Bb feb 2010 -
The population biology
of Common Sandpipers
in Britain
T. W. Dougall, P. K. Holland and D. W. Yalden
Abstract The population biology of Common Sandpipers
Actitis hypoleucos has
been studied, especially by colour-ringing breeding adults, at two sites, in the Peak
District and in the Scottish Borders. Adults are usually site-faithful, males more so
than females, contributing to a good apparent survival rate (72% and 67%,
respectively). Some, at least, return to breed at one year old, but usually not to the
site where they hatched. The population in Britain seems to be in slow decline,
most obviously indicated by a contraction along the edges of its range, which
results especially from poor recruitment of young birds. This does not seem to be
due to poorer breeding success, but it is uncertain whether it is caused by a subtle
effect of climate change, change in quality of stopover sites on migration, or
changes in wintering habitat. Since we don't know precisely where British birds
spend the winter, the last possibility is especially hard to evaluate.
or Wood Lark
Lullula arborea, but not suffi-
The Common Sandpiper
Actitis hypoleucos is
ciently abundant to benefit from mass
one of those ‘in between' birds – not rare
studies, like the Blue Tit
Cyanistes caeruleus
enough to have attracted devoted individual
or Robin
Erithacus rubecula. Unlike many
attention, like the Osprey
Pandion haliaetus
waders, it does not flock in large numbers to
British Birds 103 • February 2010 • 100–114
Common Sandpipers in Britain
fields, estuaries or the Bancd'Arguin, it is not censused bymoorland bird counts, andwhen on migration does not
get ringed in any great
numbers. There is a littleinformation from the Water-
ways Bird Survey on popula-
tion trends and the species
seems to have been targeted
by only one PhD study (Mee
2001). Our own, essentiallyspare-time, efforts have beenan attempt to fill in some of
the gaps. In reviewing whatwe have learnt, we inevitably
also highlight how much
remains unknown.
Study sites and
Fig. 1. Sketch map of the Ashop–Alport study site and the nearby
reservoirs in the Upper Derwent Valley, Peak District (areas of
Our efforts started with a
woodland shown in green, water features in blue; also shown is the
survey of numbers in the Peak
A57 between Manchester and Sheffield) (from Dougall
et al. 2005).
District in 1977–79, combinedwith a colour-ringing study ofbirds along a 10-km stretch of
the Rivers Ashop (‘Snake
Pass') and Alport (Holland
etal. 1982a,b). Of about 200pairs then breeding in the
Peak District, 20–22 pairs
occupied these valleys, withanother 45–50 pairs nearby on
the Ladybower-Derwent-
Howden (LDH) Reservoirchain. The Ashop flows intothe western arm of LadybowerReservoir, about 2 km down-
stream of the study site; thewhole forms part of theUpper Derwent catchment.
The colour-ringing study con-
tinues to the present day,giving 32 years of data (PKH,
then DWY), but has also beenextended to the reservoirchain. Thinking that a com-parative study would be
revealing, we started a parallel
investigation on the riversLeithen Water (8.5 km) and
Fig. 2. Sketch map of the Borders study site (woodland in green,
Heriot Water (6 km) in the
water features in blue; also shown is the B709 road fromEdinburgh to Innerleithen) (from Dougall
et al. 2005).
Moorfoot Hills, Borders, in
British Birds 103 • February 2010 • 100–114
Dougall et al.
1993 (TWD), and this also continues
returning migrants than the Peak District
(Dougall
et al. 2005; Pearce-Higgins
et al.
(hereafter referred to as Ashop) study area.
2009). This study site, and our knowledge of
On the other hand, the LDH reservoir chain
the species, benefited from an intensive PhD
has a shoreline of 30 km. This must also
study in 1998–99 (Mee 2001). The two
provide a good target for returning sand-
stretches of river are about 6 km apart, and
pipers, but is too extensive to be studied as
together perhaps provide a larger target for
intensively as the rivers. The whole shoreline
has been surveyedtwice (once in May,once in June) eachyear since 1992, andone or two ex-Ashopbirds are invariablypresent.
Adults are usually
caught on the riversin short, single-panelmist-nets set withinthe
usually targetingparticular individ-uals, and birds areringed with 3–4Darvic colour rings,as well as a BTO
metal ring, so that
individuals are sub-
29. An upper stretch of Glentress Water, near Blackhopebyres, in the
sequently recognis-
Borders study area. This shows the meandering river and the wide shingle
able in the field. In
banks that develop in the bends, producing favoured feeding sites for
Common Sandpipers
Actitis hypoleucos.
area, potential terri-tories are checkedweekly through Mayand June, sometimesinto mid July, for thepresence of colour-ringed birds. Nestsare rarely located,and the adults aresecretive
incubation, but oncethe chicks hatch,their parents becomevery vocal (‘alarm'),revealing roughlywhere their chicksare. By backing off,or using a car as a
hide, it is sometimes
possible to watch
30. A lower stretch of the Borders study site, on Glentress Water near
adults returning to
Whitehope. This is good habitat for Common Sandpipers
Actitis hypoleucos,
brood young chicks.
with wide shingle beds, cover for chicks in the boulders, and with theadvantage of the nearby road, allowing a car to be used as a hide; May 2009.
Every effort is made
British Birds 103 • February 2010 • 100–114
Common Sandpipers in Britain
to ring these, with a single(‘scheme') colour ring anda BTO metal ring. Ashopbirds receive a white ringon the right tibia as thescheme mark, while thosearound the reservoirs get alight blue ring on the lefttibia. The hope was thatrecruits to the Ashop fromthe reservoirs would bequickly recognised. In theBorders, chicks as well asadults were formerly fittedwith individually recognis-
able combinations, but in
more recent years both
31. ‘Walk-in' traps set at the waterline of Ladybower Reservoir,
chicks and adults have
May 2009. Two traps are set, facing in opposite directions, and with
scheme colours applied to
boulders or other debris to mask the outline of the trap, and
the right leg only: red on
mealworms on the bait trays as an added attraction.
tibia, orange over BTO on tarsus for adults;
quickly (and noisily) pair up, usually in their
orange on tibia, red over BTO on tarsus for
old territories, and have eggs by mid May. It
chicks. So far, neither Peak District nor Moor-
seems as though they fly into their territories
foot Hills birds have been seen in the recip-
very quickly (rather than trickle slowly up
rocal study area, but a juvenile ringed on
through the country, although there are some
autumn passage in Lancashire on 4th August
sightings in south and southwest England,
2001 (by PKH) was retrapped as a breeding
and in Wales, of colour-ringed Borders
bird south of Peebles, just outside the Moor-
birds). This impression is heightened by the
foots study area, on 29th May 2003. The age
fact that they get back to their territories in
of the chicks (and hence their hatch date) can
the Borders about a week earlier than those
be estimated from their bill length, and
in the Peak District. Young birds arrive back
weight relative to age gives an estimate of
5–10 days later, on average, than the old
condition (Holland & Yalden 1991b). Sexing
hands. Territories are usually 150–200 m long
Common Sandpipers is not easy, though
when there are neighbours to constrain
females are generally slightly larger (means
them, and are generally along stretches of
54 g v. 49 g; Mee 2001). The greater weights of
shingle shore with some cover for the chicks
females during egg-laying are revealing (they
(e.g. undercut shores, boulders, taller vegeta-
can weigh up to 82 g), and with colour-ringed
tion). Females typically lay a clutch of four
birds courtship behaviour can be helpful in
eggs (mean 3.65) in a nest on a grassy bank a
determining the sex of individuals. However,
few metres back from the water, but some-
both adults incubate, brood chicks and
times on shingle islands or at the top of stony
defend their territory against neighbours
reservoir shores; occasionally the nest may be
(sometimes making the mistake of chal-
over 100 m from the waterside. Incubation is
lenging their own returning mates!).
shared, but males sit mainly overnight and
Although the male usually follows behind the
females during daylight hours (Mee 2001).
female while she is feeding up prior to egg-
The eggs hatch from mid May (Borders)
laying, and signals to her with ‘wings-up', dis-
or the last few days of May into early June,
playing the strongly patterned underwing, the
but new recruits are later and pairs will
same display is used as a threat by both sexes.
produce a replacement clutch if the first islost, stretching the hatch dates to as late as
7th July. At hatching, chicks weigh around
Older Common Sandpipers reappear on
9.5 g and have a bill length of about 10 mm.
their breeding grounds in mid to late April,
Chicks are encouraged from the nest by their
British Birds 103 • February 2010 • 100–114
Dougall et al.
.Yalden. W
32. A typical Common Sandpiper
Actitis
hypoleucos nest, with a clutch of four eggs; this
33. A late, replacement nest, in growing
one, in a patch of Bracken
Pteridium aquilinum in
Bracken
Pteridium aquilinum, Ashop Valley,
the Ashop study area, 15th May 1984, belonged
9th June 1982; another of Red-Red's successful
to the long-lived male ‘Red-Red'.
parents, and for their first few days usually
this age to fledging: their camouflage, ability
frequent damp areas – rushes, streams trick-
to run and hide, and to dive and swim if sur-
ling over reservoir shores – where small
prised near deeper water, gives them an
insects are readily available. Mortality at this
ability to dodge most predators (and
stage is high, around 25% (Yalden & Dougall
ringers!). Detailed observations suggest that
2004), despite both parents being present as
they are thermally independent by then,
guards. Young chicks need brooding every
though older chicks may still be brooded in
seven minutes or so, and for about seven
rain and at night. From about seven days old,
minutes each bout. By seven days, the chicks
the chicks may be closely attended by only
are mobile, weigh around 16 g and have a bill
one parent, the other going off to feed nearby
length of 14 mm; most (75%) survive from
(though still alert to intruders). By their third
week (the chicks usuallyfledge in 19 days), oftenonly one parent is leftguarding the chicks, andusually stays beyondfledging for another weekor so. When one parentleaves early, it is usually,but not always, the female.
Partial desertion is morelikely with later-hatchingbroods (Mee 2001).
Clearly, their territory
is ver y important toCommon Sandpipers;both adults defend itagainst their neighbours.
Moreover, it is demon-
strably the stretch of
shoreline that they are
34. A brood of three young Common Sandpiper
Actitis hypoleucos
defending: the four neigh-
chicks, about four days old and about 20 m away from their knownnest-site; June 2009, Ladybower Reservoir.
bours will feed together
British Birds 103 • February 2010 • 100–114
Common Sandpipers in Britain
comfortably in nearbypasture, but renew ter-ritorial aggressionwhen they return tothe river. An analysisof habitat suggeststhat they need an areaof shingle, and theadults spend muchtime patrolling thewater's edge in searchof food. However,their diet contains aneven mix of aquaticprey (caddisfly (Tri-choptera),
(Ephemer optera) andstonefly (Plecoptera)larvae) and terrestrial
prey (click beetles
35. A young Common Sandpiper Actitis hypoleucos chick, probably only a
(Elateridae), carabids
day or two old, still with a short (10 mm) bill; Whiteholme Reservoir,Yorkshire, June 2008.
earth worms (Lumbricidae)) (Yalden 1986),
and low recruitment. The Ashop had 20–22
so reliance on the water's edge to provide a
pairs in 1977–80. However, the severe late
food supply is not the whole story. The most
April (24th–26th) snowfall of 1981 apparently
prolonged territorial fight we have observed
surprised, and killed, a number of returning
(27 minutes) involved two pairs guarding
adults; the population dropped to 14 pairs
chicks, on 18th June 1989. One pair, dis-
that year, and recovered only slowly (by one
turbed by an angler, led its chicks into the
or two pairs a year) up to 1988 (Holland &
neighbouring territory, and to start with all
Yalden 1995). Another cold, late spring in
four birds were fighting (three of them were
1989 then caused another sharp decline, from
colour-ringed) (Yalden 1992a). One male
which the population has never properly
then led his chicks back, while his mate con-
recovered. Although there were 15 pairs in
tinued fighting for another 11 minutes. Such
2006, only five pairs were present in 2007 and
observations suggest that the most important
2008. Averaged over 28 years, the population
roles of the territory are toprovide the chicks with a
feeding area and good cover.
Territorial fights are certainlymore prolonged when chicks
are also present than early inthe season, when the briefest of
challenges is usually sufficient
to maintain territorial integrity.
The early years of study in thePeak District, and the early
years of the Waterways Bird
Survey (WBS), suggested a
Fig. 3. The long-term decline of the Ashop–Alport Common
stable population, as might be
Sandpiper Actitis hypoleucos population from 1977 to 2009, and
expected for a relatively long-
the fluctuating population in the Borders study site from 1993
lived species with high survival
British Birds 103 • February 2010 • 100–114
Dougall et al.
has declined by 59%. This matches what has
Survival rates, breeding success
happened more generally on the streams
around the Peak District, and the national
We have tried hard to understand the basis of
trend revealed by WBS and BBS (Breeding
the national decline, and the contrast in for-
Bird Survey), which suggests a 25% decline
tunes between the Peak District and Borders
over eight years (www.bto.org/birdtrends/
populations. Originally (1970s and 1980s)
wcrcomsa.htm). Over the shorter run of data
there was a strong correlation between sur-
from the Borders, there seems to have been
vival rates and minimum temperatures in
no decline, though much variability. Even
late April: fewer adults returned in cold
more puzzling, the LDH reservoir population
springs. Simple apparent survival rates
near the Ashop also shows no such decline.
(resighting of the previous year's colour-
There were 49 (±8) pairs in the early 1980s
ringed adults, on average 72% for males and
and 51 (±11) pairs in the 1990s and 2000s;
67% for females) declined between 80% and
indeed the highest count ever was as recent as
47% as late April mean minimum tempera-
2007, when 84 pairs were logged. In the
ture went down from 5.6 to 1°C (Holland &
Moorfoots, along a discontinuous 10.13 km,
Yalden 2002a). The variation in population
monitored annually between 1993 and 2007,
this produced matched changes in the North
the number of territories ranged between 14
Atlantic Oscillation (NAO); warm wet
in 2000 and 31 in 1993, and the percentage
winters (high NAO, perhaps snowier) are bad
which hatched at least one chick ranged
for sandpipers (Forchammer et al. 1998).
between 48% (in 2004) and 82% (in 2003)
However, in the 1990s, this relationship no
(Dougall et al. 1999; Dougall unpubl.).
longer fitted, though survival rates were still
While estimating the national population
just as variable, as were April temperatures
is fraught with difficulties (see below), it is
(Dougall et al. 2004). Not only did the
easier to map the breeding distribution.
Borders population not show the long-term
Between the two breeding atlases (1968–72
decline, but the considerable variations in
(Sharrock 1976) and 1988–91 (Gibbons et al.
populations there from year to year were not
1993)), the breeding range contracted, partic-
correlated with those in the Ashop popula-
ularly along the fringes of the species' range.
tion (Pearce-Higgins et al. 2009). If anything,
An apparent net loss of 14% of its breeding
survival rates were slightly higher in the
range in Great Britain and a substantial,
declining Ashop population than in the
though perhaps overestimated, 54% of its
Borders population, although the difference
Irish range suggests a considerable decline
was not significant.
(Yalden 1993). It will be fascinating to see
Breeding success is hard to evaluate pre-
what the third atlas reveals.
cisely; because well-grown chicks are so good
at running and hiding from any
threat, whether ringer or pred-ator, counts of older young or
fledglings are invariably underes-
timates. One calculation suggeststhat perhaps 35% of the actual
fledglings were not recorded(Yalden & Dougall 2004), another
that 30% were missed (Holland &
Yalden 1994). However, the fact
that their guarding parents are so
vocal suggests that at least the
Fig. 4. This figure shows the slight negative correlation of
success of breeding attempts can
annual adult survival for the Ashop birds with winter North
be reliably counted: a territory
Atlantic Oscillation (NAO) values (yellow circles denote first-
that has ‘alarming' parents over
year birds, i.e. new recruits, red circles represent older
three or four weeks has surely got
adults) (from Pearce-Higgins et al. 2009). A higher NAO valueindicates a warmer, wetter winter in western Europe, but
at least one chick through to
cooler, drier conditions in NW Africa.
fledging. On that basis, there is no
British Birds 103 • February 2010 • 100–114
Common Sandpipers in Britain
evidence that breeding success has been
mate at least once (Holland & Yalden 1994).
poorer in the Ashop study area than in the
The converse pleasure is of discovering
Borders study area over the decade or more
that individual birds which have failed to
that both have been studied. What is clear,
return to their territories have moved else-
however, is that recruitment of ‘new' birds, as
where (emigrated) rather than died. Obvi-
well as returns of fledglings, is very much
ously, this does not happen often. One 1980
higher in the Borders than in the Peak Dis-
female that had been presumed dead in the
trict. Only one of the 99 chicks and fledglings
1981 spring was found in 1982, about 20 km
ringed in the Ashop study area over the 1990s
away to the southwest in the Goyt Valley,
returned to breed there, but 49 of 421 ringed
where she returned in 1983 and 1984. In
in the Borders did so. Over the same period,
1990–93, ten birds from the supposedly site-
while 68 adults were recruited to the Ashop
faithful Ashop population contributed
population, there were 172 new adults ringed
further to the decline there by moving to
in the Borders (Dougall et al. 2005). The
LDH. One, having bred successfully along the
attempt to identify the source of new recruits
Ashop in 1992, as a new recruit, returned
to the Ashop population did find four birds
there briefly in April 1993, but in June 1993
bearing LDH light blue rings, including what
was at Derwent Reservoir, guarding a family,
has become the record for longevity. A 13-
as he was in 1994, 1995, 1996 and 1997. This
day-old chick ringed as NV54164 at Lady-
example highlights one of the mysteries of
bower Reservoir on 21st June 1992 turned up
this species. What are returning birds,
as a breeding male 7 km away in one of the
whether old hands or new recruits, looking
upper territories on the Ashop in May 1993,
for when they get back? If they were heading
with an unringed female. Both were caught
for familiar habitat, we might expect riverine
and individually colour-ringed, but their
birds to return to rivers and reservoir birds to
breeding attempt failed that year, and the
return to similar shorelines. Obviously, estab-
female was never seen again. The male didn't
lished birds return to their familiar territory,
return to the Ashop, but in May 1994 was
and get back to claim it as quickly as possible.
seen three times with an unringed female at
Often, they also meet their old mate there,
Ladybower, about 3 km south of where he
and experienced birds are more likely to
hatched. He was seen back in that territory
breed successfully together (Mee 2001).
every year to 2007, being last recorded on
Rarely, females get back before their mate,
19th June 2007; moreover, judging by his
and may then pair with an already estab-
‘alarming' behaviour in June, he bred suc-
lished (but different) male. Are birds that
cessfully in 11 of his 14 years at Ladybower,
move sites ones whose mate fails to return?
and at 15 years old was much the oldest
Are they looking to set up territories near
Common Sandpiper that we or anyone else
other, already established, birds or pairs, on
has recorded.
the basis that their presence signifies a suit-
This sort of resighting is one of the
able site? If the latter is true, it might explain
delights of studying this species. The birds
the geographical retreat from peripheral
are highly site-faithful (and if they were not,
parts of the range, in the Peak District cer-
the apparent survival rates would be much
tainly, but also in northeast Scotland and in
poorer). An early calculation was that 89% of
Ireland, as revealed by Gibbons et al. (1993).
returning males and 78% of returning
It is possible that some of the apparent con-
females came back to the same or an adjacent
traction in range was actually a consequence
territory (Holland & Yalden 2002a). So, in
of less thorough surveying of the thinly pop-
late April, checking the rivers to see who is
ulated periphery of the species' range in
safely back, and with whom they are paired,
1988–91, especially in Ireland, although local
is a special pleasure. With an average survival
accounts confirm losses in many areas.
rate of about 80%, adult life is likely to lastthree years, so the chances of both members
of a pair returning are only about 50%. Put
Given that chicks wear at least a scheme ring,
another way, a bird that holds territory for
any that return to our study areas are con-
three years is statistically ‘bound' to get a new
spicuous, and targeted (albeit not always suc-
British Birds 103 • February 2010 • 100–114
Dougall et al.
perhaps a large proportion, ofthem. This is an aspect of ourstudies where sightings fromothers would be especially valu-able.
Migration and wintering
This section deals with our biggest
uncertainties! There are ringing
recoveries of British birds moving
south in Britain, and from France,
Spain, Portugal and Morocco, in
July–October. Similarly, there are
recoveries of returning birds,
through Morocco, Spain and
France in March–April. These
include recoveries/resightings of
36. A well-grown Common Sandpiper Actitis hypoleucos chick,
our own birds in southern
within about two days of fledging (bill length 19.7 mm, wing
England, France, Iberia and
length 79 mm, but underweight at 27.5 g), showing remnants
Morocco. None of our birds has
of chick down on the nape and tail. The red ring above the
been seen in November–February,
BTO metal ring identifies this as a chick from the Borders
and, more surprisingly, there has
study site; Leithen Water, June 2005.
been just one recovery from West
cessfully) for recapture, identification and
Africa of the 19,000+ Common Sandpipers
full colour-ringing. This gives us a measure
ringed in Britain. This was of a bird ringed at
of natal dispersal – the distance between
Abberton Reservoir, Essex, in July 1964 and
hatching and establishment of a breeding ter-
recovered in Guinea-Bissau in September
ritory in later years – but this is clearly going
(though birds ringed at Abberton are mostly
to be an underestimate, since we are more
Scandinavian birds on passage, as nine other
likely to detect a ringed new recruit within
recoveries in Scandinavia testify). The nearest
our study areas than farther afield. Of 36
thing to a recover y in West Africa of a
Borders chicks that returned later to breed,
British-bred bird involves an unfortunate
they were between 0 and 39 km from where
chick ringed at Grassington, Yorkshire, on
they hatched, and at a median distance of 6
2nd June 1963. Its remains were recovered on
km. It is probably no coincidence that this is
20th November 1963 from the radiator of an
roughly the distance between the two parts,
aircraft that landed at Moscow airport after a
Heriot Water and Leithen Water, of the study
journey from Accra (Ghana) via Conakry
site. In the Peak District, the 35 returning
(Guinea), Bamako (Mali) and Belgrade. The
chicks were between 0.7 and 138 km away,
likelihood is that this young bird had been
with a median of 3.3 km, which is, similarly,
caught up in Guinea or Mali; this is roughly
roughly the distance between the Ashop and
where we would predict British birds to be in
the LDH reservoirs (Dougall et al. 2005).
winter on the basis of the SSW trend shown
How biased are these two figures? We have
by recoveries of Common Sandpipers ringed
had a few exchanges between the Goyt Valley,
elsewhere in Europe (Poland, Germany,
20 km to the southwest, and the Upper
Russia) and recovered in West Africa
Derwent, and there is a weak correlation,
(Holland & Yalden 2002b).
with a two-year lag, between population
Common Sandpipers move south in
changes in the Ashop population and those
summer very quickly. Our study sites are
in the Goyt. The most distant return, at 138
usually already thinly populated by early July,
km, was in north Wales. If our returns are
and typically empty before the end of the
spread so far, it is obvious that a search con-
month. Some have been recovered in Por-
centrated within 10 km or so of the sites of
tugal and Spain as early as mid July, and we
ringing is going to miss a proportion,
suspect that failed breeders may leave for
British Birds 103 • February 2010 • 100–114
Common Sandpipers in Britain
Africa very quickly. One Peak District bird
at St Abbs Head, Borders (Dougall & Yalden
was recovered, already back in Morocco, on
2007). These August falls are presumably of
15th June 1979. On the other hand, we also
Scandinavian birds on passage, but the July
have recoveries (and sightings of colour-
flocks are likely to be from more local
ringed birds) into August of sandpipers still
breeding areas.
in Britain, and we suspect that juveniles are
A small minority of Common Sandpipers
in less of a hurry to reach Africa.
remain in Britain overwinter. We know little
Over a period of 12 years, PKH succeeded
about these, either. On the basis that, in birds
in fitting colour rings to 60 full-grown birds
generally, those summering farthest north
near the mouth of the River Lune. It was
winter farthest south, whereas those in the
thought that these might be a sample of the
middle of the range move relatively little, we
100 pairs or their offspring that breed higher
would expect that Scandinavian birds overfly
up the Lune catchment. Yet none of these was
Britain. Birds wintering in Britain are more
ever seen on territory there in later years, nor
likely to be part of our summering popula-
were any of 38 birds ringed on those
tion. The chances of spotting one of our
breeding areas ever seen at the mouth.
colour-ringed breeding birds still in Britain
However, one was seen later breeding in the
in winter seem slight, but would be a worth-
Borders (see above) and another in
while target for those with telescopes or
Northumberland. Evidently, the mouth of
digiscoping equipment. If some of the
the Lune is a feeding-up site for birds from
Common Sandpipers overwintering in
the Borders. The mean weight of these birds
Britain could be caught and colour-ringed,
was 66 g, but the six heaviest weighed 80, 80,
they are perhaps more likely to yield a
81, 81, 82 and 84 g. Others have also reported
resighting in summer. Birds regularly
birds fattening up to over 80 g at sites away
hunting along reservoir shores can some-
from their breeding grounds (e.g. Brown
times be caught in walk-in traps or small
spring-traps (Dougall & Yalden 2008). Trap-
Relatively large concentrations of
ping may also confirm that wintering birds
migrating birds sometimes occur in autumn,
are site-faithful to their wintering territories,
for example 100 at Carron Valley Reservoir,
as suggested by one caught on Southampton
Upper Forth, on 7th July 1984; 64 at Endrick
Water on 6th January 1974 that was caught
Mouth, Clyde, on 25th July 1977 and 14th
there again on 18th March 1975. The ringing
August 1978; and 83 roosting at the River
expeditions to Djoudj, Senegal, in the early
Dumfries & Gal-loway, on 25th July2003. Occasionallythere are ‘falls' inthe Northern Islesand along the eastcoast of Scotland,most
during 18th–22ndAugust 2001, whenthere were 15 onFair Isle; seven onNorth Ronaldsay;50 on the Isle ofMay; 23, 11, nine
and eight at foursites in the East
Fife; 37. An adult (female?) Common Sandpiper Actitis hypoleucos executing a
seven at Skateraw,
‘double wing-up', showing the striking underwing pattern. This is both a
Lothian; and eight
threat and a courtship display. Whiteholme Reservoir, Yorkshire, April 2007.
British Birds 103 • February 2010 • 100–114
Dougall et al.
1990s showed that Common Sandpipers
(51%) were recorded (Dougall et al. 2005).
ringed there had a higher return rate (9/65 or
However, given the median natal dispersal
14% after one year) than any of the other
distance of 6 km, and a range up to 39 km, it
waders, implying that they were indeed site-
is not known if the ‘missing' 49% were in fact
faithful (Sauvage et al. 1998). Limited obser-
recruited elsewhere, or if mortality rates are
vations suggested that they established
higher than we assumed. Moreover, we must
territories of about 200 m in length along the
expect that first-year survival rates vary
banks of the Senegal River, and defended
widely, as they do for adults, with weather
them against neighbours, but there is scope
and other conditions, but we know nothing
for considerable work on the species' winter
of this. Colour-ringing studies of wintering
birds, comparing adult and first-year survival
We have estimated that around 54% of
rates, would be interesting. Although they
newly hatched chicks survive to fledging
reach adult size by the time they fledge,
(Yalden & Dougall 2004), and we have a good
juvenile sandpipers have more conspicuous
set of estimates for adult survival (more
barring on their primary coverts, so they
strictly, adult return rates) from resighting
would still be recognisable as young birds
our colour-ringed birds (see above). Because
when they reach Africa.
fledglings rarely return to their natal sites, at
Returning birds, as we remarked above,
least in the Peak District, we have no clear
seem to return quickly to their territories.
estimate of the survival rate of first-year birds
However, this is clearly a stressful time. In
over their first winter. When the Ashop pop-
addition to the suspected, and recorded,
ulation was stable, we could create a balanced
mortality in 1981, recoveries in other years
life table by assuming that 57% of young
are quite frequent in late April/early May. A
birds survived their first winter. Using that
conspicuous example is NV82841, ringed as a
assumed survival rate, we might have
chick on 8th June 1996 at Garvald Lodge,
expected 96 recruits to the Borders popula-
Heriot Water. He was not seen again until
tion from the 1993–2002 cohorts; and 49
2002, when he was breeding successfully, at
David Hutton
38 & 39. Two splendid portraits of Common Sandpiper Actitis hypoleucos ‘Light blue//BTO,
orange/black, photographed on migration at Seaswood Pool, Nuneaton, Warwickshire, on 15th April
2008. It had been ringed as a newly fledged juvenile at Ladybower Reservoir on 10th July 2002.
It was seen in its territory at the northeastern end of Derwent Reservoir in June 2004, June 2005,
May 2007 and June 2008, and was retrapped there on 28th May 2009.
British Birds 103 • February 2010 • 100–114
Common Sandpipers in Britain
Whitehope, 6 km to the south, and he was
rivers. A major problem, of course, is to
there again in 2003 and bred successfully
translate observed linear densities (per km of
with the same mate. On 22nd April 2004,
river or shoreline) into sensible figures for
though, he was found dead at Devizes, Wilt-
areas. Among a variety of estimates, we sug-
shire. His mate returned safely, took a new
gested that 1.6 pairs per monad (1-km
mate, was unsuccessful in breeding and was
square) was the most plausible (Dougall et
never seen again.
al. 2004), while the BBS suggested that 2% ofthe monads in England and Wales but 15%
Estimating the size of the British
of those in Scotland were occupied; there are
few data for Ireland, but we assumed that
Given our interest in the densities of birds in
5% of monads were occupied. The total
our study areas, it was a natural progression
occupancy for Britain & Ireland is then
to attempt to estimate the size of the total
19,500 monads, with a breeding population
breeding population of Britain & Ireland.
of 30,622 pairs, twice the number estimated
In the New Atlas (Gibbons et al. 1993), it was
in Gibbons et al. (1993). Moreover, a recal-
suggested that the population of a hectad
culation of the appropriate figure for the
(10-km square) might be 15 pairs, implying
1968–72 breeding atlas (Sharrock 1976) sug-
a population of 15,000 pairs in Great Britain
gests a population then of 39,600, and thus a
and 2,500 pairs in Ireland. This figure was in
decline of 23% between the two breeding
part suggested by our earlier estimate of the
atlases. A decline of 23% is also suggested by
Peak District population (11 pairs per hectad
the WBS, albeit between 1989 and 1999
averaged over 18 hectads), tempered by
(Dougall et al. 2004). Using the same
some acknowledgment of the fact that the
methodology, we estimated the nationally
species was clearly most abundant in Scot-
important Scottish population to be in the
land. Since the 1980s, many more surveys
range of 17,000–24,000 pairs, most probably
have been published, including both local
around 19,000 pairs (Dougall & Yalden
tetrad atlases and particular surveys of
David Hutton
British Birds 103 • February 2010 • 100–114
Dougall et al.
Causes of the decline
tering birds) and near our two study sites in
Our attempts to explain the decline of the
Britain, produced evidence of some effect of
national and Ashop populations by studying
climatic variation (NAO) on adult survival
the population dynamics of the Ashop birds
rates, and declining adult survival rates
and contrasting these with patterns in the
explained the overall population decline;
Borders have not been especially fruitful.
however, there was no evidence that any
Adult survival rates and breeding success
long-term trend in climate was responsible
seem comparable between the two areas
for the decline of the Ashop population
(Dougall et al. 2005). There is clearly a major
(Pearce-Higgins et al. 2009). Increased agri-
discrepancy in recruitment rates between the
cultural use of the wintering grounds or the
two areas, but we are not clear what underlies
stopover sites on migration might be
this. Possibilities include the poorer survival
affecting return rates. In Britain, it is evident
of first-winter birds in West Africa and
that the decline is most marked on smaller
poorer survival on migration. If there is
streams and rivers. Moreover, this retreat
poorer survival of young birds, is this a con-
from small streams has been proceeding for a
sequence of climatic changes? Alternatively,
century or more (Holloway 1996). Are these
are new recruits put off by finding poorer
sites more affected by agricultural change
breeding habitat on the rivers when they get
than larger sites? The species is certainly sus-
back so that, perhaps, they take territories on
ceptible to recreational disturbance, from
reservoirs instead? These ideas merit some
both anglers and picnickers (Yalden 1992a),
and this is regularly cited as a harmful factor.
A recent attempt to correlate changes in
The pair described earlier that led their
population size from year to year with
chicks into their neighbours' territor y,
climatic variation, both in Africa (for win-
inducing a prolonged fight, did so because an
angler was standing in the middleof their own territory. Conversely,the species responds quickly to theprovision of secure sites (islands,fenced areas). However, in the PeakDistrict, reservoir shores suffermore human intrusion than thestreams, so the cause-effect correla-tion is weak.
One interesting possibility is
that climate change might beaffecting the populations of somefreshwater prey. Many of thecommon stream larvae prefercolder water (Durance & Ormerod2007). Stream temperatures wouldbe much more sensitive to fluctu-ating weather than reservoirwaters, which are buffered by theirlarge volume. Stream invertebratesare also more exposed to changesin management regimes on neigh-bouring farmland. A regular
hazard of mist-netting at dusk usedto be getting dor-beetles Geotrupescaught in the nets; it was quite
usual to get three or four in an
40. An adult Common Sandpiper Actitis hypoleucos carrying
just a BTO metal ring – against the grey legs, these are easily
evening whereas now there are not
missed; Whiteholme Reservoir, Yorkshire, June 2008.
that many in a season. Are the cow-
British Birds 103 • February 2010 • 100–114
Common Sandpipers in Britain
pats now inhospitable to beetle larvae
sandpipers do migrate in long hops, it might
because of the use of ivermectin vermifuges
explain why relatively few are seen in July in
on cattle? Many smaller invertebrates also
southern England as they migrate south
emerge from dung, and are eaten by sand-
(although more are seen in August, these are
pipers. Reduced sheep numbers mean that
probably Scandinavian birds). There are
vegetation is now longer along some stretches
major uncertainties with this scenario. Some
of Heriot Water; chicks are harder to locate,
evidence from elsewhere in Europe suggests
but they might also be less able to feed freely.
that Common Sandpipers accumulate little
In places, the shingle is becoming overgrown,
or no weight at stopovers (e.g. Meissner
so that too is less suitable feeding habitat.
1997, Arcas 2001). Is this a strategy that
An entirely different scenario is suggested
varies among populations, or age groups?
for the wintering habitat by recent surveywork in West Africa. In some places at least,
wintering Common Sandpipers frequent
We have learnt much about our favourite
mangrove swamps and roost communally,
bird in over 30 years of study. What began as
though it is not certain whether they also
a very local concern (how many Common
defend territories while feeding. Trolliet &
Sandpipers are there in the Peak District and
Fouquet (2004) suggested that 15,000 feed in
how is the population balanced?) has
mangrove areas and another 8,000 on the
expanded to a second study (do Borders
mudflats of Guinea; this could be a substan-
birds vary the same way?), and to national
tial proportion of the British population, if
concerns (how many breed, how is the popu-
Guinea is where they winter. Mangroves are
lation changing?). Now we are faced with
highly productive, but also subject to intense
problems on a flyway scale. We suspect that it
pressure from coastal development, for fish
is this wider population that is declining, not
farming, timber, and agriculture. Apparently,
just the Ashop or British ones (Sanderson et
Common Sandpipers exploit the abundance
al. (2006) suggested a 49% decline across the
of fiddler crabs Uca in this habitat. Therefore,
Western Palearctic). Greater understanding
this habitat is highly susceptible to human
of the species will require a co-ordinated
intervention and could be that in which most
international effort. We urgently need to
of our birds overwinter. However, it is
locate the wintering areas and habitats of the
unlikely to be affected by winter rainfall or
bulk of our birds. If there are routine
temperature, as would be expected if the
stopover points during migration, these too
Sahel region (as at Djoudj) is their main win-
need to be established. The attachment of
tering area. Thus our failure to detect any
satellite or radio tags would be one, albeit
obvious relationship between year-on-year
expensive, way to do this (cf. the studies of
population change and weather variation in
White-rumped Sandpiper Calidris fuscicollis;
West Africa (Pearce-Higgins et al. 2009)
Harrington 1999). An alternative would
might fit with such a wintering habitat.
involve routine studies on the wintering
Migration remains an area of major
areas, similar to ours on the breeding sites.
uncertainty and concern. Fat-free mass seems
Given the numbers of ringers and other
to be about 40 g (Baccetti et al. 1992). If our
ornithologists visiting exotic locations, as
birds regularly accumulate 40 g of fat (as
well as dedicated expeditions and collabora-
implied by birds weighing over 80 g in July),
tive studies with local birdwatchers in West
and have an average wingspan of 350 mm, we
Africa, we hope that someone will take up
can calculate that they should have a flight
this challenge.
range of 6,000 km in still air (Pennycuick1989). This would easily take them to the
Senegal River mouth in one hop, farther if
We must thank the estates and farms on whose land
they exploit tailwinds. If this is optimistic,
we have worked for their permission to do so, and
two major refuelling stops in, say, Morocco
those who have helped with our fieldwork (as detailed
and Senegal would enable an easier migra-
in, for example, Dougall et al. 2005). We also thank thephotographers for the accompanying images. Sightings
tion; if such sites exist, they need to be identi-
of colour-ringed birds away from our study sites can
fied so that they can be protected. If
be reported to [email protected]
British Birds 103 • February 2010 • 100–114
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T. W. Dougall, 38 Leamington Terrace, Edinburgh EH10 4JLP. K. Holland, 32 Southlands, East Grinstead, West Sussex RH19 4BZD. W. Yalden, High View, Tom Lane, Chapel-en-le-Frith, High Peak SK23 9UN
British Birds 103 • February 2010 • 100–114
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Standards in Genomic Sciences (2014) 9:821-839 Genome analysis ofstrain GrollT a highly versatile Gram-positive sulfate-reducing bacterium Jan Kuever1, Michael Visser2, Claudia Loeffler3, Matthias Boll3, Petra Worm2, Diana Z. Sousa2, Caroline M. Plugge2, Peter J. Schaap4, Gerard Muyzer5, Ines A.C. Pereira6, Sofiya N. Parshina7, Lynne A. Goodwin8,9, Nikos C. Kyrpides8, Janine Detter9, Tanja Woyke8, Patrick Chain8,9, Karen W. Davenport8,9, Manfred Rohde10, Stefan Spring11; Hans-Peter Klenk11, Alfons J.M. Stams2,12
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