Atlantic Puffin Fratercula arctica
Status; International importance;
Population
estimates; Distribution; Annual abundance/
productivity; Phenology/diet/survival

Description
The following was adapted from original text by Mike P.
Harris and Sarah Wanless in
Seabird Populations of Britain and Ireland (with
permission from A&C Black, London).
The Atlantic puffin is the most instantly recognisable and
popular of all North Atlantic seabirds. It breeds in the North
Atlantic and the adjacent Arctic Ocean, with strongholds in Iceland
and Norway, with around 10% of the population breeding around the
British Isles, where it is the second most abundant breeding
seabird. Atlantic puffins are pelagic and we are still largely
ignorant of where they spend their time away from the colony. Those
from north western Britain disperse widely outside the breeding
season, as far as Newfoundland in the west and the Canary Islands
to the south and even into the Mediterranean as far east as Italy.
In contrast, most of those from colonies in eastern Britain
remain within the North Sea though in recent decades some have
dispersed as far as the Bay of Biscay.
Atlantic puffins typically nest underground in burrows dug in
the soil of offshore islands, but where such habitat is
sparse they nest among boulder screes or at low densities in
cracks in sheer cliffs. The species is highly colonial and most
colonies occur where the nesting birds are safe from mammalian
predators. However, during the breeding season a colony can appear
deserted during the middle of the day since most birds are either
down burrows or out at sea feeding. At other times awe-inspiring
numbers can be seen standing on the slopes, bobbing around on the
sea or flying in vast wheels over the colony. Chicks are fed on
small fish that the adult carries cross-wise in its beak. In the
UK the commonest prey is the lesser sandeel, followed by
sprat, herring and a wide range of small juvenile gadoid fish. Fish
are caught by underwater pursuit, usually several at a
time.
Conservation status
Atlantic puffin is currently identified as a conservation
priority in the following:
Amber listed in Birds
of Conservation Concern 3 (2009 update)
(further information on Conservation Designations for UK Taxa)
Amber listed in
Birds of Conservation Concern in Ireland 2 (2007
update)
International importance
*AOB = Apparently Occupied Burrows
The UK population figure (rounded to the nearest hundred) was
derived from data in Mitchell, P.I., Newton, S.F.,
Ratcliffe, N. and
Dunn, T.E. (eds.) 2004. Seabird Populations of Britain
and Ireland. Poyser, London.
This was also the source of figures for the Biogeographic and World
populations.
UK population estimates and change 1969-2002 (census
data)
During Seabird 2000 and the SCR Census surveyors were able
to count apparently occupied burrows (AOB) at most colonies. Here
the main source of error is the misclassification or overlooking of
burrows. However, for logistical reasons complete coverage is
impossible to achieve at many colonies. In such situations the
density of burrows must be determined in sample plots, the area of
the colony estimated and the measures combined to get an estimated
population size. Where birds nest under boulders, in mixed colonies
with Manx shearwaters, in completely inaccessible places or at low
densities along stretches of cliffs (mainland colonies), counts of
burrows are impractical. In these cases surveyors must count
individual birds attending the colony. Such counts can be highly
variable both within and between days, and the lack of any obvious
factor influencing attendance, means that such counts are of rather
limited value in assessing breeding numbers, but they do at least
give some idea of colony size. April and May are the prime months
for counting individual birds before substantial numbers of
immatures begin to attend colonies.
For the calculation of total populations, some arbitrary
decisions had to be made to allow the combination of counts of
individuals and AOBs. The practice of assuming that one
individual corresponded to one AOB was applied to counts from
all three national censuses. This approximation may well result in
a serious underestimate of the number of AOBs. However, the overall
estimates of the SCR Census and Seabird 2000 should be broadly
comparable. In both surveys 83% of the total population
estimates came from counts or estimates of AOBs. Furthermore, in
the SCR Census 65% of the counts of individual birds came from the
preferred counting months compared to 73% during Seabird
2000.
| |
Operation Seafarer
(1969-70)
|
Seabird Colony Register
(1985-88)
|
Seabird 2000
(1998-2002)
|
| UK Population estimate (AOB*) |
424,318 |
488,763 |
580,714 |
| % change since previous census |
N/a |
+15 |
+19 |
*AOB = Apparently Occupied Burrows
For census results for individual countries and Ireland, the
Channel Islands and the Isle of Man see under relevant
sections below.
Distribution/abundance
The Seabird
2000 census provides the most comprehensive recent
assessment of the distribution and abundance of breeding seabirds.
Numbers of Atlantic puffin found in different regions,
and a map showing where colonies are found and how large they are
is provided in the Seabird 2000 Atlantic
puffin results page (PDF, 2.0 mb).
An
interactive map is available on the NBN Gateway, where you can
filter to display only the Seabird 2000 data. For more
recent, but less comprehensive, coverage view the distribution on
the NBN with all available contributing datasets.
The locations sampled during the annual
Seabird Monitoring Programme provide some information on
distribution and are accessible via the Seabird Monitoring Programme
online database.
Annual abundance and productivity by geographical area
Breeding abundance
The logistical difficulties in monitoring Atlantic puffin
colonies means that few data are collected annually and that which
is collected tends to be from the smaller colonies; these are
usually counts of individual adult birds in attendance at
breeding sites. For a variety of factors counts of individuals
can vary quite markedly between years compared with counts of
apparently occupied burrows (AOB). The UK Atlantic puffin
population increased until at least Seabird 2000
(for unknown reasons), and possibly beyond, as counts of AOBs from
two of the largest colonies (Farne Islands in NE England and Isle
of May in SE Scotland) even greater numbers in 2003. However, a
substantial decline at these two colonies was then recorded between
2003 and 2008/9. It is not known whether these decreases are
representative of the UK as a whole. The return rate of
adult puffins breeding on the Isle of May was very low in 2007
and 2008 (see under ‘Return rates and survival rates’), which
explains, at least in part, the population decrease at this colony.
The reasons for low return rate in these years are unknown.
Productivity

Figure 1: Trend in UK productivity (no. of
chicks fledged per pair) of Atlantic puffin 1986-2010.
Based on SMP data; view the methods
of analysis (PDF 158 kb).
Productivity has fluctuated throughout
the recording period (Figure 1) but appears to have been generally
lower in the years since the late 1990s contributing to a declining
trend overall. Productivity was at its lowest in 2007 due to
food shortage plus unprecedented rainfall flooding burrows; these
two factors combined also resulted in the low productivity values
recorded in 1998 and 2004.
Population estimates and change 1969-2002 (census data)
| |
Operation Seafarer
(1969-70)
|
Seabird Colony Register
(1985-88)
|
Seabird 2000
(1998-2002)
|
| Population estimate (AOB*) |
410,011 |
438,101 |
493,042 |
| % change since previous
census |
N/a |
+7 |
+13 |
*AOB = Apparently Occupied Burrows
Breeding abundance

Figure 1: Abundance of Atlantic
puffin at three colonies in Scotland, 1986-2010.
The logistical difficulties in monitoring Atlantic puffin
colonies means that few data are collected annually. National
census data indicate that the number of Atlantic puffins in
Scotland had increased since Operation Seafarer in 1969/70 by 20%
up to Seabird 2000. Figure 1 (above) shows changes at the largest
colonies between the Seabird Colony Register and Seabird 2000.
However, of the three colonies shown above only the Isle of May has
been surveyed since that time; here numbers continued to climb
before a decline was recorded between 2003 and 2009. The
return rate of adult puffins breeding on the Isle of May was
very low in 2007 and 2008 (see under ‘Return rates and survival
rates’), which explains, at least in part, the population decrease
at this colony. The reasons for low return rate in these years are
unknown.
Productivity

Figure 2: Trend in productivity (no. of
chicks fledged per pair) of Atlantic puffin in Scotland,
1986-2010. Based on SMP data; view the methods
of analysis (PDF 158 kb).
Productivity has fluctuated throughout
the recording period but appears to have been generally lower
in the years since the late 1990s, contributing to a declining
trend overall. Productivity was at its lowest in 2007 due to
food shortage and unprecedented rainfall flooding burrows;
these two factors also resulted in the low productivity values
recorded in 1998 and 2004.
Population estimates and change 1969-2002 (census data)
| |
Operation Seafarer
(1969-70)
|
Seabird Colony Register
(1985-88)
|
Seabird 2000
(1998-2002)
|
| Population estimate (AOB*) |
8,616 |
36,868 |
75,734 |
| % change since previous
census |
N/a |
+328 |
+105 |
*AOB = Apparently Occupied Burrows
Breeding abundance

Figure 1: Abundance of Atlantic
puffin on Coquet Island, 1986-2010.
Atlantic puffin numbers at colonies in England increased
since Operation Seafarer when 8,616 AOB were counted. The Seabird
Colony Register recorded four times this number and by Seabird
2000 numbers had doubled again. The largest colonies are both in
the north-east on the offshore islands of the Farnes and
Coquet. The number of AOB on Coquet decreased by
approximately one-third immediately after Seabird 2000 but had
recovered by 2008. On the Farne Islands, a far larger colony than
Coquet (so monitored less frequently) a 24% decline was
recorded between 2003 (55,674 AOB) and 2008 (36,835
AOB).
Productivity

Figure 2: Trend in productivity (no. of
chicks fledged per pair) of Atlantic puffin in England,
1986-2010. Based on SMP data; view the methods
of analysis (PDF 158 kb).
The productivity of Atlantic puffins at
English colonies has been very variable over time. Overall, there
is no clear trend. Food shortage plus unprecedented rainfall
flooding burrows resulted in low productivity in
1998 and 2007.
Population estimates and change 1969-2002 (census data)
| |
Operation Seafarer
(1969-70)
|
Seabird Colony Register
(1985-88)
|
Seabird 2000
(1998-2002)
|
| Population estimate (AOB*) |
4,255 |
11,116 |
10,328 |
| % change since previous
census |
N/a |
+161 |
-7 |
*AOB = Apparently Occupied Burrows
Breeding abundance

Figure 1: Abundance of Atlantic
puffin on Skomer based on counts of individuals attending
the colony in spring, 1986-2010.
The number of Atlantic puffins in Wales increased by
161% between Operation Seafarer and the Seabird Colony
Register, with little change thereafter up to Seabird 2000. Few
Welsh colonies of any size are monitored on an annual basis.
However, on Skomer, there are regular counts undertaken of
adults attending the colony in spring. For a variety of
factors these counts can vary quite markedly between years
compared with counts of apparently occupied burrows
(AOB). However, the trend in the Skomer
population appears to be upward in general with no hint of the
declines recorded at some of the large Scottish and English
colonies.
Productivity

Figure 2: Trend in productivity (no. of
chicks fledged per pair) of Atlantic puffin in Wales,
1986-2010. Based on SMP data; view the methods
of analysis (PDF 158 kb).
Compared with data collected in Scotland and England
the productivity of Atlantic puffins in Wales is less
variable. There appear to be no
years where productivity has been badly affected
by factors such as extreme weather events or food
shortages.
Population estimates and change 1969-2002 (census data)
| |
Operation Seafarer
(1969-70)
|
Seabird Colony Register
(1985-88)
|
Seabird 2000
(1998-2002)
|
| Population estimate (AOB*) |
1,436 |
2,678 |
1,610 |
| % change since previous
census |
N/a |
+86 |
-40 |
*AOB = Apparently Occupied Burrows
Breeding abundance
In Northern Ireland, Atlantic puffin numbers increased by 86%
between Operation Seafarer and the Seabird Colony Register but then
fell by 40% by the time of Seabird 2000.
Counts carried out on Rathlin Island, which holds
some 98% of the national population, indicate a further
large decline has occurred since Seabird 2000; 1,579 AOB were
recorded in 1999 but only 731 AOB in 2007.
Productivity
No systematic data on productivity have been collected for
Atlantic puffins in Northern Ireland as part of the SMP.
Population estimates and change 1969-2002 (census data)
| |
Operation Seafarer
(1969-70)
|
Seabird Colony Register
(1985-88)
|
Seabird 2000
(1998-2002)
|
| Population estimate (AOB*) |
26,553 |
17,435 |
19,641 |
| % change since previous
census |
N/a |
-34 |
+13 |
*AOB = Apparently Occupied Burrows
Breeding abundance
The number of Atlantic puffins in the Republic of Ireland
declined by 34% between Operation Seafarer and the Seabird Colony
Register but then increased by 13% by the time
of Seabird 2000. The status of the species in the country since
then is largely unknown. On Great Skellig a count of individuals in
early July 2010 recorded 2,170 compared to 4,000 at the same time
of year in 1999. The difference in these figures may not be
indicative of a decline as counts at this time of year can be even
more variable than counts in spring due to a wide variety of
factors e.g. adult attendance at the colony, differences between
breeding seasons.
Productivity
No systematic data on productivity have been collected for
Atlantic puffins in the Republic of Ireland as part of the SMP.
Population estimates and change 1969-2002 (census data)
| |
Operation Seafarer
(1969-70)
|
Seabird Colony Register
(1985-88)
|
Seabird 2000
(1998-2002)
|
| Population estimate (AOB*) |
27,989 |
20,113 |
21,251 |
| % change since previous
census |
N/a |
-28 |
+6 |
*AOB = Apparently Occupied Burrows
Breeding abundance
Overall, numbers of Atlantic puffins throughout Ireland fell by
28% between Operation Seafarer and the Seabird Colony Register with
little further change recorded during Seabird 2000. Few sizeable
colonies are monitored in the region with any frequency so data
since the last national census is sparse. Please refer to the
entries for the two individual countries.
Productivity
No systematic productivity data have been collected on
Atlantic puffins throughout Ireland as part of the SMP.
Population estimates and change 1969-2002 (census data)
| |
Operation Seafarer
(1969-70)
|
Seabird Colony Register
(1985-88)
|
Seabird 2000
(1998-2002)
|
| Population estimate (AOB*) |
82 |
93 |
85 |
| % change since previous
census |
N/a |
+13 |
-9 |
*AOB = Apparently Occupied Burrows
Breeding abundance
The Isle of Man has had a small but fairly stable population of
Atlantic puffins since Operation Seafarer in 1969/70. Numbers have
ranged from 82-93 pairs over the three censuses. There are no more
recent counts to indicate the species' current
status.
Productivity
No systematic data on the productivity of Atlantic puffins on
the Isle of Man have been collected as part of the SMP.
Population estimates and change 1969-2002 (census data)
| |
Operation Seafarer
(1969-70)
|
Seabird Colony Register
(1985-88)
|
Seabird 2000
(1998-2002)
|
| Population estimate (AOB*) |
1,116 |
335 |
311 |
| % change since previous
census |
N/a |
-70 |
-7 |
*AOB = Apparently Occupied Burrows
Breeding abundance
Atlantic puffins nesting on the Channel Islands have
declined greatly since Operation Seafarer recorded over 1,100
pairs. Numbers during the Seabird Colony Register and Seabird 2000
were similar having fallen by 70% prior to the 1985-88 census. No
data on the status of the species have been received since Seabird
2000.
Productivity
Few systematic data on the productivity of Atlantic
puffins on the Channel Islands have been collected as part of the
SMP.
UK phenology, diet, survival rates
Phenology
No systematic data on phenology (timing of life-cycle events)
have been collected as part of the SMP.
Diet
Our understanding of puffin diet comes from data collected
at a few geographically dispersed key sites. Because diet is
likely to vary from site to site, and given few sites are
monitored , caution should used in drawing wider
geographical conclusions from these data.
Sandeels (Ammodytes spp.), an energy-rich shoaling
fish, comprise a large proportion of the diet of puffins
at two sites where diet is studied. The proportion of
sandeels fluctuated; at Fair Isle in Shetland (Figure 4) it
declined over the monitoring period (especially the 'large'
sizeclass), though on the Isle of May in SE Scotland (Figure 3)
there was no clear trend. However, the size and energy
content of sandeels caught by and available to puffins on the
Isle of May declined over the period 1973-20021, and in some years (such as 2004)
the energy content of sandeels was found to be far lower than
would be expected for their size and coincided with very poor
breeding success for puffins and other sandeel-feeders2. The mean mass of food loads of
puffins breeding at Fair Isle declined from the mid 1990s (Figure
5) and may have contributed to low breeding success during those
years. The causes for the change in the size of sandeels around the
Isle of May are probably due to climate
change rather than from the effects of
fishing.
Between 2004 and 2008, snake pipefish Entelurus
aequoreus appeared in the diets of many seabirds around the
UK3 -including puffins
during 2006-2007. The energy content of this bony fish, which
before the early 2000s was very rare in UK waters, is
very low4. Snake
pipefish, therefore did not provide an alternative energy
source during a time when puffin's usual prey
species (sandeels and sprats) were scare. Indeed, pipefish can
choke chicks when fed to them in quantity. The reason for the
appearance of snake pipefish is uncertain, but is not
thought to be related to climate change.

Figure 3. Percentage of sandeels (by weight) in
the diet of young puffins at the Isle of May (SE
Scotland), 1987-2010.

Figure 4. Composition of fish loads brought to
puffin chicks at Fair Isle (Shetland) 1986-2010.

Figure 5. Mean mass (g) of fish loads brought
to puffin chicks at Fair Isle (Shetland), 1986-2010.
Return rates and survival rates
Important notes on interpretation: Estimation
of Atlantic puffin adult return rate and survival
rate is currently only undertaken at two sites within the
Seabird Monitoring Programme -The Isle of May in the Firth of
Forth, SE Scotland and Skomer Island, Pembrokeshire, SW
Wales. Return rates are based on sightings of individually
colour-ringed birds and are calculated as the proportion of marked
birds present in year one that is seen in the
following year. Because not every adult alive is seen each
year, return rates for 2010 presented here for Isle of May
need to be treated as minimum estimates of survival of birds seen
in 2009. In contrast, survival estimates - as presented here
for Skomer - do take into account birds that are not seen one
year but which re-appear in following years.
Return rate of Atlantic puffins breeding on the Isle of May
(Figure 6) has shown no consistent trend, though declined between
the mid 1990s and the mid 2000s; however, it fell to extremely low
levels in 2007 and 2008, which was the likely cause of a population
decline on the island noted between 2003 and 2008. Atlantic puffins
breeding on Skomer (Figure 7) show a significant downward trend in
survival, but without the sharp declines in 2007 and 2008 that were
seen on the Isle of May. Little is currently known of the causes of
changes in survival rates of puffins, though recent evidence
suggests a shift in overwintering range of Isle of May breeders
from the North Seas into the east Atlantic, possibly reflecting
deteriorating conditions in the North Sea5.

Figure 6. Annual return rate of Atlantic puffin
breeding on the Isle of May (SE Scotland), 1987-2010.

Figure 7. Annual survival rate of Atlantic
puffin breeding on Skomer (SW Wales), 1987-2009.
References
1 Wanless, S., Wright, P. J., Harris, M. P.
& Elston, D. A. 2004. Evidence for decrease in size of lesser
sandeels Ammodytes marinus in a North Sea aggregation over
a 30-yr period. Mar. Ecol. Prog. Ser. 279: 237-246.
2 Wanless, S., Harris, M. P., Redman, P. &
Speakman, J. R. 2005. Low energy values of fish as a probable cause
of a major seabird breeding failure in the North Sea. Mar.
Ecol. Prog. Ser. 294: 1-8.
3 Harris, M. P., Beare, D., Toresen, R., Nøttestad, L.,
Kloppmann, M., Dörner, H., Peach, K., Rushton, D. R. A.,
Foster-Smith, J. & Wanless, S. (2007). A major increase in
snake pipefish (Entelurus aequoreus) in northern European
seas since 2003: potential implications for seabird breeding
success. Marine Biology 151:973-983.
4 Harris, M.P., Newell, M., Daunt, F.,
Speakman, J., & Wanless, S. (2007). Snake pipefish
Entelurus aequoreus are poor food for seabirds.
Ibis, 150, 413-415.
5 Harris, M.P., Daunt, F., Newell,
M., Phillips, R.A. and Wanless, S. 2009. Wintering
areas of adult Atlantic puffins Fratercula arctica from a North Sea
colony as revealed by geolocation technology.
Marine
Biology
DOI 10.1007/s00227-009-1365-0
Partners
Data have been provided to the SMP by the generous contributions
of its partners, other organisations and volunteers throughout
Britain and Ireland. Partners to the SMP are: BirdWatch Ireland;
The British Trust for Ornithology; Centre for Ecology and
Hydrology; Countryside Council for Wales; Department of
Agriculture, Fisheries and Forestry (Isle of Man); Department of
Environment, Heritage and Local Government (Republic of Ireland);
States of Guernsey Government; JNCC; Manx Birdlife; Manx National
Heritage; The National Trust; National Trust for Scotland; Natural
England; Northern Ireland Environment Agency; The Royal Society for
the Protection of Birds; Scottish Natural Heritage; Seabird Group;
Shetland Oil Terminal Environmental Advisory Group; Scottish
Wildlife Trust. More about the SMP partners >>
Puffin image appears courtesy of Ian Rendall ©, is subject to
international copyright law and may not be reproduced in any form
whatsoever.