Fridtjof Mehlum
Summer distribution of
seabirds in northern
Greenland and Barents Seas
NORSK POLARINSTITUTT
OSLO 1989
Fridtjof Mehlum
Summer distribution of
seabirds in northern
Greenland and Barents Seas
NORSK POLARINSTITUTI
OSLO 1989
Cover page: Fulmar Fulmarus glacialis.
Drawing by Viggo Ree
ISBN 82-90307-53-5
Printed April 1989
IPil�G[]�1
II LI
Printed in Grea! Britain by
Page Bros (Norwich) Ltd
Norsk Polarinstitutt,
Rolfstangveien 12,
1330 Oslo Lufthavn,
Norway
Contents
Abstract ........................
5
Introduction ...................
7
Material and methods ...........................................
7
Cruises and areas covered ............................................................................................................................
7
Recording methods .................................................................................................................................... .
7
Distribution and abundance maps ..................................................................................................................
8
Oceanography and sea-ice conditions ................................................................................................................
8
Results
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
13
Fulmar Fulmarus glacialis ............................................................................................................................
14
Black Guillemot Cepphus grylle ....................................................................................................................
16
Puffin Fratereula arctiea ...................................... ......................................................................................
18
Little Auk A Ile alle.....................................................................................................................................
20
Briinnich's Guillemot Uria lomvia ......................................................
22
Common Guillemot Uria aalge......................................................................................................................
24
Kittiwake Rissa tridactyla... . .. . . ... . .. .. .. . . . . . . . . . . . . . . .. ... ... ... ... .. . .... . .. ... ... .. ... ... .. ... ... .. .. ... ... .. ... . . . . . ... .... . .. .... .. . . . . . . . . . . .
26
Ivory Gull Pagophila eburnea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
...................................... ...............
28
Ross' Gull Rhodostethia rosea............................................... ................ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
30
Sabine's Gull Larus sabini.................................................... .......................................................................
32
Glaucous Gull Larus hyperboreus..................................................................................................................
34
Great Black-backed Gull Larus marinus .........................................................................................................
36
Arctic Tern Sterna paradisaea .... .. .......................... .......................................................................................
38
Arctic Skua Stercorarius parasiticus.. . . . . . . . . .. . . . . . . . . . . .. ... . .. ... ... .... . ... .. . . . . . . .. ... ... .. . . . . . . .. .. . . . . . . ..... ... .......... .. ... . .. .. . . . . ..
40
Long-tailed Skua Stercorarius longicaudus .. . . . . .. . . . . .. . . . . . . ... ... ..... ... . . . . . . . . ... ... .. ... ... .. .. ... ..... ... ..... ....... ... .. ... ... ... ... .. .
42
Pomarine Skua Stercorarius pomarinus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
44
Great Skua Stercorarius skua . . . . . . .. . . . . . . . . ... . .. . . . . .. ... ... ... ... ... ..... .. . . . ... . . . . . . .. . . . . . . . . ..... ... .. . . . . . . .. .. . . . . . . . . .. . . . . . . . . . . . . .. ... .
46
Common Eider Somateria mollissima ................... ..... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
48
Long-tailed Duck Clangula hyemalis ............................................. .................................................................
50
Discussion.... ... ... ... ... . . . . . . . . . . . . . . ... . . . . . . . . . . . .. . ... . . . ... ... ... ... ... . . . ... .......... ... . . . . . ..... ... ..... . . . . . . . . . . . . . .. .......... ... . .. .. . . . . . . ... .
54
Acknowledgements... ... . . . . .. . . . . ... . .. . . .. . ... . . . . ... ... ... ... . . . . . . ... ... ....... ... . . . . . . .. ... ... .. ... ... .. . . . . . . . . . . . . . . . . .. ....... ... ... . . . . . . . . . . . ..
55
References ...... .. . ... ... . ... ..... . ... . . . . . . . . . . . . . . . ... . . . . .. . . . . .. ... ... ... . .. .. ... ... .. . . . . . . ..... ... .. . . . . .. .. . . . . . . . . ... . . . . . ............ ... ..... ... . .. .
55
Abstract
Line transects of seabirds were conducted in the seas around Svalbard during five summer seasons ( 198G-1984). The transects were
made on board Norwegian, Swedish and West-German research vessels covering the area from Northeast Greenland in the west
to Frans Josef Land in the east. The seabird observations were made along a total transect length of 26,700 km and are grouped
into 10-minute watches. Because the transects had no fixed width, absolute densities of birds per area unit could not be obtained.
A computer programme was made to generate distribution and abundance maps for each seabird species. The abundance was
given as mean number of birds recorded in each block (0.2 degrees north-south, l degree east-west) where registrations were
made. The abundances werc grouped into eight categories. A separate map was compiled showing the total observation time
(effort) in each block. The total number of individuals of each seabird speeies was also computed.
Three species, the Little Auk A lle alle, the Fulmar Fulmarus glacialis, and the Kittiwake Rissa tridactyla, dominated in numbers
during the transects and accounted for 86% of the total number of seabirds observed. Briinnich's Guillemot Uria lomvia accounted
for another 9% of the birds. A total of 567, 154 birds of 22 species was recorded. Distinct distribution patterns were obtained for
each species. Distribution and abundance maps are presented for the 19 most numerous speeies.
Fridtjof Mehlum, Norsk Polarinstitutt,
p.a.
Box 158, N-J330 Oslo Lufthavn, Norway.
Introduction
The seas around the Svalbard archipelago are rich
in biological resources. The large biomass of fish
and pelagic invertebrates is exploited by seabird
populations of many species. The Svalbard area
and the whole Barents Sea area are inhabited by
one of the largest and most important seabird
communities in the northern hemisphere (Nor­
derhaug et al. 1977; Croxall et al. 1984). The
living resources of the seas around Svalbard are
exposed to severai threats. Extensive fisheries
take place in these seas , especially in the Barents
Sea. Management offish stocks and offshore drill­
ing activities will probably be of main importance
for the fate of the marine ecosystems in these
arctic areas. Seabirds are important elements in
the marine ecosysterns , but their functions in arc­
tic ecosystems are not yet understood in full
detail.
Researchers at Norsk Polarinstitutt have be en
working for severai years on the mapping of the
distribution and abundance of seabirds in the
Svalbard area. Census es of breeding colonies
have been made all around the archipelago (Nor­
derhaug et al. 1977; Jepsen & Mobæk 1983;
Mehlum & Fjeld 1987). In 1980 we initiated a
programme to map the pelagic distribution of
seabirds in the seas around Svalbard. Before 1980
only a few ship-based censuses between the coast
of Northern Norway and the southern point of
Spitsbergen had been published (Voisin 1970;
Byrkjedal et al. 1976; Hansen 1978). Brown
(1984) conducted a comprehensive seabird census
during the spring of 1982 in the Greenland ,
Norwegian and western Barents Seas, applying
the same methods as used by the Canadian Wild­
life Service in Canadian waters (Brown et al.
1975). In 1980 we cooperated with the Swedish
'Ymer'-80 expedition in mapping the pelagic
distribution of seabirds (Edelstam 1981; Hjort et
al. 1987). The 'Ymer' is an icebreaker able to
cope with heavy multiyear ice. This cooperation
gave the opportunity to obtain data from seas far
to the north , which could not be reached by
smaller research vessels. Similarly, in 1984 we
cooperated with the Alfred Wegener Institut fur
Polarforschung, and were able to use the German
icebreaker 'Polarstern' in heavy ice-covered
waters in the northern parts of the Fram Strait.
The study of the pelagic distribution and abun­
dance of seabirds in these areas has two main
objectives. The first is to give a general description
of the occurrence of the different seabird species
at different times of the year throughout these
seas. The other is to try to explain the patterns
of abundance of seabirds by the presence of other
living organisms and different physical environ­
mental factors. Additionally, the data will help
to increase our knowledge of the biology and
migration of arctic seabirds.
In the present paper the summer distributi on
of different seabird species from five consecutive
years 1980-1984 is presented in the form of rela­
tive abundance maps. In this descriptive pres­
entation no thorough analytical discussion of the
distribution patterns has been made. These mat­
ters will be dealt with in future papers.
Material and methods
Cruises and areas covered
This report includes seabird observations col­
lected during the summer season 1980-1984. The
following cruises were conducted: 3 July-26 Sep­
tember 1980 with the Swedish icebreaker 'Ymer' ,
22 July-6 August 1980 with the Norwegian sealer
'Norvarg' , 28 July-31 August 198 1 , 22 July-31
August 1982 and 19 July-l September 1983 with
the Norwegian Polar Research Institute research
vessel 'Lance' , 20 July--4 August 1984 with the
West German ice-breaking research vessel
'Polarstern'. One to four observers participated
on each cruise. All of the cruises were multi­
disciplinary , but fishing activities , which usually
attract some seabird species , were never involved.
The total seabird observation time during these
cruises was 2,006 hours , corresponding to a tran­
sect length of 26,700 km.
Surveys were made in the northern Greenland
Sea and the Fram Strait , the northern and north­
western Barents Sea as well as north of Svalbard
into the Arctic Ocean. The actual are as covered
by the cruises and the total observation effort
(number of lO-minute watches in each geo­
graphical subunit, see below) are shown in Map
1. The range of the distribution maps is 200W to
500E and 74° to 83°N.
Recording methods
Seabirds at sea have be en recorded in many dif­
ferent ways. Most counts have been done from
ships, but in recent years aircraft have also been
7
used to some extent. Ships and aircraft both have
their advantages and disadvantages. To be able
to compare seabird census results from different
parts of the world, obtained by many observers,
it is important to have standardized recording
methods. In recent years most observers have
used lO-minute observation periods as recording
unit (Brown et al. 1975; see Tasker et al. 1984 for
review).
In our studies all seabirds seen from the moving
ship were recorded. The transect width was un­
limited, but in practice most birds were seen
doser than 300 m from the ship, and in a sector
180° ahead of the moving ship. The observer was
located outdoors on the deck above the steering
house , and the watches were of 2-4 hours dur­
ation. During most cruises two or more observers
were on board , and watch was usually kept around
the dock as long as the ship was moving, except
for periods with rough seas , fog , and from mid­
August onwards during the dark hours. The
observed birds were separated in to different cat­
egories of age , behaviour, and sex (if possible).
Data on weather, sea-ice and geographical posi­
tions were also induded. Navigational data were
obtained regularly from the ship's satellite navi­
gation systems, usually every hour and whenever
the ship's heading was changed. All data were
recorded on a special form and later transferred
to a computer file.
RELATIVE ABUNDANCE:
(MEAN NUHBER SEEN PER 10 MINUTES)
-
o
NONE
()
�
0.1 - 0.3
a
10.0 - 30.0
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0.3
1.0
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30.0
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1. O - 3.0
•
t10RE THAN 100.0
3.0
-
10.0
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Fig. 1. The abundance of birds is indicated by the mean number
of individuals observed per lO·minute watch within a unit block
(O.2°NjS x lOEjW). An open circle indicates no observations
(see text).
abundance patterns. Longitudes west of Green­
wich are indicated by negative values.
Some of the calculations involved need to be
described. Between fixed geographical positions
we calculated the positions for eve ry 10-minute
observation sub-interval. This was done by cal­
culating 1) the distance between two neighbouring
fixed positions , 2) the sum of all observation sub­
intervals (in minutes) when the ship was moving
between the two fixed positions , 3) the movement
per min ute of the ship, and by linear interpolation
for determining the position of each lO-minute
sub-interval.
The observation effort in each 0.2° x 1° block
Distribution and abundance maps
was then determined byfirst calculating the obser­
A special FORTRAN computer programme vation time in each block according to the pre­
(MAPPER) was developed for statistical tre at­ viously calculated movement per minute between
ment and for the drawing of abundance maps two fixed positions. Divided by 10, this gives the
for each species. For the presentation of species effort (number of lO-minute watches). A simil ar
abundance maps we have lumped the obser­ procedure was followed for calculating the abun­
vations into a num ber of blocks with each entry dance of observed birds in each block. For the
corresponding to 0.2°N/S and 1°E/W. For each purpose of this paper all age groups and behav­
block the observation effort, i.e. the number of ioural categories are combined.
lO-minute watches, and the mean number of birds
of different species seen per lO-minute period,
was calculated. These values of relative abun­
dance are represented by symbols on the distri­ Oceanography and sea-ice
bution maps by eight categories (Fig. 1). A special conditions
projection was made in order to obtain the same
size of all 0.2°N/S x 1°E/W degree blocks The oceanographic conditions in the Barents Sea
(squares). The abundance categOlies are in are dominated by the southwesterly ftow of cold
accordance with those used by Brown et al. (1975) arctic water from the Polar Basin, which meets a
in eastern Canadian waters. Circular symbols branch of the warmer North Atlantic Current
filled with black according to relative abundance ftowing northeast along the Bjørnøya Channel to
were applied for easy visual perception of the form a frontal system in the area southeast of
8
Atlantic water
- - - � Mixed water
Polar water
,
" - - ","
... ...... Polar front, varying considerably
Polar front, varying slightly
Fig. 2. Prevailing surface currents in the ocean areas around Svalbard. The 'polar fronts' indicated on the map suggest the
approximate positions on the surface of the division bctween temperate, saline Atlantic water and cold, less saline water of polar
origin.
Svalbard (Fig, 2), The Greenland Sea is char­
acterized by the large outftow of co Id arctic water
from the East Greenland Current to the west , and
the warmer West-Spitsbergen Current ftowing
northwards along the coast of Spitsbergen, The
Barents Sea is a shallow sea (most parts are less
than 300 m deep) and characterized by severai
banks like Storbanken , Sentralbanken (150200 m) and Spitsbergenbanken (50-100 m), The
Greenland Sea, however, except for a shelf
(70 km wide on the Svalbard and 200-250 km
wide on the Northeast-Greenland side) with
depths between 100 m and 500 m is a deep sea
area with depths of more than 3,000 m,
The northern parts of the study area are usually
covered by sea-ice, Most of the ice in the Barents
Sea is forrned locally, but there is also a variable
inftux of multiyear ice transported from the Arctic
Ocean through the passage between Svalbard and
Frans Josef Land, Winter ice may als o be
imported from the Kara Sea, Normally the ice in
the Barents Sea melts during the summer, but in
some years part of it survives and intermingies
with the new ice forrned during the following
winter. Most of the sea-ice in the Greenland Sea
is multiyear ice transported by the strong East
Greenland lee Drift Stream from the Arctic
Ocean, However, large quantities of ice are
forrned locally during the cold season. Measure­
ments indicate that 15-40% of a given area may
9
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be covered by new ice (Vinje & Finnekåsa 1986) .
The southward extent of the sea-ice varies con­
siderably both seasonally and interannually. Mini­
mum ice coverage is normally observed in August.
In Map 2 the August southern sea-ice limits for
each of the five years 1980-1984 are presented.
A detailed account of sea-ice conditions in this
region is given by Vinje (1985) and Vinje &
Finnekåsa (1986) .
As can be se en from Map 2 the area infiuenced
by the West-Spitsbergen Current was generally
free of ice during the summer. In fact it is normally
free of ice all year round because of the warm
northbound currents (Vinje & Finnekåsa 1986) .
The ice-border in the Greenland Sea was also
rather stable from year to year, while the southern
ice-limit in the Barents Sea was much more vari­
able between each of the five years.
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Results
A total of 22 species of seabirds, including two species of waterfowl (Common Eider Somateria
mollissima and Long-tailed Duck Clangula hyemalis), was observed during the cruises. A total of
567,154 birds was recorded. The species are listed in Table 1 , ranked according to their relative
abundance. The Little Auk Alle alle, the Fulmar Fulmarus glacialis, and the Kittiwake Rissa tridactyla
accounted for 86% of the total number of birds seen. A fourth species , Briinnich's Guillemot Uria
lomvia, accounted for another 9%.
In the following section distribution maps are given for the 19 most abundant species. A survey of
species diversity is presented in Map 23.
Table l. Summary table of all seabirds seen during the cruises 1980-1984.
Mean
Species
Little Auk Alle alle
Fulmar Fulmarus glacialis
Kittiwake Rissa tridactyla
Briinnich's Guillemot Uria lomuia
Ivory Gull Pagophila eburnea
Black Guillemot Cepphus grylle
Glaucous Gull Larus hyperboreus
Ross' Gull Rhodostethia rosea
Pomarine Skua Stercorarius pomarinus
Arctic Tern Sterna paradisaea
Puffin Fratereula arctiea
Long-tailed Skua Stercorarius longicaudus
Common Eider Somateria mollissima
Arctic Skua Stercorarius parasiticus
Common Guillemot Uria aalge
Great Skua Stercorarius skua
Long-tailed Duck Clangula hyemalis
Sabine's Gull Larus sabini
Great Black-backed Gull Larus marinus
Storm Petrel Hydrobatidae indet.
Gannet Sula bassana
Manx Shearwater Puffinus puffinus
Tot. no.
no· /km
184,364
163,534
138,486
51,857
14,500
4,088
3,877
1,949
1,304
1,139
866
553
206
172
171
26
26
16
15
3
1
1
6.9
6.1
5.2
1.9
0.5
0.2
0.1
<0.1
<0.1
<0.1
<0.1
<0.1
<0.1
<0.1
<0.1
<0.1
<0.1
<0.1
<0.1
<0.1
<0.1
<0.1
Mean
no· / lO-min
15.3
13.6
11.5
4.3
1.2
0.3
0.3
0.2
0.1
<0.1
'<0.1
<0.1
<0.1
<0.1
<0.1
<0.1
<0.1
<0.1
<0.1
<0.1
<0.1
<0.1
13
Fulmar Fulmarus glacialis
Breeding distribution.
The Fulmar is the on ly procellariiform species breeding within the area
covered by the censuses. The breeding colonies are widespread over most of Svalbard , including
Bjørnøya (Norderhaug et al. 1977) . It also breeds in Frans Josef Land (Golovkin 1984) , and some
small colonies are known from Northeast Greenland (Pedersen 1942; Evans 1984) . In Svalbard the
Fulmar nests in bird cliffs along the coast as well as 011 inland nunataks (Løvenskiold 1964) , and is
se en ne ar the colonies in early spring. Egg laying starts in the second half of May (Rahn et al. 1984) ,
while hatching occurs in July.
-
Pelagic distribution.
The map shows that the species is abundant in most of the pelagic regions
covered. It is the most uniformly distributed species involved in this study. The lowest abundance
was registered in the ice-covered areas off Northeast Greenland and north of Frans Josef Land.
However, it was one of the most abundant species in and around the Nordøstvandet polynya along
the coast of Northeast Greenland (together with the Ivory Gull Pagophila eburnea).
The Fulmar is known to be attracted by ships and may follow them for hours. The observers were
aware of this problem and tried to avoid repetitive counts of followers during the lO-minute counts.
Fulmars and other procellariiforms are known to have long foraging ranges. No exact data exist on
the location of foraging areas for the Svalbard nesting birds. The Fulmar is a surface feeder and in
these waters has a varied diet composed of squid, fish , pelagic crustaceans and polychaetes (Mehlum
& Gjertz 1984; Gjertz et al . 1985 ) .
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15
Black Guillemot Cepphus grylle
Breeding distribution. The Black Guillemot is a common breeder all along the coasts of Svalbard ,
usually in single pairs or in small colonies. It seems to be most abundant along the northern coasts.
The species is also a common breeder in Frans Josef Land (Golovkin 1984) but is rather scarce in
Northeast Greenland (Evans 1984) . During the breeding season it is known to stay doser to the shores
than the other auk species. It is also the only Atlantic alcid species with a dutch of two. This means
that it must have ready access to an abundant food supply dose to the breeding areas. During the
breeding season the Black Guillemot, unlike other Atlantic alcids , mainly prey upon bottom-dwelling
fishes and invertebrates in shallow waters (Cramp 1985) . In Svalbard the eggs are normally laid in
late June or early July, and the young appear on the water in late August (Løvenskiold 1964) .
-
Pelagic distribution. The distribution map shows that this species is mainly confined to ice-covered
waters, and that it is fairly abundant not only in coastal waters but als o far out at sea . The highest
relative abundances were recorded east of Kong Karls Land , at 79°N. Most individuals observed
pelagically are thought to be non-breeders and immatures. The majority of the individuals collected
in the Barents Sea (Mehlum & Gjertz 1984; Gjertz et al. 1985) were moulting birds, indicating that
the pack-ice zone , at least in the northern Barents Sea, is an important moulting area for the Black
Guillemot.
The Black Guillemot is also known to inhabit the pack-ice zone in other geographical areas , and
can be found in areas with open water in the Arctic all year round.
Food sample analyses have shown that the birds in the pack-ice prey mainly on arctic cod and
different crustaceans associated with the subsurface of the sea-ice (Mehlum & Gjertz 1984; Gjertz et
al. 1985) .
-
16
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Puffin Fratereula arctiea
Breeding distribution. Within the study area this species breeds mainly on the western and northern
coasts of Svalbard and on Bjørnøya, while it is absent from Frans Josef Land and Northeast Greenland
(Golovkin 1984; Evans 1984) . Except for the Common Guillemot Uria aalge this is the least abundant
auk species in the Svalbard area (exeluding Bjørnøya) , and no large breeding colonies like those
known from regions farther south (like Røst of the Lofoten Islands , Northern Norway) are found.
One reason for this may be the lack of suitable nesting places. In more temperate regions nest burrows
are dug out in grassy slopes. The perrnafrost in high arctic areas prevents the Puffins from digging
out nesting burrows, so they have to nest in appropriate crevices in eliffs and scree for nesting. Few
details are known of the bre eding biology of Svalbard Puffins .
-
Pelagic distribution. The pelagic distribution map refiects the limited distribution o f breeding colonies
in the area. Most of the transects were made within the breeding period of the Puffin. In Svalbard
Puffin chicks normally leave the colonies in late August or in September (Løvenskiold 1964) . Assuming
that part of the pelagically observed birds were adults on feeding trips for their young, the results
show that the travel distance between the nest and the feeding area may exceed 100 km off northwest
Spitsbergen.
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18
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Little Auk Alle alle
Breeding distribution. This typical high arctic species is known to bre ed in very large numbers in
Svalbard, especially along the western coast of Spitsbergen (Løvenskiold 1964) . It is also an abundant
species in Frans Josef Land (Golovkin 1984) , while no large colonies are known in Northeast
Greenland north of Scoresby Sound (Evans 1984) . In Svalbard the species is mainly associated with
screes made up by 'Hecla Hoek' rocks. These types of rock are confined to the western parts of
Svalbard. The Little Auk is less numerous in the eastern parts of the archipelago , but is abundant at
Hopen, �jørnøya and Tusenøyane. The latter area is a group of fiat islands where the Little Auk
nests among boulders . The breeding colonies are occupied later than the colonies of cliff nesting auk
species, since the screes normally become snow-free later in the season . Egg-laying usually starts in
late June or early July , while the young be gin to leave the colonies in mid-August (Stempniewicz
1980) .
-
As shown on the distribution map , the eastern parts of the Greenland Sea are
of major importance for this species. There is an almost abrupt disappearance of the species as one
moves to the northwestern parts of the Greenland Sea. Also important are the regions of the Barents
Sea between Kong Karls Land in eastern Svalbard, and Frans Josef Land. It is unclear whether the
pelagic concentrations of Little Auks in these eastern regions include birds belonging to the nesting
populations in the territories of the Soviet Union (Frans Josef Land and Novaj a Zemlj a) , or if they
are Svalbard nesting birds. This problem can be investigated by biometrical studies. Most birds
observed in this area are probably non-breeding immatures.
It is known from ringing recoveries that some Little Auks breeding on the western coast of
Spitsbergen are wintering southwest of Greenland (Norderhaug 1967; Anker-Nilssen & Jensen 1981) .
The total wintering area of the Spitsbergen Little Auks i s not known , but probably most of the birds
winter along the ice-edge from southwest of Greenland to the central Greenland Sea. The observations
by Brown (1984) of Little Auks along the ice-edge in the central Greenland Sea in March 1982 also
support this. The winter quarters of eastern Svalbard birds are unknown .
Large nesting colonies of Little Auks are found in the Hornsund area, Spitsbergen. That part of
the Greenland Sea which is off Hornsund, was not covered by the censuses . We assurne that high
abundances would also be recorded in these waters.
Pelagic distribution.
20
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Briinnich's Guillemot Uria lomuia
Breeding distribution.
This cliff-nesting auk is very numerous in the Svalbard area. The total
population is probably in the order of one million pairs (Mehlum unpublished) . The largest breeding
colonies are concentrated on Bjørnøya, Hopen, in the Storfjorden and Hinlopenstretet areas, and on
the western coast of Spitsbergen (Norderhaug et al. 1977) . It is also very numerous on Novaj a Zemlja
(one million pairs) and less abundant in Frans Josef Land (100,000 pairs) (Golovkin 1984) . It is typical
of this species, as well as of the Common Guillemot Uria aalge and the Razorbill Alca torda, that the
young leave their nesting ledges before they are able to fly. The chick is accompanied by a parent
when it starts its swimming migration from its breeding place to the open sea towards the winter
quarters. Comprehensive ringing has demonstrated that the waters southwest of Greenland are an
important wintering area for Briinnich's Guillemots breeding in Western Spitsbergen (Norderhaug et
al. 1977) . Nothing is known about the winter quarters of Briinnich's Guillemots breeding in other
parts of Svalbard . The species arrives in the early spring at the breeding ledges (in Hornsund 1985 on
3 April, C. Lydersen & I. Gjertz pers. comm . ) . The main egg-laying period is in June (Rahn et al.
1984) , and the chicks dep art from the colonies in the first half of August.
-
The pelagic distribution map reflects the breeding distribution around both
Svalbard and Frans Josef Land. The species is observed further to the west in the Greenland Sea than
the Puffin Fratereula arctiea, which may indicate a longer foraging radius during breeding. But many
of these birds could als o be non-breeders , or birds which have started their SW migration early.
The species was found both in open and ice-covered waters. Studies on feeding ecology (Mehlum
& Gjertz 1984; Gjertz et al. 1985 ; Lydersen et al. 1985 ; Mehlum unpublished) have shown that in the
Svalbard area it feeds mainly on the pelagic amphipod Parathemisto libellula and arctic cod Boreogadus
Pelagic distribution.
saida.
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SUMMERS
Common Guillemot Uria aalge
This species breeds in large numbers on Bjørnøya (Franeker & Luttick 1981),
but only about two hundred pairs are documented as breeders farther north in Svalbard (Norderhaug
1974; Kempf & Sittler 1988). The largest known colony in Svalbard except for Bjørnøya is the one
at Fuglehuken , the northernmost point of the large island Prins Karls Forland off the west coast of
Spitsbergen. Scattered pairs have been observed elsewhere on the western coast of Spitsbergen . It
may also breed at Hopen, but this still has to be documented. The Common Guillemot is not reported
breeding in Frans Josef Land (Golovkin 1984) or in Northeast Greenland (Evans 1984).
Breeding distribution.
-
As se en on the distribution map , the Com mon Guillemot is only documented
to have been seen in are as dose to Bjørnøya. The Common Guillemot overlaps in breeding distribution
with the more northerly distributed Briinnich's Guillemot Uria lomvia. Because these two species are
difficult to distinguish at sea, some Common Guillemots may have be en overlooked and incorrectly
identified as the more abundant Briinnich's Guillemot. The existing data indicate that most Common
Guillemots forage in the are as to the south of 75°N during late summer. In contrast, the Briinnich's
Guillemot was observed in abundance in the area between Bjørnøya , Hopen and Storfjorden (Map
Pelagic distribution.
7).
24
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Map 8. Common GuiLLemot
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25
Kittiwake Rissa tridacyla
Breeding distribution. - The Kittiwake belongs to the group of the most abundant breeding seabird
species in the area covered. Breeding colonies are distributed on cliffs all along the coasts of the
Svalbard archipelago . It is also common in Frans Josef Land (Golovkin 1984) but very scarce or
absent in Northeast Greenland north of Scoresbysund (Hjort et al. 1983; Evans 1984). In Svalbard
the Kittiwakes occupy their nesting cliffs in April , and egg-laying usually starts in early June . One or
twa eggs are laid in each clutch , and the young leave the nests in August. Immature birds are often
seen foraging in the fiords and other coastal are as during summer (Mehlum unpublished) .
Pelagic distribution. The general pattern obtained by the surveys is a wide distribution and high
relative abundance over most of the study area. However, a distinctly reduced abundance was
observed in the northwest Greenland Sea, comparable to the distribution pattern of the Little Auk
Alle alle and Briinnich's Guillemot Uria lomvia. Kittiwakes are attracted to ships, and this has to be
considered when comparing the relative abundance maps with maps of other species which are not
attracted. Their abundance may have been somewhat overestimated. Additionally, some of the birds
following the ship for severai lO-minute periods may have been recorded severai times.
The chicks of the Kittiwake usually stay in their nests until mid-August (Løvenskiold 1964). Time­
budgets at Kongsfjorden, Spitsbergen (Mehlum unpublished; Gabrielsen & Mehlum 1989) and at
Hopen (Gabrielsen et al. 1987) show that the adults may have very long shifts of attending the nest.
The normal pattern was 12-24 hours between each time the parents changed guarding the chicks at
the nest. The birds can fly considerable distances during that many hours. The high abundance
recorded in the central part of the Greenland Sea may suggest that feeding ranges can be more than
200 km for individuals nesting on the western coast of Spitsbergen. However, these birds could also
be non-breeders.
The Kittiwake was found both in open waters and in ice-covered areas, except in most of the
northwestern part of the Greenland Sea. Stomach analyses (Mehlum & Gjertz 1984; Gjertz et al.
1985) and direct observations show that Kittiwakes mainly eat arctic cod Boreogadus saida in the ice­
covered waters, but that they also prey on various crustaceans (mainly amphipods) . Farther south ,
e.g. around Hopen, they mainly depend on capelin Mallotus villosus in the summer (Gabrielsen et
al. 1987).
-
26
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Map 9. Kittiwake
27
Ivory Gull Pagophila eburnea
Breeding distribution. This gull is generally confined to High Arctic breeding places. In Svalbard it
is known to breed in scattered colonies at some nunataks on Spitsbergen, but most of the known
breeding colonies are in the easternmost part of the archipelago . In this area, where the characteristic
pointed nunataks of Spitsbergen are missing, the Ivory Gull breeds in small numbers on bird diffs
or, occasionally, on flat ground. The species is fairly abundant as a breeding bird in Frans Josef Land
(P. S. Tomkowich pers. comm. ) and it probably breeds in small numbers in most of High Arctic
Greenland (Evans 1984) . Very little is known about the breeding sch edule of this species in Svalbard
(Løvenskiold 1964) .
-
Pelagic distribution. The pelagic distribution map shows dearly the Ivory Gull's dose association
with sea-ice . Its main distribution is north of 79°N. The map als o shows the high relative abundance
of the species , even if the calculated values may be influenced by its habit of being attracted to ships.
The Ivory Gull is the bird which is recorded in highest abundance in waters off NE Greenland , on
the coast of which severai breeding colonies are known (Håkansson et al. 1981; Hjort et al. 1983) .
The extreme northern pelagic distribution may be determined by the species' feeding habits. Our
studies (Mehlum & Gjertz 1984; Gjertz et al. 1985) confirm earlier studies of the species' varied diet.
It feeds on arctic cod and different crustaceans associated with the sub-surface of the sea-ice . Their
food is caught either by plunging into the water or pecking food items from the ice surface . Another
important food source is the remains of seal carcasses and blubber left by polar bears.
-
28
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l1ap 10. Ivory Gull
29
Ross' Gull Rhodostethia rosea
Breeding distribution. Outside the breeding are as very little is known about Ross' Gull. The main
breeding groups are located in the tundra and swampy northern taiga lowlands between the Jana and
Kolyma rivers in Eastern Siberia (Andreev & Kondratev 1981). Isolated nesting are as have also been
found in Tajmyr (Pavlov & Dorogov 1978) and in the Caun Lowland (Andreev & Kondratev 1981).
Single breeding records are reported from Svalbard (Løvenskiold 1964), Northern Greenland (Kampp
& Kristensen 1980; Hjort 1980) and arctic Canada (MacDonald 1978; Chartier & Cooke 1980).
-
SeveraI old records of Ross' Gulls exist from ice-covered areas north of Svalbard
and Frans Josef Land . Ross (Parry 1828), Nansen (Collett & Nansen 1900), and Andree (L6nnberg
1931) observed severaI individuals during the summer months July and August. The importance of
these are as to Ross' Gull was confirmed by ornithologists on the 'Ymer'-expedition in 1980 (Meltofte
et al. 1981).
Additional observations of Ross' Gulls have been made on cruises conducted since the 'Ymer'­
expedition , but the distribution pattern is similar to that reported by Meltofte et al. (1981). The data
obtained during the 'Ymer'-expedition are included in the distribution map presented here . Ross'
Gull was encountered in the ice-covered areas north of Nordaustlandet, Svalbard , south of Victoria
Island and north of Frans Josef Land. West of Svalbard the species was widely distributed in the
central and northwestern parts of the Greenland Sea, even if the relative abundance in each block
was low. In most squares in the Greenland Sea area the species was recorded in abundance category
0. 1-0.3 individuals per lO-minute period, while high er abundances were recorded north and east of
Svalbard. Ross' Gull, like the Ivory Gull , was restricted to ice-filled waters in the northern part of
the study area.
Pelagic distribution.
30
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Sabine's Gull Larus sabini
Breeding distribution. Sabine's Gull is mainly a Low Arctic species and has an almost circumpolar
breeding range , but with large gaps along the Eurasian coast of the Arctic Ocean. An account of its
breeding distribution is given by Blomqvist & Elander (1981) and Abraham ( 1986) . It breeds in single
pairs or in small groups , almost exclusively in colonies of Arctic Terns Sterna paradisaea. It is a rare
bird in the Svalbard area and only a few records of nesting have been reported (Løvenskiold 1964) .
However, observations indicate that Sabine's Gull may have nested at several localities in Svalbard
in recent years (Norsk Polarinstitutt data base , unpublished) . A few breeding localities are also known
in High-Arctic Greenland (Blomqvist & Elander 1981; Hjort et al . 1983; Håkansson et al. 1988) .
-
As shown on the pelagic distribution map, Sabine's Gull is a very uncommon
seabird in the waters between Northwest Greenland and Frans Josef Land. The observations are
scattered all over the study area, except for the southernmost parts southeast of Svalbard . A total of
16 birds was recorded in 11 different squares during the five years.
Pelagic distribution.
32
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33
'
Glaucous Gull Larus hyperboreus
The Glaucous Gull is a common breeding species all over Svalbard, ineluding
Bjørnøya. It is also widespread in Northeast Greenland (Evans 1984) and occurs in Frans Josef Land.
Its status in the latter area is unknown, however (Golovkin 1984) . This species nests in association
with typical bird-eliff species, and with island colonies of eiders and geese . It preys both on the eggs
and chicks of other bird species and utilizes a wide variety of marine food items. It will often also
feed on garbage around human dwellings (Larsen 1965; Mehlum & Gjertz 1984; Lydersen et al. 1985).
Breeding distribution .
-
Pelagic distribution. The species is widely distributed in the pelagic regions studied. The highest
abundance was recorded elose to the western coast of Spitsbergen. Farther west in the Greenland
Sea, the numbers deelined, and it was almost absent from the waters off Northeast Greenland . The
Glaucous Gull was distributed both in ice-covered waters and in are as with open waters (such as in
the northwestern Barents Sea and along the western coast of Spitsbergen) .
-
34
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Great Black-backed Gull Larus marin us
Breeding distribution. This speeies now breeds in single pairs at several localities along the western
coast of Spitsbergen, but the total population probably amounts to less than 50 pairs. It has gradually
expanded and increased in numbers during the last three decades (Norderhaug 1983). It also breeds
at Bjørnøya in small numbers (Franeker & Luttick 1981), and is abundant along the coasts of the
southern parts of the Barents Sea (Cramp & Simmons 1983). The Great Black-backed Gull is not
known to breed in Northeast Greenland (Evans 1984) and Frans Josef Land (Golovkin 1984).
-
The distribution of observations of the Great Black-backed Gull during the
censuses was very seattered, and the abundance very low. A total of only 15 individuals was recorded .
All records were from east o f 15°E . The results indicate that the speeies may b e encountered
fortuitously all over the Barents Sea area.
Pelagic distribution.
36
-
(f)
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Map 14 . Great B lack-backed Gu l l
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37
Arctic Tern Sterna paradisaea
Breeding distribution. The Arctic Tern has a circumpolar breeding distribution, and is a common
breeding species all over Svalbard , induding the easternmost and northernmost islands (Løvenskiold
1964) . It breeds on fiat tundra along the coast and on small coastal islands . The species is also
widespread along the coasts of Northeast Greenland (Evans 1984) and breeds in Frans Josef Land
(Golovkin 1984) . The Arctic Tern usually arrives in Svalbard by the end of May or early June , and
leaves the breeding area in late August or September (Løvenskiold 1964) .
-
Pelagic distribution. The Arctic Tern was the 10th most numerous species in the censuses . As the
distribution map shows , this species is almost missing in the northernmost parts of the area concerned .
I t was fairly common dose t o the shores , but was also recorded i n mid-ocean regions i n the Barents
and Greenland Seas . In the Greenland Sea it seems to be associated with the marginal ice-zone . Some
observations were also made off the coast of Northeast Greenland , probably of birds breeding along
the adjacent coast.
-
38
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Arctic Skua Stercorarius parasiticus
Breeding distribution. - The Arctic Skua is known as a solitary breeding species in most coastal are as
in the Svalbard archipelago , but with its lowest abundance in the northeastern parts, where it nests
on the tundra (Løvenskiold 1964) . It is reported to be absent as a breeder in Northeast Greenland
north of 76°N (Evans 1984) . It is a scarce breeder in Frans losef Land (Demente'ev & Gladkov 195 1 ) .
Pelagic distribution. - The Arctic Skua was less abundant i n the investigated area than both the
Pomarine and Long-tailed Skuas Stercorarius pomarinus and S. longicaudus, despite the fact that it is
the only abundant breeding skua species in Svalbard. The observations were scattered over large parts
of the study area, but the species was lacking in the northwestern parts of the Greenland Sea. Except
for records in two blocks north of Frans losef Land it was als o absent from are as east of 40oE . This
indicates that the breeding population in Svalbard probably feeds mainly in coastal areas during the
months luly-September.
40
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Long-tailed Skua Stercorarius longicaudus
Breeding distribution. The Long-tailed Skua breeds in Svalbard in very low numbers and only a few
nests have been found along the western coast of Spitsbergen (Løvenskiold 1964). It also breeds over
most of the northern and eastern parts of Greenland (Evans 1984), but it is not recorded as a breeding
bird in Frans Josef Land. It is scarce in Novaja Zemlja and on the Murmansk coast but more abundant
in the inland tundra zones in northern Fennoscandia and the USSR (Cramp & Simmons 1983).
-
The distribution map of this species distinguishes between two important
geographical areas of abundance . One stretches southeastwards from the northeastern coast of
Greenland to the middle of the Greenland Sea, the other is in the central to northwestern parts of
the Barents Sea. This distribution pattern is probably influenced by the presence of migratory birds,
the Greenland Sea birds probably originating from Greenland and migrating southwards to the
Atlantic. The Barents Sea birds may be individuals originating from arctic USSR and migrating
westwards through the Barents Sea, before turning southwards into the Atlantic. However, it cannot
be ruled out that the Barents Sea birds belong to the same general population as the Greenland Sea
birds, and that a large part of them are indeed non-breeders spending the summer in pelagic areas
(cf. the Pomarine Skua Stercorarius pomarinus) .
Pelagic distribution.
42
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Map 1 7 . Long-tai led Skua
43
Pomarine Skua Stercorarius pomarinus
Breeding distribution. The Pomarine Skua is not a common breeding bird in Svalbard, and the only
breeding record is from Tusenøyane in 1984 (Løvenskiold 1964; Bentz 1986). The closest known
breeding localities are in the northern parts of Novaja Zemlja (Cramp & Simmons 1983). It do es not
breed in' either Northeast Greenland or Frans Josef Land, but it do es breed in most arctic parts of
the USSR and North America bordering the Arctic Ocean (Muus et al . 1981; Cramp & Simmons
-
1983).
Pelagic distribution. Even though this species is not known to bre ed regularly within the study area,
it was the most abundant of all the skua species recorded during the censuses. The largest con­
centrations of birds were encountered in the northwestern part of the Barents Sea (southeast of
Edgeøya and Hopen) . Another concentration was recorded along the ice-edge north of Spitsbergen.
The species was also recorded in abundance in the waters between Nordaustlandet, Kvitøya and Kong
Karls Land.
The individuals observed are probably non-breeding and migratory birds from the USSR, which
utilize the rich food resources of the Barents Sea and the marginal ice-zone before migrating
southwards through the northern Atlantic to the wintering areas (Demente'ev & Gladkov 1951;
Løvenskiold 1964). September is given as the main departure period from the breeding range
(Demente'ev & Gladkov 1951). They were often seen chasing Kittiwakes Rissa tridactyla and stealing
their food.
-
44
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Map 1 8 . Pomarine Skua
�
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45
Great Skua Stercorarius skua
Breeding distribution. Probably less than 10 pairs of Great Skuas breed yearly along the western
coasts of Spitsbergen and on the island of Hopen. On Bjørnøya the breeding population has apparently
increased sharply since the species' establishment around 1970, to about 20 pairs in 1980 (Vader 1980;
Franeker & Luttick 1981). The first reported breeding in Svalbard (except for Bjørnøya) was in 1976
(Larsen 1977) . No breeding localities of Great Skuas outside Svalbard are known north of Iceland
and Northern Norway. Recoveries of ringed individuals prove an immigration from British colonies
(Vader 1980; R. Barrett pers. comm. ; Mehlum unpublished) .
-
Pelagic distribution. Our observations of Great Skuas were confined to the northwestern part of the
Barents Sea and waters west and north of Spitsbergen (north to about 81°N) . During the years of
immaturity the north Atlantic subspecies of the Great Skua is known to have a wide marine range,
extending north to the Greenland Sea, especially birds in their third year (Cramp & Simmons 1983) .
Most of the individuals observed are probably immatures and non-breeders. Many non-breeding
individuals are also regularly seen along the Svalbard coasts during summer, and up to 100 birds may
be observed at Bjørnøya (Vader 1980) .
-
46
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Map 1 9 . Grea t Skua
47
Common Eider Somateria mollissima
The Common Eider breeds abundantly on small islands and in coastal areas
all along the coasts of Svalbard. The largest breeding concentrations are found on small islands along
the western coast of Spitsbergen (Løvenskiold 1964) . It also breeds in East Greenland north to
Nordostrundingen (81°N) (Muus et al. 1981 ; Hjort et al. 1983 ; Håkansson et al. 1988) and in Frans
Josef Land (Cramp & Simmons 1983) . The Eiders arrive in Svalbard waters in the spring as soon as
there are coastal areas with open water. They are usually waiting at the ice-edge outside the fiords
and coasts until their nesting islands are free of snow and surrounding sea-ice . Breeding normally
starts in June.
Breeding distribution.
-
Pelagic distribution. The distribution map do es not reftect the true distribution patterns of the Eider
in this region. Most Eiders gather in shallow water dose to the shores after the breeding season before
leaving Svalbard in the late autumn (Løvenskiold 1964; Karlsen & Mehlum 1986) . The map shows
that the speeies can also be encountered in small numbers farther out at sea, but no observations
were recorded in mid-ocean areas (no records west of 9°E and east of 300E) .
Only four recoveries of ringed Svalbard Eiders exist, of which one is from Iceland and the other
three from the coasts of Northern Norway (Mehlum unpublished) . The main wintering area is probably
the coast of Northern Norway (Røv & Parker unpublished) .
-
48
'"
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Map 20 . Common Eider
49
Long-tailed Duck Clangula hyemalis
Breeding distribution. The Long-tailed Duck breeds in small numbers on the coastal tundra all over
Svalbard except in the northeastern parts (Løvenskiold 1964) . It is also present as a breeder in
Northeast and North Greenland (Muus et al. 198 1 ) , but not in Frans Josef Land (Cramp & Simmons
1977) .
-
Pelagic distribution. This speeies is known to be more pelagic than the Eider Somateria mollissima
outside the breeding season, and it winters in northern waters (Cramp & Simmons 1977) . The pelagic
habits are reflected in the distribution map given for this speeies. The total number of individuals
observed was only 26, recorded in four different blocks. Three of these blocks were in the northern
pack-ice zone . The low number of birds observed indicates that the speeies is not common in the
waters investigated. Even though the total number of birds obsenied was low , the distribution pattern
was quite different from the other duck speeies induded in this paper, the Eider, which was confined
to areas doser to the coasts of Svalbard.
-
50
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Map 21 . Long-tai led Duck
51
Map 22. Summa ry map of all speeies.
52
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53
Discussion
The census data from the five years 1980-1984
are combined in the distribution maps presented
in this paper. This procedure will mask differences
of abundance and distribution between years.
However, the main purpose of the paper is to
present the average distribution patterns for the
five studied summer seasons in order to identify
pelagic areas generally important to seabirds dur­
ing the months July-September. Distribution
maps for each of the five years are not presented
since only parts of the total area were censused
each year, and because the census effort in each
block was limited.
As seen from the distribution maps seabirds
are abundant in most parts of the study area.
An abundance map of all species combined is
presented in Map 22. Large concentrations were
recorded in the Greenland Sea off the western
coast of Spitsbergen, in the waters north of Spits­
bergen north to the pack-ice in the Arctic Ocean,
in the northernmost part of the Barents Sea
between Svalbard and Frans Josef Land, and in
the northwestern part of the Barents Sea towards
the southeastern parts of Svalbard. Although the
highest numbers of seabirds were recorded in
these waters, the individual seabird species
showed distinct distribution patterns. These dif­
ferences in distribution may be infiuenced by the
location of breeding colonies, migration patterns
and feeding habits. The number of species
observed in each block (l°N/S and 5°E/W ) is
shown in Map 23 . The rather low abundances in
the western part of the Fram Strait are also
refiected in a generally low species diversity in
this area. An exception is, to a certain degree,
the Nordøstvandet polynya south of Nord­
ostrundingen, Greenland's easternmost point.
Three species, the Black Guillemot, Ross' Gull,
and the Ivory Gull , were closely associated with
ice-filled waters in the northern parts of the study
area. This pattern is in agreement with reports
from other regions (Brown & Nettleship 1981;
Blomqvist & Elander 1981). The abundance of
Black Guillemots more than 200 km from the
nearest shore, however, is very different from the
near shore distribution of this species in more
temperate regions (Cramp 1985 ) . Their pelagic
distribution in arctic areas is probably related to
the availability of crustaceans and fish associated
with the sea-ice.
54
Our seabird observations may be biased in sev­
eral ways, so the maps have to be viewed with
some caution. First of all some species are more
easily overlooked than others. This group
includes the a\cids, with their dark plurnage which
is not easily detected on the sea surface, especially
in rough sea. They also dive for food and may not
be present on the sea surface when the ship is
passing by. Other species, like fulmars and gulls
(except for Ross' Gull) , are attracted by ships and
may follow them for hours. There is always the
possibility that individuals of these species may
be recorded severai times, and during more than
one lO-minute watch. Differences in weather con­
ditions, such as vi sibilitY wind and waves, may
also infiuence the detectability of the birds. The
ship probably attracts relative ly more birds when
it moves slowly through more or less heavy ice
than when it is in open water, and this will arti­
ficially increase the figures from the ice-covered
areas as compared with those from ice-free
waters.
In the maps presented here the abundance cate­
gories in each block may be infiuenced by the
types of bi as mentioned above . In addition , the
results in some blocks may not be representative
because of low observation effort (small number
of 10-minute watches) .
It has to be emphasized that in this study the
pelagic abundances of seabirds are presented as
relative values (i. e . the mean num ber of indi­
viduals observed per 10-minute watch) . The tran­
sect width was unlimited, and therefore absolute
numbers of seabirds seen per square unit could
not be computed. In other recent studies a fixed
transect width, usually 300 m, has be en applied
(see Tasker et al. 1984 for review) . The capacity
of shipboard surveys to provide reasonable values
of the absolute sizes of seabird populations at sea
is questioned by Brown (1986, p. 3). He argues
that 'most shipboard observations are made in
the course of limited transects that cover only a
very small fraction of a marine area, and do not
even cover all the habitats within it' . I agree with
Brown that given the limitations of the capacity
of the methods applied during the censuses, the
goal should be to refine the measurements of
relative variations in numbers of seabirds. These
relative abundances of different species may then
be compared with each other and interpreted in
terms of the occurrence of physical and biological
oceanographic phenomena.
,
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