Bird Study

ISSN: 0006-3657 (Print) 1944-6705 (Online) Journal homepage: https://www.tandfonline.com/loi/tbis20

Evaluating the Breeding Bird Survey for producing

national population size and density estimates

Stuart E. Newson , Rick J.W. Woodburn , David G. Noble , Stephen R. Baillie &
Richard D. Gregory

To cite this article: Stuart E. Newson , Rick J.W. Woodburn , David G. Noble , Stephen R. Baillie
& Richard D. Gregory (2005) Evaluating the Breeding Bird Survey for producing national population
size and density estimates, Bird Study, 52:1, 42-54, DOI: 10.1080/00063650509461373

To link to this article: https://doi.org/10.1080/00063650509461373

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Bird Study (2005) 52, 42–54

Evaluating the Breeding Bird Survey for producing

national population size and density estimates

STUART E. NEWSON1*, RICK J.W. WOODBURN1, DAVID G. NOBLE1, STEPHEN R. BAILLIE1

and RICHARD D. GREGORY2
1BritishTrust for Ornithology, The Nunnery, Thetford, Norfolk, IP24 2PU, UK and 2Royal Society for the
Protection of Birds, The Lodge, Sandy, Bedfordshire, SG19 2DL, UK

Capsule The BBS has potential for producing better estimates of habitat-specific densities and population
sizes for many UK bird populations than those available previously.
Aims To examine the use of the Breeding Bird Survey (BBS) in providing unbiased national population
and habitat-specific density estimates of British birds.
Methods Line transect data collected by volunteers in 1998 from 2287 1-km squares across the UK were
analysed using distance sampling methods to calculate habitat-specific density and abundance estimates.
For each species, the habitat-specific decline in detectability with distance from a transect line was
modelled and applied at a regional level to incorporate variation in sampling intensity in different areas
of the country.
Results National population and density estimates calculated here were at a magnitude expected for at
least seven species in this study. However, national population size estimates were higher than expected
for Starling Sturnus vulgaris, House Sparrow Passer domesticus, Blackbird Turdus merula, Greenfinch
Carduelis chloris, Jackdaw Corvus monedula, Whitethroat Sylvia communis, Woodpigeon Columba
palumbus and Linnet Carduelis cannabina and lower than expected for Skylark Alauda arvensis,
Dunnock Prunella modularis, Song Thrush Turdus philomelos and Corn Bunting Miliaria calandra. These
differences are likely to be related to differences in sampling design and survey methods. For example,
Starling, House Sparrow, Blackbird, Jackdaw, Greenfinch and Wood Pigeon, which have considerable
populations in urban areas, were undoubtedly underestimated by the Common Birds Census (CBC). The
counts of species that flock during the breeding season, or are not strongly territorial (e.g. Linnet, Jackdaw
and Wood Pigeon) could be biased if detectability is strongly related to flock size. National population
estimates of Skylark and Corn Bunting are lower than those based on CBC, but higher, or similar, to
targeted national surveys of these species. Possible reasons for the differences between these estimates
are considered.
Conclusions This study highlights the strength of the BBS over previous data sources in producing
national estimates of density and abundance at the habitat and national level. More research on the sex
ratio and status of birds counted during surveys, and on the reliability of the detectability functions derived
from distance sampling is needed to improve the interpretation of population estimates derived from BBS
data.

The British Trust for Ornithology (BTO)/Joint Nature
Conservation Committee/Royal Society for the
Protection of Birds Breeding Bird Survey (BBS) was
introduced in 1994 to monitor population trends of a
broad range of breeding birds in the UK. Annual
population indices are calculated for all species where
there is sufficient coverage, and population trends are
*Correspondence author. Email: stuart.newson@bto.org
reported annually (Noble et al. 2001, Baillie et al.
2002). In order to make the maximum use of the data
collected, these analyses use all bird registrations.
However, through the recording of birds in different
distance categories, the BBS was also designed to
enable an assessment of variation in detectability and
hence to derive estimates of density and population
size.
For most common and widespread species in the UK,

© 2005 British Trust for Ornithology

 Using BBS to estimate density and population size 43

the New Atlas of Breeding Birds in Britain and Ireland: ulations mainly occur on farmland and for which there
1988–91 (Gibbons et al. 1993) provides the most recent is sufficient coverage to produce annually updated
estimates of density and national population size. population trends.
However, these estimates have a number of limitations.
The Atlas uses the mean density of bird territories on
METHODS
woodland and farmland Common Birds Census (CBC)
plots surveyed in 1989 to derive habitat-specific density
Data collection
and hence population size estimates for these habitats.
Because the CBC allows volunteers to select their own We use BBS data for 1998 during which fieldwork was
plots in mainly farmland and woodland, there are too undertaken on 2287 1-km squares. Full details of the
few plots to calculate density estimates for habitats survey design can be found elsewhere (Gregory et al.
other than woodland and farmland, so density and pop- 1996, Gregory & Baillie 1998). In brief, volunteers
ulation size estimates for these other habitats are based were provided with squares following a stratified
on observed densities in woodland and farmland. CBC random sampling design. Initially, the number of
coverage is also biased to the south and east of Britain, squares allocated to each of the BTO’s 83 regions
although there is some attempt to correct for this in the (roughly counties or groups of counties) was a fixed
calculation of population size. A major advantage of the proportion of the number of potential volunteers in the
BBS over the CBC is that it employs a formal sampling region, estimated using BTO membership information.
design with 1-km survey squares of the National Grid Within each region, squares are selected randomly and
selected at random, within each of 83 regions of the UK allocated to volunteers through a network of voluntary
and is stratified by observer density to allow participa- regional organizers (ROs). Fieldwork involves three
tion to vary across regions without introducing bias into visits by a volunteer observer to each survey square.
the analysis. The first is to determine two parallel ideal 1-km line
Secondly, an analysis of BBS and CBC data since
1989 shows that the long-term declines reported for
Table 1. A list of species in the analyses and information of their
many widespread farmland birds (Noble et al. 2001) status within the UK.
have continued over this period of time. This suggests
that national population estimates such as those in Population
Stone et al. (1997) based on data from the last Breeding change
Species Scientific name Code 1970–98 (%)1
Bird Atlas (Gibbons et al. 1993), regardless of its
limitations, are no longer appropriate, and need to be Kestrel 2,5 Falco tinnunculus K. –17
updated. This is particularly true for species included in Grey Partridge 2,4,6 Perdix perdix P. –82
the national Biodiversity Action Plan (BAP) Lapwing 2,5 Vanellus vanellus L. –52
Wood Pigeon 2 Columba palumbus WP 201
programme, such as Grey Partridge, Skylark, Song Turtle Dove 2,4,6 Streptopelia turtur TD –77
Thrush, Tree Sparrow, Linnet, Corn Bunting and Reed Skylark 2,4,6 Alauda arvensis S. –52
Bunting. Table 1 lists all the scientific names. Blackbird 3 Turdus merula B. –29
Relative habitat-specific densities were previously Song Thrush 3,4,6 Turdus philomelos ST –55
Whitethroat 2 Sylvia communis WH 34
estimated for eight species monitored by the BBS Jackdaw 2 Corvus monedula JD 148
during 1995 (Gregory & Baillie 1998). The percent- Starling 2,4 Sturnus vulgaris SG –58
ages of the British populations of sparrows, finches and Dunnock 3,5 Prunella modularis D. –21
buntings (10 species) in broad habitat classes were also House Sparrow 4 Passer domesticus HS –58
Tree Sparrow 2,4,6 Passer montanus TS –87
estimated from BBS data for 1996 (Gregory 1999). Greenfinch 2 Carduelis chloris GR –2
However, neither of these studies explores the uncer- Linnet 2,4,6 Carduelis cannabina LI –38
tainties that may bias estimates of this type, or presents Bullfinch 3,4,6 Pyrrhula pyrrhula BF –40
absolute population size estimates. Yellowhammer 2,4 Emberiza citrinella Y. –43
Reed Bunting 2,4,6 Emberiza schoeniclus RB –52
We extend these studies to examine the potential of Corn Bunting 2,4,6 Miliaria calandra CB –85
BBS data for producing unbiased estimates of habitat-
specific densities and population size for the UK. We 1Data from Common Birds Census (CBC). 2A farmland species indi-
use as examples, data from 20 species that commonly cator as per Gregory et al. (1999). 3A woodland species indicator
as per Gregory et al. (1999). 4Red-listed in Gregory et al. (2002).
use farmland. Species (shown in Table 1) were selected 5Amber-listed in Gregory et al. (2002). 6Priority species in the UK
to cover all widespread species of birds whose pop- Government’s Biodiversity Action Plan.

© 2005 British Trust for Ornithology, Bird Study, 52, 42–54

44 S.E. Newson et al.

transects through the square and two 1-km line
transects that will actually be surveyed. These actual
line transects should follow the ideal as closely as
possible, but allow for minor deviation due to impracti-
calities for the observer caused by land features or other
obstacles. The habitats of each 200-m section along
each of the ideal and actual line transects were also
recorded on this first visit using a hierarchical coding
system (Crick 1992).
Early morning bird counts are carried out on the
second and third visits, between early April and mid-
May and between mid-May and late June respectively.
It is recommended that the two visits should be at least
4 weeks apart. Counts begin at 06:00–07:00 hours
(GMT) so that they coincide with maximum bird
activity but avoid concentrated song activity at dawn.
Observers record all birds that they see or hear as they
walk along their transect routes within each 200-m
section in one of three distance categories from the line
(0–25 m, 25–100 m, 100 m or more) or in flight. Flying
birds obviously associated with the square (e.g. display
flight of Skylark) are assigned to the appropriate
distance band rather than recorded as in flight.
Distances are estimated at right angles to the transect
line. Juvenile and immature birds are recorded in the
field, but as far as possible they are excluded from the
computerized data and subsequent analyses. Thus, the
density and population estimates calculated in this
paper relate to breeding, or potentially breeding adults
only.
Distance sampling analysis
Distance sampling software developed by Buckland et al.
2001 (DISTANCE, Version 3.5; Thomas et al. 1998) was
used to produce estimates of density and population size
using BBS data. This technique models the decline in
detectability of birds with distance from the transect line
in order to include an estimate of undetected individuals
while estimating real density and hence population size
of the species of interest. As recommended by Buckland
et al. (2001), birds recorded in the final distance band
(100 m or more) were excluded from the analyses, as
counts within an unbounded category are difficult to
interpret. Birds in flight were also excluded. Count data
from the two remaining bands (0–25 m, 25–100 m) were
fitted with a half-normal detection function in all cases,
as recommended for binomial data of this kind
(Buckland et al. 2001).
For each species, the decline in detectability with
distance was modelled for each of 20 broad habitat
classes (Table 2). This was done because one might

Table 2. Habitat classes used in the analyses. Habitat classes are based on Crick (1992).

Code Name Description

WOOD1 Broadleaved woodland Broadleaved woodland

WOOD2 Coniferous woodland Coniferous woodland
WOOD3 Mixed woodland Broadleaved and coniferous (min. 10% of each)
SCRUB Scrub Young regenerated woodland, downland scrub, heath scrub, young coppice, young
plantation and clear-felled woodland (<5 m in height)
GRASS1 Dry semi-natural grassland Chalk downland, grass moor (unenclosed), grass moor mixed with heather and other dry
grassland
GRASS2 Wet semi-natural grassland Machair, water meadow, grazing marsh, reed swamp, saltmarsh and other open marsh
HEATH Heath Dry heath, wet heath, mixed heath and breckland
BOG Bog Bog, drained bog and bare peat
FARM1 Improved grass farmland Grass that has been regularly treated with fertilizer and distinguished by its bright colour,
lush growth and even texture
FARM2 Unimproved grass farmland Grass that has not been treated with fertilizer but is usually grazed or mown and may be
rank and neglected
FARM3 Mixed grass/tilled farmland Mixture of grassland and tilled farmland
FARM4 Tilled farmland Arable farmland including standing crop, plough and bare earth awaiting cultivation
FARM5 Other farmland Orchard and other farmland not specified above
URBAN Urban human sites Land relating to human habitation in a city or town
SUBURB Suburban human sites Land relating to human habitation on the outskirts of a city or town
RURAL Rural human sites Land relating to human habitation in the countryside, e.g. a village or hamlet
WATER Freshwater bodies Freshwater pond, lake, reservoir, stream, river, canal and ditch
COAST Coastal Marine open shore, inlet, cove, loch, estuary, brackish lagoon and open sea
ROCK Inland rock Cliff, scree/boulder slope, limestone pavement, other rock outcrop, quarry, mine
spoil/slag heap and cave
MISC Miscellaneous Sites not categorized above

© 2005 British Trust for Ornithology, Bird Study, 52, 42–54

 Using BBS to estimate density and population size 45

expect large differences in detectability between Table 3. A list of the UK level 1 NUTS (Nomenclature of Territorial
Units for Statistics) regions of the European Community used in the
habitats. For example, birds are likely to be detected at
analyses. The table also includes details of the number of squares
greater distances from the transect line in open country surveyed and the land-area in each region.
than they are in woodland or scrub. Habitat-specific
detectability curves were then applied to count data Squares Area % UK
Code Name surveyed (km2) by area
from BBS squares assigned to one of 12 NUTS
(Nomenclature of Territorial Units for Statistics) 71 North England 94 15 853 6.2
regions of the European Community (see Table 3) to 72 Yorkshire/Humberside 138 15 178 6.0
calculate a regional estimate of density for each habitat 73 East Midlands 154 16 112 6.3
74 East Anglia 130 13 091 5.1
and, by doing so, controlling for broad differences in
75 Southeast England 573 28 317 11.1
sampling intensity in different areas of the country. 76 Southwest England 295 24 408 9.6
BBS count data were pooled across bird recording visits 77 West Midlands 174 12 800 5.0
(early and late) to produce a mean density estimate 78 Northwest England 152 7 653 3.0
79 Wales 192 21 657 8.5
over these two visits. Counts were assumed to be a
7A Scotland 296 85 024 33.4
collection of sightings of individual birds rather than 7B Northern Ireland 82 13 991 5.5
taking the approach of modelling the detectability of 7C Channel Islands 7 261 0.1
flocks (see later for discussion of these assumptions). Total 2287 254 345 100
Although fitting habitat-specific detection curves is
preferable to the fitting of a single curve, this proved
difficult where there were few observations of birds in were calculated using a bootstrap resampling procedure
particular habitats. Since these species were considered of 400 iterations (Crowley 1992). For each iteration,
very rare in these habitats (e.g. Yellowhammer in urban squares were randomly resampled within region from a
habitat), it was considered appropriate to remove these list of squares in the original data with replacement. The
data (17 of 400 habitat–species combinations, habitat data and bird count data for the new sample of
Appendix) from the analyses. The resulting estimates squares forms the new data set of the iteration. The
of density and abundance were considered to be much entire process of analyses that was used to calculate the
more robust because of these exclusions. density and population estimates detailed above, is
Habitat-specific population estimates were calcu- repeated for each new sample to produce estimates by
lated by multiplying habitat-specific density estimates habitat across the UK and by NUTS region. We calcu-
for each NUTS region by the area represented by that lated 95% confidence intervals of the original estimates
habitat in each region. The area of each habitat within from the 400 new density and population size estimates
each NUTS region was calculated from the proportion for each habitat and for the total UK estimates. We cal-
that each habitat comprised of the total area of each culated 95% confidence intervals in a similar way for
region (see Table 3). The habitat data that the observer density and population size estimates by NUTS regions,
would have recorded had they walked the ideal transect but these are not presented here.
line through the square were used in these calculations Previous estimates of national population size in the
as this represents the true random sample of the region last Breeding Atlas or targeted national surveys were
(Field & Gregory 1998). based on the number of territories or pairs. To provide
National population estimates were calculated as the a broadly comparable measure of population size to the
sum of regional habitat-specific population estimates. numbers of adults estimated here, previous estimates
On the rare occasion that a habitat recorded by the based on territories or pairs are doubled.
ideal transect across all BBS squares was not surveyed
by the actual transect, a mean density estimate for that
RESULTS
habitat across NUTS regions was substituted as this
represents our best estimate for that habitat class. A
National population estimates
mean density of each species in each habitat across the
UK was estimated by dividing the habitat-specific National population size estimates with 95% confi-
national population estimates by the estimated total dence intervals for species derived from BBS data are
area of that habitat in the UK. Regional estimates of summarized in Table 4. We also present population size
density and population size are not considered here. estimates for these species from the literature. Of the 20
Confidence intervals for the estimates for each species species in this study, estimates for eight species

© 2005 British Trust for Ornithology, Bird Study, 52, 42–54

46 S.E. Newson et al.

Table 4. Comparison between previous national population size estimates (number of territories/pairs x 2) and new national population esti-
mates for 1998 calculated here with bootstrapped 95% confidence intervals. Figures are rounded to the nearest 10 individuals.

Estimates based on last Breeding Atlas

Species unless otherwise stated New estimates 95% CI

Kestrel 100 0001 77 420 62 040–156 130

Grey Partridge 300 000 265 040 229 630–746 320
Lapwing 370 000–476 000 436 310 325 390–574 290
Lapwing 265 450 2,3 436 310 325 390–574 290
Wood Pigeon 5 100 000 7 895 750 7 312 790–8 464 750
Turtle Dove 150 000 69 150 54 100–377 330
Skylark 4 000 000 2 649 650 2 350 650–2 939 730
Skylark 1 475 560–2 155 350 4 2 649 650 2 350 650–2 939 730
Blackbird 8 800 000 9 836 130 9 292 440–10 400 800
Song Thrush 1 980 000 1 233 160 1 124 520–1 351 030
Whitethroat 1 320 000 1,575,390 1 429 87–1 814 250
Jackdaw 780 000 2 504 340 2 211 950–3 016 970
Starling 2 200 000 9 231 150 8 181 740–10 458 580
Dunnock 4 000 000 3 350 010 3 138 410–3 632 590
House Sparrow 5 200 000–9 200 000 12 973 830 11 813 040–14 384 650
Tree Sparrow 220 000 283 980 218 490–760 400
Greenfinch 1 060 000 3 034 410 2 788 710–3 323 630
Linnet 1 040 000 2 258 410 2 015 950–2 658 200
Bullfinch 380 000 392 910 359 790–727 640
Yellowhammer 2 400 000 2 185 070 1 964 690–2 429 790
Reed Bunting 440 000 335 030 284 560–478 890
Corn Bunting 320 000 113 140 82 850–155 100
Corn Bunting 15 960–23 3205 113 140 82 850–155 100

1Gibbons et al. (1993) – based on an informed guess; 2Wilson & Browne (1999); 3Wilson et al. (2001); 4Browne et al. (2000); 5Donald &
Evans (1995).

(Kestrel, Grey Partridge, Lapwing, Turtle Dove, Tree
Sparrow, Bullfinch, Yellowhammer and Reed Bunting)
gave estimates whose confidence intervals encom-
passed the estimate based on the last Breeding Atlas.
Estimates for eight species (Wood Pigeon, Blackbird,
Whitethroat, Jackdaw, Starling, House Sparrow,
Greenfinch and Linnet) were larger than Atlas
estimates, whilst estimates for four species (Skylark,
Dunnock, Song Thrush and Corn Bunting) were
smaller than Atlas estimates. Population size estimates
for Lapwing, Skylark and Corn Bunting, for which
targeted surveys had been carried out in recent years,
were consistently smaller than Atlas estimates, but in
the case of Lapwing and Corn Bunting were still
considerably larger than estimates based on targeted
surveys of these species.
Habitat-specific density and population estimates
Of 22 330 200-m transect sections surveyed in 1998,
12 646 (57%) belonged to one of five farmland habitat
classes (Table 5). By taking the variation in coverage
between different NUTS regions into account, the
total estimated area of farmland in 1998 in the UK, as
defined by Crick (1998), was approximately 129 692
km2, (equivalent to over 50% of the total land area).
Habitat-specific density and population size
estimates for each of the twenty species in this study are
shown in Tables 6 & 7 respectively. We found that
more than 60% of Turtle Dove, Dunnock, White-
throat, Greenfinch and Linnet and over 90% of Grey
Partridge, Yellowhammer, Tree Sparrow, and Corn
Buntings occurred on farmland (Table 7 & Fig. 1).
Whilst Corn Bunting and Grey Partridge showed a very
high preference for arable land, Turtle Dove, Tree
Sparrow, Whitethroat, Linnet and Yellowhammer were
found on a number of farmland habitats, with scrub and
rural and urban habitats also holding important popu-
lations of these species. Although farmland was found
to hold important populations of all 20 species in this
study, the importance of rural, suburban and urban
habitats for a number of species is obvious. Whilst an
estimated 10% of the UK was classified as containing
habitat associated with humans (rural, suburban or
urban habitats), a large proportion (17–62%) of the
populations of Blackbird, Song Thrush, Dunnock,
Greenfinch, Starling, House Sparrow, Wood Pigeon
and Jackdaw were found in this broad habitat class.

© 2005 British Trust for Ornithology, Bird Study, 52, 42–54

 Using BBS to estimate density and population size 47

Table 5. Number of 200-m transects surveyed within each habitat class and the estimated area of each habitat in the UK (with bootstrapped
95% confidence intervals).

Name Transects surveyed Total habitat in UK (km2) 95% CI % UK by area

Broadleaved woodland 1125 10 715 9 577–11 954 4.2

Coniferous woodland 771 14 436 11 757–16 953 5.7
Mixed woodland 558 5 679 4 766–6 535 2.2
Scrub 487 6 929 5 502–8 365 2.7
Dry semi-natural grassland 860 13 681 11 442–16 102 5.4
Wet semi-natural grassland 164 1 850 1 297–2 522 0.7
Heath 1268 29 261 25 712–33 568 11.5
Bog 212 4 688 3 278–6 477 1.8
Improved grass farmland 4798 53 827 50 543–56 786 21.2
Unimproved grass farmland 1518 16 799 14 734–18 631 6.6
Mixed grass/tilled farmland 759 7 350 6 380–8 323 2.9
Tilled farmland 5259 49 356 46 446–52 779 19.4
Other farmland 312 2 360 1 873–2 956 0.9
Urban human sites 761 5 716 4 638–6 796 2.2
Suburban human sites 1670 12 988 11 371–14 697 5.1
Rural human sites 1068 9 205 8 302–10 240 3.6
Freshwater bodies 414 4 994 4 082–6 135 2.0
Coastal 163 2 094 1 343–2 963 0.8
Inland rock 121 2 107 1 443–3 000 0.8
Miscellaneous 42 309 173–461 0.1
Total 22 330 254 345 – 100

DISCUSSION is if a significant proportion of observers deviate from

Limitations and assumptions of BBS data and the
chosen approach
Producing reliable population estimates for wild bird
species is important for a number of reasons. In
addition to providing information for those interested
in the numbers of a species and for directing academic
research, population estimates are widely used for
setting conservation priorities and are important for
influencing policy and promoting the conservation of
birds. Within a country, the population status and
trends of a species may be examined by comparing
current population estimates with earlier estimates, as
in the third assessment of the population status of birds
in the UK (Gregory et al. 2002). Population estimates
may also be used to determine the national or inter-
national importance of a particular site or to compare
the status and trends of a species in one country to
those in other countries within the species’ range. This
approach has been used in Europe to identify species of
European conservation concern to prioritize those
species in most need of conservation action (Tucker &
Heath 1994).
There is no reason to believe that the BBS violates
the main assumptions of distance sampling and there-
fore our estimates are reliable (see Buckland et al. 2001
for a discussion). However, one potential source of bias
their ideal transect route due to land features to follow
linear or edge features. In this situation, an apparently
rapid fall in detectability may be related to the absence
of the critical feature for that species at greater
distances (e.g. hedgerows), which might inflate the
resulting density estimates. Moreover, recording birds
in two distance bands (0–25 m and 25–100 m) means
that it is not possible to test the data for violations of
these assumptions or to examine quantitatively the fit
of the model. It would therefore be useful to collect
more detailed distance and habitat data from a sample
of BBS sites for each species, to look for possible
violations of our assumptions. The resulting detection
functions could then be used to improve the precision
and confidence in existing estimates.
Our estimates of population size refer to the recorded
number of individuals, specifically adults. The assump-
tion that BBS observers correctly determine the age of
all birds cannot be tested, but data included in
our analysis might include some juveniles for early
breeding, mainly resident, species, especially during the
late visit (mid-May to late June). This would inflate
estimates for these species. Conversely, for migratory
species such as the Whitethroat which arrives in the
UK relatively late in the spring, numbers on the first
visit (April to mid-May) may be less than present later
in the season. Given these potential biases, it is

© 2005 British Trust for Ornithology, Bird Study, 52, 42–54

48 S.E. Newson et al.

Table 6. Habitat-specific density estimates of each species within the UK with bootstrapped 95% confidence intervals in parentheses. Units:
birds/km2 quoted to one decimal place for the estimate and to the nearest whole number for confidence intervals. Habitat and species
codes are explained in Tables 2 & 1 respectively.

Species density estimate (birds/km2)

K. P. L. WP TD S. D. B. ST WH

WOOD1 0.2 0.3 0.1 64.5 0.3 0.9 11.5 57.1 11.9 3.5
(0–5) (0–6) (0–0) (55–75) (0–1) (0–1) (9–14) (51–65) (8–14) (2–5)
WOOD2 0.1 0 0.6 22.7 0 0.8 7.3 12.7 5.9 0.5
(0–0) (0–0) (0–1) (17–30) (0–0) (0–1) (5–10) (10–17) (4–8) (0–1)
WOOD3 0.1 0.3 0.1 79.2 0.1 1.3 7.9 42.7 9.7 2.1
(0–7) (0–5) (0–0) (53–113) (0–0) (1–3) (5–10) (35–50) (7–13) (1–4)
SCRUB 0.2 0.8 0.3 29.7 0.2 2.7 14.4 31.1 7.1 9.5
(0–4) (0–28) (0–4) (21–39) (0–0) (1–5) (9–20) (23–40) (5–10) (6–13)
GRASS1 0.1 0.2 6.9 3.9 0 28.9 2.3 5.4 1.1 1.5
(0–4) (0–6) (2–14) (2–7) (0–0) (21–36) (1–4) (3–8) (1–2) (1–2)
GRASS2 0.2 0.3 10.8 13.6 – 19.1 13.2 22 3.4 8.6
(0–1) (0–1) (4–19) (7–25) (–) (7–33) (7–24) (12–32) (1–7) (3–15)
HEATH 0.5 0 1.9 0.9 0 16.6 0.8 1.2 0.6 0.5
(0–1) (0–0) (1–4) (0–2) (0–0) (12–21) (0–1) (1–2) (0–1) (0–1)
BOG – 0 0 1.7 0 22.6 1.5 2.1 2.4 0
(–) (0–0) (0–0) (0–6) (0–0) (12–35) (0–26) (0–5) (0–6) (0–0)
FARM1 0.3 1 1.3 25.2 0.1 5.6 16.7 39.6 4.9 5.6
(0–1) (0–1) (1–2) (22–28) (0–0) (4–7) (15–19) (37–43) (4–6) (5–7)
FARM2 0.3 1.6 4.1 25.3 0.2 11.6 10.5 31.4 3.4 4
(0–0) (1–26) (2–8) (19–34) (0–0) (8–16) (8–13) (27–36) (2–5) (3–5)
FARM3 0.2 1.4 1.1 50.2 0.2 9.7 23.7 53.1 7.2 14.4
(0–1) (0–3) (1–2) (39–68) (0–0) (7–13) (19–28) (45–63) (4–11) (10–19)
FARM4 0.4 3.1 1.8 46.1 0.7 18.9 16.6 32.5 2.8 15.3
(0–1) (2–4) (1–3) (41–53) (0–1) (16–22) (15–19) (29–36) (2–4) (13–17)
FARM5 0.3 2.2 1 52.6 2 5.7 24.9 68.3 10.8 9.2
(0–9) (0–106) (0–2) (33–78) (0–4) (3–9) (16–37) (51–92) (5–18) (5–17)
URBAN 0.2 0 0.1 37.1 0 0.5 15.7 103.1 6.6 0.9
(0–7) (0–0) (0–0) (28–48) (0–0) (0–1) (11–22) (83–128) (4–10) (0–2)
SUBURB 0.2 0.1 0.1 46.9 0.2 0.6 23 132.1 9.7 2
(0–0) (0–0) (0–2) (40–54) (0–0) (0–1) (19–27) (120–145) (8–11) (1–3)
RURAL 0.6 0.1 0.5 55.1 0.5 3.5 29.1 103.8 14.6 4.3
(0–1) (0–0) (0–1) (48–63) (0–1) (2–6) (25–34) (95–114) (12–17) (3–6)
WATER 0.2 0.3 1 41 2 1.9 13.8 43.3 6.8 8.4
(0–0) (0–1) (0–2) (29–56) (1–59) (1–3) (9–20) (32–55) (4–10) (5–13)
COAST 0 0 0.5 4.7 0 2.1 16.1 46.6 1.3 15.6
(0–0) (0–0) (0–1) (1–14) (0–0) (1–5) (2–32) (13–84) (0–3) (3–34)
ROCK 0.4 – 0 6.8 0 3.2 2.6 12.6 0.1 2
(0–12) (–) (0–0) (1–23) (0–0) (1–7) (0–27) (3–32) (0–0) (0–89)
MISC 0.4 0 4 8.3 0 19.4 – 23.9 – 0
(0–1) (0–0) (0–9) (1–153) (0–0) (3–35) (–) (7–44) (–) (0–0)

continued

remarkable that the estimates are so similar to those
from the last Atlas. As an indication of the level of
bias, we can compare the counts of Whitethroat from
the early and late visits. During 1998, 2127
Whitethroat were recorded during the early visit
compared with 2651 during the late visit. Data from
both visits were used in the analyses, meaning that the
resulting estimates are likely to be intermediate
between those calculated if early or late visit data only
were used. In the future it may be more appropriate to
use data from early visits for resident species and late
visits for migrant species and perhaps both visits for
‘intermediate’ species. Counts for intermediate species
such as Collared Dove Streptopelia decaocto may not be
biased in either of the two visits. Alternatively, it may
be most appropriate to treat data in relation to the
behaviour of each species, although this needs to be
examined and decisions made at the species level.
The BBS does not ask observers to record sexes sepa-
rately, indeed for many species examined in this study,

© 2005 British Trust for Ornithology, Bird Study, 52, 42–54

 Using BBS to estimate density and population size 49

Table 6 continued.

Species density estimate (birds/km2)

JD SG HS TS GR LI BF Y. RB CB

WOOD1 12.1 7.8 6.3 – 8.7 2 2.8 3 0.4 –

(7–19) (5–11) (4–10) (–) (6–12) (0–5) (2–4) (2–5) (0–6) (–)
WOOD2 2.6 0.3 3.5 – 3.7 0.9 1.6 0.9 0.1 0
(0–9) (0–1) (0–19) (–) (2–7) (0–6) (1–3) (0–2) (0–2) (0–0)
WOOD3 9 2.9 7.6 – 6 1.2 5.5 1.4 0.1 0
(5–15) (1–5) (3–14) (–) (3–10) (0–7) (3–9) (1–2) (0–0) (0–0)
SCRUB 2 9.4 6.6 0 6.6 9.9 5.3 6.4 2.1 0
(1–4) (4–17) (3–13) (0–0) (4–9) (5–16) (2–10) (4–10) (1–4) (0–0)
GRASS1 2.4 3.8 1.4 0 0.8 3.2 0.7 1.3 2.3 –
(1–4) (2–7) (0–3) (0–0) (0–2) (1–6) (0–2) (1–2) (1–5) (–)
GRASS2 5 13.9 12.5 0 7.7 12.4 – 1.6 17.3 0
(2–12) (6–29) (2–29) (0–0) (2–15) (5–23) (–) (0–5) (9–27) (0–0)
HEATH 0.4 1.3 0 0 0.4 3.6 0.2 0.7 0.5 0
(0–1) (0–3) (0–1) (0–0) (0–4) (1–9) (0–5) (0–1) (0–1) (0–0)
BOG 0.7 0.4 0.2 0 0.1 0 0.5 0 1.4 0
(0–20) (0–22) (0–0) (0–0) (0–0) (0–0) (0–19) (0–0) (0–29) (0–0)
FARM1 13.6 39.5 37.6 2 11.7 10.6 1.6 8 1 0.1
(11–17) (31–50) (32–43) (1–4) (10–14) (8–13) (1–2) (6–9) (1–2) (0–0)
FARM2 15.3 38.2 35.9 2.4 9.7 8.7 1.5 5.8 2.1 0.1
(10–23) (22–60) (24.7–47.4) (1–6) (7–12) (6–12) (1–2) (4–8) (1–4) (0–0)
FARM3 11.9 30.6 60.6 2.5 19.7 17.4 1.5 24.4 1.4 0.2
(7–18) (22–40) (44–77) (1–28) (14–27) (11–26) (1–3) (18–30) (0–4) (0–0)
FARM4 5.7 13.7 26.2 1.6 11.3 17.1 1.3 24.8 1.7 2
(4–8) (11–17) (21–32) (1–3) (9–13) (14–20) (1–2) (22–27) (1–2) (1–3)
FARM5 7.5 44.9 72.1 3.8 26.4 19.5 2.6 10 0.3 0.2
(3–14) (28–60) (47–100) (0–142) (18–36) (8–38) (1–46) (6–18) (0–1) (0–1)
URBAN 18.4 223.3 345.3 0 25.5 5.4 0.6 – 0.2 0
(10–29) (167–288) (275–440) (0–0) (19–34) (3–10) (0–4) (–) (0–1) (0–0)
SUBURB 26.9 214 324.4 0.1 40.1 5.4 1.2 0.2 0.1 0.2
(15–41) (176–261) (276–378) (0–1) (34–46) (3–8) (1–20) (0–0) (0–1) (0–0)
RURAL 34.7 99.7 194.8 2.2 53.1 10.6 2.8 5.9 0.8 –
(26–44) (79–132) (167–226) (1–4) (45–61) (7–15) (2–4) (4–9) (0–2) (–)
WATER 6.9 30.7 25.3 1.4 8.6 7 2.9 4 7.1 0
(3–13) (16–45) (15–36) (0–37) (6–14) (3–15) (1–33) (2–7) (3–12) (0–0)
COAST 8.1 17.2 33.3 0 5.6 3.8 – 2.2 1.9 0.6
(0–32) (5–43) (7–81) (0–0) (1–12) (1–10) (–) (1–5) (0–6) (0–2)
ROCK 8.4 0 – 0 1.4 1.7 0.3 2 – 0
(1–133) (0–0) (–) (0–0) (0–31) (0–42) (0–6) (0–73) (–) (0–0)
MISC 1.7 25.9 45 0 – 2.1 0 24.4 0.7 6.1
(0–4) (0–1395) (2–188) (0–0) (–) (1–5) (0–0) (3–59) (0–2) (2–17)

it would not be possible to distinguish sex in the field.
However, where there is a large difference in detect-
ability between the sexes, the resulting figures could
underestimate the true density and population size for
that species. In the case of the Skylark for
example where singing males are highly detectable, but
females much less so, the resulting population size
estimates may be close to the number of adult males
only. We also assume that BBS data are a collection of
sightings of individuals. It could be argued that the BBS
is designed to monitor breeding birds, which for the
majority of species tend not to flock at this time of year.
However, if there is a considerable difference in
detectability between different flock sizes, this could
bias detection functions and hence the resulting
estimates. A study that examines sex-specific detect-
ability and importance of flocking is required to
examine the extent to which these bias the resulting
estimates.
Comparison with existing estimates
In the past, there has been a tendency for population
size estimates to be reported with little or no discussion

© 2005 British Trust for Ornithology, Bird Study, 52, 42–54

50 S.E. Newson et al.

Table 7. Habitat-specific population estimates of each species within the UK with bootstrapped 95% confidence intervals. Units: 1000
birds quoted to one decimal place for the estimate and to the nearest whole number for confidence intervals. Habitat and species codes are
explained in Tables 2 & 1 respectively.

Species population estimate (thousands of birds)

K. P. L. WP TD S. D. B. ST WH

WOOD1 1.6 3 1.1 691.3 2.9 9.7 122.9 611.8 127.5 37.1
(1–49) (0–69) (0–3) (590–821) (1–5) (7–13) (96–147) (528–703) (99–155) (24–55)
WOOD2 1.4 0 8.8 327.4 0.4 12 105.3 183.2 84.4 7.5
(0–2) (0–0) (0–12) (231–431) (0–1) (6–19) (65–156) (135–240) (54–116) (2–17)
WOOD3 0.7 1.6 0.4 449.9 0.8 7.2 44.8 242.4 55.2 12.1
(0–40) (0–27) (0–2) (291–712) (0–2) (4–16) (31–60) (187–294) (40–74) (6–23)
SCRUB 1.4 5.7 1.9 205.9 1.1 18.5 99.5 215.7 49.4 65.6
(0–25) (1–187) (0–30) (152–272) (0–2) (8–34) (64–135) (164–264) (33–68) (43–92)
GRASS1 1.9 2.4 94.6 53.7 0 394.7 31.3 73.2 15 20.5
(1–8) (0–73) (32–178) (30–90) (0–0) (264–530) (15–51) (49–104) (7–26) (10–33)
GRASS2 0.4 0.6 20 25.1 – 35.4 24.4 40.7 6.3 15.8
(0–1) (0–2) (7–37) (13–43) (–) (11–72) (11–42) (20–62) (2–14) (5–28)
HEATH 15.5 0 55.4 26.2 0.2 484.9 22 35.7 16.7 14.4
(0–24) (0–0) (21–108) (9–51) (0–0) (339–661) (10–38) (16–65) (7–30) (0–24)
BOG – 0 0 7.7 0 106.1 6.8 9.6 11.1 0
(–) (0–0) (0–0) (1–24) (0–0) (49–161) (1–128) (2–23) (1–30) (0–0)
FARM1 18 50.9 72.2 1355.9 5.8 299.3 899 2132.6 261.8 302
(11–29) (25–79) (49–100) (1180–1524) (2–12) (220–384) (809–1014) (1939–2349) (218–314) (247–371)
FARM2 4.5 27.4 68.2 425.7 3.5 194 176.7 526.8 56.8 67.5
(3–7) (12–42) (26–129) (319–563) (1–7) (123–264) (140–216) (448–604) (40–75) (48–87)
FARM3 1.7 10.2 7.7 368.8 1.3 71.5 174 390.4 52.6 106.2
(1–5) (3–23) (4–12) (272–498) (0–3) (47–100) (134–216) (323–469) (33–82) (74–144)
FARM4 19.1 154.4 89.5 2273.4 32.1 934.5 819.8 1604.8 136.6 755.8
(13–28) (116–197) (61–133) (1990–2650) (21–40) (793–1081) (712–923) (1429–1772) (108–169) (655–852)
FARM5 0.6 5.2 2.4 124.1 4.7 13.4 58.8 161.2 25.5 21.6
(0–20) (0–222) (0–6) (72–192) (1–10) (7–24) (35–92) (106–239) (11–47) (10–39)
URBAN 1.2 0 0.4 212 0 2.6 89.6 589.2 37.4 5
(0–43) (0–0) (0–2) (147–283) (0–0) (0–3) (56–128) (431–779) (22–58) (2–11)
SUBURB 2 1.2 1.8 609.6 2.3 7.3 298.7 1715.7 125.7 26
(1–5) (0–4) (1–23) (498–729) (0–5) (3–14) (241–364) (1458–2014) (97–152) (14–42)
RURAL 5.5 1.2 4.9 507.5 4.2 32.4 268.2 955.4 134.2 39.3
(3–10) (0–4) (2–11) (429–592) (2–8) (18–56) (220–317) (835–1089) (108–162) (29–52)
WATER 1 1.3 4.8 204.8 9.9 9.2 69 216.4 34.1 42.1
(0–2) (0–3) (2–10) (141–284) (3–314.9) (4–15) (43–108) (156–273) (18–52) (24–68)
COAST 0 0 1 9.9 0 4.3 33.6 97.5 2.7 32.7
(0–0) (0–0) (0–2) (2–31) (0–0) (1–10) (4–74) (25–201) (1–7) (6–73)
ROCK – – 0 14.2 0 6.6 5.5 26.5 0.2 4.1
(–) (–) (0–0) (2–48) (0–0) (2–15) (0–61) (6–62) (0–1) (0–195)
MISC 0.1 0 1.2 2.5 0 6 – 7.4 – 0
(0–0) (0–0) (0–3) (0–42) (0–0) (1–12) (–) (2–15) (–) (0–0)
Sum 77.4 265 436.3 7895.7 69.2 2649.6 3350 9836.1 1233.2 1575.4

continued

as to the limitations of the data or analyses on which
they are based. Obviously the greater number of differ-
ing population estimates that are ‘in use’, the greater
the potential confusion, and the potential damage to
the credibility of population estimates in the wider
public domain. By highlighting the limitations and
uncertainties in the data and analyses presented here, it
is hoped that the reader is not only aware of these
issues, but that through this awareness steps are taken
to address these before further estimates of this type are
adopted for general use.
Habitat-specific density estimates presented here are
similar to those in Gregory & Baillie (1998) and
Gregory (1999), suggesting population size estimates, if
they were to be derived from these studies would
be similar to those presented here. However, this
comparison is not particularly informative, because
these studies are all based on BBS data, use similar

© 2005 British Trust for Ornithology, Bird Study, 52, 42–54

 Using BBS to estimate density and population size 51

Table 7 continued.

Species population estimate (thousands of birds)

JD SG HS TS GR LI BF Y. RB CB

WOOD1 129.9 83.1 67.4 – 93.4 21.6 30.2 31.8 4.3 –

(72–207) (53–123) (39–107) (–) (66–130) (5–58) (18–42) (17–49) (0–69) (–)
WOOD2 37.6 4.3 50.6 – 52.9 12.8 23.3 13.2 0.8 0
(7–125) (1–13) (5–253) (–) (25–93) (1–91) (9–42) (5–26) (0–30) (0–0)
WOOD3 51.2 16.5 43.2 – 34.3 6.8 31.1 8.2 0.3 0
(28–85) (8–29) (15–80) (–) (20–55) (2–30) (16–53) (3–14) (0–1) (0–0)
SCRUB 13.7 65.4 45.4 0 45.6 68.8 36.8 44.6 14.3 0
(6–25) (28–122) (18–92) (0–0) (26–64) (36–106) (14–66) (26–66) (5–28) (0–0)
GRASS1 33.1 52.2 18.6 0 11.1 43.1 9.9 17.5 31.3 –
(17–55) (29–94) (5–37) (0–0) (5–19) (18–80) (1–29) (9–28) (12–64) (–)
GRASS2 9.3 25.7 23.2 0 14.2 22.9 – 3 31.9 0
(4–22) (12–52) (3–59) (0–0) (3–26) (8–41) (–) (1–8) (15–53) (0–0)
HEATH 11.3 37.6 0.5 0 11.7 103.8 6.7 19 14.8 0
(3–33) (8–101) (0–22) (0–0) (2–115) (35–249) (3–139) (4–37) (4–29) (0–0)
BOG 3.2 2 0.7 0 0.5 0 2.2 0 6.8 0
(0–77) (0–108) (0–2) (0–0) (0–1) (0–0) (0–79) (0–0) (1–141) (0–0)
FARM1 731.1 2127.2 2021.5 108.7 630 570.8 85.6 429.4 51.4 5.3
(568–895) (1600–2706) (1728–2342) (60–194) (525–728) (450–701) (60–115) (341–514) (31–82) (4–20)
FARM2 256.2 641 603.3 39.5 162.8 146.6 25.1 97.6 35.1 1.1
(170–390) (383–975) (427–793) (8–109) (118–204) (103–201) (15–38) (66–137) (18–61) (0–4)
FARM3 87.3 224.5 445.5 18.3 144.6 127.9 11.2 179.4 10.6 1.3
(52–133) (160–304) (313–585) (6–188) (99–201) (83–198) (6–20) (132–234) (3–28) (0–3)
FARM4 279.1 674.9 1294.1 80.2 558.1 841.9 64.9 1224.7 83.9 99.2
(184–399) (525–859) (1021–1565) (46–127) (453–669) (687–1009) (44–98) (1064–1365) (58–115) (64–134)
FARM5 17.7 105.9 170.2 9.1 62.3 46 6.1 23.6 0.7 0.6
(8–31) (64–146) (106–252) (0–327) (37–96) (17–97) (1–118) (12–46) (0–2) (0–1)
URBAN 105.1 1276.5 1973.7 0.1 145.9 31 3.4 – 1.1 0
(55–173) (903–1692) (1450–2637) (0–0) (96–208) (15–58) (1–22) (–) (0–3) (0–0)
SUBURB 349.9 2778.9 4212.8 1.2 520.4 70.2 15.8 2.1 0.9 2.6
(191–565) (2117–3513) (3326–5292) (0–16) (419–635) (39–110) (8–275) (1–5) (0–10) (0–3)
RURAL 319.1 918.1 1793.4 20 489.1 97.2 25.8 54.7 7.5 –
(237–412) (712–1202) (1511–2083) (7–37) (412–566) (67–137) (16–38) (34–85) (2–17) (–)
WATER 34.7 153.3 126.1 6.9 43 34.7 14.2 20 35.3 0.1
(13–63) (82–223) (74–185) (0–169) (25–75) (14–75) (5–15) (10–35) (17–63) (0–0)
COAST 16.9 36.1 69.7 0 11.6 8 – 4.6 4 1.1
(0–66) (9–101) (11–183) (0–0) (2–35) (2–19) (–) (1–10) (1-13) (0-4)
ROCK 17.6 0 – 0 2.9 3.6 0.6 4.3 – 0
(1–289) (0–0) (–) (0–0) (0–54) (1–78) (0–10) (0–162) (–) (0–0)
MISC 0.5 8 13.9 0 – 0.6 0 7.5 0.2 1.9
(0–1) (0–435) (0–46) (0–0) (–) (0–1) (0–0) (1–21) (0–1) (1–6)
Sum 2504.3 9231.1 12973.8 284 3034.4 2258.4 392.9 2185.1 335 113.1

analyses and the timing of data collection spans a
relatively short period (1995–98), in which there is
likely to be little population change.
One assessment of the feasibility of using the BBS to
derive densities and population size estimates for
species in this study, is to compare the estimates
reported here with independent population estimates
in the Breeding Atlas based on CBC data for 1989 and
targeted national surveys for Lapwing, Skylark and
Corn Bunting. Because CBC farmland plots are
believed to be representative of lowland farmland in
the south and east of Britain (Fuller et al. 1985), it
might be expected that CBC estimates for farmland
specialists such as Corn Bunting, Grey Partridge, Tree
Sparrow, Yellowhammer, Whitethroat, Linnet and
Turtle Dove occurring predominantly in lowland
England might be least affected by geographical and
habitat bias.
This study suggests that at least 70% or more of all
Corn Bunting, Grey Partridge, Tree Sparrow,
Yellowhammer, Whitethroat, Linnet and Turtle Dove
occur in farmland habitat (Fig. 1), the majority of

© 2005 British Trust for Ornithology, Bird Study, 52, 42–54

52 S.E. Newson et al.
Other
100
Proportion of population (%)
75
50
25
0
CB P. TS Y. WH LI TD D. WP S. K. L. JD RB GR BF B. ST SG HS
Figure 1. Estimated proportions of the total British populations of 20 species in this study occurring within five major habitat classes in 1998.
Other habitat here includes SCRUB, HEATH, BOG, WATER, COAST, ROCK and MISC as defined in Table 2. A key to species codes is given
in Table 1.
which are found in lowland in the south and east of
Britain. Assuming this to be correct and adjusted for
temporal change in these species populations between
the Atlas and this study (see Baillie et al. 2002), species
estimates reported from BBS were similar to those
expected from previous studies, except those for Linnet
and Corn Bunting (Table 4). The national population
estimate for Linnet in this study is considerably higher
than that based on the last Breeding Atlas.
Linnets may travel several kilometres to communal
foraging sites (Cramp & Perrins 1994) and because
communal foraging groups are likely to be more
detectable than individuals, this could upwardly bias
estimates using distance sampling and BBS data if
detectability is strongly related to flock size.
In contrast, the national population size estimate for
Corn Bunting here is considerably lower than in the
Breeding Atlas, but considerably higher than an esti-
mate produced by Donald & Evans (1995) as a result of
the 1993 national survey. Donald & Evans (1995)
suggested that the Breeding Atlas overestimates
abundance of Corn Bunting because of a regional bias
in the data towards regions subsequently shown to
support the highest densities of birds. The reason why
the estimate here is higher than in the national survey
is not known, but might be explained by the fact that
this, and other targeted studies discussed here, assume a
full count is made of all individuals present, whilst
our BBS study corrects for individuals present but
undetected using the distance-sampling approach
employed here.
© 2005 British Trust for Ornithology, Bird Study,
Grass Wood Human Farm
Species
Species in our BBS analyses producing similar popu-
lation size estimates to those expected from the Atlas
include Kestrel, Song Thrush, Dunnock, Bullfinch and
Reed Bunting. However, the previous population
estimate for Kestrel in the Breeding Atlas was based on
an informed guess of density and hence population size
(Gibbons et al. 1993), so we can probably have greatest
confidence in the estimate presented here for this
species. Although Song Thrush, Dunnock, Bullfinch
and Reed Bunting all occur in a number of habitats
poorly covered or not covered by the CBC, this study
suggests that densities in habitat classes other than
farmland or woodland are either similar to densities in
these habitats or are very different in rare habitat only.
Using estimates for the other habitat category based on
densities in woodland and farmland probably explains
the similarity between estimates in the Atlas and this
study. Population estimates of Skylark from a national
survey in 1997 (Browne et al. 2000) are very similar to
the estimates presented here (Table 4), although both
are considerably lower than the Atlas estimates. There
are a number of reasons why the CBC methodology
may suggest a high population size for this species. The
1997 national survey of Skylark, and probably the BBS,
effectively record singing males only. The mapping of
territories by the CBC over a large number of visits
(10–12) is therefore likely to provide a better estimate
of the real number of Skylark territories than the
national survey (four visits) and our use of BBS data
(two visits) even if the maximum count over visits is
used, as in the national survey.
52, 42–54

This study effectively uses the mean counts over two
visits and will almost certainly underestimate the true
population. This point needs to be examined before
BBS data can be used to routinely generate wide-scale
density and population estimates. However, the CBC
methodology may still overestimate population size for
two main reasons. The method by which data are
extracted from CBC maps may bias towards higher
densities of birds because territories lying partly outside
a CBC plot boundary will still be recorded as one
territory rather than a proportion of that sighting
within the plot. The CBC tends to use field boundaries
as the plot edge, so the bias may be greatest for
hedgerow specialists, but may affect all species to some
degree. Also the fact that the CBC is concentrated in
the southeast of Britain, where Browne et al. (2000) has
shown there to be the highest densities of Skylark in
Britain, is likely to increase the estimated number of
Skylarks across the country. In the case of the Lapwing,
the population size estimate here was higher than the
national survey of this species carried out in 1998
(Wilson & Browne 1999, Wilson et al. 2001) but simi-
lar to the estimates in the last Breeding Atlas.
The reason for the difference between the national
survey and estimate here is not known, although there
is the possibility that the national survey based on a
single survey visit could have missed a small proportion
of nesting birds present. Residential areas were found to
support an estimated 54% of Starlings and 62%
of House Sparrows (Fig. 1), which is likely to be the
result of the high availability of nest holes and feeding
opportunities in this environment (Crick et al. 2002).
This may explain the considerably higher new popula-
tion size estimates for these species compared with past
estimates based on CBC plots, which under-surveyed
human habitats and, as Table 6 demonstrates, these
species occur at particularly high densities within these
habitats. For a similar reason, population estimates for
other species that occur at high densities in human
habitats (Blackbird, Greenfinch, Jackdaw and probably
Wood Pigeon) are likely to have been underestimated
by the CBC methodology.
Conclusions
The approach described here demonstrates that the
BBS has the potential to produce more robust estimates
of habitat-specific densities and population sizes for
many species than those that have been previously
available. While single-species surveys may provide the
best estimates for individual species there is a need for a
Using BBS to estimate density and population size 53
multi-species approach to provide such information for
large numbers of bird species. The BBS methodology
represents an improvement on earlier multi-species
approaches based on CBC data, such as Atlas estimates,
because it takes explicit account of geographical and
habitat-specific variation in population densities and
sampling intensity. For these reasons we should have
greatest confidence in estimates derived from BBS data,
although there are still a number of limitations and
uncertainties related to this approach. Probably the
main difficulty with the BBS is the interpretation of the
number of adults counted which cannot readily be
connected to the number of breeding pairs as has been
used previously. However, results of earlier studies based
on territory mapping may also be difficult to interpret.
More research on the sex ratio and status of birds
counted by BBS surveys, and on the reliability of the
detectability functions derived from BBS distance sam-
pling data is needed.
ACKNOWLEDGEMENTS
We are grateful to the thousands of volunteers who have
contributed to the Breeding Bird Survey, which provided the
data on which these analyses are based. The BBS is jointly
funded by the British Trust for Ornithology, the Joint Nature
Conservation Committee (on behalf of the Countryside
Council for Wales, English Nature, Scottish Natural Heritage
and the Department of the Environment for Northern
Ireland) and the Royal Society for the Protection for Birds. We
thank Jeremy Wilson, Peter Vickery and John Marchant for
helpful comments on an earlier draft of this paper
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(MS received 20 April 2003; revised MS accepted 27 February 2004)

APPENDIX

Habitat–species combinations for which there were too few obser-

vations of birds to fit habitat-specific detection curves and so were
removed from the analyses.

Species Habitat

Kestrel BOG
Grey Partridge ROCK
Turtle Dove GRASS2
Dunnock MISC
Song Thrush MISC
House Sparrow ROCK
Tree Sparrow WOOD1, WOOD2 & WOOD3
Greenfinch MISC
Bullfinch GRASS2, COAST
Yellowhammer URBAN
Reed Bunting ROCK
Corn Bunting WOOD1, GRASS1, RURAL

© 2005 British Trust for Ornithology, Bird Study, 52, 42–54