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58 © British Birds 98 • February 2005 • 58-72
ABSTRACT The Turtle Dove Streptopelia turtur has declined, in terms of both
range and abundance, throughout much of northern Europe over the last 30
years. Until recently, the species was relatively poorly studied in the modern
agricultural landscape in Britain. Concerns over its conservation status
prompted an intensive research programme to gather information on its
breeding ecology, identify the causes of the recent decline and make
recommendations to aid its recovery and conservation.Turtle Doves require
tall, overgrown bushes for nesting and short weed-rich areas for feeding, but
agricultural intensification has markedly reduced the availability and suitability of
these habitats. Over the last 40 years,Turtle Doves have switched from foraging
in ‘natural habitats’ to those created by humans, and their diet is now primarily
seeds from cultivated plants.The number of chicks fledged per pair per year
is today almost half that in the 1960s, while autumn migration has become
significantly earlier.Many management options to help to conserve the species
can be achieved within the current UK agri-environment policy framework.
Studies of
West Palearctic birds:
Turtle Dove
Stephen Browne and Nicholas Aebischer
Ray Scally
BB Feb 2005 21/1/05 1:07 pm Page 58
59British Birds 98 • February 2005 • 58-72
Studies of West Palearctic birds: Turtle Dove
23. Turtle Dove Streptopelia turtur,Austria, May.
Gordon Langsbury
The Turtle Dove Streptopelia turtur is a
summer migrant to Britain and Europe,
but perhaps most people in Britain
would associate the bird with the second day of
Christmas rather than high summer, thanks to
the popular carol The Twelve Days of Christmas.
This subversive, pro-Catholic hymn was sup-
posedly written at a time when it was illegal to
make direct reference to Catholicism, and the
‘two Turtle Doves’ are thought to represent the
Old and New Testaments. Historically, human
interest in the Turtle Dove was associated with
the bird’s faithfulness, and because it was an
emblem of hope and peace. The writings of
Saint Jerome (c. AD 320-420) and of Alain of
Lille, in his twelfth-century work The Complaint
of Nature, state that a Turtle Dove would not
seek another mate should its original partner
die. Indeed, it was thought that if a Turtle Dove
lost its mate then it too would soon die. It was
not until the eighteenth century that the species
became more noted for its migratory abilities,
being referred to in at least two of Gilbert
White’s letters. Today, however, most bird-
watchers in southeast England and much of
Europe would associate its purring calls with
warm summer evenings, rather than love or
peace. In recent years, the calls of this small,
attractive and once-common dove have been
heard less frequently and its population decline
and range contraction in Britain and
throughout Europe prompted The Game Con-
servancy Trust, with support from English
Nature and the World Wide Fund for Nature
(UK), to establish a dedicated research pro-
gramme focused on the species’ breeding
ecology. In this paper, we outline the biology of
the species based on the results of a number of
studies undertaken across Europe. We also
present the findings and recommendations of
our intensive four-year study, undertaken
between 1998 and 2001, to show how the
breeding biology of the species has changed
over the last 40 years.
Range and distribution
The Turtle Dove is a small, slim member of the
Columbidae, with a thin neck, a protruding
head and deep chest, giving it a pigeon-like
form, but with a long, wedge-shaped tail and
swept-back wings. The sexes are broadly similar,
but females are slightly smaller than males and
somewhat less brightly coloured. Four races of
Turtle Doves occur in the Western Palearctic,
with nominate turtur found throughout much
of Europe and western Asia. Slightly smaller
and paler, with more white on the belly, S. t.
arenicola occurs across North Africa (including
the Balearics) and into the Middle East. The
two remaining races are much more restricted
geographically, with S. t. hoggara (also smaller
and paler than nominate turtur, with broader
BB Feb 2005 21/1/05 1:07 pm Page 59
edging on the wing-coverts and on the tail)
inhabiting the central Saharan mountains, and
S. t. rufescens (more distinct than the other
races, being smaller and more richly rufous-
coloured) in Egypt and northern Sudan
(Cramp 1985; Gibbs et al. 2001). Both turtur
and arenicola are true migrants, spending the
winter (late October to late March) chiefly in
the Sahel region of Africa, roughly between
10°N and 20°N. The other two races are
thought to be sedentary or undertake only
small migratory movements (Cramp 1985).
In the breeding season (early May to mid
August), the range of turtur extends from the
Mediterranean to all but the most northern
areas of Europe. In Britain, at the northwest
edge of the species’ breeding range, it is a
lowland bird, favouring warm, dry conditions
and avoiding the higher ground and rainfall of
the west and north (Gibbons et al. 1993). Con-
sequently, Turtle Doves are concentrated in the
most intensively farmed, and now predomi-
nantly arable, areas of Britain.
Population trends in Britain and Europe
Survey and census work organised by the BTO
suggests that numbers of Turtle Doves in
Britain declined by 70% between 1968 and 1999
(Baillie et al. 2002), with a 25% contraction in
range between the 1968-1972 and 1988-1991
breeding Atlases (Gibbons et al. 1993). This
sharp decline commenced in 1979, following a
lengthy period of apparent increase and range
expansion from at least the mid nineteenth
century (Spencer 1965; Duckworth 1992; Hol-
loway 1997). Some local declines were apparent
from the 1950s (Goodwin 1989), but trends
from census work suggest that the period of
general increase lasted until about 1978/79
(Marchant et al. 1990). A few local studies
showed that numbers of Turtle Doves fluctu-
ated in a pattern similar to the national trend
(Hongell & Saari 1983; Miller 1992). From the
late 1970s, however, a sustained decline in the
breeding population in Britain became
apparent, and this continues today (Baillie et al.
2002) (fig.1). The decline has been so severe
that the majority of the British population is
now restricted to southern and eastern counties
of England and, by extrapolation from the
1988-91 census and subsequent rates of decline,
probably numbered only about 30,000 pairs in
2001. These population trends are of such
concern that Turtle Dove is now recognised as a
‘Red List’ species of conservation concern in the
UK (Gregory et al. 2002), and the UK Govern-
ment has placed it on the list of priority species
considered by the UK Biodiversity Action Plan.
One of the recommendations of the UK Species
Action Plan for the Turtle Dove is to undertake
an autecological research project to identify the
cause ofthe recent decline, in order to form a
recovery plan (Anon 1998).
This pattern of range expansion followed by
recent decline has been repeated elsewhere in
Europe, with evidence of local downturns being
noted from the 1950s onwards (Holzwarth
1971; Kraus et al. 1972; Cederwell 1978;
Hongell & Saari 1983; Bijlsma 1985; Yeatman-
Berthelot & Jarry 1995). Although census and
60 British Birds 98 • February 2005 • 58-72
Studies of West Palearctic birds: Turtle Dove
Fig. 1. Index of abundance of Turtle Doves Streptopelia turtur in Britain from 1963 to 1998, derived from
BTO Common Birds Census data.The index (solid line) and the 95% confidence limits (dashed lines) are shown.
63 65 67 69 71 73 75 77 79 81 83 85 87 89 91 93 95 97 99
CBC Index
2
1.5
1
0.5
0
BB Feb 2005 21/1/05 1:07 pm Page 60
migration data are often rudimentary or non-
existent, it is believed that, on a European scale,
the decline of the Turtle Dove occurred from
the mid 1980s onwards, particularly in western
Europe (Tucker & Heath 1994; Heath et al.
2000). Turtle Dove is now a Category 3 species
of European conservation concern (SPECs): a
‘species whose global populations are not con-
centrated in Europe, but have an Unfavourable
Conservation Status in Europe’ (Heath et al.
2000).
Previous research on the Turtle Dove
Until recently, the only major ecological study
of the Turtle Dove in Britain was completed in
the early 1960s (Murton et al. 1964; Murton
1968). In 1996, a pilot study was undertaken (by
The Game Conservancy Trust, funded by
English Nature) to collect basic information on
the ecology of the Turtle Dove and to test the
methodology for a more detailed study (Calla-
dine et al. 1997). The factors causing its recent
decline, in a modern agricultural environment
which has changed considerably since the 1960s
(Grigg 1989), and the stages of its life history at
which those factors operate, were therefore
unknown before a study undertaken by The
Game Conservancy Trust in the late 1990s
(Browne & Aebischer 2001; Browne 2002).
More widely, the breeding biology of the
Turtle Dove has been investigated in Bulgaria
(Nankinov 1994a, Nankinov 1994b), (the
former) Czechoslovakia (Pikula & Beklova
1984), France (Genard 1989), Germany
(Holzwarth 1971; Kraus et al. 1972), The
Netherlands (Bijlsma 1985), Portugal (Dias &
Fontoura 1996; Dias et al. 1996), Spain (Peiró
1990), Sweden (Cederwell 1978), and in the
southern Urals (Kotov 1974). Other studies
have looked at specific aspects of Turtle Dove
biology, including diet (Garzón 1974; Kiss et al.
1978; Jimenez et al. 1992; Dias & Fontoura
1996) and habitat requirements (Aubineau &
Boutin 1998). Turtle Dove migration was docu-
mented by, for example, Ash 1956, Rappe 1965,
Marchant 1969, Bourne & Beaman 1980,
Mountfort 1981, Genard 1989, Nankinov 1994b
and Aebischer 2002, and the biology of the
species on its wintering grounds in Africa by
Morel & Morel 1979, Morel 1985 and Morel
1987.
Migration
The Turtle Dove is the only British member of
the pigeon family to undertake a long-distance
migration, travelling up to 4,000 km twice a
year between breeding and wintering grounds.
The species’ annual migration, combined with
its exclusively granivorous diet, sets it apart
from all other farmland bird species in Britain.
After breeding, Turtle Doves begin to leave
Britain in August, with the last birds departing
61British Birds 98 • February 2005 • 58-72
Studies of West Palearctic birds: Turtle Dove
24. Turtle Doves Streptopelia turtur are found in a variety of habitats on their wintering grounds in western Africa,
including scrub, open woodlands,savannah and agricultural land;the open scrub with low-density cattle
browsing shown here (in The Gambia) would be a typical backdrop for encountering the species in winter.
Stephen Browne
BB Feb 2005 21/1/05 1:07 pm Page 61
in October and November. Their migration
route takes them through western France,
central and western Iberia, along the Atlantic
coast of Morocco, and through Mali and
Senegambia into western Africa. The main
migration period is between mid August and
late October, when most birds arrive in western
Africa. Migration is thought to be mainly noc-
turnal, but there is a diurnal component at
times (Guyomarc’h 1998). Turtle Doves tend to
migrate in groups of 5-30 individuals, but
flocks of over 100 are reported regularly; large
congregations, of up to 1,000, can occur in
food-rich areas, such as fields of ripe sunflowers
Helianthus (Aebischer 2002). Spring migration
commences during February-March, when
huge flocks of many thousands of birds can
occur in Mali and Senegambia (Curry 1974;
Morel & Morel 1979, 1988). Turtle Doves arrive
in Britain from mid April through to June, with
the main influx during May (see Aebischer 2002
for an overview).
In recent years, the autumn migration of
Turtle Doves in the UK has taken place eight
days earlier, on average, than in the 1960s
(Browne & Aebischer 2003a); concomitantly,
their breeding season is now some 12 days
shorter, on average, than 40 years ago (fig. 2).
For some bird species, it has been shown that a
recent increase in average temperatures, attrib-
uted to global warming, has altered the timing
of migration, but this relationship has not been
established for Turtle Doves (Tryjanowski et al.
2002; Browne & Aebischer 2003a) and it is
likely that factors other than temperature (e.g.
day length) may trigger the start of migration
for this species (Lofts et al. 1967). One conse-
quence of this lack of response in migratory
behaviour to increasing global temperature is
that, like some other migrant species, such as
Pied Flycatcher Ficedula hypoleuca (Both &
Visser 2001), the timing of the Turtle Dove’s
breeding season and food supply may now be
asynchronous. Studies have shown that the
fruiting of various plants now occurs earlier, in
response to increasing temperature (e.g.
Walther et al. 2002), and it is possible that the
peak period of Turtle Dove breeding is now out
of phase with the peak in food availability, a sit-
uation which may have contributed to the
species’ decline in recent years (Browne &
Aebischer 2003a).
Hunting
Little accurate information is available on the
numbers of Turtle Doves taken by hunters (see
Boutin 2001). The Turtle Dove is a traditional
quarry species in many European countries,
particularly those bordering the Mediterranean,
and hunting is legal in seven EU countries in
autumn: Austria, France, Greece, Italy, Malta,
Portugal and Spain. All these countries operate
an open season, and although national or
regional legislation controls the exact opening
date of that season, EU regulations ensure that
this is not before 15th August. This prevents
resident breeding populations from being tar-
geted specifically, and it is primarily migrating
62 British Birds 98 • February 2005 • 58-72
Studies of West Palearctic birds: Turtle Dove
Fig. 2. Length of residence of Turtle Doves Streptopelia turtur in Britain during the period 1963 to 2000.
The values represent the difference between mean arrival and departure dates at six bird observatories in
south and east England. Redrawn from Browne & Aebischer (2003a).
63 65 67 69 71 73 75 77 79 81 83 85 87 89 91 93 95 97 99
Length of residency (days)
130
120
110
100
90
80
BB Feb 2005 21/1/05 1:07 pm Page 62
birds which are shot. A bag limit per hunter (i.e.
the number each hunter can shoot legally per
day) is set in Greece, Portugal and Spain. In
addition, Turtle Doves are hunted illegally in
France during the spring. Although accurate
data on the numbers hunted each year do not
exist, and estimates are difficult, the total
annual hunting bag for the EU is thought to be
of the order of 2-4 million birds (Boutin 2001);
it is not possible to assess the long-term trend
of the EU hunting bag. In Morocco, the esti-
mated bag is up to 140,000 birds, and because
the open season runs from mid June to early
August, most are of the race arenicola. Hunting
of Turtle Doves is not permitted in their win-
tering areas, and information is so limited that
the numbers of any taken illegally cannot be
estimated.
Overwinter ecology
Little is known about the ecology of Turtle
Doves on their wintering grounds (see Cramp
1985 for overview), certainly in recent times.
The overwintering range of the migratory
turtur and arenicola is thought to extend from
Senegambia and southern Mali south to the
Gulf of Guinea, and from the Atlantic coast
east to Cameroon and Nigeria. Although areni-
cola probably occurs in the east of this range,
with turtur in the west, there appears to be con-
siderable overlap. Both races are known to
occur in Senegal and Mauritania during winter.
They are found in a variety of habitats,
including acacia scrub, open woodlands,
savannah plains and agricultural land. Large
flocks have been recorded feeding on wheat
fields, harvested rice fields and on quaysides
where peanuts are being loaded (Barlow et al.
1997). They eat a wide variety of seeds from
wild and cultivated plants. Huge flocks, of up to
a million birds, most likely to be arenicola,have
been recorded at food sources, water and at
roost, typically in acacia scrub or waterlogged
woodland. Turtle Doves tend to drink at first
light or at sunset, when the largest congrega-
tions are usually seen. Although they appear
able to feed during the intense heat of the day,
they will take shelter in shade under trees and
bushes during extreme heat (Curry 1974); they
often associate with Vinaceous Doves Strep-
topelia vinacea while foraging or resting (Morel
& Morel 1988). Recent poor rainfall in the
Sahel, deforestation, scrub clearance and
improvements to agriculture in western Africa
are thought to have contributed to the decline
of the species (Marchant et al. 1990; Jarry
1995), although this is unsubstantiated by any
recent studies.
Habitat use on the breeding grounds
The habitat requirements of Turtle Doves away
from the nest-site have been little studied. In
general landscape terms, Turtle Doves occur in
a range of open lowland habitats which are
interspersed with hedges, scrub and small
woods (e.g. Kraus et al. 1972, Bijlsma 1985).
They avoid open areas such as heathland and
also large tracts of extensive woodland, but
occur in young plantations and managed
(thinned) woodland (e.g. Genard 1989). Turtle
Doves appear to favour warm areas, being
found in those parts of Germany where the
mean daily maximum July temperature is at
least 17°C (Kraus et al. 1972), and in Britain
where the equivalent temperature is at least
19°C (Norris 1960).
An analysis of30 BTO Common Birds
Census (CBC) plots showed that, by 1995, the
mean density of Turtle Doves was 0.8 territories
per km2on 15 farmland plots and 10.1 terri-
tories per km2on 15 woodland plots (Browne et
al. 2004). Browne & Aebischer (2004) recorded
farmland densities of 4.1-4.6 territories per km2
in Suffolk and 0.3-0.5 territories per km2in Lin-
colnshire. Densities of 6.5 territories per km2in
woodland and over 40 territories per km2in
63British Birds 98 • February 2005 • 58-72
Studies of West Palearctic birds: Turtle Dove
25. Turtle Dove Streptopelia turtur,
Austria, June 2000.
Olaf Lessow
BB Feb 2005 21/1/05 1:07 pm Page 63
scrub have been recorded in northeast Essex
(C. F. Mason pers. comm.). Other reported den-
sities for Turtle Doves in Britain are 1.4 territo-
ries per km2on farmland and 2.2 in woodland
during 1968-72 (Sharrock 1976); corresponding
figures of 0.6 and 2.6 territories per km2during
1988-91 (Gibbons et al. 1993); and 1.6-2.1 terri-
tories per km2on farmland in eastern England
(Mason & Macdonald 2000). Other studies
across the species’ breeding range have shown
that Turtle Dove density varies from 1.4 to 30.0
pairs per km2in a range of wooded and farm-
land habitats (Holzwarth 1971; Kraus et al.
1972; Bijlsma 1985; Genard 1989; Dias & Fon-
toura 1996).
In Britain, it appears that suitable woodland
areas support densities of Turtle Dove terri-
tories up to 6.5 times higher than on farmland.
This probably reflects the species’ propensity to
nest in overgrown bushes, with woodland areas
being used predominantly for nesting and the
arable areas for feeding. Studies by Fuller &
Moreton (1987) and Hinsley et al. (1995)
showed that use of woodland by Turtle Doves
was governed by nest-site selection. Turtle
Doves were associated with shrubby vegetation,
avoided dense woodland and did not hold terri-
tories in coppice woodland until five years after
coppicing, when mean tree height was 5.6 m
and canopy cover was 66%. Within woodland
habitats, Turtle Doves appear to favour scrub
rather than pure woodland stands (Mason &
Macdonald 2000). Woodland area is also
important, with the minimum required being
around 0.2 ha (Mason 2001).
On farmland, based on information
extracted from 15 farmland CBC plots, density
was related to the amount of suitable nesting
habitat on each plot (Browne et al. 2004). As the
length of hedgerow and length of woodland
and scrub edge decreased within farmland,
Turtle Dove density also decreased (fig. 3).
The home range of British Turtle Doves is
highly variable (on average occupying some
83.5 ± 23.9 ha at one site and 497.3 ± 22.5 ha at
another site), depending on habitat availability,
and ranging from 0.3 ha to 1,130 ha (Browne &
Aebischer 2003c). Information from radio-
tracking suggests that Turtle Doves choose their
territories in a non-random way. Cropped
arable habitats were avoided and wooded ones
most preferred, relative to availability. The latter
reflects the species’ behaviour of loafing near
the nest-site or in tall trees and hedges, which is
also where most territorial behaviour is con-
ducted.
Social behaviour
Turtle Doves are monogamous and may form
pairs which last for more than one breeding
season (Cramp 1985). Some pairs are formed
during migration but, in most cases, it would
seem that the males arrive on the breeding
grounds first, begin to deliver their continuous
purring calls immediately and attract females
quickly. Males generally call most frequently
early in the morning and in the evening, and
from favoured song-posts (Calladine et al.
1999). The calls may be followed by a display
flight, which is usually triggered by the arrival
64 British Birds 98 • February 2005 • 58-72
Studies of West Palearctic birds: Turtle Dove
Fig. 3. Relationship between the amount of suitable nesting habitat and the density of Turtle Doves Streptopelia
turtur on 15 BTO farmland CBC plots during 1965-1995. Redrawn from Browne et al. (2004).
-2.00 -1.50 -1.00 -0.50 0.00 0.50 1.00 1.50 2.00
Change in the availability of suitable nesting habitat
Change in Turtle Dove density
0.25
0.20
0.15
0.10
0.05
0.00
-0.05
-0.10
-0.15
BB Feb 2005 21/1/05 1:07 pm Page 64
of another Turtle Dove, apparently to further
attract females or discourage males. The display
flight consists of a steep, upward flight, with
rapid wingbeats producing wing-clapping, fol-
lowed by a long, slow, circular descending glide
with wing- and tail-feathers outstretched,
returning the bird to its original take-off point.
If joined by a female, the calling male will also
undertake a bowing display, in which he fluffs
out his feathers, particularly around the throat,
bobs his head up and down and takes small,
two-footed jumps towards the female.
Pairs tend to aggregate in suitable nesting
habitat and adjacent nests can be as close as 3 m
(Cramp 1985). Further nesting attempts are
made close to the first nest, often in the same
bush and occasionally on top of the old nest.
Breeding activities, including nest-building,
incubation and tending the young, are under-
taken by both parents, although the male
appears to take on more of these activities than
the female. During the breeding season, birds
tend to roost near their nests, although occa-
sionally they roost communally. Soon after
breeding, young and adults disperse and do not
form family groups. Consequently, Turtle Doves
are usually observed singly or in pairs on their
breeding grounds, although small flocks may
form around food sources.
Breeding ecology
The breeding season begins upon arrival on the
breeding grounds in late April, with the peak of
nesting activity being in early June. Turtle
Doves defend only a small area immediately
around the nest-site, but utilise a much larger
area for foraging. The size of these defended
territories around the nest ranges from 0.75 to
8.90 ha, and they contain a higher proportion
of woodland, grassland and other non-crop
habitats, and a lower proportion of cropped
habitats, than expected by chance (Browne &
Aebischer 2004). The majority of nests are
found in hedgerows and scrub, with some
within young plantations and woodland, and in
isolated bushes; most are in thorny bushes, but
also in conifers, elder Sambucus and fruit trees
(Browne & Aebischer 2004; Browne et al. 2005).
The nest is usually a small platform of thin
sticks, built by the female with the occasional
assistance of the male. Sometimes other mater-
ials, such as thin wire and plastic, are utilised
too.Many nests are built with climbers as an
integral part of the nest (Aubineau & Boutin
1998; Browne & Aebischer 2004), and it is
believed that these provide additional support
to the nest structure. Nests are usually 0.1-20.0 m
above ground level, the majority being 1-3 m
high (Browne & Aebischer 2004). Nest height
65British Birds 98 • February 2005 • 58-72
Studies of West Palearctic birds: Turtle Dove
26. During the breeding season,Turtle Doves Streptopelia turtur will use a range of open lowland habitats.
Most nests are found in hedgerows and scrub,with smaller numbers in young plantations and woodland,
and in isolated bushes. Nests are typically in thorny bushes,but conifers,elder Sambucus and fruit trees
are also favoured;dense woodland is generally avoided.This photo also shows the type of broad,
weedy-edge habitat which is important for feeding.
Stephen Browne
BB Feb 2005 21/1/05 1:07 pm Page 65
varies according to habitat; for example, nests in
conifers tend to be higher as the lower levels of
conifers are generally unsuitable. The majority
of nests are located within 1 m of the edge of
the bush or tree.
Clutches vary from one to three eggs, with
two being typical. Turtle Doves may produce up
to three successful broods in a season, but may
nest more than three times if a brood is lost and
a replacement clutch laid. Incubation is under-
taken by both sexes and lasts for 14 days.
Nestlings are cared for by both parents and
leave the nest after 15 days. Birds are able to
breed in their second calendar-year. About half
the eggs laid are successful, with the remainder
being predated, deserted or failing to hatch
owing to infertility (Murton 1968; Cramp 1985;
Browne & Aebischer 2004).
The likelihood of various outcomes of
modern-day nesting attempts is different from
that during the 1960s (Murton 1968; Browne &
Aebischer 2004). During Murton’s study, 6% of
nests were abandoned, 56% of nests were lost to
predation and 39% fledged young, compared
with 15%, 37% and 47% respectively during the
late 1990s. In other words, abandonment rates
were lower and predation rates higher in the
1960s, but in the 1990s hatching success was
higher, yet fledging success was lower. Overall,
the success of an individual breeding attempt
appeared to be approximately the same in the
1990s as in the 1960s.
In recent years, the breeding productivity of
Turtle Doves has fallen dramatically. In the
1960s, Murton (1968) found that, on average,
Turtle Doves laid 2.9 ± 0.1 clutches per year,
which produced on average 1.6 ± 0.1 fledged
young per pair. Today, each pair produces 2.1 ±
0.3 clutches and 1.3 ± 0.2 young fledge per pair
(Browne & Aebischer 2004). This decline in
productivity is a result of fewer breeding
attempts and is related to a contraction of the
breeding season (see above and fig. 2); for
example, in the 1960s, 24% of nesting attempts
were started in August compared with just 5%
during the 1990s (Browne & Aebischer 2004; fig
4). This, in turn, links in with the species’
markedly earlier autumn migration from
Britain (discussed above), and implies that the
reduced number of nesting attempts and earlier
cessation of breeding identified by Browne &
Aebischer (2004) is likely to apply on a wide
geographical scale. Furthermore, an analysis of
approximately 1,900 BTO Turtle Dove Nest
Record Cards has shown that there has been no
discernable difference in any aspect of the
species’ breeding ecology over the last 40 years
(Browne et al. 2005). It is clear that the reduc-
tion in the number of nesting attempts per bird
per year has caused the reduction in breeding
performance.
Diet on the breeding grounds
Early studies identified seeds from wild plants
(weeds) as the main component of Turtle Dove
diet, with seeds of locally cultivated crops
making up the remainder. The seeds from two
species, Common Fumitory Fumaria officinalis
and Common Chickweed Stellaria media,were
particularly important (Murton et al. 1964). In
66 British Birds 98 • February 2005 • 58-72
Studies of West Palearctic birds: Turtle Dove
Fig. 4. Proportion of nests started in each half-month period of the Turtle Dove Streptopelia turtur breeding
season in the early 1960s (black bars) and the late 1990s (white bars). Redrawn from Browne & Aebischer (2004).
May 2 Jun 1 Jun 2 Jul 1 Jul 2 Aug 1 Aug 2
% of nests started
30
25
20
15
10
5
0
BB Feb 2005 21/1/05 1:08 pm Page 66
areas where cultivated seeds were taken, this
usually coincided with harvest time. Today,
seeds from cultivated plants (chiefly wheat and
rape) are the main constituents of Turtle Dove
diet, and comprise, on average, 69% of the seeds
identified in the faecal samples of nestlings, and
60% ofthe seeds taken by adults (Browne &
Aebischer 2003c; fig. 5). Forty years ago, the
percentages were 23% and 5% respectively
(Murton et al. 1964). It is likely that these dif-
ferences are real and do not reflect different
methodologies.
The foraging behaviour of Turtle Doves is
now markedly different from that recorded in
the 1960s, when they fed predominantly on
clover leys, hay fields and stooked wheat
(Murton et al. 1964). Today, they feed in a range
ofdifferent habitat types, but principally those
which include spilt grain: in farmyards, animal
feed and non-arable habitats. In almost all
cases, birds recorded on crops are feeding on
the weed strip around the edge of fields, weedy
areas that sprayers have missed, on crops that
have limited herbicide use (e.g. daffodils) and
on stubbles after harvest (Browne & Aebischer
2003c). Recent changes in agricultural practices
have removed or reduced many of the feeding
opportunities available in Murton’s day. At the
same time, increased use of herbicides and fer-
tilisers and more efficient screening procedures
have greatly reduced the abundance and diver-
sity of weeds and weed seeds throughout the
farmed environment, compared with 40 years
ago (O’Connor & Shrubb 1986).
Turtle Doves not only utilise different habi-
tats for feeding nowadays, but also undertake
relatively large foraging trips. On average, Turtle
Doves travelled 0.5-1.5 km per trip during the
1990s, but some feeding sites were located up to
10 km from the nest (Browne & Aebischer
2003c). Such relatively long trips probably
reflect a lack of suitable feeding opportunities
closer to nesting areas.
The greatest influence on these dietary
changes is likely to be the spatial and temporal
availability of food during the breeding season.
The seeds from crops are generally most widely
available immediately before, during and after
harvest, at the end of the Turtle Dove’s breeding
season. Earlier availability is dependent on spilt
grain, which usually occurs during its transport
from storage. When cereal prices are high, farm-
based grain stores are emptied earlier than
during times when prices are low. Conse-
quently, in some years, spilt grain may be avail-
able to Turtle Doves during the breeding
season, in other years it may not, and this
depends solely on market pressures. Cultivated
seed availability is not only limited and variable
temporally, but also spatially. Whereas a weed-
rich farming landscape would provide food for
Turtle Doves throughout their home range,
there may be only one or two patches of spilt
grain across a wide geographical area, forcing
birds to travel large distances between nesting
and feeding habitats.
Reasons for the species’ decline in Britain
Habitat use and feeding ecology
Hedgerow and scrub removal has long been a
67British Birds 98 • February 2005 • 58-72
Studies of West Palearctic birds: Turtle Dove
Fig. 5. Proportion of weed seeds in the diet of adult and nestling Turtle Doves Streptopelia turtur in the early
1960s (black bars) and the late 1990s (white bars). Error bars show ±1 standard error. Redrawn from
Browne & Aebischer (2003c).
Adult Nestling
% of weed seeds in diet
100
80
60
40
20
0
BB Feb 2005 21/1/05 1:08 pm Page 67
feature of agricultural intensification (Pollard et
al. 1974; Barr et al. 1993), and remaining hedges
are often intensively managed. Most of these are
cut annually (Macdonald & Johnson 2000), cre-
ating hedgerows which are usually too low and
not wide enough for the Turtle Dove’s needs.
The territory distribution of Turtle Doves
appears to be linked to availability of suitable
nesting habitat, primarily bushes and trees in
scrub, hedgerows, woodland edge, etc. Both the
quantity and the quality of suitable habitat have
declined in the modern farming landscape.
As described above, Turtle Doves now
occupy relatively large home ranges, undertake
long foraging trips and make little use of
natural feeding sites, being much more depen-
dent on (for example) spilt and stored grain
and animal feed. The results from experimental
work suggest, however, that, at the local scale,
food provided by humans does not appear to
improve breeding densities or success (Browne
& Aebischer 2002). Diet has changed as well as
feeding behaviour, from weed seeds to culti-
vated seeds (in broad terms). Although it
appears that diet does not have a direct effect on
breeding success, it seems likely that all the
factors outlined here will combine to have a
detrimental impact on the species.
Breeding ecology
Our data suggest that the reduction in length of
the Turtle Dove’s breeding season means that
the number ofnesting attempts would now be
expected to be at least 20% lower than in the
1960s. The observed reduction in the number
of clutches each pair lays is 45%, from 2.9 in the
1960s to 1.6 in the 1990s. In turn, the number
of young that successfully fledge per pair has
dropped from 2.1 in the 1960s to 1.3 in the
1990s. This reduction in breeding output would
lead to a per annum decrease of approximately
17% in the British breeding population of
Turtle Doves if replicated throughout its British
range. The ultimate cause of the reduction in
breeding output is not known, but it is possible
that changes in the spatial and temporal avail-
ability of food and the associated shift in diet
may affect adult body condition. Birds may
struggle to achieve and maintain suitable body
condition throughout the breeding season and
as a result may finish breeding earlier, although
this remains unproven (Browne & Aebischer
2003b).
Hunting
The limitations of the available data make it dif-
ficult to assess the importance of hunting. For
example, a worst-case scenario, using the lowest
European population and highest bag estimates
(Hagemeijer & Blair 1997; Boutin 2001) and a
breeding productivity of 1.3 fledged young per
pair (Browne & Aebischer 2004), could predict
68 British Birds 98 • February 2005 • 58-72
Studies of West Palearctic birds: Turtle Dove
27. First-winter Turtle Dove Streptopelia turtur,Scilly, October 2003.
Gary Bellingham
BB Feb 2005 21/1/05 1:08 pm Page 68
up to 40% of the autumn population being
killed, but the comparable best-case scenario
(highest population and lowest bag estimates)
predicts that only 4% is killed. Until accurate
information on the numbers of Turtle Doves
being hunted is available, it is impossible to
make a scientifically sound statement on the
contribution of hunting to the decline of the
Turtle Dove.
Changes on overwintering areas
For some migrants that overwinter in sub-
Saharan Africa, population declines have been
linked with drought on the wintering grounds
(e.g. Winstanley et al. 1974, Cowley 1979,
Marchant et al. 1990, Peach et al. 1991).
Decreased rainfall, rising temperature and other
changes linked to desertification in the Sahel,
perhaps linked to global warming, may indeed
have important consequences for Turtle Doves
breeding in Europe. Nonetheless, the decline of
the Turtle Dove did not immediately follow the
severe drought of 1968/69, when many other
species that winter in the Sahel suffered a severe
population crash (Marchant et al. 1990).
Indeed, Turtle Doves increased during the 10-15
years following the 1968/69 drought, and it was
not until 1980 that the population began to
decline (Marchant et al. 1990). Unlike the
species that were badly affected by that drought,
the Turtle Dove is solely granivorous; it is pos-
sible that its food supply was less severely
affected and it may have adapted to become
more mobile, and exploit new food sources,
such as rice and peanuts, that arose from
improved agriculture in the region (Cramp
1985; Marchant et al. 1990; Barlow et al. 1997).
In addition, forest and scrub clearance in over-
wintering areas, for agriculture and to provide
fuel, has also been cited as a possible cause of
the Turtle Dove’s decline (Jarry 1994), particu-
larly through the removal of suitable roosting
sites. The impact of this is likely to be relatively
small, however.
Common factors causing Turtle Dove and other
farmland bird declines
Recent studies have highlighted a number of
ecological changes that have affected Turtle
Doves and other farmland species alike. The
loss of nesting habitat, primarily hedges and
scrub,is thought to restrict the territory distrib-
ution of Grey Partridge Perdix perdix (Rands
1986), Red-backed Shrike Lanius collurio (Van-
69British Birds 98 • February 2005 • 58-72
Studies of West Palearctic birds: Turtle Dove
28. Edge habitats are now crucial for Turtle Doves
Streptopelia turtur in the modern agricultural
landscape. During the breeding season, most birds
recorded feeding on cropped land are concentrated
on the strip of weeds around the edge of fields,
weedy areas that sprayers have missed,farm tracks
or on crops that have limited herbicide use.
Stephen Browne
hinsberg 2000) and Yellowhammer Emberiza
citrinella (Bradbury & Stoate 2000) as well as of
Turtle Dove. A reduction in breeding perfor-
mance, brought about by a falling number of
nesting attempts per pair, has consequences for
the population size of Song Thrush Turdus
philomelos (Thomson & Cotton 2000) and
Linnet Carduelis cannabina (Moorcroft &
Wilson 2000), as well as of Turtle Dove. The
decrease in food availability, attributable either
directly or indirectly to pesticide use, has been
shown to affect Grey Partridge (Potts 1986), Sky
Lark Alauda arvensis (Donald & Vickery 2000),
Red-backed Shrike (Vanhinsbergh 2000), Yel-
lowhammer (Bradbury & Stoate 2000) and
Corn Bunting E. calandra (Brickle & Harper
2000), as well as Turtle Dove. Such findings
make it difficult to separate the decline of the
Turtle Dove, and of many other farmland bird
species, from changes in land management
associated with recent agricultural intensifica-
tion, although the potential negative effects of
hunting and changes on the overwintering
grounds cannot be dismissed.
Conservation measures
The Turtle Dove UK Species Action Plan (Anon
BB Feb 2005 21/1/05 1:08 pm Page 69
1998) had the following objectives and targets:
•In the short term, halt or reverse the decline
in numbers of the Turtle Dove by 2003 so
that the Breeding Bird Survey (BBS) index is
at least at 1996 levels;
•In the long term, see a sustained recovery in
numbers so that the BBS index is at least
50% higher than 1996 levels by 2008.
The following recommendations will help to
deliver these targets in the UK. Although tail-
ored towards Turtle Doves, these are also likely
to help other farmland wildlife.
Management options
Nesting habitat
Turtle Doves have specific requirements for the
types of tree and bush in which they nest, and
management needs to recreate suitable
hedgerows and patches of woodland or scrub,
and ensure that intensively managed hedgerows
are restored and replanted. Hedges should be
allowed to become overgrown with climbers
and reach a minimum height of 4.5 m and
width of 3 m.
Food availability
Turtle Doves like to feed amongst short, sparse
vegetation with a predominance of seed-rich
weedy plants. Management needs to encourage
patches of arable plants and provide alternative
sources of seed. Within or adjacent to arable
fields, areas should be tilled annually each
autumn and receive limited herbicide applica-
tions (only those targeted specifically at perni-
cious agricultural weeds) to allow the
establishment of weed-rich areas, with low open
vegetation cover. Alternatively, or additionally,
such areas can be established by sowing seed
mixes. Existing herb-rich grassland should be
protected and managed for a late hay crop (cut
after 15th July). These grasslands should be
encouraged by de-intensifying the use of
improved grassland or by arable reversion.
Post-harvest cereal, rape and pea stubbles
should be left until at least the end of August to
allow good pre-migratory feeding. Supplemen-
tary food, such as waste grain or tailings, is best
provided close to suitable nesting habitat.
Policy options
The current framework of agri-environment
policies allows many of the conservation man-
agement recommendations for Turtle Doves to
be incorporated within modern agricultural
systems. Ongoing reform of the Common Agri-
cultural Policy, scheduled for 2005, should
encourage farmers to manage land in a more
environmentally friendly fashion. This will
allow the introduction of compulsory cross-
compliance, whereby a farmer receives agricul-
tural subsidy (e.g. Arable Area Payments)
conditional on compliance with environmental
standards; payments may be reduced or can-
celled if standards are not met. Future policy
also needs to address those areas which are cur-
rently not adequately covered, including the
widespread management of arable field
margins, the re-establishment of arable fields in
pasture-dominated areas, the planting and
management of scrub and the provision of sup-
plementary food.
Acknowledgments
The study undertaken by The Game Conservancy Trust
was funded by The Game Conser vancy Trust, English
Nature, WWF-UK and, in part, by The Habitat Research
Trust. We thank all landowners, farm managers and their
gamekeepers for allowing access to their estates and
farms. Alastair Burn, Julie Ewald, Jim Fowler, Phil Grice and
Becky Halahan provided useful advice during the study.
Mike Armitage, Sarah Coe,Victoria Edward,Abigail Knight,
Stephen Murphy, Lynsey Robinson, Blair Urquhart,
Nicholas Watts and Georgios Yfantis assisted with the
fieldwork. We thank all the BTO bird ringers, CBC and
Nest Record Card surveyors and bird-observatory staff
for their efforts over the last 50 years in collecting some
of the data used here. We are also grateful for help from
the Bird Observatories Council (BOC), in particular Peter
Howlett and Dick Loxton, and the BTO, in particular the
Ringing Unit staff, Humphrey Crick, John Marchant and
Richard Thewlis.
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72 British Birds 98 • February 2005 • 58-72
Studies of West Palearctic birds: Turtle Dove
Stephen Browne and Nicholas Aebischer
The Game Conservancy Trust, Fordingbridge, Hampshire SP6 1EF
BB Feb 2005 21/1/05 1:08 pm Page 72