Seabirds in the Marine Wilderness of the
Western North Atlantic
Falk Huettmann
Abstract—Presenting the largest habitat in the world, patterns
and processes of oceans beyond the inshore area are poorly investigated. Seabirds are among the best researched marine creatures,
but their research at sea still presents the last frontier in field
ornithology. The Programme Inté gré de recherches sur les oiseaux
pé lagiques (Integrated Research Program on Seabirds, or PIROP)
database of the Canadian Wildlife Service is one of the largest data
sets on pelagic seabird distribution. In concert with biotic and
abiotic environmental factors, this data set presents a unique
opportunity to study how marine animals interact across national
borders (Canada, Greenland, and the United States) with the
unconstrained forces of nature in the huge marine area of the
Northwest Atlantic, from the Gulf of Maine to the Canadian Arctic.
This Marine Wilderness offers a mosaic of several “hot spots,”
including threats such as dependence on ocean currents driven by
the global climate, as well as those normally related to human
activities such as overfishing, oil pollution, and traffic. This paper
presents results from an international, multidisciplinary 3-year
research project that used advanced tools such as Geographic
Information Systems (GIS), remote sensing, international governmental long-term databases, spatial statistics, the Internet, modeling, Landscape Ecology approaches, and seabird surveys at sea.
Implications for seabird conservation and management are
shown also, emphasizing that knowledge of Marine Wilderness is
insufficient.
Presenting the largest habitat in the world, oceans cover
more than two-thirds of the globe, and the vast areas far
away from land can truly be considered as Marine Wilderness. Marine Wilderness in the western North Atlantic,
defined here as remote areas at sea far away from shore, lies
in international (Canada, Greenland, and the United States)
waters and is not fully researched and monitored, nor does
it enjoy proper and consistent protection. Coastal areas,
here defined as the waters within 5 km of the shore, are
easier to access and enjoy better monitoring. Seabirds are
among the best researched marine creatures, but seabird
research at sea still presents the last frontier in field ornithology. The only information about seabirds in the Marine
Wilderness of the western North Atlantic stems from surveys, based on vessels of opportunity, carried out by the
In: Watson, Alan E.; Aplet, Greg H.; Hendee, John C., comps. 2000.
Personal, societal, and ecological values of wilderness: Sixth World Wilderness Congress proceedings on research, management, and allocation, volume II;
1998 October 24–29; Bangalore, India. Proc. RMRS-P-14. Ogden, UT: U.S.
Department of Agriculture, Forest Service, Rocky Mountain Research Station.
Falk Huettmann is Ecologist, Atlantic Cooperative Wildlife Ecology Research Network, University of New Brunswick, P.O. Box 44555, Fredericton,
NB, Canada E3B 6C2, email: k9wk@unb.ca
USDA Forest Service Proceedings RMRS-P-14. 2000
Programme Inté gré de Recherches sur les Oiseaux Pé
lagiques (Integrated Research Program on Seabirds or
PIROP). This paper is based on a 3-year study by the author,
investigating the importance of the Marine Wilderness for
seabirds using PIROP, environmental long-term data sets
from a variety of sources, and Geographic Information Systems (GIS).
Description of the Study Area and
Marine Wilderness ______________
The study area (fig. 1) of this research project is located in
the western North Atlantic, from the Gulf of Maine to the
Canadian Arctic (Brown 1986; Brown and others 1975;
Diamond and others 1993). Brown and others (1975) present
an “oceanographical zonation” for this “Seascape”
(Huettmann and Diamond 1998c) based on Ashmole (1971),
Dunbar (1968), and Salomonsen (1965, 1972), and divides
the water bodies into high arctic, low arctic, boreal, and cool
subtropical without further temporal or vertical reference. A
classification based on biodiversity criteria and indicators
was not carried out for the study area (Angel 1993; Gaskin
1995).
There are nine major surface currents and streams in the
study area, such as the Gulf Stream and the Labrador
Current (Backus and Bourne 1987; Drinkwater 1996;
Greenberg and Petrie 1988; Salomonsen 1965; Stommel
1965). Due to a funnel effect (Defant 1961), areas with fast
currents can be found in the study area, for example, Flemish Pass, Strait of Belle Isle, and Grand Manan Channel.
Salinity increases with distance to the coast.
The occurrence and locations of fronts and eddies are well
described for the southern study area and they are among
the richest and most productive regions in the study area
(Fournier 1978; Loder and Greenberg 1986; Mann and
Lazier 1991; Olson and Backhus 1985; Wiebe 1977). There
are major banks in the study area, some of which play a role
in global fisheries: Grand Banks, Labrador Banks, St. Pierre
Banks, and the Scotian Shelf. All of these banks are relatively
close to the coast and, therefore, considered part of the
Continental Shelf. Other distinctive features of the study
area are deep sea channels and deep sea basins. The study
area has few sea mounts and no active underwater volcanoes.
Being one of the most important fishing grounds in the
world, the southern study area is heavily used by the
international fishing fleet and, therefore, divided into management zones by the North Atlantic Fishing Organization.
Due to the relevance of the western North Atlantic as a
traditional fishing ground, an oil resource, a mining resource, and until very recently, as a strategically important
area for submarines, the political setup of the study area is
quite complex.
237
Figure 1—Map showing locations of observed seabirds in PIROP, 1967-1992, and the seabird
species richness for the study area.
Agnew (1993) discusses the Davis Strait/Labrador Sea as
one of three regions with high sea ice and atmospheric
variability. The region from the Middle Atlantic Bight to the
Grand Banks exhibits the highest inter-annual variability
of sea surface temperature anywhere in the North Atlantic
Ocean (Cayan 1986 in Petrie and Drinkwater 1993;
Drinkwater 1996; Petrie and others 1992). One reason for
this variability is the connection of the study area with the
global climate and global seas. However, the biologically
important Cold Intermediate Layer is relatively stable. One
of the biggest environmental changes in this marine ecosystem, a near surface event known as the “Great Salinity
Anomaly,” occurred throughout the southern study area
from approximately 1969 to 1972 (Angel 1993; Mertz and
Myers 1994). Furthermore, Mann and Drinkwater (1994)
and Drinkwater (1996) described the North Atlantic Oscillation (NAO) and big variations of the NAO Index occurring
from 1900 to 1986. Severe and overall changes can be
expected for the study area in the event of global warming
(Mann and Lazier 1991; Montevecchi and Myers 1997). In
particular, the southern section of the study area is threatened by overfishing (Backus and Bourne 1987; Wilson and
Addison 1984 for Gulf of Maine) (do Carmo Gomes 1993;
Harris 1998 for Grand Banks) and contamination by radionuclides, chemical and thermal effluents from the Point
Lepreau Nuclear Power Plant (Nelson and others 1998). It
is also threatened by industrial sources, heavy metals, and
sea floor contamination in the Bay of Fundy/Gulf of Maine
238
(Backus and Bourne 1987; Braune 1987; Braune and Gaskin
1987; Gaskin 1987; Loring 1981, 1982).
The biggest threats, oil exploration and the transportation of oil and cargo, can be found in the southern highly
populated areas (Backus and Bourne 1987; Nettleship 1997).
Traffic already accounts for considerable mortality to endangered North Atlantic right whales (Eubalaena glacialis)
(Brown and others 1995; Gaskin 1987) and poses significant
threats of oil spills to wintering auk populations (Green and
Hirons 1993; Lock and others 1994).
The arctic regions have not been fully investigated for
threats or for their potential as protected areas (Beckmann
1995). Although the need for protected areas at sea has been
emphasized by a variety of researchers and agencies
(Nettleship 1997; Shackell and Willison 1995), there are no
regions at sea within the study area that have any true, yearround protection status.
Data Sets ______________________
Integrated Research Program on Seabirds
Initiated by the Canadian Wildlife Service in 1966 for
monitoring seabirds at sea, the Integrated Research Program on Seabirds (PIROP) database originally consisted of
vessel-based 10-minute seabird counts in the western North
Atlantic (Brown 1986; Brown and others 1975; Lock and
USDA Forest Service Proceedings RMRS-P-14. 2000
others 1994). This program also includes aerial survey data
and a large amount of occasional observations of sea mammals and birds other than seabirds, such as waterfowl and
passerines (Lock and others 1994). The PIROP database is
fully digitized (dBASE, relational database) and includes
software for analytical GIS work and convenient data entry
(Huettmann and Lock 1997). It was recently extended with
the addition of the U.S. Manomet Seabird data for the Gulf
of Maine area (Powers and others 1979). With 26 years
(1966 to 1992) of seabird-at-sea observations, PIROP is the
largest data set on this topic (Diamond and others 1993;
Lock and others 1994) and is available for the scientific
community.
Environmental Data Sets
For this study, 20 environmental data layers were used for
GIS overlays in combination with PIROP within the study
area (table 1) (Huettmann 1997a). All data sets used are
available for the scientific community free of charge and
were derived from satellite images, and official oceanographic, meteorological, and governmental data sources.
Data transfer was accomplished via the Internet (ftp, WWW,
and email) or sometimes digital media (floppy disks, zip
disks, and CD-ROMs); the Internet option proved to be the
best, most convenient way to locate, present, and transfer/
download data and subsets from large public databases
across platforms.
Table1—Overview of the data sets used to investigate Seabirds in the Marine Wilderness.
No.
Name of data set
Seabirds
1
PIROP
Atmospherical
2
Atmospheric
temperature at
880 mbar
3
Standard temperature
atmospheric at
880 mbar
4
Wind speed
5
Air pressure at
sea level
6
Air temperature
7
Wind when seabird
observation was done
Geographical
8
Sea depth
9
Slope of sea floor
10
Aspect of sea floor
11
Distance from coast
12
Distance from
a seamount
Water
13
Sea surface
temperature
14
Water temperature
at 30 m depth
15
Sea surface salinity
16
Water salinity at
30 m depth
17
Salinity difference
surface - 30 m depth
18
Temperature difference
surface - 30 m depth
19
Sea state when seabird
observations were done
Additional Data Sets
20
ECNASAP groundfish
21
Seabird colonies
Data source
Data type
Units
Bedford Institute for Oceanography,
Environment Canada, Canadian
Wildlife Service, Manomet Bird
Observatory
Continuous
Seabirds within a
10-minute block
1966-1992
Pathfinder (NOAA)
Continuous
Kelvin
April 1987December 1988
Pathfinder (NOAA)
Continuous
Standard deviation
April 1987December 1988
COADS (NOAA)
COADS (NOAA)
Continuous
Continuous
Meter/second
Mbar
1854-1993
1854-1993
COADS (NOAA)
PIROP
Continuous
Discrete
Kelvin
Classified
1854-1993
1966-1992
ETOPO5
SPANS-GIS from ETOPO5
SPANS-GIS from ETOPO5
SPANS-GIS
SPANS-GIS from ETOPO5
Continuous
Continuous
Discrete
Discrete
Discrete
Meters
Degrees
Classified
Distance bands
Distance bands
1988
1988
1988
1998
1996
WOA (NOAA)
Continuous
Degrees Celsius
1948-1988
WOA (NOAA)
Continuous
Degrees Celsius
1948-1988
WOA (NOAA)
WOA (NOAA)
Continuous
Continuous
Parts per million
Parts per million
1948-1988
1948-1988
SPANS-GIS based on WOA (NOAA)
Continuous
Parts per million
1948-1988
SPANS-GIS based on WOA (NOAA)
Continuous
Degrees Celsius
1948-1988
PIROP
Discrete
Classified
1966-1992
Department of Fisheries
and Oceans, Dartmouth
Lock and others 1994, Cairns 1989
Continuous
KG biomass,
species richness
Breeding pairs
1970-1994
USDA Forest Service Proceedings RMRS-P-14. 2000
Continuous
Time cover
1989, 1994
239
Seabirds in Summer _____________
Migration ______________________
The distribution of seabirds at sea is most easily monitored in summer, due to relatively mild weather conditions
and ongoing colony-based research. Most of the seabird
colony locations in the study area are well known (Boertmann
and others 1996; Cairns and others 1989; Nettleship 1980).
However, systematic sampling of the huge oceanic ecosystem is rarely attempted, and opportunistic sampling, such
as PIROP, is the source of most information (Brown 1986).
The temporal extent and timing of the breeding season for
seabirds in the study area is mostly a function of latitude;
Diamond and others (1993) define a northern and a southern
breeding regime (borderline at approximately 52o N). During the summer, breeding seabirds are attached to their
breeding colonies due to the need for providing their chicks
with food (Huettmann and Diamond 1997b; Weimerskirch
1998). Most of the seabird species in the Canadian North
Atlantic have up to 40 percent nonbreeders in their population (Diamond and others 1993), which could theoretically
be found dispersed all over the Atlantic because they are not
constrained in their movements by the duty to provide for
their chicks. Breeding in the southern hemisphere, Wilson’s
storm petrels (Oceanites oceanicus) as well as greater
(Puffinus gravis) and sooty shearwaters (P. griseus) are the
only seabird species that occur in great numbers in the study
area without actually breeding there.
All seabirds in the North Atlantic migrate. Normally,
their migration covers many hundreds of kilometers and is
affected by foraging, breeding, molting, flight energetics,
and the travel between winter and summer grounds (Furness
and Burger 1988; Newton and Dale 1996; Pennycuick 1987).
The importance of the Marine Wilderness for migration, for
resting areas, and as a flyway for North Atlantic seabirds is
not well known; these topics are difficult to investigate in the
field, and most knowledge consists of rough descriptions and
arrows on maps (Braune 1987; Furness 1987; Powers 1983;
Tuck 1961). The only quantitative information on seasonal
seabird migration stems normally from recoveries of birds
banded in colonies during summer (Brewer and others, in
preparation) or, to a lesser degree, from terrestrial and atsea observations (Stenhouse and Montevecchi 1996). Satellite transmitters offer the best data on movement patterns,
but to date have been applied to larger seabirds only and not
in the area covered by PIROP (Prince and others 1992).
Seabirds in Winter _______________
Research about seabirds at sea presents “the last frontier
in field biology” (Brown 1981 in Thurston 1990). The importance of this statement can be shown by the marginal
scientific knowledge about seabird distribution during winter months (November through February) in the offshore
region of the North Atlantic (Brown 1986; Nettleship and
Birkhead 1985; Powers and others 1979). Significant seabird information for this time of the year (for example, their
distribution within the Marine Wilderness) is scarce (Gaston
and McLaren 1990; Huettmann and others, in preparation).
So far, it is believed that most pelagic seabird populations,
such as thick-billed murres (Uria lomvia) and Atlantic
puffins (Fratercula arctica), simply disperse in winter over
the vast Northern Atlantic (Nettleship and Birkhead 1985).
The small amount of existing information for this time of the
year includes some feeding studies (Elliot and others 1990;
Falk and Durinck 1993) and movement patterns (Donaldson
and others 1997) derived from work in Greenland, the
“Turr” (mostly thick-billed murres and common murres
Uria aalge that hunt off Newfoundland), polynyas (Brown
and Nettleship 1981), and ice cover (Orr and Parsons 1982).
Large data gaps for winter also exist in PIROP. Therefore,
transferring knowledge derived from seabirds found in comparable areas elsewhere offers perhaps the only way to learn
more on how seabirds cope with winter in the Marine
Wilderness (see Divoky 1979 for the Pacific) (see Ainley and
others 1993; Hunt and Nettleship 1988 for the Antarctic)
(see Durinck and Falk 1996; Joiris 1983 for the North
Atlantic).
240
Methods _______________________
For analyzing seabird distribution during winter and
breeding season, the PIROP database was queried with
FoxPro (Siegel 1995) for the particular months. Only seabird
species that had sufficient observations were included; resulting data were then imported and visualized in a GIS
format. I am using SPANS-GIS (Huettmann 1998;
Huettmann and Diamond 1998a; INTERAC TYDAC 1995)
to present bird observations in the study area, to create
surfaces for environmental data layers (Huettmann 1998),
and to overlay different data layers per month (Huettmann
1997a) for further statistical analysis in SPLUS (StatSci
1995). Logistic regression and Classification and Regression
Trees were used to test for significant results (StatSci 1995).
Seabird queries for the environmental data overlays were
filtered for “standardized seabird counts” following Tasker
and others (1984). To track seabird movements during the
year, locations of observed juveniles and molting seabirds in
PIROP were queried and then analyzed per month in SPANSGIS.
Results and Discussion __________
Seabirds in Summer
Due to their limited feeding range, breeding seabirds in
the study area show a very coastal distribution concentrated
around their seabird colonies (Huettmann and Diamond
1998b). For most of the breeding seabirds, such as northern
gannet (Sula bassanus), black-legged kittiwake (Rissa
tridactyla), northern fulmar (Fulmarus glacialis), herring
gull (Larus argentatus), great black-backed gull (L. marinus),
Iceland gull (L. glaucoides), glaucous gull (L. hyperboreus),
thick-billed murre, common murre and Atlantic puffin, high
concentrations occur on the west and north shores of the St.
Lawrence region, Labrador coast, east Newfoundland coast,
USDA Forest Service Proceedings RMRS-P-14. 2000
northern Hudson Bay, West Greenland, and the Canadian
High Arctic. These are areas that are normally free of ice in
the pre-breeding season (Huettmann and Diamond 1998a).
For seabird species included in the investigation of the
environmental data overlay with PIROP for the breeding
season (black-legged kittiwake, common murre, thick-billed
murre, northern fulmar, Atlantic puffin, northern gannet),
results show a complex situation with several factors determining their distribution and colony locations. However,
depth, distance to coast, sea floor aspect, and meteorological
parameters such as sea level pressure, wind speed, and air
temperature are significant factors for the seabird distribution in summer. Nonbreeding birds were observed either
close to these colonies (most likely in order to obtain mates
for breeding and to occupy potential breeding sites) or far
away from any colonies. We assume that the latter birds are
normally young individuals that disperse from their original
colonies to look for potential mates elsewhere or to feed
opportunistically in areas where food is plentiful and easily
accessed. During the breeding season, northern fulmars
(likely nonbreeding individuals from the whole northern
Atlantic) can be found on the Labrador Shelf (Hatch and
Nettleship 1998). Baird (1994) presents that the Grand
Banks are of international importance for black-legged kittiwakes year round. Nonbreeding northern gannets were
observed in the Grand Manan region. Our data show an
abundance of greater shearwaters in the southern study
area from mid-May through July (breeders) and June through
November (nonbreeders) carrying out a trans-equatorial
migration from the southern hemisphere for molting (Brown
1988).
During summer, the Marine Wilderness is not used by
most seabirds to the same extent as the coastal zones. Even
nonbreeding birds, with the ability to range freely, were
most often found in offshore regions of the Continental Shelf.
Seabirds in Winter
Environmental data suggest that the winter season must
be divided into early (November and December) and late
winter (January and February). The Davis Strait is still
open in early winter but is covered with sea ice in December,
driving most arctic birds into southern waters such as the
Labrador Sea, Grand Banks, Scotian Shelf, and Gulf of
Maine. This is also supported by observations indicating
that the marine environment changes substantially from
the early winter season to the late winter season (Braune
1987 for early winter plankton peak in the Bay of Fundy;
Elliot and others 1990 for shift of seabird prey).
The environmental data overlays with seabirds in PIROP
for the winter season were done for the following species:
black-legged kittiwake, dovekie (Alle alle), northern fulmar,
great black-backed gull, herring gull, white gulls (Iceland
gull, glaucous gull, ivory gull Pagophila eburnea, Ross’s gull
Rhodostethia rosea), and large auks (thick-billed murre,
common murre, Atlantic puffin, razorbill Alca torda). Results show that mostly sea floor aspect, distance to coast, and
the classical oceanographical factors such as water temperature, salinity, and wind speed determine seabird distribution in winter. The data clearly indicate a pelagic distribution
for northern fulmars and black-legged kittiwakes. Great
USDA Forest Service Proceedings RMRS-P-14. 2000
black-backed gulls and herring gulls were found offshore,
but are mainly coastal (Root 1988). Large auks were observed offshore, but the winter hunt on “Turrs” and, in
particular, findings from winter surveys off Grand Manan
for wintering razorbills indicate a very high density of
wintering birds in inshore (less than approximately 5 km
from land) coastal waters (Huettmann and others, in preparation; Root 1988). Root (1988) indicated that the Grand
Manan and Passamaquoddy Bay area also hosts the largest
numbers of black-legged kittiwakes observed during Christmas bird counts on the North American continent. Iceland
gulls were mostly found in coastal areas southwest of Newfoundland, Cape Breton, south of Nova Scotia, and around
the Magdalen Islands. Root (1988) reports that observations
of Iceland gulls were high at the extreme southeastern point
of New Brunswick. In terms of their distribution, Iceland
gulls can, therefore, be labeled as coastal birds in winter.
These results suggest that, in terms of numbers and
concentrations of seabirds occurring in the study area during winter, the coastal waters may be more important than
suggested earlier, at least for the southern study area such
as the Bay of Fundy, southern Nova Scotia, and the Gulf of
Maine. Ice-free coastal zones in the study area present an
important wintering ground for most seabird species, where
they share their habitat with other waterbirds, such as
waterfowl, grebes, and loons (Root 1988). I conclude that
coastal zones must present a richer habitat for seabirds in
winter than the ice-free Marine Wilderness far offshore.
Post and Pre-Breeding Migration
Seabird migration can be characterized as a directed
movement between summer and winter grounds. Although
migration strategies for seabirds are not that well researched,
at least some resting grounds during migration are roughly
known. My results indicate that juvenile thick-billed murres
do not show a rapid southward movement after leaving the
colonies. Their migration flyway seems to follow the coastline and shelf edges, instead of crossing the open sea directly
(except for the Davis Strait off the Cumberland Peninsula).
Most of the northern gannets leave the breeding colonies in
the St. Lawrence estuary and off Newfoundland completely
by September, and then return in mid-April. Greater shearwaters arrive in the study area beginning in mid-May,
followed by juvenile birds during the next months. They can
be found in the coastal waters off southern Newfoundland
feeding on capelin (Mallotus villosus) and have been observed dispersing as far north as Greenland. After July, they
make full use of the whole North Atlantic. Breeding birds
seem to leave the study area starting in July; remaining
birds, likely nonbreeders, can be found in the area until
December. Northern fulmars were observed in the study
area year-round. Hatch and Nettleship (1998) indicate that
birds in North America generally do not show a directed
movement, except perhaps birds from the High Arctic.
However, due to the presence in the study area of birds from
the whole North Atlantic, movement patterns can be detected from changing numbers on the banks. Black-legged
kittiwakes were found all over the study area, with high
densities in the Gulf of Maine region in winter. Herring gulls
show a clear removal from the St. Lawrence area during
241
winter and a southward movement for wintering to the Gulf
of Maine and beyond. Great black-backed gulls exhibit the
same wintering pattern as herring gulls, but breed farther
north. Iceland gulls move between winter and summer
grounds along the coastal shelf edge and ice edge. Glaucous
gull movements indicate a more pelagic wandering than
Iceland gulls.
General
The distribution and biology of seabirds that are rare,
difficult to determine, and hard to see or count at sea still
presents a challenge, and their conservation status and
biology remain unknown in the study area. Jaegers (Stercocarius parasiticus, S. pomarinus, S. longicaudus), and Cory
(P. diomedea) and Manx’s shearwaters (P. puffinus) occur in
the study area, but even basic biological knowledge is lacking. The world’s largest known nesting colony of Leach’s
storm-petrels (Oceanodroma leucorhoa) can be found in the
study area on Baccalieu Island on the Grand Banks
(Sklepkovych and Montevecchi 1989), but their feeding
grounds, wintering grounds, and movement patterns are not
known. Schneider and Heinemann (1996) indicate the importance of the southern study area for two other seabird
species from the southern hemisphere (Wilson storm petrel
and sooty shearwater), but details on distribution, population, and biology for these species are only conjecture.
Conclusion _____________________
The importance of the Marine Wilderness for seabirds is
not known nor fully monitored. The results, so far, indicate
that coastal areas, and to a lesser degree the Continental
Shelf, are key zones for seabirds during the winter (such as
the Bay of Fundy, southern Nova Scotia, and the Gulf of
Maine) and during the breeding season (almost the entire
coastline of the study area) (see fig. 1). The shelf edge is used
by most seabirds for activities such as feeding, migrating, or
roosting (nonbreeders). For the study area, I conclude that
the Marine Wilderness presents a relatively poor and homogenous environment for seabirds (see also Dunbar 1968
for the arctic; Wiebe 1977 for mosaics in boreal waters)
where environmental conditions can be extreme and prey is
difficult to catch. Shallow but remote areas at sea, such as
the extending Continental Shelf, are used by truly pelagic
seabird species, such as black-legged kittiwakes and northern fulmars, and for a relatively short period of time by
greater shearwaters, herring gulls, and great black-backed
gulls.
Besides other basic biological parameters in winter, habitat use of the Marine Wilderness is not fully known for the
Atlantic puffin, thick-billed and common murre, jaegers,
manx and Cory’s shearwater, and Leach and Wilson’s storm
petrel. During the migration season, deep-water sections of
the Marine Wilderness are used by seabirds like the glaucous gull, Iceland gull, and great black-backed gull to connect
winter and summer grounds. I recommend that a specially
designed, interdisciplinary seabird survey program be
conducted for the Marine Wilderness (with consideration of
the area’s international nature) to learn more about this
huge and important habitat.
242
References _____________________
Agnew, D. 1993. Simultaneous winter sea-ice and atmospheric
circulation anomaly patterns. Atmosphere-Ocean. 31(2): 259-280.
Ainley, D. G.; Ribic, C. A.; Spear, L. B. 1993. Species-habitat relationships among antarctic seabirds: a function of physical or biological factors? The Condor. 95: 806-816.
Angel, M. V. 1993. Biodiversity in the pelagic ocean. Conservation
Biology. 4(4): 760-772.
Ashmole, N. P. 1971. Seabird ecology and the marine environment.
In: Farner, D. S.; King, J. R., eds. Avian Biology. New York, NY:
Academic Press: 223-286.
Backus, R. H.; Bourne, D. W. 1987. The Georges Bank. Woods Hole,
MA: MIT Press. 593 p.
Baird P. 1994. Black-legged kittiwake (Rissa tridactyla). In:
Poole, A.; Gill, F., eds. The birds of North America, No. 92.
Philadelphia, PA: The Birds of North America Inc. 27 p.
Beckmann, L. 1995. Marine conservation in the Canadian Arctic.
In: Shackell, N.; Willison, J. H. M., eds. Marine protected areas
and sustainable fisheries. Science and Management of Protected
Areas Association. Wolfville, Nova Scotia: 227-235.
Boertmann, D.; Mosbech, A.; Falk, K.; Kampp, K. 1996. Seabird
o
o
colonies in western Greenland (60 -79 30' N. lat.), NERI Tech.
Rep. 170. Roskilde, Denmark: National Environmental Research
Institute. 148 p.
Braune, B. M. 1987. Seasonal aspects of the diet of Bonaparte’s gulls
(Larus philadelphia) in the Quoddy region, New Brunswick,
Canada. The Auk. 104: 167-172.
Braune, B. M.; Gaskin D. E. 1987. Total mercury levels in Bonaparte
gulls (Larus philadelphia) during autumn molt in the Quoddy
region, Brunswick, Canada. Archive of Environmental Contamination and Toxicology. 16: 539-549.
Brewer, D.; Diamond T.; Woodsworth, E. [In preparation]. Canadian bird banding atlas. Ottawa, ON, Canada: Canadian Wildlife
Service.
Brown, R. G. B. 1986. Revised atlas of Eastern Canadian seabirds.
Ottawa, ON, Canada: Canadian Wildlife Service. 110 p.
Brown, R. G. B. 1988. The Wing-molt of fulmars and shearwaters
(Procellariidae) in Canadian Atlantic waters. The Canadian
Field-Naturalist. 102(2): 203-208.
Brown, R. G. B.; Nettleship, D. 1981. The biological significance of
polynyas to arctic colonial seabirds. In: Stirling, I.; Cleator, H.,
eds. Polynyas in the Canadian Arctic. Occasional Paper Number.
Ottawa, ON, Canada: Canadian Wildlife Service: 59-65.
Brown, R. G. B.; Nettleship, D. N.; Germain, P.; Tull, C. E.; Davis,
T. 1975. Atlas of eastern Canadian seabirds. Ottawa, ON, Canada:
Canadian Wildlife Service. 219 p.
Brown, M.; Allen, J. M.; Kraus S. 1995. The designation of seasonal
Right Whale Conservation Areas in the waters of Atlantic Canada.
In: Shackell N.; Willison, J. H. M., eds. Marine Protected Areas
and sustainable fisheries. Wolfville, NS, Canada: Science and
Management of Protected Areas Association: 82-90.
Cairns, D. K.; Montevecchi, W. A.; Threlfall, W. 1989. Researcher’s
guide to Newfoundland seabird colonies. Occasional Papers in
Biology, No.14. St. John’s, NF, Canada: Memorial University of
Newfoundland. 34 p.
Defant, A. 1961. Physical oceanography. Vol I. Oxford, UK: Pergamon
Press. 729 p.
Diamond, A. W.; Gaston, A. J.; Brown, R. G. B. 1993. Studies of highlatitude seabirds. 3. A model of the energy demands of the
seabirds of eastern Arctic Canada. In: Montevecchi, W. A., ed.
Canadian Wildlife Service Occasional Paper No. 77. Ottawa, ON,
Canada. 39 p.
Divoky, G. J. 1979. Sea ice as factor in seabird distribution and
ecology in the Beaufort, Chukchi and Bering Seas. In: Bartonek,
J. C.; Nettleship, D. N., eds. Conservation of marine birds of
northern North America. Washington, DC: U.S. Department of
the Interior, Fish and Wildlife Service: 9-17.
do Carmo Gomes, M. 1993. Predictions under uncertainty. St.John’s,
Newfoundland: Institute of Social and Economic Research. 240 p.
Donaldson, G. M.; Gaston, A. J.; Chardine J. W.; Kampp, K.;
Nettleship, D. N.; Elliot, R. D. 1997. Winter distributions of thickbilled murres from the eastern Canadian Arctic and western
Greenland in relation to age and time of year. Occasional Paper
Number 96. Ottawa, ON: Canadian Wildlife Service. 26 p.
USDA Forest Service Proceedings RMRS-P-14. 2000
Drinkwater, K. F. 1996. Atmospheric and oceanic variability in the
northwest Atlantic during the 1980’s and early 1990’s. Journal of
Northwest Atlantic Fisheries Science. 18: 77-97.
Dunbar, M. J. 1968. Ecological developments in polar regions: a
study in evolution. Englewood Cliffs, NJ: Prentice-Hall. 119 p.
Durinck, J.; Falk, K. 1996. The distribution and abundance of
seabirds off southwestern Greenland in autumn and winter 19881989. Polar Research. 15(1): 23-42.
Elliot, R. D.; Ryan, P. C.; Lidster, W. W. 1990. The winter diet of
thick-billed murres in coastal Newfoundland waters. Studies in
Avian Biology. 14: 125-138.
Falk, K.; Durinck, J. 1993. The winter diet of thick-billed murres,
Uria lomvia, in western Greenland, 1988-1989. Canadian Journal of Zoology. 71: 264-272.
Fournier, R. O. 1978. Biological aspects of the Nova Scotia shelfbreak
fronts. In: Bowman, M. J.; Esaias, W. E., eds. Oceanic fronts in
coastal processes. Marine Sciences Research Center. New Orleans, LA: Springer Verlag: 22-35.
Furness, R. W. 1987. The skuas. Calton, UK: T. & A. D. Poyser.
363 p.
Furness, R. W.; Burger, A. E. 1988. Effects of energy constraints on
seabird breeding at high latitudes. In: Quellet, H., ed. Proceedings XIX International Ornithological Congress 1986. Ontario,
Canada: University of Ottawa Press: 1205-1217.
Gaskin, D. E. 1987. Updated status of the right whale, Eubalaena
glacialis, in Canada. Canadian Field-Naturalist. 101(2): 295-309.
Gaskin, D. E. 1995. Marine biodiversity monitoring. Department of
Zoology, University of Guelph, Ontario. 120 p.
Gaston, A. J.; McLaren, P. L. 1990. Winter observations of black
guillemots in Hudson Bay and Davis Strait. Studies in Avian
Biology. 14: 67-70.
Green, R. E.; Hirons, G. J. M. 1993. The relevance of population
studies to the conservation of threatened birds. In: Perrins, C. M.;
Lebreton, J. D.; Herons, G. J. M., eds. 1993. Bird population
studies: relevance to conservation and management. Oxford
University Press, UK: 594-637.
Greenberg, D. A.; Petrie, G. D. 1988. The mean barotropic circulation on the Newfoundland Shelf and Slope. Journal of Geophysical Research. 93(C12): 15,541-15,550.
Harris, M. 1998. Lament for an ocean. Toronto, Ontario: McClelland
& Stewart. 342 p.
Hatch, S. A.; Nettleship, D. N. 1998. Northern fulmar (Fulmarus
glacialis). In: Poole, A.; Gill, E., eds. The birds of North America,
Vol. 361. Philadelphia, PA: The Birds of North America, Inc.
32 p.
Huettmann, F. 1997a. Birds at sea: linking long-term monitoring
data for seabirds with oceanographic data. In: The ecological
monitoring and assessment network report on the third national
science meeting: proceedings; 1997 January 21-25; Saskatoon,
Saskatchewan, Canada: 54-57. [Online] EMAN website: http://
www.cciw.ca/eman-temp/reports/publications/nm97_birds/
intro.html.
Huettmann, F. 1997b. [Personal communication]. Selected issues
for the successful application of a Geographic Information System in ornithology: some considerations for working with
georeferenced data. GIS Workshop for the Society of Canadian
Ornithologists; 7-10 August 1997; Peterborough, ON, Canada.
Huettmann, F. 1998. An ecological GIS research application for the
Northern Atlantic—The PIROP database software, environmental data sets and the role of the Internet/WWW. In: Riekert, W.F.; Tochtermann K., eds. “Hypermedia im Umweltschutz” Proceedings of Deutsche Gesellschaft fü r Informatik (GI) and
Forschungsinstitut fü r anwendungsorientierte Wissensverarbeitung (FAW) Ulm. Umwelt-Informatik aktuell; Bd.17,
Metropolis Verlag/Marburg: 213-217.
Huettmann, F.; Diamond, A. W. 1998a. Seabird surveys and selected environmental data sets in the Bay of Fundy: findings and
conclusions from monthly ferry transects (St. John-Digby-St.
John). Proceedings of EMAN 97 Conference Bay of Fundy; 1997
November; St.Andrews, New Brunswick: 85-92.
Huettmann, F.; Diamond, A. W. 1998b. Characterizing, modeling
and predicting locations of seabird colonies in the Davis Strait:
using the PIROP database, GIS and environmental data to
evaluate the suitability of marine breeding habitats for Arctic
seabirds. Proceedings of environmental prediction workshop;
1998 February 17-19; Halifax, NS. Environment Canada: 86-94.
USDA Forest Service Proceedings RMRS-P-14. 2000
Huettmann, F.; Diamond, A. W. 1998c. “Seascape” ecology: applying
terrestrial concepts of marine systems. Poster at the International Association for Landscape Ecology Conference; 1998 March
17-21; East Lansing, MI.
Huettmann, F.; Lock, A. R. 1997. A new software system for the
PIROP database: data flow and an approach for a seabird-depth
analysis. International Council for the Exploration of the Sea.
Journal of Marine Science. 54: 518-523.
Huettmann, F.; Dalzell, B.; Diamond, A. W.; Lock, T.; MacIntosh, K.;
Nettleship, D. [In preparation]. Winter distribution of razorbills
Alca torda and other auks in the lower Bay of Fundy, New
Brunswick.
Hunt, G. L.; Nettleship, D. N. 1988. Seabirds of high-latitude
northern and southern environments. In: Ouellet, H., ed. Proceedings XIX International Ornithological Congress 1986. University of Ottawa Press, ON: 1143-1155.
INTERAC TYDAC 1995. International SPANS workshop. Technical Demonstrations Notes; 1995 August 30-September 1; Ottawa,
ON, Canada. 281 p.
Joiris, C. 1983. Winter distribution of seabirds in the North Sea: an
oceanographical interpretation. Le Gerfaut. 73: 107-22.
Lock, A. R.; Brown, R. G. B.; Gerriets, S. H. 1994. Gazetteer of
marine birds in Atlantic Canada. Halifax, NS: Canadian Wildlife
Service. 137 p.
Loder, J. W.; Greenberg, D. A. 1986. Predicted positions of tidal
fronts in the Gulf of Maine region. Continental Shelf Research.
6(3): 397-414.
Loring, D. H. 1981. Potential bioavailability of metals in eastern
Canadian estuarine and coastal sediments. Rapport Pour la Ré
union de la Conseil International de la Exploration de la Mer
(Report from the International Council for the Exploration of the
Sea). 181: 93-101.
Loring, D. H. 1982. Geochemical factors controlling the accumulation and dispersal of heavy metals in the Bay of Fundy. Canadian
Journal for Earth Sciences. 19: 930-943.
Mann, K. H.; Drinkwater, K. F. 1994. Environmental influences on
fish and shellfish production in the Northwest Atlantic. Environmental Review. 2: 16-32.
Mann, K. H.; Lazier, R. N. 1991. Dynamics of marine ecosystems.
Boston: Blackwell Scientific Publications. 390 p.
Mertz, A.; Myers, R. A. 1994. The ecological impact of the Great
Salinity Anomaly in the northern Northwest Atlantic. Fisheries
and Oceanographers. 1: 1-14.
Montevecchi , W. A.; Myers, R. A. 1997. Centurial and decadal
oceanographic influences on changes in northern gannet populations and diets in the northwest Atlantic: implications for climate
change. International Council for the Exploration of the Sea.
Journal of Marine Science. 54: 608-614.
National Oceanic and Atmospheric Administration. 1996. Five
minute gridded earth topography data. [Online] Http://
edcwww.cr.usgs.gov/glis/hyper/guide/etopo5
National Oceanic and Atmospheric Administration. 1997. Live
access to climate data. [Online] Http://ferret.wrc.noaa.gov/fbin/
climate_server
Nelson, R. W. P.; Ellis, K. M.; Smith J. N. 1998. Long-term trends in
radionuclide distributions in the vicinity of a CANDU nuclear
generating station. Radiation Protection Dosimetry. 75(1-4):
71-76.
Nettleship, D. N. 1980. A guide to the major seabird colonies of
eastern Canada: identity, distribution and abundance. Canadian
Wildlife Service “Studies on Northern Seabirds” Manuscript Rep.
97: 1-133.
Nettleship, D. 1997. [Personal communication]. Ecosystem disturbance and seabirds in crisis: Eastern and Atlantic Canada.
Invited keynote oral paper presentation for WWF Canada’s
Atlantic Canada Endangered Species Technical Workshop; 1997
March 21; Dartmouth, NS, Canada. Bedford Institute of Oceanography.
Nettleship, D.; Birkhead, T. 1985. The Atlantic Alcidae. London:
Academic Press. 584 p.
Newton, I.; Dale, L. 1996. Relationship between migration and
latitude among west European birds. Journal of Animal Ecology.
65: 137-46.
Olson, D. B.; Backus, R. H. 1985. The concentrating of organisms at
fronts: a cold-water fish and a warm-core Gulf Stream ring.
Journal of Marine Research. 43: 113-137.
243
Orr, C. D.; Parsons, J. L. 1982. Ivory gulls, Pagophila eburnea, and
ice edges in Davis Strait and the Labrador Sea. Canadian FieldNaturalist. 96: 323-8.
Pennycuick, C. J. 1987. Flight of seabirds. Croxall, J. P., ed.
Seabirds: feeding ecology and role in marine ecosystems. Cambridge, UK: Cambridge University Press.
Petrie, B.; Drinkwater, K. 1993. Temperature and salinity variability on the Scotian Shelf and in the Gulf of Maine 1945-1990.
Journal of Geophysical Research. 98(C11): 20079-20089.
Petrie, B.; Loder, J. W.; Lazier, J.; Akenhead, S. 1992. Temperature
and salinity variability on the eastern Newfoundland Shelf: the
residual field. Atmosphere-Ocean. 30: 120-139.
Powers, K. 1983. Pelagic distributions of marine birds off the
northeastern United States. U.S. Department of Commerce,
National Oceanographic and Atmospheric Administration (NOAA)
Technical Memorandum NMFS-F/NEC-27. 201 p.
Powers, K. D.; Pittman, B. G. C.; Burrell, G. C. 1979. Distribution of
marine birds on the Mid- and North-Atlantic. U.S. Outer Continental Shelf. Volume Technical Progress Report, September
1978-August 1979. Manomet Bird Observatory, U.S. Department of Energy, Ecological Research Division, Office of Health
and Environmental Research, Office of Environment. 120 p.
Prince, P. A.; Wood, A. G.; Barton, T.; Croxall, J. P. 1992. Satellite
tracking of wandering albatrosses (Diomedea exuland) in the
south Atlantic. Antarctic Science. 4(1): 31-36.
Root, T. 1988. Atlas of wintering North American birds, an analysis
of Christmas bird count data. Chicago: University of Chicago
Press. 312 p.
Salomonsen, F. 1965. The geographical variation of the fulmar
(Fulmarus glacialis) and the zones of marine environment in the
North Atlantic. Auk. 82: 327-355.
Salomonsen, F. 1972. Zoogeographical and ecological problems in
Arctic birds. Proceedings of the XVth International Ornithological Congress 1970: 25-77.
Schneider, D. C.; Heinemann, D. W. 1996. The state of marine bird
populations from Cape Hatteras to the Gulf of Maine. In: Sherman,
244
K.; Jaworski, N. A.; Smayda, T. J., eds. The Northeast Shelf
ecosystem: assesment, sustainability, and management. Cambridge, MA: Blackwell Science: 197-216.
Shackell N.; Willison, J. H. M., eds. 1995. Marine protected areas
and sustainable fisheries. Wolfville, NS, Canada: Science and
Management of Protected Areas Association. 300 p.
Siegel, C. 1995. Mastering FoxPro 2.6 -Special Edition-. San Francisco: Sybex. 981 p.
Sklepkovych, B. O.; Montevecchi, W. A. 1989.The world’s largest
known nesting colony of Leach’s storm petrels on Baccalieu
Island, Newfoundland. American Birds. 43(1): 38-42.
StatSci. 1995. SPLUS guide to statistical and mathematical analysis. Version 3.3. Seattle: MathSoft Inc. 450 p.
Stenhouse, I.; Montevecchi, A. W. 1996. Winter distribution and
wrecks of little auks (Dovekies) Alle A. Alle in the northwest
Atlantic. SULA. 10(5): 219-228.
Stommel, H. 1965. The Gulf Stream, a physcial and dynamical
description. 2nd edition. Los Angeles: University of California
Press. 248 p.
Tasker, M. L.; Hope Jones, P.; Dixon, T.; Blake, B. F. 1984. Counting
birds at sea from ships: a review of methods employed and a
suggestion for a standardized approach. The Auk. 101: 567-577.
Thurston, H. 1990. Tidal life, a natural history of the Bay of Fundy.
Willowdale, ON: Firefly Books. 240 p.
Tuck, L. M. 1961. The murres: their distribution, populations and
biology—a study of the genus Uria. Canadian Wildlife Monograph Series No.1. St. John’s, NF. 260 p.
Weimerskirch, H. 1998. How can pelagic seabird provision its chick
when relying on a distant food resource? Cyclic attendance at the
colony, foraging decision and body condition in sooty shearwaters. Journal of Animal Ecology. 67: 99-109.
Wiebe, P. H. 1977. Rings of the Gulf Stream. Scientific American.
246(3): 60-70.
Wilson, R. C. H.; Addison, R. F. 1984. Health of the northwest
Atlantic. Dartmouth, NS, Canada: Environment Canada and
Department of Fisheries and Oceans. 174 p.
USDA Forest Service Proceedings RMRS-P-14. 2000