function both of both the probabilities of encountering species and the
sampling effort. CMR methods can be used on species information to directly estimate the
total number of species present (Nichols & Conroy 1996). This is accomplished simply by
treating species as individuals, and deriving capture histories for each species by observing
presence/absence. This is completely analogous to the presence-absence data distribution
normally used to estimate demographic parameters using CMR. Data collected in this way
can be analyzed using CMR, resolving some of the sampling problem inherent in most
studies of species richness. Furthermore, they allow us to use the formal CMR statistical
framework to test specific hypotheses about changes in richness over time. In fact, the
range of uses of this approach can readily be extended to permit estimation of parameters
associated with community dynamic, such as rate of change in richness, local extinction
rates, and local colonization rates.
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WILDLIFE ECOLOGY RESEARCH CHAIR
B. Other Initiatives
One of the key requirements of a major research program is the development of an
intellectual framework for ideas and projects. To this end, we have attended a series of
symposia, seminars and workshops to disseminate our ideas outlined above. These
activities include:
i) The third annual Western Sandpiper workshop, held 7 November, 1995, in Victoria,
B.C. The workshop continued the work of co-ordinating research and discussing
ideas. Dr. Bruno Ens from The Netherlands was for the second time a participant, and
20 other researchers from four countries took part. Tony Williams also organized and
moderated a shorebird symposium at this meeting.
ii) Members of the Chair played a major role in hosting the Joint Conference of the
Pacific Seabird Group and the Colonial Waterbird Society, held in Victoria,
November 8-11, 1995.
iii) 8th North American Arctic Goose Conference and Workshop, 9 -14 January 1995.
iv) Drs. E. Syroechkovsky and V. Baranyuk, sponsored to visit the 8th North American
Arctic Goose Conference for further discussions on Wrangel Island Goose Research.
v) The Senior Chair represented NSERC for an assessment of ACWERN, a similar
cooperative venture to our Chair involving CWS, NSERC and Universities. It is
located in Atlantic Canada.
vi) Our outdoor aviaries attached to the Animal Care Facility at SFU now house Western
Sandpipers and Dunlin. These provide us with a powerful experimental model to
investigate a) migration physiology and b) foraging behaviour, prey selection etc. in
shorebirds.
vii) The Senior Chair became a Council Member of Bird Studies Canada, an NGO
established to coordinate bird studies across Canada and to coordinate research
initiatives among amateurs and professionals. He attended the inaugural meeting in
Long Point Ontario in Oct. 1995, and organised the second meeting in Delta, BC, in
April 1996. He is a member of the Program Planning Committee.
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C. Community Involvement
i)
ii)
iii)
iv)
v)
vi)
vii)
Attendance by the senior Chair, and presentations by Pippa Shepherd and Eric Reed,
at the Public Advisory Committee on Boundary Bay. This was to provide input to the
policy making for the new Boundary Bay Wildlife Management Area.
Presented talks to several naturalist groups throughout the years about the work of
the Chair.
gave interviews with CBC and BCTV.
served on the Biodiversity panel of FRBC.
established, with the Surrey and White Rock Naturalists, a monthly survey of the
water birds of the western Boundary Bay.
became an executive member of the Surrey and White Rock Naturalists.
Evan Cooch established the GooseRef Bibliography System, an indexed reference
database of papers devoted to geese available on the World Wide Web.
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WILDLIFE ECOLOGY RESEARCH CHAIR
D. Funding
Our total funding has increased considerably since the establishment of the Chair. Almost
all of this increase has been due to the acquisition of funds from outside sources (see
Figure 1.) Our forecast budget for 1996/97 is more than $1,000,000.
1. Research Funding Obtained (1995)
Applicant
Source of Funding
Amount
(per annum)
Fraser River Action Plan
(M. Sewell, P. Shepherd, R. Vadas)
$58,000.00
Graduate Research Fellowships
(B. Carter, S. Cullen, G. Robertson, B. Sandercock, D. Schamel)
$23,050.00
Clowater, J.
NSERC Graduate Scholarship
$15,000.00
Cooch, E.
Polar Continental Shelf Project
"Banks Island Snow Goose Project"
$6,000.00
Cooch, E.
Arctic Goose Joint Venture
"Banks Island Snow Goose Project"
$21,000.00
(first of 3 yrs)
Cooch, E.
/M. Samuel
/J. Takekawa
Ducks Unlimited
“Metapopulation dynamics of the
Banks Island-Wrangel Island Lesser Snow Goose Complex
$25,000.00
(2nd of 4 yrs)
Cooch, E.
Environment Canada
“Fraser-Skagit Snow Goose”
$3,800.00
Cooke, F.
Forest Industry Donations
“Demography of Marbled Murrelets”
$44,500.00
Cooke, F.
NSERC CRD Matching Funds
“Demography of Marbled Murrelets”
$89,000.00
Cooke, F.
BC Ministry of Forests
"Population Status Of Marbled Murrelets"
$6,500.00
Cooke, F.
Science Council of B.C. - FRBC
“Nesting Habitats, Abundance and Ecology of Marbled
Murrelets on the B.C. Coast”
$165,000.00
Cooke, F.
Science Council of B.C. - FRBC
“Effects of Forest Management on Biodiversity of Riparian
Ecosystems in Interior Forests of the Cariboo-Chilcotin”
$40,000.00
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THIRD ANNUAL REPORT - MARCH 1996
Applicant
Source of Funding
Amount
(per annum)
Cooke, F.
IWP - Ducks Unlimited
“Waterfowl in Becher’s Prairie:
Waterbirds, Grazing and Disturbance”
$20,000.00
Cooke, F.
IWWR
“Winter Philopatry of Harlequin Ducks”
$12,500.00
Cooke, F.
J.L. Baillie Memorial Fund
$2,500.00
Cooke, F.
NSERC Individual Operating Grant
$100,000.00
(fifth of 5 yr)
Lank, D.B.
Challenge Grant
$1,100.00
Lank, D.B./
F. Cooke
Skills Now Student Summer Works Program
$5,253.00
Lank, D.B.
NSERC Individual Operating Grant
"Genetic Polymorphism in Male Mating Strategy."
$17,500.00
(first of 4 yrs)
O'Hara, P.
Northern Studies Training Program
"Niche Overlap in Sympatric Sandpipers."
$3,100.00
Sandercock, B.
Northern Studies Training Program
"The Adaptive Significance of Clutch Size in Shorebirds"
$5,000.00
Sandercock, B.
President’s Ph.D. Research Stipend
Schamel, D.
H.R. MacCarthy Bursary
$2,324.00
Shepherd, P.
NSERC Graduate Scholarship
$17,400.00
Warnock, N.
United States Fish and Wildlife Service
“Nongame Migratory Bird Program, San Francisco Bay
Program, Coastal Marine Bird Program”
$34,000.00
Warnock, N.
National Fish and Wildlife Foundation
“Conservation of Western Sandpipers along the Pacific Flyway”
$61,200.00
Williams, T.
NSERC Individual Operating Grant
"Reproductive Investment in Wild Birds"
$34,000.00
(second of 4 yrs)
Williams, T.
NSERC Equipment Grant
"Reproduction and Migration in Birds:
Individual Variation in Physiological Traits."
$30,226.00
$4,850.00
TOTAL:
3RDAR8DOC (23-Jan-98)
$847,803.00
Page 29
WILDLIFE ECOLOGY RESEARCH CHAIR
2. Research Funding Obtained * or Applied For (1996)
Applicant
Source of Funding
Amount
(per annum)
Graduate Research Fellowships*
(C. Betancourt-Lougheed, S. Cullen, C. Guglielmo,
A. MacCharles, P. O’Hara, E. Reed, G. Robertson,
B. Sandercock, D. Schamel)
$40,650.00
Bertram, D.
Pacific Seabird Group*
$4,700.00
Cooch, E.
Polar Continental Shelf Project*
"Banks Island Snow Goose Project"
$20,000.00
Cooch, E.
Arctic Goose Joint Venture*
"Banks Island Snow Goose Project"
$26,000.00
(3rd of 4 yrs)
Cooch, E.
/M. Samuel
/J. Takekawa
Ducks Unlimited*
“Metapopulation dynamics of the
Banks Island-Wrangel Island Lesser Snow Goose Complex”
$25,000.00
(3rd of 4 yrs)
Cooke, F.
Forest Industry Donations
“Demography of Marbled Murrelets”
$44,500.00
Cooke, F.
NSERC CRD Matching Funds
“Demography of Marbled Murrelets”
$89,000.00
Cooke, F.
Science Council of B.C. - FRBC*
“Nesting Habitats, Abundance and Ecology of Marbled
Murrelets on the B.C. Coast”
$190,000.00
(first of 5 yrs)
Cooke, F.
IWP - Ducks Unlimited*
“Waterfowl in Becher’s Prairie:
Waterbirds, Grazing and Disturbance”
$26,000.00
Cooke, F.
Science Council of B.C. - FRBC*
“Effects of Forest Management on Biodiversity of Riparian
Ecosystems in Interior Forests of the Cariboo-Chilcotin”
$212,375.00
(first of 5 yrs)
Cooke, F.
IWWR
“Winter Philopatry of Harlequin Ducks”
$19,300.00
Cooke, F.
J.L. Baillie Memorial Fund
$2,500.00
Cooke, F./
E. Cooch
Vancouver International Airport
$98,000.00
Cooke, F.
NSERC Individual Operating Grant*
$79,000.00
(first of 4 yrs)
Page 30
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THIRD ANNUAL REPORT - MARCH 1996
Applicant
Source of Funding
Amount
(per annum)
Cooke, F./
D. Lank
NSERC Equipment Grant*
Vehicle-mounted radio telemetry system.
$15,160.00
Knechtel, H.
Anne Vallee Scholarship
$1,500.00
Lank, D.B./
F. Cooke
Skills Now Summer Works Program
$7,361.00
Lank, D.B.
NSERC Individual Operating Grant*
"Genetic Polymorphism in Male Mating Strategy."
$17,500.00
(2nd of 4 yrs)
Sandercock, B.
Northern Studies Training Program*
"The Adaptive Significance of Clutch Size in Shorebirds"
$4,560.00
Schamel, D.
United States National Parks*
$5,000.00
Schamel, D.
UAF Natural Resources Fund*
$21,000.00
Schamel, D.
Steel Memorial Scholarship
$15,000.00
Shepherd, P.
Fraser River Action Plan*
$10,000.00
Shepherd, P.
NSERC Graduate Scholarship*
$17,400.00
Shepherd, P.
American Ornithologists Union
$1,500.00
Shepherd, P.
Society of Canadian Ornithologists
$1,000.00
Shepherd, P.
John K. Cooper Foundation
$1,000.00
Warnock, N.
United States Fish and Wildlife Service*
“Nongame Migratory Bird Program, San Francisco Bay
Program, Coastal Marine Bird Program”
$28,800.00
Williams, T.D.
NSERC Individual Operating Grant*
"Reproductive Investment in Wild Birds"
$34,000.00
(3rd of 4 yrs)
Williams, T.D./
LBendell-Young
Environmental Science & Technology Alliance of Canada
“Determining the ecological viability of constructed wetlands.”
$48,000.00
(first of 2 yrs)
TOTAL:
3RDAR8.DOC (23-Jan-98)
$1,105,806.00
Page 31
WILDLIFE ECOLOGY RESEARCH CHAIR
E. Publications
We continue to publish actively in 1995, and increasingly our publications reflect the
activities which have taken place since we arrived in BC. However since most of our focus
is on long term studies, publications relevant to these long term goals will occur with
increasing frequency in later reports.
1. Refereed Publications - 1995/96 or in press.
Beauchamp, W.D., R.R. Koford, T.D. Nudds, R.G. Clark & D.H. Johnson. (in press)
Long-term declines in nest success of prairie ducks. Journal of Wildlife Management.
Beauchamp, W.D., T.D. Nudds & R.G. Clark. (in press) Declines in nest success of ducks
with and without predator management. Journal of Wildlife Management.
Bertram, D.F. (1995) The roles of introduced rats and commercial fishing in decline of
Ancient Murrelets on Langara Island, B.C. Conservation Biology 9: 865-872.
Bertram, D.F. and D.W. Nagorsen. (1995) Introduced rats on the Queen Charlotte
Islands: implications for seabird conservation. Canadian Field-Naturalist 109: 6-10.
Bertram, D.F. (1996) Size-dependent predation risk in larval fishes: mechanistic inferences
and levels of analysis. Fishery Bulletin 94: 371-373.
Bertram, D.F., C.F.J. Welham & R.C. Ydenberg. (in press) Flexible effort in breeding
seabirds: Adjustment of provisioning according to nestling age and mass. Canadian
Journal of Zoology.
Bertram, D.F., T. J. Miller & W.C. Leggett. (in press) Individual variation in growth and
development during the early life stages of winter flounder. Fishery Bulletin.
Brault, S., S. Boyd, F. Cooke & J. Takekawa. (1995) Population models as tools for
research cooperation and management: the Wrangel Island snow geese. Trans. 59th
North American Wildlife and Natural Resources Conference 79-90.
Butler, R.W., F.S. Delgado, H. de la Cueva, V. Pulido & B.K. Sandercock. (in press)
Migration routes of the Western Sandpiper. Wilson Bulletin.
Cooch, E.G., D.B. Lank & F. Cooke. (in press) Intraseasonal variation in the development
of sexual size dimorphism in a precocial bird. Journal of Animal Ecology.
Cooch, E.G., D. B. Lank, R.J. Robertson & F. Cooke. (in press) Offspring sex ratio
change in a precocial bird: growth, survival and parental experience in a changing
environment. Journal of Animal Ecology.
Ganter, B. (in press) Site tenacity and movements of staging Barnacle Geese. Ardea.
Ganter, B. & F. Cooke (in press) Pre-incubation feeding activities and energy budgets of
snow geese: can food on the breeding grounds influence fecundity? Oecologia.
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Ganter, B., F. Cooke, P. Mineau. (in press) Long-term vegetation changes in a snow
goose nesting habitat. Canadian Journal of Zoology.
Guglielmo, C.G., W.H. Karasov, & W.J. Jakubus. (1995) Nutritional quality of winter
browse for ruffed grouse. Journal of Wildlife Management 59: 427-436.
Guglielmo, C.G., W.H. Karasov, & W.J. Jakubus. (in press) Nutritional costs of a plant
secondary metabolite explain selective foraging in an avian herbivore, the ruffed
grouse. Ecology.
Hitchcock, C. & Gratto-Trevor, C. (in press) Diagnosing a shorebird local population
decline with a stage-structure population model. Ecology.
Hugie, D.M. & D.B. Lank. (in press) The resident’s dilemma: a female-choice model for
the evolution of alternative male reproductive strategies in lekking male ruffs
(Philomachus pugnax). Behavioural Ecology.
Iverson, G.C., S. Warnock, R. Butler, M.A. Bishop & N. Warnock. (1996) Spring
migration of western sandpipers along the Pacific coast of North America: a telemetry
study. Condor 98: 10-21.
Jakubus, W.J., C.G. Guglielmo, W. H. Karasov & C. Vispo. (1995) Sodium balance in
ruffed grouse. Canadian Journal of Zoology. 73:
Lank, D.B., C.M. Smith, O. Hanotte, T.A. Burke & F. Cooke. (1995) Genetic
Polymorphism for alternative mating behaviour in lekking male ruff, Philomachus
pugnax. Nature 378: 59-62.
Mulder, R.S., T.D. Williams, & F. Cooke. (1995) Dominance, brood size and foraging
behaviour during brood-rearing in the Lesser Snow Goose: an experimental study.
Condor.
Pradel, R., E.G. Cooch & F. Cooke. (1995) Transient animals in a resident population:
local emigration or heterogeneity. Journal of Applied Statistics.
Reed, J.M., N. Warnock & L.W. Oring (Eds.) (in press) Conservation and management of
shorebirds in the western Great Basin of North America. International Wader
Studies.
Reed, J.M., N. Warnock & L.W. Oring. Censusing shorebirds in the western Great Basin
of North America. In Conservation and management of shorebirds in the western
Great Basin of North America. (J.M. Reed, N. Warnock & L.W. Oring, Eds.).
International Wader Studies (in press).
Robertson, G.J. (in press) Factors affecting nest site selection and nesting success in the
Eider Somateria mollissima. Ibis
Robertson, G.J. (1995) Annual variation in common eider egg size: effects of
temperature, clutch size, laying date and laying sequence. Canadian Journal of
Zoology. 73: 1579-1587
Robertson, G.J., M.A. Owen & R.F. Rockwell. (1995) Responses of snow geese to
artificially induced hatching asynchrony and increased clutch size. Auk 112-790-794.
3RDAR8.DOC (23-Jan-98)
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WILDLIFE ECOLOGY RESEARCH CHAIR
Robertson, G.J., F. Cooke, R.I. Goudie & W.S. Boyd. (in press) Within year fidelity of
Harlequin Ducks to a moulting and winter area. Proc. Seaduck Sym., PSG/CSW
conference.
Samuel, M.D., D.R. Goldberg, D.J. Shadduck, J.I. Price & E.G. Cooch. (in press) Avian
Cholera Bacterium Isolated from Healthy Lesser Snow Goose. Journal of Wildlife
Diseases.
Sandercock, B.K. (in press) Removal of Eggshells at hatch by Western and Semipalmated
Sandpipers. Condor.
Sandercock, B.K. & C.L. Gratto-Trevor. (in press) Patterns of local survival in
Semipalmated Sandpipers (Calidris pusilla) breeding at La Perouse Bay, Manitoba.
Ibis.
Sandercock, B.K. (in press) The effect of manipulated brood size on parental defence in a
precocial bird, the willow ptarmigan. Journal of Avian Biology.
Sandercock, B.K. & Pederson. (in press) The effect of renesting ability and nesting
attempt on egg size variation in willow ptarmigan. Canadian Journal of Zoology.
Stock, M., F. Hofeditz, K. Mock, B. Pohl. (1995) Impact of aircraft and tourism on
behaviour and salt marsh use of Brant geese in the Wadden Sea (in German). Corax
16, 63-.
Szekely, T., & T.D. Williams. (1995) Costs and benefits of brood desertion in female
Kentish Plover Charadrius alexandrinus. Behavioural Ecology and Sociobiology.
37: 155-161.
Syroechkovsky, E.V., F. Cooke & W.J.L. Sladen. (1995) Population structure of the
lesser snow geese of Wrangel Island, Russia. Ecoscience.
Viallefont, A., E.G. Cooch & F. Cooke. (1995) Estimation of trade-offs with capturerecapture models: a case study on the lesser snow goose. Journal of Applied
Statistics.
Viallefont A., F. Cooke & J.D Lebreton. (1995) Age-specific costs of first time breeding.
Auk.
Warnock, N. & S. Scharzbach. (1995) Incidental kill of Dunlin and Killdeer by strychnine.
Journal of Wildlife Diseases 31: 566-569.
Warnock, N., G.W. Page & L. E. Stenzel. (1995) Non-migratory movements of Dunlin on
their California wintering grounds. Wilson Bulletin 107: 131-139.
Warnock, N. & R.E. Gill, Jr. (1996) Dunlin (calidris alpina). In The Birds of North
America No. 203 (A. Poole & F. Gill, Eds.). Philadelphia: The academy of Natural
Sciences; Washington, D.C.: The American Ornithologists’ Union.
Warnock, N. & L.W. Oring. (1996) Nocturnal nest attendance of Killdeers: More than
meets the eye. Auk 113: 502-504.
Warnock, N., J.M. Reed & L.W. Oring. (in press) Synthesis: shorebirds in the arid
western Great Basin of North America. In Conservation and management of
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THIRD ANNUAL REPORT - MARCH 1996
shorebirds in the western Great Basin of North America. (J.M. Reed, N. Warnock &
L.W. Oring, Eds.) International Wader Studies.
Warnock, N (in press) Local and regional differences in habitat utilization by Dunlins
(Calidris alpina) as revealed by radiotelemetry: conservation implications. In
Shorebird Ecology and Conservation in the Western Hemisphere (P. Hicklin et al.
editors). International Wader Studies.
Williams, T.D. (1996) Intra- and inter-individual variation in reproductive effort in
captive-breeding zebra finches, Taeniopygia guttata. Canadian Journal of Zoology.
74: 85-91.
Williams, T.D. (in press) Variation in reproductive effort in female zebra finches
(Taeniopygia guttata) in relation to nutrient-specific dietary supplements during egglaying. Physiological Zoology.
Williams, T.D. & E.G. Cooch. (1996) Egg size, temperature and laying sequence: why do
snow geese lay big eggs when its cold? Functional Ecology 10: 112-118.
Williams, T.D., C. Jeffs, K.A. Murray & S. Choudhury. (1996) Intraclutch egg-size
variation in the Barnacle Goose Branta leucopsis: An egg removal experiment. Ibis.
Williams, T.D., M.J.J.E. Loonen & F. Cooke. (1995) Fitness consequences of parental
behaviour in relation to offspring number in a precocial species: the lesser snow
goose. Auk 111, 563-572.
2. Other publications, book reviews, etc.
Cooke F. (1994) Are Chickens really birds? Condor 96: 246-247. (this ms. was omitted
from the previous report)
Cooke F. 1995. The Snow Geese of the Fraser Delta. Marshnotes Fall 95: 4-6.
Cooke F. 1996. Waterbird Surveys of Boundary Bay, BC. Part 1. Discovery 25: 30-34.
Cooke F. (in press) Waterbird Surveys of Boundary Bay, BC. Part 2. Discovery.
Toochin, R. (1995) A Great Crested Flycatcher at Triangle Island, British Columbia - first
provincial record. Birders Journal. Volume 4, Number 3, pp 106-109
3. Publications Submitted
Bertram, D.F. & R.R. Strathmann. (submitted to Ecology) Effects of maternal and larval
nutrition on developmental plasticity and growth rates of planktotrophic larvae of
echinoids.
Black, J.M., E.G. Cooch, M.J.J. Loonen & M. Owen. (submitted to Journal of Animal
Ecology) Geographical variation in body size in a barnacle goose metapopulation: the
effect of temporal and spatial scale.
3RDAR8.DOC (23-Jan-98)
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WILDLIFE ECOLOGY RESEARCH CHAIR
Cooke, F., G.J. Robertson, R.I. Goudie & W.S. Boyd. (submitted to Condor) Moult and
the basic plumage of male Harlequin Ducks (Histrionicus histrionicus).
Gowans, B., G.J. Robertson & F. Cooke. (submitted to Wildfowl) Behavior and
chronology of pair formation in Harlequin Ducks.
Kopachena, J. & B. Falls. (submitted to Evolution) Differential allocation of reproductive
effort as a stabilizing mechanism in the maintenance of polymorphism in Whitethroated Sparrows.
Sandercock, B.K. (submitted to J. Field Ornith.) Methods for determining the chronology
of nesting events in Western and Semipalmated Sandpipers.
Sandercock, B.K. (Submitted to Oecologia) Incubation capacity and clutch size
determination in two calidrine sandpipers: a test of the four-egg threshold.
Shepherd, P. (submitted to Conservation Biology) Effects of a commercial bloodworm
harvest on Semipalmated Sandpipers (Calidris pusilla) and their prey in the Bay of
Fundy Hemispheric Shorebird Reserve.
4. Books Published in 1995
Cooch, E.G., R. Pradel & N. Nur (1995) A practical Guide to Mark-recapture analysis
using SURGE. Publication as a textbook by CNRS (France). approx. 125 pp.
Cooke, F., R.F. Rockwell & D.B. Lank (1995) The Snow Geese of La Perouse Bay:
Natural Selection in the Wild. Oxford: Oxford University Press. 289 pp.
Williams, Tony (1995) The Penguins: Spheniscidae, Bird Families of the world Series.
Oxford. Oxford University Press. 295 pp. (Japanese translation to be published 1996.)
3. Papers Presented at Conferences in 1995
Brault, S. & E.G. Cooch: Demographic effects of density changes in Lesser Snow Geese.
8th North American Arctic Goose Conference and Workshop. Albuquerque, New
Mexico.
Buenrostro, A. & H. de la Cueva. Age and sex distribution of Western Sandpiper,
Calidrus maurii at Estera Punta Banda. Baja California, Mexico. Cooper
Ornithological Society CIPAMEX joint meeting. La Pas BIS, Mexico.
Cooch, E.G.: Overgrazing in Arctic: An example in the Lesser Snow Goose. New Mexico
State University, Department of Fish and Wildlife.
Cooch, E.G., D.B. Lank & F. Cooke: Old genes in new bodies: do shrinking snow geese
reflect evolutionary change? Society for the Study of Evolution. McGill University,
July, 1995.
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Cooch, E.G., D.B. Lank, R.F. Rockwell & F. Cooke Body size and breeding propensity
in Snow Geese: do runts wimp out? 8th North American Arctic Goose Conference
and Workshop. Albuquerque, New Mexico.
Cooch, E.G., D.B. Lank, R.J. Robertson & F. Cooke: The cost of being larger: natural
selection on sexually dimorphic growth in Lesser Snow Geese. 8th North American
Arctic Goose Conference and Workshop. Albuquerque, New Mexico.
de la Cueva, H. Aerodynamic of migration of the Western Sandpiper, Calidris maurii.
Cooper Ornithological Society CIPAMEX joint meeting. La Pas BIS, Mexico.
Gowans, B. Behaviour and chronology of pair formation in Harlequin Ducks. 3rd
Harlequin Duck Working Group meeting.
Guglielmo, C. & T.D. Williams. Variability in digestive systems of migrating Western
Sandpipers: allometry and effects of sex, age and time of year. Pacific Seabird
Group/Colonial Waterbird Meeting. Victoria, BC.
Lank, D.B., D.M. Smith. Genetic Polymorphism for Alternative Male Mating Strategies
in the Ruff (Aves:Scolopaciae): Support From Pedigree Data. Society for the Study of
Evolution. McGill University.
Robertson, G.J. Sources of egg size variation in eiders. Pacific Ecology Conference,
Oregon Institute of Marine Biology.
Robertson, G.J. Within season moulting and wintering site philopatriy of Harlequin
ducks. IWWR student/staff symposium. Oak Hammock Marsh, Manitoba. PSG/CWS
joint conference. Victoria.
Robertson, G.J. Moult chronology and the timing of pairing in Harlequin ducks. Pacific
Ecology Conference, Friday Harbour Labs.
Robertson, G.J. Moulting and wintering philopatry in Harlequin Ducks at White Rock,
British Columbia. 3rd Harlequin Duck Working Group meeting.
Sandercock, B.K. Survival estimates for semi-palmated sandpipers. Pacific Ecology
Conference. Coos Bay, Oregon.
Sandercock, B.K. Breeding biology of western and semi-palmated sandpipers. Western
Sandpiper Workshop. CWS/PSG Joint Meeting, Victoria.
Sandercock. B.K. Incubation ability in calidrine shorebirds: a test of the four-egg
threshold. Pacific Ecology Conference. Friday Harbour, Washington.
Sandoval Y. & H. de la Cueva. Nutritional status of migratory Western Sandpipers,
Calidris maurii. Cooper Ornithological Society CIPAMEX joint meeting. La Pas
BIS, Mexico.
Savard, J.L., A. Reed, A. Morrier, L. Lesage, E. Reed & Y. Aubry. Time budget of surf
scoter broods. Pacific Seabird Group/Colonial Waterbird Meeting. Victoria, BC.
Shepherd, P. The winter ecology of Dunlin Calidris alpina in the Fraser River Delta.
Fraser River Action Plan Research Workshop and Ecological Monitoring and
Assessment Network Workshop. Vancouver.
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WILDLIFE ECOLOGY RESEARCH CHAIR
Warnock, N. Nocturnal incubation in Killdeer. Cooper Meeting. San Diego, California.
Warnock, N. Migration strategies of radiomarked Western Sandpipers along the Pacific
Flyway. Pacific Seabird Group/Colonial Waterbird Meeting. Victoria, BC.
Warnock, N. Spring migration ecology of Western Sandpipers as revealed by
radiotelemetry, 1995. International Waterfowl Ecology Symposium and Wader Study
Group. Aveiro, Portugal.
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F. Personnel
1. Board of Directors
Name
Dr. Larry Dill
Dr. Robert Elner
Dr. Chris Pharo (CWS alt.)
Dr. Nico Verbeek (SFU alt.)
Dr. Brian Wilson
Dr. Ron Ydenberg
Position
Professor
Head of Ecosystem Research
Head PWRC
Professor
Regional Director, ECB, DOE
Professor
Affiliation
SFU
CWS
CWS
SFU
CWS
SFU
2. Research Team
i. Faculty and Research Associates
Name
Dr. Fred Cooke
Dr. Tony Williams
Dr. Evan Cooch
Dr. Dov Lank
Dr. Doug Bertram
Position
Senior Chair
Associate Chair
Assistant Professor
Assistant Professor
Research Associate
ii. Research Group
Postdoctoral Fellow
Dr. Mary Sewell
Dr. Robert Vadas
Dr. Nils Warnock
Ph.D. Student
Chris Guglielmo
Greg Robertson
Brett Sandercock
Pippa Shepherd
Douglas Schamel
M.Sc. Student
Brian Carter
Julian Christians
James Clowater
Sean Cullen
Oliver Egeler
Hugh Knechtel
Andrea MacCharles
Laura Jones
Irene Manley*
Patrick O'Hara
Eric Reed
Debbie van de Wetering
Others
Barbara Sherman
Wendy Beauchamp
Connie Smith
Barbara Pohl
Uli Steiner
* supervised by Dr. Alton Harestad.
3RDAR8.DOC (23-Jan-98)
Page 39
WILDLIFE ECOLOGY RESEARCH CHAIR
3. Workshops/Meetings
i. Attended by Chair Representatives
Title
Date
Location
Participants
8th North American Arctic Goose
Conference
01/95
Albuquerque, NM
F. Cooke
E. Cooch
B. Ganter
Pacific Seabird Group
01/95
San Diego, CA
I. Jones
H. Knechtel
Pacific Ecology Conference
02/95
Coos Bay, OR
B. Sandercock
G. Robertson
Pacific Ecology Conference
03/95
Bamfield, BC
E. Reed
Western Society of Comparative
Endocrinologists
03/95
Seattle, WA
T. Williams
Society for the Study of Evolution
07/95
Montreal, Que.
D. Lank
IWWR Symposium
08/95
Oak Hammock Marsh,
MB
G. Robertson
BC Seabird Group Annual Meeting
09/95
SFU
W. Beauchamp
D. Bertram
F. Cooke
10th IWRB Waterfowl Ecol. Conf.
09/95
Aveiro, Portugal
N. Warnock
BC Field Ornithologists Annual
Meeting
09/95
Saanich BC
F. Cooke
Bird Studies Canada Meeting
10/95
Long Point, Ont.
F. Cooke
PSG/CWS Joint Meeting
11/95
Victoria, BC
W. Beauchamp
D. Bertram
J. Clowater
F. Cooke
C. Guglielmo
G. Robertson
D. Lank
B. Sandercock
D. Schamel
P. Shepherd
H. Knechtel
I. Manley
N. Warnock
SFU/UBC Ecology Retreat
10/95
Squamish, BC
B. Sandercock
G. Robertson
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THIRD ANNUAL REPORT - MARCH 1996
ii. To be attended by Chair Representatives
Workshop/Meeting Title
Date
Location
Participants
Pacific Ecology Conference
02/96
Friday Harbour, WA
B. Sandercock
G Robertson
Fraser River Action Plan Research Workshop
02/96
Vancouver, BC
P. Shepherd
D. Lank
Harlequin Duck Working Group
03/96
White Rock, BC
G. Robertson
B. Gowans
F. Cooke
BC. Wildlife Meetings
03/96
108 Mile House
F. Cooke
Ecological Monitoring and Assessment
Network Workshop
03/96
Vancouver, BC
P. Shepherd
Cooper Ornithological Society Meeting
04/96
San Diego, CA
D. Lank
N. Warnock
International Avian Endocrinology Meeting
04/96
Lake Louise, AB
T. Williams
Brant Festival
04/96
Parksville, BC
F. Cooke
B. Sandercock
E. Reed
Bird Studies Canada
04/96
Ladner, BC
F. Cooke
American Ornithologist's Union
07/96
Boise, Idaho
F. Cooke
G. Robertson
International Behavioral Ecology Congress
09/96
Canberra, Australia
F. Cooke
D. Lank
Evolution and Behaviour of Birds. Festschrift
Meeting in Honour of Prof. J. Kikkawa
09/96
Brisbane, Australia
F. Cooke
D. Lank
3RDAR8.DOC (23-Jan-98)
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WILDLIFE ECOLOGY RESEARCH CHAIR
III. APPENDIX I. MISSION STATEMENT (JANUARY 1995 - F. COOKE)
The mandate of the Chair is to foster high quality, graduate level research in the field of
wildlife ecology, with specialisations in behavioural, population and physiological ecology.
The central concept is to provide an interface between the mission-oriented research and
management policies of the Canadian Wildlife Service and the more curiosity-driven
research of the university community. By encouraging this interface, the government
agencies would gain access to a broader base of scientific information for government
policy and decision making and university researchers would benefit from a broader
practical application for the more theoretical ideas which their disciplines generate.
With these perspectives in mind, I have formulated the mission of the Chair
i) by defining how the mandate of the Chair fits into the broader objectives of the CWS,
Pacific and Yukon Region and
ii) by considering the theoretical framework of wildlife ecology and what questions the
Chair is in position to focus on most effectively.
A. Conservation Goals, The CWS Perspective
To conserve and protect wildlife and habitat, managers require scientific advice based on
an understanding of population dynamics and ecosystem functioning. If we can discover
the processes involved in population dynamics of particular species in their environment
then we can begin to understand the dynamics of those ecosystems in which they live.
Under the Migratory Birds Convention and Canada Wildlife Acts, the CWS is
required to protect and maintain migratory bird populations through protection of
habitats, policy decisions, public forums, influence and research. Such an allembracing mandate requires focused objectives and cooperation with other research and
management agencies. From the perspective of CWS, Pacific and Yukon Region, 3 major
ecotypes are defined.
i) Marine and Coastal,
ii) Wetlands and Freshwater
iii) Forest and Grasslands.
Detailed definitions and understanding of priority ecosystems (e.g. Strait of Georgia) is the
mandate of the Wildlife Ecology Division, with whom we will coordinate for advice on the
relationship between our target species and the ecosystems in which they occur.
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1. Habitats
When the Chair was established it was agreed that the research emphasis would be on
birds of the Marine and Coastal environment. Specific ecosystems are as follows:
i) Coastal Salt and Freshwater Marshes
ii) Mudflats, Sandy and Rocky Shorelines
iii) Agricultural Lands in the Fraser Lowlands
iv) Inshore waters of the Fraser Delta and the Strait of Georgia
v) Open Sea
vi) Off-shore Islands
2. Bird Species
Each of these environments provide food and shelter for many of Canada's waterfowl,
shorebirds and seabirds, the taxa which will be the main research focus of the Chair. Our
aim is to focus on a few of the important target bird species for detailed research with the
objective of understanding not only the demographic processes but through them, the
dynamics of the ecosystems of which the birds are a part.
The choice of species is based on our assessment of their importance to the ecosystems
where they occur, their likely importance as species of conservation concern and their
suitability as research organisms. Additionally we felt it important to cover year round
residents, winter visitors, passage migrants, summer residents, and species which breed in
one area and winter in another.
The key to conservation of birds is understanding the demographic processes which
influence population numbers. Ultimately we are interested in the factors that influence
reproduction and survival, and for this reason it is important to have the ability to examine
these factors throughout the life cycle of the birds. We have therefore chosen species
amenable to direct or indirect study throughout the annual cycle.
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The species of birds which we plan to concentrate on, their seasonal status, and the
environments in which they primarily occur in BC are listed below:
Species
Seasonal Status
Habitat
Snow Goose
winter
Trumpeter Swan
winter *
American Widgeon
winter *
Harlequin Duck
resident
(different habitats)
resident
(different habitats)
winter and transient
salt-marshes
agricultural fields
salt-marshes
agricultural fields
mud flats
agricultural fields
rocky shores, inshore waters,
mountain streams
inland ponds and open sea
Barrow's Goldeneye
Brant
Western Sandpiper
Dunlin
Grebes sp.
Cassin's Auklet
Rhinoceros Auklet
Tufted Puffin
Pigeon Guillemot
Marbled Murrelet
transient
winter
resident
(different habitats)
summer
resident
(different habitats)
resident
(different habitats)
resident
resident
(different habitats)
* also breeds in interior parts of province
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mudflats
inshore waters
mudflats
mudflats
inshore waters and
inland lakes and ponds
open sea
off-shore islands
open sea, inshore waters,
off-shore islands
open sea
off-shore islands
inshore waters, rocky shores,
off-shore islands
inshore waters,
old growth forest
THIRD ANNUAL REPORT - MARCH 1996
3. Integrated Population Approach
We take as a useful model the approach to bird conservation pioneered in United
Kingdom by the British Trust for Ornithology (BTO ), referred to as Integrated Population
Monitoring (IPM). It identifies the following stages.
Stage 1. Is there a mechanism in place to detect conservation problems associated with
bird populations? This would usually be associated with disturbing changes in
population size due to anthropogenic effects on the environment of the birds,
Stage 2. If disturbing changes are recognized, can one identify the stages in the life-cycle
that are affected? If survival is involved, does it affect all age groups equally? If
reproduction is involved, at what stage - egg production, fledgling success or
recruitment? At what level does the problem lie?
Stage 3. If the stage of the life cycle that is affected has been identified, can one identify
the causal components that influence it? For example, if over-winter mortality is high,
is this due to changes in food availability, predator pressure (including hunting
pressure), contaminant levels or other factors?
Stage 4. Can remedial steps be taken? What management advice can we give?
IPM provides us with a logical framework for bird conservation but still presents a
formidable task. Fortunately there is already considerable background information
available on most of the species we have chosen through the work of the CWS. For
example, the production of the book "Birds of British Columbia" represents an important
initiative by CWS as do the various programs, such as Christmas counts, breeding bird
surveys, nest record schemes that involve the public, particularly bird watchers, in the
monitoring function.
The role of the Chair in this program relates more to stages 2 and 3, the demographic
aspects of the IPM and in this we hope to work closely with scientists of the Wildlife
Ecology Division, using a team approach. The specifics of this approach will be pointed
out under the individual research projects.
The scientific approaches to IPM fall under three headings:
i. Population Ecology,
ii. Behavioural Ecology and
iii. Physiological Ecology
although in reality there are no such sharp divisions between these groups.
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i. Population Ecology
Our general aim will be to expand existing projects or set up new field research projects
that monitor variability in reproductive and survival performance of the species of
concern. To investigate variation in reproduction will entail several seasons of field work
on reasonably large samples of nesting birds. Where possible, we intend to embark on a
program of individually marking birds, and to collect blood, tissue and feather samples
from the birds to assess genetic variability and contaminant levels. The aim of the research
is to identify the environmental factors responsible for variation in reproductive
performance and to detect long term demographic trends. To examine variation in survival
rates, we will use individually marked birds and Capture-Mark-Recapture (CMR) models
to assess variability of local and age-specific survival, immigration and emigration. Where
possible we will build on existing data-bases of banded birds.
These data will be used for population modelling to help us understand both the
demographic processes and the sensitivity of populations to environmental perturbations.
The systems ecologist to be hired by CWS will be essential for this aspect of the research.
ii. Behavioural Ecology
Behavioural ecology seeks to understand the natural behaviour of animals including
foraging, distribution, daily and annual movements, reproductive behaviour and life
history. The central tenet of the discipline is that all these behaviours evolved as a result of
identifiable and measurable factors acting on the lifetime reproductive success of
individuals. Behavioural ecology developed from the cross-fertilization between long-term
field studies of animals in their natural habitats, theoretical insights into the evolution of
behaviour, and the successful conduct of quantitative, experimental work in both
laboratory and field. Using these approaches, much progress has been made over the past
few decades in understanding animal behaviour.
We can use behavioural ecological approaches to help understand why animals use their
environment as they do, aiming to more than simply describe. For example, we may be
able to describe the dispersion of a species over available feeding sites, but to be able to
predict the outcome of a change resulting from , for example, development, we need to
investigate the basis of the distribution. Behavioural ecology has developed a number of
tools, including dynamic programming and games theory to help investigate this basis, and
identify the ecological constraints under which birds find themselves.
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iii. Physiological Ecology
Ecological constraints are more readily understood if we have some knowledge of the
underlying physiological processes, essential for maintenance, reproduction and survival.
The appointment of a physiological ecologist to the Associate Chair position will fill a gap
that is evident in most other demographic studies. By looking at intra-specific variability
of physiological processes we hope to discover the responses of birds to environmental
perturbations such as might be imposed by environmental contaminants for example. In
this way we hope to interface with the Ecosystem Health Section, Wildlife Ecology
Division.
Physiological ecology is concerned with how animals function, whereas population and
behavioural ecology seek to explain why animals function the way they do. In the past,
these two questions have often been considered to be independent, and physiological
ecologists have been accused of showing only that 'animals live where they can'. Although
it is now becoming increasingly apparent that the ecology and evolution of any species can
not be considered in isolation from it's physiology - and in fact that these two approaches
are complementary - this continues to be a weakness of many other demographic studies.
At the most general level, the efficiency of energy and nutrient intake, its processing,
allocation and expenditure is critical to the survival and reproductive success of individuals
and populations. Within-species, individual variation in physiological traits can be as
marked as that in ecological, behavioural or morphological traits, although the causes and
consequences of this variation are poorly understood. An understanding of the
mechanisms underlying this intra-specific variability in physiological traits is essential in
determining the constraints that might limit the ability of individuals, populations and
species to respond to chemical and biological changes in the environment.
Reproduction and migration represent two major aspects of the life cycles of most birds,
and in particular of those species that have been targeted as being of primary importance
in this study (colonially-breeding seabirds and migratory shorebirds and waterfowl). These
will therefore form the focus of the work of the Associate Chair. Reproductive output is
dependent on breeding propensity and breeding effort (the number and size of eggs laid)
and these vary markedly between individuals. Understanding the physiological basis of egg
production, and egg-size variation, in wild birds will represent one of the two main goals
of this project. Egg production has been demonstrated to be the phase of the life-cycle of
birds that is most sensitive to environmental change (particularly with regard to pollution
and habitat alteration). Secondly, efficient utilisation of food resources and control of body
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reserves is essential for the successful completion of migration and for over-winter
survival. Again, it is known that there is marked intra-specific variation, for example, in
the rate of fat deposition between individuals. The second goal of this project will be to
investigate the physiological basis of this variation in relation to possible constraints
operating on diet choice, digestive capabilities and fat storage.
B. Academic Goals, The SFU Perspective
1. Population Ecology
What are the major academic issues in population ecology, which the Chair can contribute
to the discipline as a whole? We identify two large areas where the projects outlined here
can contribute. Firstly, although many long term bird population studies have investigated
variation in reproduction, much less has been done in terms of variation in survival. This is
partly due to the fact that more effort and resources are required to collect the data, and
partly because until recently the necessary mathematical models have not been available.
Recently however, new software packages have become available and we have spent
considerable time learning these new techniques with the view to applying them to long
term bird-banding data. Having pioneered this with the La Perouse Bay data set, we are
now in a position to adapt it for other data sets, both those already in existence and those
which we generate during the mandate of the Chair. One of the academic goals of the
Chair therefore will be to develop a Canadian centre where the measurement of survival
variation can be measured and assessed. To this end, we have hired Dr. E. Cooch whose
mathematical and computer skills make him ideal for this work. As well, Dr. R. Pradel,
who helped to develop the Survival General Estimation (SURGE) program for
investigating survival variation has joined our team as a post-doctoral fellow (PDF). We
also plan to hold workshops to teach these methods and make them more readily available
for Canadian academics and wildlife biologists.
The SURGE and other recent methodologies are also valuable for examining patterns of
philopatry in birds and will be used for a theoretical and practical exploration of the whole
question of philopatry in birds. Although some general predictions about natal and
breeding philopatry can now be made, there is very little understanding about the
processes involved in winter and migration philopatry. Many applied questions depend on
an understanding of the rules that determine whether a bird will return to a previously used
location. Often it is important to know whether the populations of birds in an area are
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residents, transients or a mix of the two. Turnover rates of birds within populations are
also important in assessing impact of birds on their environment. Both these questions
ultimately depend on understanding the principles of philopatry away from the breeding
ground. We hope to concentrate on these questions as a unifying theme for our population
work.
2. Behavioural Ecology
Although population ecology and behavioural ecology have been closely tied during their
histories, fundamental assumptions remain unique to each. In particular, population
models consider neither adaptive adjustments in behaviour nor the possibility of
behavioural evolution in making predictions. Yet there is every reason to believe that these
are fundamentally important processes.
The discipline of behavioural ecology as a whole has set, as a major goal, the integration
of behavioural and population theory into a unified whole. Several major works of the past
decade have been devoted to considering the ecological consequences of adaptive
behaviour, but so far, the real progress has been very limited.
The Chair will be able to make a major contribution in this area because of the crossfertilization between large and active groups in both population and behavioural ecology.
The aim is to understand intra-specific life history variation, and to use that as a basis for
population predictions, rather than the reverse, which is the existing procedure. This
would constitute a major and important advance, and we plan to be major contributors in
this area.
3. Physiological Ecology
The principal academic goal of research in Physiological Ecology will be to gain an
understanding of the significance of, and mechanisms underlying, individual and agespecific expression of physiological traits. As a model system, the work will initially
investigate the physiological basis of reproductive investment in egg-laying and the
functional significance of egg-size variation in birds. Specifically, the project will
concentrate on three main aspects: 1) energetic costs of egg production, 2) the
relationship between egg-size and offspring fitness, and 3) mechanisms and constraints in
egg-laying females. The approach taken will be to conduct experimental work under
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WILDLIFE ECOLOGY RESEARCH CHAIR
more-or-less controlled conditions, for example using captive breeding birds or perhaps a
nest-box breeding population, and then to test ideas and hypotheses generated in this way
on selected free-living species of particular interest to the Chair. A similar approach would
be taken toward the study of the physiological basis of, and possible constraints on, diet
choice, digestive capabilities and fat deposition. This work would concentrate on
migratory shorebirds, probably Dunlin or western sandpipers, and the aim would be to
establish a population of these birds in captivity.
These two areas of research are linked by the common goals of understanding
1) individual differences in the patterns of resource acquisition and expenditure, and
2) the adaptive significance of individual variation in physiological traits in free-living
birds, this in turn allowing a better understanding of their likely responses to
environmental change.
4. General Points
In addition to the specific themes outlined above, we hope that the projects which we
undertake will lead to a more general understanding of avian demographic processes
which will have applicability to the conservation problems associated with both lesser
known and endangered species. Often the detailed study necessary for conservation action
is not possible if the species is rare and sensitive to disturbance and conservationists have
no option but to extrapolate from more widespread and better studied species. For
example, the work with Western Sandpipers may provide valuable insights into the
management and protection of the endangered Piping Plover.
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