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The seabird wreck in the Bay of Biscay and South-Western Approaches in 2014: A review of reported mortality

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Between December 2013 and February 2014, a series of storm events occurred in areas of the North Atlantic frequented by migratory seabirds. Prolonged exposure to sustained storm conditions was followed by an unprecedented level of seabird mortality, apparently due to starvation, exhaustion and drowning. A total of 54,982 wrecked birds was recorded along European coastlines of the North-East Atlantic over the winter; 94% of which were dead. The majority of birds found were recorded on the French coastline (79.6%), and the most impacted species was the Atlantic Puffin Fratercula arctica (53.5%). In this paper, we describe the conditions surrounding this wreck event and report the numbers of wrecked and stranded seabirds by combining reports from multiple affected countries.
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SEABIRD 29 (2016): 22–38
22
Reported mortality in the 2013/14 NE Atlantic seabird wreck
The seabird wreck in the Bay of Biscay
and Southwest Approaches in 2014: A
review of reported mortality
Tim I. Morley1*, Annette L. Fayet2, Helene Jessop3, Paul Veron4, Merlin Veron5,
Jacquie Clark6and Matt J. Wood7
* Correspondence author. Email: t.i.m-eee09@hotmail.co.uk
1Alderney Wildlife Trust, Slades, 48 Victoria Street, Alderney, GY9 3TA,
Channel Islands;
2Department of Zoology, University of Oxford, Oxford OX1 3PS, UK;
3RSPB, Keble House, Southernay Gardens, Exeter, Devon, EX1 1NT, UK;
4Mermaid Cottage, Mannez Lighthouse, Alderney, GY9 3YJ, Channel Islands;
5Le Grand Verger, Rue Des Marais, Vale, Guernsey, GY6 8AU, Channel Islands;
6British Trust for Ornithology, The Nunnery, Thetford, IP24 2PU, UK;
7School of Natural & Social Sciences, University of Gloucestershire,
Cheltenham GL50 4AZ, UK.
Abstract
Between December 2013 and February 2014, a series of storm events occurred in
areas of the North Atlantic frequented by migratory seabirds. Prolonged exposure
to sustained storm conditions was followed by an unprecedented level of seabird
mortality, apparently due to starvation, exhaustion and drowning. A total of 54,982
wrecked birds was recorded along European coastlines of the northeast Atlantic
over the winter; 94% of which were dead. The majority of birds found were
recorded on the French coastline (79.6%), and the most impacted species was the
Atlantic Puffin Fratercula arctica (53.6%). In this paper, we describe the conditions
surrounding this wreck event and report the numbers of wrecked and stranded
seabirds by combining reports from multiple affected countries.
Introduction
Seabird wrecks occur when large numbers of dead, injured or exhausted seabirds
wash up on coastlines with no obvious cause of death (Birkhead 2014). Such
events are extremely hard to quantify because most mortality occurs at sea. Even
when birds do wash ashore they often do so in inaccessible locations, such as at
the bottom of steep cliff shorelines. Despite these challenges, it is important to
attempt to quantify the magnitude of wreck events and put them into context
with other wrecks to inform long-term studies of seabird population dynamics,
because they can have large impacts on seabird populations (Votier et al. 2005,
2008; Mesquita et al. 2015).
Between December 2013 and February 2014 a succession of extreme and persistent
weather events generated such severe conditions that a number of seabird species,
usually wintering in the open ocean, were adversely affected. Although no individual
23
SEABIRD 29 (2016): 22–38
Reported mortality in the 2013/14 NE Atlantic seabird wreck
Table 1. Known survey effort within affected areas.
Country Date Location Distance Source
United Kingdom 22–23 February Accessible coastline 1,984 km Schmitt 2014
throughout the UK
excluding the southwest
England region*
Channel Islands: Regular surveys 6.51 km of 77.18 km total Broadhurst &
Alderney from 10 February accessible Morley 2014
to 22 March coastline
Channel Islands: 22 February Accessible 96 km total C. Veron, pers. comm.
Guernsey to 8 March coastline
Channel Islands: 16–23 February, Accessible Variations in personnel G. Young &
Jersey 2–9 March coastline make total distance C. Sellares,
incalculable pers. comm.
France Six weekends from French Atlantic Surveys conducted Farque et al.
1 February to coastline across 2,773.7 km but 2014
9 March 2014 not all was accessible
Spain Some areas regular, Atlantic and Varying effort in C. Torrell,
others intermittent northern Spanish different locations pers. comm.
coastlines makes distance
incalculable
Portugal 14–27 March 12.7 km of accessible 22.7 km total T. van Nus,
coastline between pers comm.
Sao Jacinto and Torreira,
northwest Portugal
* SW region could not be surveyed on the designated days due to weather
storm was an exceptional event, the clustering and persistence of the storms was
highly unusual (Slingo et al. 2014). The storms occurred from the first week of
December 2013, through January and into early February 2014 causing record wind
gusts (> 60 knots) and rainfall in the UK (Slingo et al. 2014). A notable feature of
the storms was the long peak wave period and high wave height, resulting in waves
carrying a large amount of energy causing substantial damage to northeast Atlantic
coasts; presumably conditions at sea for wintering seabirds must have been similarly
extreme. Conditions also included an unusually strong North Atlantic jet stream and
a prolonged series of storm events, with winds gusting in excess of 100 mph; the
worst recorded for a century (Slingo et al. 2014).
Tracking studies have shown that several seabird species including Atlantic Puffin
Fratercula arctica, hereafter ‘Puffin’ (Guilford et al. 2011; Jessopp et al. 2013; Fayet
et al. 2016), Common Guillemot Uria aalge, hereafter ‘Guillemot’ (Stone et al.
1995), Northern Gannets Morus bassanus, hereafter ‘Gannet’, from the UK and
Norway (Veron & Lawlor 2009; Fort et al. 2012), and Black-legged Kittiwakes Rissa
tridactyla, hereafter ‘Kittiwake’, from Western Europe (Frederiksen et al. 2012),
would have been in the Celtic Sea, Bay of Biscay and/or western English Channel
SEABIRD 29 (2016): 22–38
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Reported mortality in the 2013/14 NE Atlantic seabird wreck
during late winter (January–February), and therefore at risk of being affected by the
storm events during this period. Large numbers of dead and injured seabirds were
subsequently washed ashore over much of the northeast Atlantic coast of Europe,
as ocean currents and wind carried dead and moribund birds ashore.
This paper attempts to give a detailed report of the mortality associated with the
2013/14 seabird wreck by combining beached bird surveys from across the entire
affected area. Information will be provided on the origin of the species most
affected by the wreck using ring recovery data; note that data were available from
the UK only for this review. We also report evidence on cause of death from post-
mortem examinations of dead seabirds.
Methods
Survey of stranded seabirds: The methods used by different organisations or
volunteers in different countries were sufficiently similar to warrant comparison.
Typically, a designated route along an accessible coastal area, usually sandy beaches
but including all accessible coastline habitats, is walked by surveyors standing a
distance apart that allows the maximum amount of beach area to be covered in one
transect as they scanned the location for specimens. This methodology accords very
closely with that of the Royal Society for the Protection of Birds’ (RSPB) ‘Beached
Bird Survey’ which coincided with the 2013/14 wreck, conducted on its annual
schedule in the UK on 2223 February (Schmitt 2014). Not all surveys in all areas
of the UK (Figure 1) would have been through the RSPB scheme, due to the reactive
approach to seabird wrecks, but many of them were (Table 1).
Besides the survey effort described in Table 1, additional data were gathered from
members of the public. These records were subsequently confirmed and verified
by the relevant organisation, minimising the chances of double-counting
individuals by cross-checking against previous reports from the same location and
confirming identification from photos where available. All these data are included
in the totals given. Despite the coordinated Beached Bird Survey in parts of the
UK and France, and continued major survey effort across multiple locations
providing a considerable amount of data on the seabird wreck, the challenges
involved in collating and continuing surveys across such a large area, the varying
search effort between organisations around the period of the wreck, the discovery
of only a proportion of the fatalities occurring at sea and the varying time that
different species of seabird corpses will remain afloat, are likely to result in a
substantial underestimation of mortality.
Post-mortem examinations: To determine cause of death, staff from the Groupe
Ornithologique Normand, Alderney Wildlife Trust and Alderney Animal Welfare
conducted a necropsy workshop in Alderney on 22–23 March 2014. Subcutaneous
fat (between the feathers on the breast), fat deposits around the distal part of the
gut, curvature of pectoral muscle extending from the sternum and presence of
spume in the lungs were recorded, alongside general information on the state of the
specimens, to determine any potential cause of death (Broadhurst & Morley 2014).
25
SEABIRD 29 (2016): 22–38
Reported mortality in the 2013/14 NE Atlantic seabird wreck
Ringing recoveries: Ringing recoveries reported to the British Trust for
Ornithology (BTO) over the wreck period were analysed. Only recoveries
occurring between December 2013 and March 2014 were included. We limited
the analysis to the recoveries made in the area affected by the storms, including
the French, Spanish and Portuguese Atlantic coasts, the coast along the English
Figure 1. Location of survey areas recorded during the seabird wreck event (red line).
05001,000250
km
N
SEABIRD 29 (2016): 22–38
26
Reported mortality in the 2013/14 NE Atlantic seabird wreck
Results
Estimates of mortality: In total, 54,982 birds were found during the survey
period; of these 3,243 (5.9%) were live birds. The majority of the birds was found
in France (79.6%) and Puffins were by far the species most impacted (53.6%;
Table 2). Of the live birds retrieved, 3,014 (93%) were found in France, most likely
due to currents and proximity to this coastline during the storm events. Of these
a third subsequently died in transit, a third died in care and a third were
successfully released back into the wild (Farque et al. 2014). The only other
locations in which live birds were recorded were in Guernsey (18 Guillemots and
one Gannet), southwest England (209 birds) and Wales (one Guillemot).
Channel, and the Irish and British coast of the Celtic and Irish Seas (Figure 1).
Note, all recoveries originated from British colonies (Figure 2). Causes of death for
inland recoveries are uncertain, with factors other than the storms potentially
impacting the birds, whilst the death of many birds recovered during the time
period of the wreck was not linked to the storms (e.g. disease or shot); all these
recoveries were excluded from the results.
Figure 2. Origin colonies of ring recoveries recorded on the northeast Atlantic coast of Europe between
December 2013 and March 2014, for the four main species affected (Puffin Fratercula arctica, Guillemot Uria
aalge, Razorbill Alca torda and Shag Phalacrocorax aristotelis); colonies with more than one recovery labelled.
015030075 km
N
Lunga
Port Ban
Fair Isle
Sule Skerry
Leac Buidhe
Isle of May
Faraid Head
Calf of Man
Ailsa Craig
Sanda Island
Great Saltee
Garbh Eilean
Castle Craig
Skomer Island
Puffin Island
Porth Ysgaden
Lambay Island
Isle of Canna
Great Skellig
Ceann Ousdale
Bardsey Island Ynys Gwylan Islands
Number of recoveries
1–2
3–8
9–24
25–61
62–157
Pen y Parc
27
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Reported mortality in the 2013/14 NE Atlantic seabird wreck
Table 2. Numbers of wrecked seabirds (live strandings and dead birds combined) recorded on the northeast
Atlantic coast of Europe from January to April 2014.
Channel SW Rest of UK
Species Portugal Spain France Islands England Wales & Ireland1Total
Gannet 5 67 305 8 33 54427
Fulmar 14131 43 7 15 165
Shag 12 47 110 22 10192
Kittiwake 5 51 868 31 29 20 73 1,077
Puffin 121 277 28,745 119 115 25 42 29,444
Razorbill 9 99 1,197 468 902 234 911 3,820
Guillemot 0 1,538 11,801 441 1,311 139 713 15,943
Auk spp. 1 85 254 16 590 22950
Other seabirds 14 105 229 43 53 7 33 484
Non-seabirds 6 12 176 48 50 5 252
Unidentified 13 022762217 1,729 2,228
Total 163 2,253 43,753 1,290 3,339 657 3,527 54,982
1Region encompasses data from Ireland, Northern Ireland, Scotland, Shetland and England (except the southwest region).
2Value includes 209 live birds retrieved but species composition was not recorded, likely mostly auks.
Post-mortem examinations: Necropsy analyses conducted on a sample of 12
Guillemots in Alderney showed that 58% had emaciated pectoral muscles, 92%
had no stomach contents (no data available for the final bird), 83% had no fat on
the chest and 50% had some or no fat in the intestine, indicating that starvation
was the most likely cause of death (Broadhurst & Morley 2014). In addition,
specimens of Guillemot and Razorbill Alca torda, hereafter ‘Razorbill’, were
collected on Chesil Beach, Dorset, on 2 March 2014 (P. Read pers. comm.). Although
data on time since stranding were unavailable, the mean (± SD) masses were 451.9
± 53.9 g (n = 44) for the Razorbills and 601.5 ± 85.5 g (n = 17) for the Guillemots.
Winter weight of healthy individuals is difficult to attain, but Guillemots are likely
in the range of 900–1,200 g (Hope Jones et al. 1984). Values for Razorbill are harder
to come by still but likely higher than the average weight of the wreck individuals.
Furthermore, live birds collected in France were emaciated and extremely weak,
typically weighing less than half that of a healthy adult (O. Le Gall, pers. comm.).
The Animal Health and Veterinary Laboratories Agency (AHVLA) also necropsied 27
seabirds from the wreck, primarily Razorbills and Guillemots, all of which were
emaciated (AHVLA 2014). The presence of spume (frothy water) in the lungs was
noted by the AHVLA (AHVLA 2014) and recorded in three specimens in the
Alderney Guillemot autopsy (Broadhurst & Morley 2014).
Finally, only 10 (3.3%) out of 300 wrecked Puffins whose wings were examined were in
moult to such an extent that they would have been flightless (M.P. Harris pers. comm.).
Ring recovery data: Since the first recovery of a ringed Puffin was made in
1935/36, the BTO have recorded an average of 11.5 recoveries of ringed Puffins
washed ashore dead in the UK per winter (November to April) up until 2012/13; a
maximum of 127 recoveries were reported in the winter of 1982/83. Additionally,
SEABIRD 29 (2016): 22–38
28
Reported mortality in the 2013/14 NE Atlantic seabird wreck
recoveries of dead ringed Puffins from France and Spain would normally equal two
or three a year (Grantham & Stancliffe 2014). However, during the wreck, 631
ringed birds were recovered, including 216 Puffins, of which 205 were from France
(Tables 3 and 4). The recoveries were spread across the entire impacted area, but
Table 3. Recovery locations of ringed seabirds found wrecked on the northeast Atlantic coast of Europe
between December 2013 and March 2014, as registered by the British Trust for Ornithology.
Channel UK &
Species Portugal Spain France Islands Ireland Total
Gannet 112 6 10
Fulmar 22
Shag 50 50
Cormorant 11 9 11
Kittiwake 178
Herring Gull 77
Great Black-backed Gull 44
Lesser Black-backed Gull 21 3
Black-headed Gull 44
Puffin 15205 14 216
Razorbill 1226 1 126 156
Guillemot 6 90 3 44 143
Black Guillemot 17 17
Total 5 17 324 5 280 631
Table 4. Ringing locations of ringed seabirds wrecked on the northeast Atlantic coast of Europe between
December 2013 and March 2014, as registered by the British Trust for Ornithology.
Northern
Ireland and Isle of Rest of
Species Ireland England Scotland Wales Man Europe* Total
Gannet 441110
Fulmar 112
Shag 72427 10 50
Cormorant 2126 11
Kittiwake 21 5 8
Herring Gull 2113 7
Great Black-backed Gull 44
Lesser Black-backed Gull 12 3
Black-headed Gull 11 2 4
Puffin 5 196 15 216
Razorbill 29 72 50 5 156
Guillemot 29 74 40 143
Black Guillemot 11 6 17
Total 89 13 357 141 18 13 631
* Including data from Belgium, France, Finland, Iceland and Norway.
Figure 3 (overleaf). Colony origin of ringed auks and Shags Phalacrocorax aristotelis (for colonies contributing
more than 5 recoveries) recovered on the northeast Atlantic coast of Europe between December 2013 and
March 2014, as registered by the British Trust for Ornithology.
29
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Reported mortality in the 2013/14 NE Atlantic seabird wreck
Garbh Eilean
Skomer
Lunga
Sule Skerry
0250500125
km
N
3a) Puffin
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30
Reported mortality in the 2013/14 NE Atlantic seabird wreck
Great Saltee
Skomer
Canna
Puffin Island
Sanda
0250500125
km
N
3b) Guillemot
31
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Reported mortality in the 2013/14 NE Atlantic seabird wreck
3c) Razorbill
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Reported mortality in the 2013/14 NE Atlantic seabird wreck
Calf of Man
Lambay
Puffin Island
Ynys Gwyland
Porth Ysgaden
N
0 100 20050
km
3d) Shag
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Reported mortality in the 2013/14 NE Atlantic seabird wreck
certain UK colonies were seemingly represented in higher numbers than others
(Figures 2 and 3). Data were filtered to remove any ring recoveries during the wreck
time period that were not attributable to the storm conditions; therefore, we only
included recoveries of birds with ‘unknown’ (53.57%) or ‘natural’ (42.95%; predom-
inantly attributed to ‘violent weather’) recorded as the cause of death. Ring
recoveries of oiled birds (3.49%) were also recorded within the wreck event as
specimens from other locations showed signs of oiling without it necessarily being
the final cause of mortality (AHVLA 2014; Broadhurst & Morley 2014).
Discussion
A total of 54,982 birds was recorded during the 2013/14 northeast Atlantic seabird
wreck, 79.6% of which were found in France. This number is likely to be a large
underestimate of the final death toll, due to the limited surveying across all regions
impacted. Also, many birds that perish in storms may not be washed ashore, the
proportion of which can only be estimated by drift experiments made during the
incident (Weise 2003). However, with such a large level of mortality (94% of the
birds, of which 53.6% were Puffins, were found dead), it is important to consider
potential factors that may have affected the severity of the wreck event.
Causes of mortality: The ringing recoveries recorded by the BTO attribute 42.3%
of mortality to ‘violent weather’; this is the likely overall cause, but more specif-
ically several independent lines of evidence point to starvation as the most likely
direct explanation for the extensive mortality observed in the 2013/14 seabird
wreck. Autopsy data from Alderney (Broadhurst & Morley 2014), Chesil Beach (P.
Read pers. comm.) and the AHVLA (AHVLA 2014) all show marked levels of
emaciated body condition in analysed specimens. The recording of weight of dead
birds comes with the caveat that many corpses are found in a desiccated state,
potentially overestimating the amount of weight loss. However, the Alderney
autopsy states that 50% of specimens were fresh and 25% rather fresh, reducing
the likelihood of desiccation after death being a major factor for these results.
Although starvation is the main factor in cause of mortality, evidence of drowning,
such as spume in lungs, was also noted in some specimens (AHVLA 2014;
Broadhurst & Morley 2014).
Many factors are likely to affect the susceptibility of different species or
individuals to death from starvation or drowning. Seabirds spend a significant
amount of time and energy foraging in winter, particularly in November and
December, to regain body condition for the breeding season (e.g. Daunt et al.
2006; Fort et al. 2009), so challenging winter conditions sustained over several
weeks or, as in this case, months are likely to affect the birds’ foraging opportu-
nities with possible consequences on their future condition, survival and
reproductive ability. For example, European Shags Phalacrocorax aristotelis,
hereafter ‘Shags’, have been shown to forage less during times of high wind
speeds, with males foraging for shorter times than females which could have
consequences for sex-specific survival rates (Lewis et al. 2015), and they may
even halt diving altogether during high onshore gales (Frederiksen et al. 2008).
SEABIRD 29 (2016): 22–38
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Reported mortality in the 2013/14 NE Atlantic seabird wreck
Additionally, the continued turbid water conditions associated with storms may
create poor visibility for foraging. This is a particular issue for auks which have
less flexible foraging strategies than Shags (Watanuki et al. 2008; Cook & Burton
2010) and spend more energy on longer, deeper dives (Elliott et al. 2013; Elliott
& Gaston 2014) to pass through the turbid surface conditions, possibly causing
the energetic cost of foraging to outweigh the nutritional gain from successful
catches. The fact that the overwhelming majority (91.3%) of retrieved birds
during the 2013/14 seabird wreck belonged to the auk family supports difficult
foraging conditions as a major contributor to mortality.
In addition to considering the weather conditions, the habits of the species found
must also be considered; susceptibility may vary between pelagic and inshore
species and between long- and short-ranging species. Pelagic seabirds were the
most impacted during the wreck, accounting for 92.5% of all findings (auks alone
were 91.3%), whilst inshore species contributed just 3.3% of all findings (Table 2)
suggesting they were less susceptible to the conditions created by the storms.
Findings from a small number of Puffins carrying geolocation loggers also suggest
that the migratory patterns of some individuals may make them more susceptible
to suffer from a wreck than others (A. Fayet et al. unpubl. data). Moult stage could
be another factor adding to birds’ susceptibility to a wreck event. For example,
Puffins shed all their main wing feathers synchronously during the winter (Harris
& Yule 1977) and are then unable to fly for a few weeks (Harris 2014; Harris et al.
2014). However, the small number of moulting Puffins found suggests moult was
not an important factor in this wreck (M.P. Harris pers. comm.). Nevertheless, it
cannot be totally disregarded, given the variability in the timing of Puffin moult
(Harris et al. 2014) and the role of post-moult feather growth in the Razorbill
wreck in 2007 (Heubeck et al. 2011).
Besides differences in foraging, migratory strategies, and habitat use, the physio-
logical differences between species could also have affected mortality rates. For
instance, the partially wettable plumage of Shags (Grémillet et al. 1998) increases
mortality in times of low ambient temperature, most likely through hypothermia
(Frederiksen et al. 2008). The ability to survive through a wreck event may also be
linked to body size, with larger-bodied species retaining greater energy reserves.
Finally, metabolism could also play a role, as suggested by Hope Jones et al. (1984)
who discovered a higher percentage of Razorbills to have lower fat reserves on
autopsy than Guillemots following the 1983 seabird wreck. However, the small
samples examined of these species from the 2013/14 wreck are insufficient to
provide any further evidence on the role of metabolism, particularly as weight loss
was high for both species. A recent study of some specimens showed, however, that
high concentrations of mercury (known to increase stress) may have acted as an
aggravating mortality factor during the event (Fort et al. 2015).
Impacts of mortality: Whilst the majority of wreck birds were located in France
(Tables 2 and 3), most ringed birds were from breeding colonies in Scotland, Wales
and Ireland (Table 4). This is to be expected as most of the large seabird colonies
35
SEABIRD 29 (2016): 22–38
Reported mortality in the 2013/14 NE Atlantic seabird wreck
where BTO-licensed ringers operate are in these locations, but it does highlight
the possibility that reproductive success, recruitment or survival of birds from
these colonies were impacted by the wreck. Furthermore, whilst the scale of the
wreck event is staggering in terms of numbers, it is not the first event of this scale.
For example, in February and March 1994, 20,000–50,000 Guillemots and
3,000–5,000 Shags were reported washed ashore in emaciated states due to
storm events (Harris & Wanless 1996), and in February 1983 prolonged storms
across Western Europe resulted in 34,000 seabirds washing ashore in the
northeast UK alone with another 24,000 across Western Europe (Underwood &
Stowe 1984). Therefore, the regularity of wrecks impacting on a particular species
or colony should also be considered when trying to uncover any longer-term
effects at the population level.
In summary, the seabird wreck along the Atlantic coastline of northwest Europe
in winter 2013/14 was an extreme example of weather-induced mortality from
starvation in seabird populations. The subsequent impacts on population
numbers and productivity are likely to be detectable in multiple species and
colonies (M.J. Wood et al., unpubl data; T.R. Birkhead, pers. comm.). Whilst any
individual wreck can have significant consequences for the populations involved,
the frequency of wreck events is also likely to be an important factor. The
potential of climate change to increase the frequency and strength of storms
(Slingo et al. 2014; Fischer & Knutti 2015) can have severe implications for
seabird populations in the future, and the vulnerability of species to extreme
events needs to be accounted for in future predictions of ecological impacts of
climate change (Frederiksen et al. 2008).
Acknowledgements
We would like thank the hundreds of dedicated staff, volunteers and members
of the public across Western Europe who responded to the wreck event with
efforts to survey its impact. We would like to thank the organisations across
Europe that contributed to the response to and recording of the wreck,
including: Ligue pour la Protection des Oiseaux (LPO), RSPB, Cornwall, Devon
and Dorset Wildlife Trusts (and the Cornwall Marine Strandings Network and
DoWT staff at Chesil Beach, Dorset in particular), RSPCA, South Devon Seabird
Trust, BTO Ringing Scheme, National Trust, Natural England, SEO Birdlife,
Alderney Wildlife Trust, Durrell Wildlife Conservation Trust, States of Jersey,
States of Guernsey, other county Wildlife Trusts and Sussex Ornithological
Society. We would particularly like to thank the following people for their
specific help in supplying data and details on responses to the wreck event in
their location: Mike Harris (CEH, Scotland), Glyn Young (Durrell Conservation
Trust, Jersey), Cristina Sellares (National Trust, Jersey), Catherine Veron (La
Société Guernesiaise, Guernsey), Janice Dockerill (Environment Department,
Guernsey), Anne-Isabelle Boulon (Living Islands, Alderney), Ed Stubbings
(Warden Skomer Island, Wales), Bee Bueche (Warden Skomer Island, Wales),
Pembrokeshire Bird Group (Wales), Phil Read (Chesil Beach, Dorset), John
Walmsley (Newgale Beach, Wales), Tim van Nus (Portugal), Olivier le Gall (LPO,
France), Amélie Boue (LPO, France) and Albert Cama Torrell (Spain). We are also
grateful to Martin Heubeck, Viola Ross-Smith, Richard Sherley and two
anonymous referees, whose comments greatly improved this paper.
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... However, most often the geographic origins of impacted seabirds and the causes of their deaths remain unclear. 6 We performed the first ocean-basin scale assessment of cyclone exposure in a seabird community by coupling winter tracking data for $1,500 individuals of five key North Atlantic seabird species (Alle alle, Fratercula arctica, Uria aalge, Uria lomvia, and Rissa tridactyla) and cyclone locations. We then explored the energetic consequences of different cyclonic conditions using a mechanistic bioenergetics model 7 and tested the hypothesis that cyclones dramatically increase seabird energy requirements. ...
... Necropsies suggest that seabirds are lean after being exposed to high-intensity cyclones 6,15 and that the resulting mortality can be aggravated by mercury contamination. 16 Yet we do not know whether they starve due to abnormally high energy expenditure linked to harsh climatic conditions, 17 because of reduced foraging profitability and energy inputs, or because both these constraints synoptically affect their energy balance. ...
... 30,31 Alternatively, cyclones may shift alcids away from their preferred prey patches and into unprofitable foraging habitats. 32 Overall, even if starvation may be the main driver of seabird winter wrecks, we cannot exclude other causes of mortality, such as drowning, 6 collision with reefs and rocky coastlines (M. Baran, personal communication), or inland stranding. ...
Article
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Each winter, the North Atlantic Ocean is the stage for numerous cyclones, the most severe ones leading to seabird mass-mortality events called ‘‘winter wrecks.’’ During these, thousands of emaciated seabird carcasses are washed ashore along European and North American coasts. Winter cyclones can therefore shape seabird population dynamics by affecting survival rates as well as the body condition of surviving individuals and thus their future reproduction. However, most often the geographic origins of impacted seabirds and the causes of their deaths remain unclear. We performed the first ocean-basin scale assessment of cyclone exposure in a seabird community by coupling winter tracking data for 1,500 individuals of five key North Atlantic seabird species (Alle alle, Fratercula arctica, Uria aalge, Uria lomvia, and Rissa tridactyla) and cyclone locations. We then explored the energetic consequences of different cyclonic conditions using a mechanistic bioenergetics model and tested the hypothesis that cyclones dramatically increase seabird energy requirements. We demonstrated that cyclones of high intensity impacted birds from all studied species and breeding colonies during winter but especially those aggregating in the Labrador Sea, the Davis Strait, the surroundings of Iceland, and the Barents Sea. Our broad-scale analyses suggested that cyclonic conditions do not increase seabird energy requirements, implying that they die because of the unavailability of their prey and/or their inability to feed during cyclones. Our study provides essential information on seabird cyclone exposure in a context of marked cyclone regime changes due to global warming.
... Extreme weather conditions at sea can impair the ability of some species to forage and find enough food. This can lead to poor body condition, lower survival and can cause substantial 'wrecks' (Morley et al., 2016;Louzao et al., 2019). This is at least partly because flight (in flapping flight species) and diving are more costly at higher wind speeds (Kogure et al., 2016). ...
... Most recently, the 'Beast from the East' storm in February and March 2018 caused breeding seasons to be markedly delayed at seabird colonies on the (Masselink et al., 2016). A total of 54,982 birds were 'wrecked' along the European coastline, of which 94% were dead, apparently due to starvation, exhaustion and drowning (Morley et al., 2016). The majority of birds found were recorded on the French coastline (80%) and just over half were Atlantic puffins (Morley et al., 2016). ...
... A total of 54,982 birds were 'wrecked' along the European coastline, of which 94% were dead, apparently due to starvation, exhaustion and drowning (Morley et al., 2016). The majority of birds found were recorded on the French coastline (80%) and just over half were Atlantic puffins (Morley et al., 2016). The 2013/14 wreck provided an opportunity to test models that predict the cumulative effect of extreme wind events. ...
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EXECUTIVE SUMMARY • The seabird declines that commenced at the end of the last century have continued during the last two decades. • Further research into the causes of these declines is required if we are to fully understand the complex mechanisms operating, which are known to vary geographically. Climate change is considered to be one of the main causes of the declines. The principal mechanism is the effect of climate warming on food supply. • There is growing evidence that short-term weather conditions have an important effect, including extreme weather events. Climate models predict further warming and increased severity and frequency of extreme weather events in UK waters. • Seabirds face an uncertain future and may decline further in the coming decades, as the interacting effects of new and existing influences will pose additional challenges.
... Shortly after completing breeding, most seabirds initiate feather moult, a process with high energetic requirements and in some volant species a period of flightlessness, limiting the ability of individuals to move between locations and away from threats (Bridge 2006). Following the moult period, seabirds may experience challenging winter conditions, with individuals exposed to prolonged periods of poor weather (Morley et al. 2016), lower food availability (Osborn et al. 1984) and shorter daylight hours in which to forage (Daunt et al. 2006, Moe et al. 2021. The moult and midwinter periods are therefore key parts of the annual cycle when individuals may experience more hostile environmental conditions and/or energetic constraints (Burke & Montevecchi 2018), making their populations more vulnerable to other marine threats. ...
... Furthermore, the volume of marine litter present in the North Sea continues to increase (Gutow et al. 2018), and guillemots and razorbills are vulnerable to entanglement or ingestion (Battisti et al. 2019). Extreme weather events during the non-breeding season can cause high levels of mortality for seabirds such as auks, with multiple wrecks in our study region in recent decades (Harris & Wanless 1984, 1996, Heubeck et al. 2011, Morley et al. 2016, and it is predicted that frequencies of such events will increase under climate change (IPCC 2018). Furthermore, within our study region, there are large-scale plans for marine development of wind energy in northern European waters over the next decade, with coastal Scottish waters and the southern North Sea being key areas for development (The Scottish Government 2020, The Crown Estate 2021). ...
Article
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Migration is a widespread strategy for escaping unfavourable conditions during winter, but the extent to which populations that segregate during the breeding season aggregate during the non-breeding season is poorly understood. Low non-breeding season aggregation may be associated with higher likelihood of overlap with threats, but with fewer populations affected, whereas high aggregation may result in a lower probability of exposure to threats, but higher overall severity. We investigated non-breeding distributions and extent of population aggregation in 2 sympatrically breeding auks. We deployed geolocation-immersion loggers on common guillemots Uria aalge and razorbills Alca torda at 11 colonies around the northern UK and tracked their movements across 2 non-breeding seasons (2017-18 and 2018-19). Using 290 guillemot and 135 razorbill tracks, we mapped population distributions of each species and compared population aggregation during key periods of the non-breeding season (post-breeding moult and mid-winter), observing clear interspecific differences. Razorbills were largely distributed in the North Sea, whereas guillemot distributions were spread throughout Scottish coastal waters and the North, Norwegian and Barents Seas. We found high levels of aggregation in razorbills and a strong tendency for colony-specific distributions in guillemots. Therefore, razorbills are predicted to have a lower likelihood of exposure to marine threats, but more severe potential impact due to the larger number of colonies affected. This interspecific difference may result in divergent population trajectories, despite the species sharing protection at their breeding sites. We highlight the importance of taking whole-year distributions into account in spatial planning to adequately protect migratory species.
... The media is also reporting opportunistic species such as gulls (Larus spp.) (SGO 2019). Large scale seabird wrecks, i.e., a vast number of dead, injured or exhausted seabirds wash up ashore, are a well-known phenomenon to or major pollution events are often implicated (Newman et al. 2007, Morley et al. 2016, Tavares et al. 2016). However, the particularly smaller scale seabird wreck events may be associated with other environmentally natural causes of death such as toxins produced by: (1) marine algae, mainly and (2) bacteria such as Clostridium botulinum (Shumway et al. 2003). ...
... The media is also reporting opportunistic species such as gulls (Larus spp.) (SGO 2019). Large scale seabird wrecks, i.e., a vast number of dead, injured or exhausted seabirds wash up ashore, are a well-known phenomenon to or major pollution events are often implicated (Newman et al. 2007, Morley et al. 2016, Tavares et al. 2016). However, the particularly smaller scale seabird wreck events may be associated with other environmentally natural causes of death such as toxins produced by: (1) marine algae, mainly and (2) bacteria such as Clostridium botulinum (Shumway et al. 2003). ...
... In just the past decade, various record-breaking ocean-warming events have occurred, including the 'Ningaloo Niño' off Western Australia (Pearce and Feng, 2013) and the extreme El Niño that affected most of the Indo-Pacific in 2016 . Elsewhere, marine heatwave-related decreases in primary productivity have also been correlated with decreased survival and breeding success of the following: Atlantic puffins (Fratercula arctica), common terns (Stirna hirundo) and Cory's shearwater (Calonectris diomedea) in the Atlantic (Jenouvrier et al., 2009;Morley et al., 2016;Szostek and Becker, 2015); king penguins (Aptenodytes patagonicus) in the Southern Ocean (Le Bohec et al., 2008); and roseate terns (Sterna dougallii) in the western Indian Ocean (Monticelli et al., 2007). Other marine vertebrates including marine mammals and fish have also been negatively affected by such phenomena due to bottom-up effects of ecosystem shifts (Sydeman et al., 2015). ...
Article
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‘The Blob’, a mass of anomalously warm water in the Northeast Pacific Ocean peaking from 2014 to 2016, caused a decrease in primary productivity with cascading effects on the marine ecosystem. Among the more obvious manifestations of the event were seabird breeding failures and mass mortality events. Here, we used corticosterone in breast feathers (fCort), grown in the winter period during migration, as an indicator of nutritional stress to investigate the impact of the Blob on two sentinel Pacific auk species (family Alcidae). Feathers were collected from breeding females over 8 years from 2010 to 2017, encompassing the Blob period. Since Pacific auks replace body feathers at sea during the migratory period, measures of fCort provide an accumulated measure of nutritional stress or allostatic load during this time. Changes in diet were also measured using δ15N and δ13C values from feathers. Relative to years prior to the Blob, the primarily zooplanktivorous Cassin’s auklets (Ptychoramphus aleuticus) had elevated fCort in 2014–2017, which correlated with the occurrence of the Blob and a recovery period afterwards, with relatively stable feather isotope values. In contrast, generalist rhinoceros auklets (Cerorhinca monocerata) displayed stable fCort values across years and increased δ15N values during the Blob. As marine heatwaves increase in intensity and frequency due to climate change, this study provides insight into the variable response of Pacific auks to such phenomena and suggests a means for monitoring population-level responses to climatological variation.
Article
This is the 82nd annual report of the British Trust for Ornithology’s Ringing Scheme, incorporating the report of the Nest Record Scheme and covering work carried out and data processed in 2018. We reviewed the literature on tracking devices to assess the impact of their use on individual birds. More than 3400 studies were identified, the first dating back to the early 1960s. The ecological questions being studied varied, with migration being the most common study subject. The frequency of effects was significantly related to attachment type and the likelihood of adverse effects was related to device weight. The proportion of studies reporting adverse effects has declined over time and the research highlighted the need for systematic documentation of potential effects. NRS and CES data were used in a collaborative study which investigated mismatches between the timing of breeding for 21 British songbirds and the availability of food for their chicks. Results indicate that birds did breed later in warmer springs relative to the timing of spring; however, this did not have a significant impact on long-term breeding success. In 2018, 135 CE sites submitted data. Long-term (1984–2017) declines in abundance were observed in six migrant, one partial migrant and three resident species while long-term increases in abundance continue to be recorded in two short-distance migrants and four resident species. Seven species are exhibiting significant long-term declines in productivity; only two species, Reed Warbler Acrocephalus scirpaceus and Chaffinch Fringilla coelebs, are exhibiting a significant long-term increase in productivity. Long-term trends in survival indicate increases for 12 species and declines for eight. Compared to the five-year mean (2013–17), the abundance of 14 species decreased significantly in 2018 with five species being recorded in lower numbers than in any previous year since CES monitoring began. No significant increases in abundance were recorded. Productivity increased significantly for six migrant and four resident species in 2018 and decreased for only two species (both residents). Significant declines in survival rates were observed in 10 species in 2018 compared to the five-year mean. The number of Retrapping Adults for Survival submissions fell slightly in 2018 to 190; eight projects ran for the first time. In total, 58 species were monitored, with Pied Flycatcher Ficedula hypoleuca (23 projects), House Sparrow Passer domesticus (18), Starling Sturnus vulgaris (17), Sand Martin Riparia riparia (13) and Reed Warbler (10) the most-studied species. A total of 982 858 birds were ringed during the year, comprising 170 619 nestlings, 465 610 first-year birds, 319 955 adults (birds in their second calendar year or older) and 26 674 birds whose age could not be determined. In addition, there were 236 314 recaptures of ringed birds at or near the ringing site. In total, 54 324 recoveries (birds found dead, recaptured or resighted at least 5 km from the place of ringing) of BTO-ringed birds were reported in 2018. The Appendix highlights a selection of recoveries that have extended our knowledge of movements, as well as longevity records established during the year. A total of 38 269 NRS submissions were received in 2018, the lowest total since 2009. Mean laying dates for four species were significantly earlier than the five-year mean (2013–17) with eight of the 52 species analysed laying significantly later. Eight species displayed a significant increase in breeding success (fledglings per breeding attempt) in 2018 compared to the five-year mean, with one decreasing significantly. Permits or licences to disturb breeding birds on Schedule 1 of the Wildlife and Countryside Act 1981 (as amended), were prepared and issued to 502 ringers and nest recorders in 2018. During 2018, the use of special methods was authorised for 557 marking projects, 66 trapping projects and 52 sampling projects.
Thesis
Seabirds are particularly vulnerable to the direct and indirect effects of climate change, however little is known about those impacts outside of the breeding season. This lack of knowledge is problematic because the conditions encountered during migration and wintering strongly shape seabird population dynamics. It is therefore essential to understand the effects of climate on their winter distribution and migration routes. Linking the distribution of organisms to environmental factors is therefore a primary task benefiting from the concept of energyscapes (defined as the variation of an organism's energy requirements across space according to environmental conditions) which has recently provided a mechanistic explanation for the distribution of many animals. In this context, we have predicted the current and future winter habitats of five species representing 75% of the seabird community in the North Atlantic (Alle alle, Fratercula arctica, Uria aalge, Uria lomvia and Rissa tridactyla). To this aim, we monitored the movements of more than 1500 individuals to identify the birds' preferred habitats through resource selection functions based on the modeling of their energy expenditure and prey availability. Electronic tracking data were also overlaid with cyclone locations to map areas of high exposure for the seabird community across the North Atlantic. In addition, we explored the energetic consequences of seabird exposure to storms using a mechanistic bioenergetic model (Niche MapperTM). Finally, we examined the impact of total summer sea ice melt from 2050 on Arctic bird migration. Our analyses predict a northward shift in the preferred wintering areas of the North Atlantic seabird community, especially if global warming exceeds 2°C. Our results suggest that cyclonic conditions do not increase the energy requirements of seabirds, implying that they die from the unavailability of prey and/or inability to feed during cyclones. Finally, the melting sea ice at the North Pole may soon allow 29 species of Arctic birds to make new trans-Arctic migrations between the Atlantic and the Pacific. We also estimate that an additional 26 currently migratory species could remain in the Arctic year-round. This work illustrates how climate change could radically alter the biogeography of migratory species and we provide a methodological toolbox to assess and predict these changes by combining movement ecology and energetic physiology.
Article
This is the 81st annual report of the British Trust for Ornithology’s Ringing Scheme, incorporating the report of the Nest Record Scheme and covering work carried out and data collected and processed in 2017. Demographic data contributed to several research studies during the year, including a review of the conservation status of Raven Corvus corax and the impacts of licensed control. This quantified the extent to which non-breeding birds could be removed without impacting overall population sustainability, as well as the data permitted, but noted that a better understanding of age-specific survival rates was required. Nest Record Scheme data contributed to a large-scale collaborative study investigating the impact that temperature and rainfall have on the timing of events throughout the annual cycles of 10 000 animal and plant species across different trophic levels. The results indicated that the speed of response was greatest at low trophic levels. Future changes under a range of emission scenarios were estimated; these results indicated that advances in timing would be greatest for primary consumers. A statistical method was developed, using nest record data, to quantify nest failure, taking into account uncertainties generated during data collection. The method was applied to Blackbird Turdus merula records to compare survival rates of nests in gardens with those in the wider countryside. Survival in suburban areas was found to be higher than in urban or rural areas, and nests survived better in wetter conditions across all habitats. In order to quantify any welfare impacts of capturing wild birds, we examined the frequency with which passerines were reported to have died either whilst in the net or during handling. Analyses of capture data submitted from 2005 to 2013 indicated that the average mortality rate is around 0.1%, and that this is consistent between years. Most fatalities occurred before individuals had been extracted from the nets and juveniles appeared to be at greater risk than adults. In 2017, 135 CE sites submitted data. Long-term (1984–2016) declines in abundance were observed in four migrant, one partial migrant and one resident species and concurrent increases in abundance continued to be recorded in two short-distance migrants and four resident species. There was no change to the 10 species which exhibited significant long-term declines in productivity; only one species, Chaffinch Fringilla coelebs, exhibited a significant long-term increase in productivity. Compared to the five-year mean (2012–16), the abundance of six species increased significantly in 2017; Treecreepers Certhia familiaris were seen in greater numbers in 2017 than in any other year since CES monitoring began. Conversely, the abundance of six species fell significantly in 2017, with Chaffinch and Greenfinch Chloris chloris numbers at their lowest since CES monitoring began. Productivity increased significantly for three species in 2017 and decreased for eight species. The only significant change in survival rates between 2017 and the five-year mean was for Wren Troglodytes troglodytes, although the survival rate for Song Thrush Turdus philomelos was at its highest. The number of Retrapping Adults for Survival submissions fell slightly in 2017 to 195; 11 projects ran for the first time. In total, 58 species were monitored, with Pied Flycatcher Ficedula hypoleuca (20 projects), House Sparrow Passer domesticus (16), Starling Sturnus vulgaris (15) and Sand Martin Riparia riparia (11) continuing to be the most-studied species. Once again, over a million birds (1 020 356) were ringed during the year, comprising 165 565 nestlings, 500 580 first-year birds, 321 945 adults (birds in their second calendar year or older) and 32 266 birds where the age could not be determined. In addition, there were 242 719 recaptures of ringed birds at or near the ringing site. In total, 41 049 recoveries (birds found dead, recaptured or resighted at least 5 km from the place of ringing) of BTO-ringed birds were reported in 2017. The Appendix highlights a selection of recoveries that have extended our knowledge of movements, as well as longevity records established during the year. NRS submissions in 2017 exceeded 4 0 000 for the eighth consecutive year, with a total of 43 570 records received. Mean laying dates for 22 species were significantly earlier than the five-year mean (2012–16) with none of the 52 species analysed laying significantly later. There was a significant increase in breeding success (fledglings per breeding attempt) for two species in 2017, compared to the five-year mean, with one decreasing significantly. Permits or licences to disturb breeding birds on Schedule 1 of the Wildlife and Countryside Act 1981 (as amended), were prepared and issued to 580 ringers and nest recorders in 2017. During 2017, the use of special methods was authorised for 584 marking projects and 67 trapping projects, a 4% decrease from 2016.
Article
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Animals can be flexible in their migration strategies, using several wintering sites or a variety of routes. The mechanisms promoting the development of these migratory patterns and their potential fitness consequences are poorly understood. Here, we address these questions by tracking the dispersive migration of a pelagic seabird, the Atlantic puffin Fratercula arctica, using over 100 complete migration tracks collected over 7 years, including repeated tracks of individuals for up to 6 consecutive years. Because puffins have high flight costs, dispersion may generate important variation in costs of migration. We investigate differences in activity budgets and energy expenditure between different strategies. We find that puffins visit a range of overwintering destinations, resulting in a diversity of migratory routes differing in energy expenditures; however, they show interindividual similarity in the timings and location of major movements. We consider 3 hypothetical mechanisms that could generate this pattern: 1) random dispersion; 2) sex segregation; and 3) intraspecific competition or differences in individual quality. First, we dismiss random dispersion because individuals show strong route fidelity between years. Second, we find that sex differences contribute to, but do not account fully for, the migratory variation observed. Third, we find significant differences in breeding success between overwintering destinations, which, together with differences in foraging levels between routes, suggest that birds of different quality may visit different destinations. Taken together, our results show that dispersive migration is a complex phenomenon that can be driven by multiple factors simultaneously and can shape a population’s fitness landscape.
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The distribution of Atlantic puffins (Fratercula arctica) from Skellig Michael, south-west Ireland, was investigated using geolocation loggers between the 2010 and 2011 breeding seasons. All tracked birds travelled rapidly west into the North Atlantic at the end of the breeding season in August, with the majority undertaking transatlantic trips from Ireland to the Newfoundland-Labrador shelf. The furthest distance from the colony reached by each bird was not influenced by body mass or sex and was achieved in approximately 20 days. By October, all birds had moved back to the mid Atlantic where they remained resident until returning to the breeding colony. The most parsimonious explanation for the rapid, directed long-distance migration is that birds exploit the seasonally high abundance of prey [e.g., fish species such as capelin (Mallotus villosus) and sandlance (Ammodytes spp.)] off the Canadian coast, which is also utilised by large populations of North American seabirds at this time. Once the availability of this short-term prey resource has diminished, the tracked puffins moved back towards the north-east Atlantic. A relationship between relative abundance of puffins and zooplankton was found in all winter months, but after correcting for spatial autocorrelation, was only significant in November and January. Nevertheless, these results suggest a potential switch in diet from mainly fish during the breeding and early post-breeding periods to zooplankton over the remaining winter period. This study suggests that puffins from south-west Ireland have a long-distance migration strategy that is rare in breeding puffins from the UK and identifies a key non-breeding destination for puffins from Ireland. This has implications for the susceptibility of different breeding populations to the effects of possible climatic or oceanographic change.
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