Present Status and Trends in Population of Seabirds in Norway

by

Einar Brun[74]

University of Tromsø
Tromsø, Norway

Abstract

The most numerous seabird in Norway is the puffin (Fratercula arctica), but its current breeding population of 1.25 million pairs is slowly declining. The kittiwake (Rissa tridactyla), however, is increasing and establishing new colonies; its population now stands at 510,000 pairs. The population of the common murre (Uria aalge), the seabird species most vulnerable to human activity, was about 160,000 breeding pairs in 1964 but is now decreasing at a rate of nearly 5% per year. Of the other alcids, the razorbill (Alca torda) and thick-billed murre (Uria lomvia) show similar declines, and the black guillemot (Cepphus grylle) is maintaining a stable population. The fulmar (Fulmarus glacialis) and the gannet (Sula bassana) have both spread from the British Isles and have established a number of breeding colonies in Norway during this century. Evidently immigration of gannets is still occurring, since the observed rate of increase far exceeds the population's intrinsic rate of increase. The impact of human activity on bird mortality varies from species to species. The two most serious factors are coastal oil pollution and the use of fishing gear; direct hunting pressure accelerates the decline of murres and razorbills. Persistent toxic chemicals are not yet a serious problem in Norway.

Norway, with a coastline of more than 20,000 km, an abundance of islands, and areas of offshore upwelling, provides good conditions for a rich seabird fauna. A regional study of this seabird fauna has been undertaken as a sideline of basic marine research. Although the ultimate aim has been to evaluate the importance of seabirds in the energy flow of a marine ecosystem, a more realistic problem (given priority so far) has been to study yearly production and the dynamics behind changes in the breeding populations.

Good population estimates are of fundamental importance to studies of population dynamics. Because the available censuses of seabirds in Norway were few and largely inadequate, a long-term program was started in 1961. In the beginning, resources and assistance were very limited, and the work was concentrated on cliff-breeding seabirds, particularly the gannet (Sula bassana), fulmar (Fulmarus glacialis), kittiwake (Rissa tridactyla), razorbill (Alca torda), common murre (Uria aalge), thick-billed murre (U. lomvia), and puffin (Fratercula arctica). Until 1970, the study involved making annual censuses in the approximately 20 major colonies of cliff-breeding seabirds and mapping the distribution of the quantitatively less important colonies.

Since 1970, the Norwegian seabird program has also involved more detailed studies in some selected colonies. In these colonies, emphasis has been on investigation of yearly production and of the factors limiting this production, and evaluation of the effects of human activity on the population growth.

Material and Methods

The logistics of census operations have gradually improved from the use of slow, local transportation to the use of fast pneumatic boats and, in more recent years, seaplanes. Various census methods have been used, depending on species and circumstances.

For puffins, a method based on measurement of feeding frequency and on the number of puffins per time unit that pass a particular observation post when they return from the feeding ground was used (Brun 1971a). Kittiwakes and gannets were readily censused by a combination of photographic methods and detailed counts in sample areas (Brun 1971b). Direct counting is by far the most accurate method for razorbills, murres, and fulmars; but in the larger colonies of common murre, lack of time permitted accurate counts for only a limited proportion of the cliff. Direct counts of individuals, the egg/chick ratio, and estimates of the relative size of the censused population were used to estimate the total population of the colony.

In a colony of kittiwakes near Tromsø, environmental factors that limit breeding success, such as temperature and wind exposure, were monitored throughout the breeding season on a data recorder, and detailed measurements of temperatures on and inside the eggs have been recorded. For further information about the influence of environmental parameters on incubation rhythm and nest attendance, the presence of the male and female at a particular nest was recorded by using radioactive bands and a Geiger-Muller tube connected to a pen recorder.

In a study of the effects of human activity, egg samples of selected species were analyzed for mercury, PCB, and DDT derivates. An effort was also made to obtain figures for the mortality caused by oil pollution and fishing gear as well as by direct hunting pressure.

Results

Status and Trends of Cliff-breeding Species
Puffin (Fratercula arctica)

By far the most numerous seabird in Norway is the puffin (Fig. 1), which is the only species with a breeding population of more than 1 million breeding pairs (Tables 1, 2). In a 1964 census (Brun 1966) the total breeding population was put at 1.5 million pairs. The current figure of 1.25 million pairs includes several newly discovered colonies and some not censused in 1964; it is more accurate than the previous census for most of the 15 largest colonies which make up 99.9% of the total population. The puffin population is concentrated in Troms and Nordland (94%), with only about 3% in Finnmark.

Fig. 1. Distribution of the puffin (Fratercula arctica) in Norway. Numbers refer to localities listed in Table 2.

Kittiwake (Rissa tridactyla)

The second most numerous seabird species in Norway is the kittiwake, which dominates in a number of the larger cliff colonies. Its distribution pattern differs from that of the puffin—the main occurrence of the kittiwake population (about 63%) is in Finnmark (Table 3).

Table 1. Estimate of the numbers of seabirds breeding on the coast of Norway 1970-1974. Species are listed in descending order of breeding population size.
Species Thousands of pairs[75] Increase (+) or decline (-)
Fratercula arctica 1250 -
Rissa tridactyla 510 +
Larus argentatus (260)[76] +
L. canus (150)[76] +
Uria aalge 100 -
L. marinus (40)[76] +
Phalacrocorax aristotelis 33 +
Alca torda 30 -
Cepphus grylle 22 0
Sterna paradisaea (21)[76] -
S. hirundo (13)[76] -
Phalacrocorax carbo 12 +
L. fuscus 9[76] +
Stercorarius parasiticus (8) 0
L. ridibundus 4[76] +
Fulmarus glacialis 1.1 +
U. lomvia 1.0 -
Hydrobates pelagicus ? ?
Sula bassana 0.76 +
Oceanodroma leucorrhoa ? ?

The annual production of kittiwakes shows enormous variation, both throughout the coastline and in different years; however, at our sample stations in north Norway, the mean production in 1974 (Table 4) was more stable and was near the minimum value necessary to maintain zero population growth.

This minimum production, mx (number of females produced per breeding female), can be computed from survival rates

mx = (1-P)/1x = 0.13/0.57 = 0.23

where P is annual adult survival and 1x is survival of fledged chicks up to first breeding. Data on survival are taken from Coulson and White (1959) and from Norwegian banding recoveries.

The kittiwake has, however, established a number of new colonies, and although the local increase in some of these is spectacular, the long-term change during the last 15 years is only about 1% increase per year in northern Norwegian colonies (E. Brun, unpublished data). In southern Norway, the population has increased much more rapidly (Brun 1971c).

Common murre (Uria aalge)

The common murre (Fig. 2) has shown a considerable decrease. The most spectacular decrease is at Sør-Fugløy, where a colony of 10,000 pairs in 1940 was reduced to 4,000 pairs in 1961, to 1,100 pairs in 1966, and to only about 10 breeding pairs in 1974 (Table 5). Most of the census work was done in 1964 and 1974. The general trend in population change, as expressed by the yearly decrease or increase, has been extrapolated forward to 1974 or back to 1964 for those colonies where censuses were missing for either of these years, to enable a better comparison (Table 6). The overall decrease in Norwegian colonies of the common murre is, thus, near 5% per year; the few cases with a positive trend are based either on very small figures or on extrapolation from old, inadequate censuses.

Thick-billed Murre (Uria lomvia)

The thick-billed murre (Fig. 3) was first proved to breed in Norwegian colonies in 1964; it was then found at three localities and has since been found breeding at eight localities (Table 7). It is now fairly certain that the thick-billed murre is not a newcomer but has remained unnoticed among the common murre for generations, possibly since the original immigration of the Uria species after the last glacial period. Data are not sufficient to show whether this small population of thick-billed murres is decreasing at the same rate as the common murre.

Table 2. Status of the puffin (Fratercula arctica) in Norway (cf. Fig. 1).
Locality Year of census Number of pairs Percent of population
1. Kjør 1975 80 <0.1
2. Heglane 1970 4 <0.1
3. Ferkingstadøyene 1970 5 <0.1
4. Utsira 1970 2 <0.1
5. Utvær 1970 200 <0.1
6. Ryggsteinen 1970 2 <0.1
7. Veststeinen 1970 1,500 0.1
8. Einevarden 1970 1,500 0.1
9. Svinøy 1970 100 <0.1
10. Runde 1974 30,000 2.4
11. Saløy 1970 2 0.1
12. Sklinna 1974 2,000 0.2
13. Lovunden 1968 60,000 4.8
14. Fugløy i Gildeskål 1968 800 0.1
15. Røst 1964 700,000 55.7
16. Værøy 1974 70,000 5.6
17. Nykvåg 1967 40,000 3.2
18. Frugga 1975 5,000 0.4
19. Anda 1970 10,000 0.8
20. Bleik 1968 40,000 3.2
21. Sør-Fugløy 1968 40,000 3.2
22. Nord-Fugløy 1967 218,000 17.3
23. Loppa 1968 180 <0.1
24. Hjelmsøy 1964 20,000 1.6
25. Gjesværstappen 1973 18,000 1.4
26. Kongsøy 1966 30 <0.1
27. Syltefjord 1966 100 <0.1
28. Hornøy 1967 160 <0.1
29. Reinøy 1967 40 <0.1
Total 1,257,705 100.0
Razorbill (Alca torda)

Another colonial cliff-breeding alcid, the razorbill (Fig. 4), has a distribution pattern very similar to that of the common murre, but the individual colonies (Table 8) are, with one exception, smaller. The total breeding population was estimated at 36,000 pairs in 1966-69 (Brun 1969b); some more recent censuses show a definite decline, but data are not sufficient to estimate the overall decline in the Norwegian population. At most, the current breeding population is 30,000 pairs.

Fulmar (Fulmarus glacialis)

The fulmar is one of two species of seabirds that have spread from colonies in the British Isles and established themselves as breeding birds in Norway during this century (the other is the gannet).

Fig. 2. Distribution of the common murre (Uria aalge) in Norway. Numbers refer to localities listed in Tables 5 and 6.

Fig. 3. Distribution of the thick-billed murre (Uria lomvia) in Norway. Numbers refer to localities listed in Table 7.

Table 3. Status of the kittiwake (Rissa tridactyla) in Norway, and a comparison of distribution with that of the puffin (Fratercula arctica).
County Breeding pairs
Rissa Fratercula
Number
(thousands)
Percent Number
(thousands)
Percent
Finnmark 321 62.9 38 3.0
Troms 9 1.8 258 20.5
Nordland 72 14.1 928 73.7
Trøndelag (S, N) 1 0.2 2 0.2
Møre and Romsdal 105 20.6 30 2.4
Sogn and Fjordane 1.9 0.4 3 0.2
Rogaland 0.1 <0.1
Total 510 100.0 1,259 100.0

The fulmar began nesting in the early 1920's on Runde, the only sizeable seabird colony in south Norway, off Alesund. Further immigration of birds from the British Isles probably occurred in the first 25 years, when the population increased about 10% annually to about 350 pairs in 1947 (Valeur 1947). Since then the population increase has slowed down to about 3% annually, and the population on Runde in 1971 was about 700 pairs (Table 9). From Runde, fulmars have spread not only to a number of islands in the same region, but also much farther afield—south to Utsira (59°18'N, 4°55'E) and north to Bleik (69°3'N, 15°42'E). The total Norwegian population of fulmars in 1971 was estimated at 1,100 pairs.

Fig. 4. Distribution of the razorbill (Alta torda) in Norway. Numbers refer to localities listed in Table 8.

Fig. 5. Distribution of the gannet (Sula bassana) in Norway. Numbers refer to localities listed in Table 10.

Table 4. Annual production, mx, of the kittiwake (Rissa tridactyla) at some North Norwegian coastal localities (mx = number of females produced per female).
Locality Year Sample size (number) mx
Vedøy, Røst 1972 852 0.21
Hekkingen, Troms 1974 264 0.46
Hjelmsøy 1974 357 0.18
Jarfjord 1974 146 0.31
Total 1,619 0.25
Gannet (Sula bassana)

The gannet (Fig. 5), the most recently established and least numerous of the cliff-breeding seabirds, has the best-known population change. Like the fulmar, it was established in 1946 on Runde, and the first individuals were undoubtedly of British origin. During its entire breeding history on Runde, and also in two of the three new colonies in northern Norway established in the 1960's, the yearly increase has far exceeded the intrinsic rate of increase (Table 10); for gannets with a 50% breeding success, adult mortality of 6%, and 35% survival up to first breeding, the intrinsic rate of increase is about 2% per year. The Runde and Syltefjord colonies are naturally protected by their inaccessibility, but the colonies at Mosken and Nordmjele, which are on small islets, are both easily accessible. The Nordmjele colony, however, has been effectively protected from its start, whereas the Mosken colony has been open to visitors; this difference is probably reflected in their different breeding success and annual growth rate (Table 11). The breeding success necessary to maintain a stable population with the mortality figures given above is 34%:

mx = (1-P)/1x = 0.66/0.35 = 0.17

For equal sex ratio, breeding success is 2 times mx = 0.34.

A British ringed gannet from Ailsa Craig (55°12'N, 5°07'W) was found nesting when 4 years old in the Nordmjele colony in 1970 (Brun 1972), giving direct evidence that immigration from colonies in Great Britain (Scotland) still takes place.

Table 5. Status of the common murre (Uria aalge) in Norway (cf. Fig. 2).
Locality Last census Previous census Reference
Year No. of breeding pairs Year No. of breeding pairs
1. Utsira 1970 1 1950 10 Holgersen 1951
2. Utvær 1970 17 1948 55 Willgohs 1952
3. Veststeinen 1970 29 1950 40 Willgohs 1952
4. Klovningen 1970 35 1950 20 Willgohs 1952
5. Einevarden 1970 30 1952 25 Willgohs 1955
6. Runde 1974 6,000 1963 7,600 Brun 1969a
7. Storholmen 1970 8
8. Røst 1974 6,800 1964 9,700 Brun 1969a
9. Værøy 1974 1,750 1964 2,400 Brun 1969a
10. Nykvåg 1974 350 1966 430 Brun 1969a
11. Bleik 1974 60 1952 90 Regnell 1957
1964 75 Brun 1969a
12. Sør-Fugløy 1974 10 1940 10,000 Soot-Ryen 1941
1961 4,000 Brun 1963
1966 1,100 Brun 1969a
13. Nord-Fugløy 1967 9,000 1963 15,000 Lütken 1965
14. Loppa 1974 500 1966 800 Brun 1969a
15. Hjelmsøy 1974 70,000 1964 110,000 Brun 1965
1967 95,000 Brun 1969a
16. Gjesværstappene 1973 580 1967 750 Brun 1969a
17. Sværholtklubben 1973 20 1966 25 Brun 1969a
18. Omgangsstauran 1973 70 1967 85 Brun 1969a
19. Syltefjorden 1974 9,000 1966 12,300 Brun 1969a
20. Hornøy 1974 500 1964 730 Brun 1969a
21. Reinøy 1974 110 1964 160 Brun 1969a
22. Kjøfjord 1970 21
23. Skogerøy 1970 8 1967 6 Brun 1969a
24. Sagfjord 1970 9 1967 12 Brun 1969a
25. Kobbholmfjorden 1970 2 1967 1 Brun 1969a

Estimates of Total Seabird Population in Norway

In addition to the more detailed censuses of the cliff-breeding species dealt with so far, notes have been made on all seabirds observed during numerous flights along the Norwegian coast. Although a first attempt at putting a figure to all seabird species in Norway may be somewhat premature, it is believed that even an extrapolation combined with an educated guess is of some value until more accurate censuses covering the whole coast can be made. Although the data (Table 1) are arranged in the same way as the results from "Operation Seafarer" in the British Isles (Cramp et al. 1974), it must be stressed that the accuracy of the Norwegian figures, at least for the non-cliff-breeding birds, is far inferior to the very fine British data. The table includes data for two petrels (Hydrobates pelagicus, Oceanodroma leucorrhoa), which in Norway breed on Røst (well north of the Arctic Circle), where they have adapted to a delayed breeding season with egg laying in August because of the conflict of their nocturnal habits with the continuous daylight due to the midnight sun. Of the present population trends that are given for each species in Table 1, all auks except the black guillemot (Cepphus grylle) are decreasing, whereas the gulls, the gannets, and the fulmars are increasing.

Table 6. Population trends in colonies of the common murre (Uria aalge) in Norway. Numbers for 1964 and 1974 are, when not censused those years, extrapolated from present trends, using estimated yearly decrease or increase from all available census figures.
Locality Number of breeding pairs[77] Percentage yearly decrease (-) or increase (+)
1964 1974
1. Utsira 2 1 -12.2
2. Utvær 23 14 -5.5
3. Veststeinen 32 27 -1.6
4. Klovningen 30 39 +2.8
5. Einevarden 28 31 +1.0
6. Runde 7,438 6,000 -2.2
7. Storholmen 9 7 (-2.2)[78],[79]
8. Røst 9,700 6,800 -3.6
9. Værøy 2,400 1,750 -3.2
10. Nykvåg 453 350 -2.6
11. Bleik 75 60 -2.3
12. Sør-Fugløy 1,844 10 -68.5
13. Nord-Fugløy 13,201 3,681 -13.6
14. Loppa 900 500 -6.1
15. Hjelmsøy 110,000 70,000 -4.6
16. Gjesværstappen 853 556 -4.4
17. Sværholtklubben 27 19 -3.2
18. Omgangsstauran 94 68 -3.3
19. Syltefjorden 13,299 9,000 -4.0
20. Hornøy 730 500 -3.9
21. Reinøy 160 110 -3.8
22. Kjøfjord 21 20 (-0.4)[78],[79]
23. Skogerøy 5 12 +10.1
24. Sagfjord 16 6 -10.1
25. Kobbholmfjord 1 5 +26.0
Total 161,341 99,566 -4.9
Table 7. Status of the thick-billed murre (Uria lomvia) in Norway (cf. Fig. 3).
Locality Year No. of breeding pairs Percentage of total Uria population
1. Vedøy, Røst 1974 15 0.3
2. Værøy 1966 20 0.9
3. Hjelmsøy 1974 850 1.2
4. Gjesværstappene 1973 25 4.3
5. Syltefjord 1970 90 0.9
6. Hornøy 1966 55 8.1
7. Reinøy 1964 1 0.6
8. Kjøfjord 1970 1 4.8
Estimated number, Norway, 1974 >1,000 ca. 1.0
Table 8. Status of the razorbill (Alca torda) in Norway (cf. Fig. 4).
Locality Year No. of breeding pairs Percent
1. Kjør 1970 1 <0.1
2. Utsira 1970 25 0.1
3. Utvær 1970 16 0.1
4. Veststeinen 1970 22 0.1
5. Klovningen 1970 12 <0.1
6. Einevarden 1970 45 0.2
7. Runde 1974 2,800 9.5
8. Sklinna 1974 15 0.1
9. Lovunden 1968 8 <0.1
10. Røst 1974 3,900 13.2
11. Værøy 1974 800 2.7
12. Nykvåg 1966 250 0.8
13. Bleik 1968 28 0.1
14. Sør-Fugløy 1974 15 0.1
15. Nord-Fugløy 1967 10,000 33.8
16. Loppa 1969 750 2.5
17. Hjelmsøy 1974 7,000 23.7
18. Gjesvær 1973 2,500 8.5
19. Sværholtklubben 1973 18 0.1
20. Omgangsstauran 1973 6 <0.1
21. Kongsøy 1966 8 <0.1
22. Syltefjorden 1966 1,200 4.1
23. Hornøy 1967 65 0.2
24. Reinøy 1967 55 0.2
25. Kjøfjord 1970 9 <0.1
26. Skogerøy 1970 4 <0.1
27. Jarfjordnes 1970 3 <0.1
Total ca. 30,000

Since the coastline of Norway is about the same length as the coastline of Great Britain and Ireland, it is interesting to compare the population figures (Table 12), although the accuracy is very different. Populations of auks and gulls are similar in both areas, but the species composition is different. There are more terns in the British Isles, but skuas (Catharacta skua), shags (Phalacrocorax aristotelis), and great cormorants (P. carbo) are present in similar numbers. The most striking difference is the very small number of procelli-forms and gannets in Norway compared to Britain and Ireland, where they are almost as numerous as the gulls and the auks.

Table 9. Status of the fulmar (Fulmarus glacialis) in Norway.
County Number of localities Number of breeding pairs
Nordland 6 140
Møre and Romsdal 7 945
Sogn and Fjordane 2 11
Rogaland 2 2
Total 17 1,098
Table 10. Population increase of the gannet (Sula bassana) in Norway (cf. Fig. 5).
Colony Year established Mean yearly growth rate 1969-1974
(%)
No. of breeding pairs
1969 1970 1971 1972 1973 1974
1. Runde 1946 8.4 330 331 383 422 450 494
2. Mosken ca. 1960 5.4 50 83 77 60 62 65
3. Nordmjele 1967 83.3 7 36 65 103 127 145
4. Syltefjord 1961 14.5 28 29 44 48 51 55
Total 12.8 415 479 569 633 690 759
Yearly growth rate (%) 15.4 18.8 11.2 9.0 10.0

Discussion

Impact of Human Activity
Direct Exploitation

According to Norwegian laws, all seabirds, with the exception (for some odd reason) of the gannet and fulmar, can be hunted from 21 August to 1 March. However, only the two species of murre and the razorbill are still regularly hunted and, although no statistics support it, an estimate based on interviews with some of the hunters reveals that murres and razorbills are shot in the ratio of about 50:1. One man can shoot as many as 380 murres and razorbills during a winter season as a sideline to fishing. Although not many hunt on this scale, an absolute minimum of 5,000 murres and razorbills are killed this way each season.

A new law based on modern principles of conservation has been under consideration for several years, and this will mean an improvement. However, the speed of the decline of the auks, particularly the murres, makes it imperative to stop this hunting immediately, and it is of very little economic importance to the few who take part. Some illegal "fishing" for auks still takes place at Røst and Vaerøy, where fishnets are anchored over wooden frames outside the auk colonies at the beginning of the nesting season. At Vedøy on Røst in 1972, up to 80 murres were taken daily. Thus an estimated total of 500-700 murres were taken that year—about 5% of the breeding population on this island.

Egg collecting was important during World War II, but in these more affluent times and because of the relative inaccessibility of the auks' nests, egg collecting is now both less attractive and less important. Human disturbance of the breeding colonies, however, is gradually becoming a more serious factor.

Table 11. Comparison of annual growth rate and breeding success in two colonies of gannets (Sula bassana).
Mosken Nordmjele
Annual growth rate Breeding success (%) Annual growth rate Breeding success (%)
1969 62
1.66 5.14
1970 51 61
0.93 1.81
1971 36 46
0.78 1.58
1972 33 62
1.03 1.23
1973 50 35
1.05 1.14
1974 12 39
1969-1974 1.05 40 1.83 46
Fishing Gear

Although on a scale different from that in western Greenland, drift-net and longline fishing for Atlantic salmon (Salmo salar) outside the 19-km (12-mile) limit off the northern Norwegian coast present a serious mortality hazard to some seabirds. Reliable data exist only for the longline fisheries. In the 1969 season (with 75 effective days from mid-March to mid-June), one boat using 1,040 hooks per day caught 294 birds: 52 fulmars, 3 gannets, 43 kittiwakes, 107 murres, and 89 puffins. No razorbills were identified, but they may have been included in the murre figure. If this sample is representative, the 100 or so Norwegian boats using longlines plus about 20 Danish boats (which used 4,000-6,000 hooks per day and consequently caught more birds) would have caught roughly 10,000 fulmars, 600 gannets, 9,000 kittiwakes, 21,000 murres, and 18,000 puffins in the 1969 season. The drift-nets in Norwegian waters are reported to be less damaging to seabirds than are the longlines, but even without adding the figures from the drift-nets, the numbers are substantial in view of the size of the Norwegian breeding populations.