International Association for Ecology

The Influences of Habitat, Landscape Structure and Climate on Local Distribution Patterns
of the Nuthatch (Sitta europaea L.)
Author(s): P. E. Bellamy, N. J. Brown, B. Enoksson, L. G. Firbank, R. J. Fuller, S. A.
Hinsley and A. G. M. Schotman
Source: Oecologia, Vol. 115, No. 1/2 (1998), pp. 127-136
Published by: Springer in cooperation with International Association for Ecology
Stable URL: http://www.jstor.org/stable/4221987
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Oecologia
(1998)
Oecologia
(1998) 115:127-136

115:127-136

Springer-Verlag
1998
Springer-Verlag
1998

P.E. Bellamy N.J. Brown B. Enoksson L.G. Firbank

R.J. Fuller S.A. Hinsley A.G.M. Schotman

The influences of habitat, landscape structure and climate

on local distribution patterns of the nuthatch (Siftta europaea L.)

Received: 3 November 1997 / 22 January 1998

10-km grid square scale were related to variables deAbstract The nuthatch, Sitta europaea L., is a small
scribing climate and the amount and dispersion of
(23 g), cavity-nesting woodland bird which, since the
1970s, has been expanding its range in Britain. However, broadleaved woodland. While climate in the study area
within this range, the species is notably scarce in an area appeared suitable, models including landscape variables
suggested that the study area as a whole was unlikely to
of eastern England. This gap in the species distribution
could arise for several reasons including habitat quality,
support nuthatches. Although suitable habitat was
available, woodland in the study area appeared to be too
local landscape structure, regional landscape structure
and climate. Field surveys and logistic models of breedisolated from surrounding nuthatch populations for
ing nuthatch presence/absence were used to investigatecolonisation to be successful. This situation may change
the relative influences of habitat quality, landscape
if current increases in both national and regional popucontinue, thus increasing the number of potential
structure and climate on the prevalence of nuthatches lations
in
colonists reaching the study area.
eastern England. Field surveys of woods in the study
area indicated that habitat quality was sufficient to
support a nuthatch population. A model of habitat ocKey words Metapopulation Population viabilitycupancy in relation to local landscape structure, develDispersal Climate space Isolation
oped in the Netherlands, was applied to the study area.
The number of breeding pairs predicted for the study
area by the model was lower than expected from habitat
Intriuction
area alone, suggesting an additional effect of isolation.
However, observed numbers were even lower than those

This study uses a multiscale approach to look at the

predicted by the model. To evaluate the possible roles of
roles of (a) habitat quality within woods, (b) the pattern
climate and large-scale landscape structure on distribuof woodland in the local landscape, (c) the pattern of
tion, presence/absence data of breeding nuthatches at the woodland and climate at regional and national levels
and (d) historical factors in determining the distribution
of the nuthatch Sitta europaea L. in west Cambridgeshire, eastern England. A multiscale approach was used,
P.E. Bellamy (E) * N.J. Brown S.A. Hinsley
because factors can operate over a wide range of spatial
NERC Institute of Terrestrial Ecology, Monks Wood,
and temporal scales to determine whether or not a speAbbots Ripton, Huntingdon, Cambridgeshire, PE17 2LS, UK
cies is present in a particular location. For example,
Fax: + 44-1487-773467; e-mail: p.bellamy@ite.ac.uk
habitat quality directly influences individual survival
B. Enoksson
and reproduction. Population size and viability are in
Department of Ecology, Lund University,
turn influenced by local landscape factors, such as the
S-22362 Lund, Sweden
area of individual habitat patches, the distances between
L.G. Firbank
patches and the nature of the intervening landuse.
NERC Institute of Terrestrial Ecology, Merlewood,
Landscape structure can also be important at the reGrange-over-Sands, Cumbria, LAll 1 6UJ, UK
gional
level; exceeding a critical level of habitat fragR.J. Fuller
mentation
may affect the functional linking of habitat
British Trust for Ornithology, The Nunnery,
patches, and hence reduce the viability of the regional
Thetford, Norfolk, IP24 2PU, UK
population (With and Crist 1995; Andren 1996), or the
A.G.M. Schotman
Institute for Forestry and Nature Research, Bosrandweg total
20, amount of habitat may fall below a threshold level
PO Box 20, 6700 Wageningen, The Netherlands
sufficient to sustain a species (Enoksson et al. 1995). At

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128

the largest geographical scales, the distribution of species is also limited by climate. This complex control of
distribution is appreciated in general terms (e.g. Krebs
1978; Wiens 1989), but we are unaware of any study of a
particular species distribution that has attempted to
account for all these factors.

During research in eastern England on the effects of

habitat fragmentation on woodland bird populations
(Hinsley et al. 1995a, b, 1996a, b; Bellamy et al. 1996a,
b), it was noted that the nuthatch was largely absent
from apparently suitable woods. The national distribution of breeding nuthatches confirmed that, even though
the study region was within the potential range of the
nuthatch, few breeding pairs were found there (Gibbons
et al. 1993). The reasons for this absence could be linked
to control at any level in the hierarchy from habitat
quality, immediate landscape, regional landscape to climatic factors. In this paper, we will address each of these
possibilities in turn to help unravel the reasons why there

are so few nuthatches in the 500 km2 of west Cam-

bridgeshire.
The nuthatch is a resident bird of mature broadleaved

woodland maintaining a territory throughout the year

(Enoksson 1987, 1988; Glutz von Blotzheim and Bauer
1993). Territories are established by juveniles in their
first summer (Enoksson 1987; Matthysen 1988, 1989)
and typically occupy 1-3 ha in good habitat (Enoksson
and Nilsson 1983; Enoksson 1990; Cramp and Perrins
1993). The short time period over which dispersal occurs
results in most juveniles settling within a few kilometres
of their natal territory. Median distance was 1 km and
very few moved more than 10 km (Matthysen and
Schmidt 1987; Enoksson et al. 1995). Once a nuthatch
has bred on a territory, it is likely to remain there until
Fig. 1 Location of study area (hatched) in the U.K., showing the local
county boundaries. The main map shows the current county boundary
its death (Enoksson 1987). These life history traits sugof Cambridgeshire, divided by a dashed line into the old counties of
gest that the distribution of the nuthatch is likely to be

affected by habitat isolation.

It is known from studies elsewhere in Europe (Verboom et al. 1991; Schotman and Meeuwsen 1994) that

Huntingdonshire (west) and Cambridgeshire (east)

tributed, and the surrounding landscape, which was dominated by

intensive arable farming, had few mature trees. Of the approxithe presence of nuthatches within particular woods demately 450 woods in the selected area, 80 were visited at least once,
pends on the spatial arrangement of the habitat patches.

Several studies have found that isolated woods were less

likely to have breeding nuthatches than similar woods

closer to other suitable woods (Opdam et al. 1985;

Opdam and Schotman 1987; Enoksson et al. 1995), and

woodland fragmentation was thought to reduce
the ability of birds to find the best available habitat
(Matthysen et al. 1995).

and 65 were surveyed in detail for breeding birds in from one to

seven years during 1990-1996.

Habitat quality of individual woods

A six-point ranking of habitat quality for nuthatches was developed on the basis of tree size and species composition, shrub species
and woodland structure. Habitat quality was assessed on the basis
of the densities of nuthatches breeding in different woodland types,

e.g. a mean density of 1.0 pair ha-1 in category 5 (excellent, see

Table 1) woodland (B. Enoksson and A.G.M. Schotman, unpublished results). This system was developed to compare quality of
nuthatch territories in different countries, using a description of the
Study area
woody plant species and structure in a woodland. A simplified
description of the habitat classification is given in Table 1. Three of
the woods in the study area, including Monks Wood itself, were
The study area was in west Cambridgeshire, England, and extended
by PEB, BE, SAH and AGMS as part of this process. A
approximately 20 km north-south and 25 km east-west (Fig.visited
1).
further 62 woods were subsequently assessed by PEB and SAH, by
Some 3% of the whole area was wooded, and 20% of this woodrecording the species and diameter at breast height of a sample of
land was potentially suitable for nuthatches, including several large
between 20 and 75 trees in each wood, depending on woodland
broadleaved woods. Monks Wood (5227 N, 012 W) was the
largest wood (158 ha) in the area. The woods were patchily dis- area, as well as describing woodland structure. In woods with less

Materials and methods

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129

than 20 trees, all were recorded. These data were used to classify the
habitat quality of each wood.
The results are presented with woods divided into three size
classes, based on data describing the frequency of occupation of
woodland patches by nuthatches in the Netherlands (Opdam et al.
1995). Even if containing suitable habitat, woods of < 2 ha were
unlikely to hold breeding nuthatches in their own right, as the
maximum population size for such a small wood was only one pair.
Many, but not all, woods with suitable habitat of 2-10 ha were
expected to have breeding nuthatches, and all woods > 10 ha were
expected to have breeding nuthatches.

Occupancy of individual woods within the landscape

estimates the probability (P) of a wood being occupied by breeding

nuthatches as:

P = e[-9.96+2.841n(A+l)+1.141n(H)]/(l + e[-9.96+2.841n(A+1)+1.141n(H)]) (1)

and the expected number of breeding pairs (N) as:

N e[-4.23+1.341n(A+l)+0.451n(H)]

(2)

where A = area of wood with suitable habitat (ha) and H = area

of suitable habitat within a 3-km radius of the wood (ha) (Schotman and Meeuwsen 1994).
Woods in the study area were estimated as having suitable
habitat for breeding nuthatches if they fell into habitat categories
3-5 (Table 1). A map of the woodlands in the study area was

digitised (Fig. 2) and the parameters estimated using ARC/INFO,

Models of woodland occupancy developed by Schotman and

(Workstation ARC/INFO version 7.04 from Environmental SysMeeuwsen (1994) were applied to the woods in the study area. tems
The Research Institute Inc., Redlands, Calif.). The mean density
models were based on data from 96 woods in the Netherlands over
of nuthatches in good habitat in continuous woodland was previously estimated at 0.5 pairs ha-1 (Enoksson and Nilsson 1983).
the years 1988-1993. These data came from fragmented woodland
Therefore woods of more than 20 ha should hold at least ten pairs,
in an arable landscape as in this study, although the Dutch landa population size sufficient to persist over a period of tens of years
scape had slightly more woodland cover and a higher density of
(Pimm et al. 1988; Liu 1993). For all suitable woods > 2 ha for
hedges and tree rows. This was reflected in the original model,
which we had detailed survey data, the probability of occurrence
which had a term describing a small positive influence of the
was predicted using Eq. 1, and the number of breeding pairs estiamount of hedgerow in the landscape on occupancy by nuthatches,
mated using Eq. 2. These predictions were then compared with the
which was omitted from the present analysis because of the paucity
results of the bird surveys.
of hedges and mature trees in our study area. The revised model

a score of 0 are unlikely to be good enough for winter survival, but

Table 1 Classification of habitat suitability for breeding nua pair of nuthatches may try to breed there if a suitable hole is
thatches, with expected breeding densities. Categories 3-5 are suipresent. Breeding densities are based on data from the Netherlands
table habitat for nuthatches, 0-2 are marginal habitat. Woods with
Category Suitability Breeding density Definition of habitat
(pairs ha- 1)

5 Excellent 1.0 Big (>40 cm diameter at breast height) oaks, elms and beech and/or hazel (or
other winter food sources including e.g. feeding tables)

4 Very good 1.0 Fewer big oaks (but still some >30 cm), less beech/hazel/extra food

3 Good 0.5 Either (a) no or few big oaks, but otherwise suitably sized trees (beech, elm,
aspen, ash, birch, >30 cm), or (b) mixed forest with conifer
and deciduous trees (>25 cm)
2 Tolerable 0.2 Some suitably sized trees (>30 cm including conifers), few oaks but at least
beech, birch, or aspen, or small oaks (25-30 cm)
1 Poor 0.05 No deciduous trees over 25 cm, nestboxes essential to provide nest sites

0 Very poor 0.05 No trees >20 cm, few deciduous trees at all, nestboxes essential

Fig. 2 Distribution of woods in

the local study area: shaded ones
are thought to contain suitable
habitat for breeding nuthatches

5km
I

0'

-C. e

P, I 0
-

,.

e
.4

I C/y

9>
I

*4f -P

IL~~~~~~

x, I( I LAIhC *
I

I'. 02

I',i

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130

Effects of large-scale landscape structure and climate

on the probability of occurrence of breeding nuthatches

taining the Monks Wood study area were first removed from the
data set, and then the regression models were constructed using
1500 randomly selected squares out of the total of 2793 across
mainland Britain. The resulting models were then tested using the
The presence or absence of breeding nuthatches within 10-km grid
remaining 1293 squares (including those of the study area). The
squares in Britain for the years 1988-1991 (Gibbons et al. 1993)
models were evaluated by the degree of agreement between obwas modelled in relation to a number of climate and landscape
served nuthatch distribution (Gibbons et al. 1993) and the prevariables for each square.
dicted
probabilities of occurrence for the remaining squares. This
The landscape data were derived from the Institute of Terrestrial
was quantified using the F-value of the regression between the
Ecology Land Cover Map of Great Britain (Fuller et al. 1994),
predicted probability of occurrence and the proportion of positive
which in turn was derived from Landsat Thematic Mapper images.
records in predicted probability classes of 0.1, weighted by number
The percentage cover of broadleaved woodland (% woodland) was
of observations in each class and with no constant; slopes are not
calculated as the proportion of 25-m square pixels within each
reported, as none were significantly different from 1.
10-km square (excluding water bodies) which were classified as
Five models were selected to help distinguish between the
broadleaved woodland. The mean woodland area was calculated for

importance of climate and landscape variables: (a) with climate

each 10-km square, by assuming that contiguous broadleaved
variables only, (b) with proximate landscape variables only, (c)
woodland pixels constituted a single wood. In addition, the influwith all landscape variables (individual and surrounding squares),
ence of the surrounding landscape for each square was accounted
(d) with climate and proximate landscape variables, and (e) with all
for by estimating the total area of woodland in the eight surrounding
climate and landscape variables. Models with interactive terms
10-km squares, and counting the number of surrounding 10-km
were generated, but they did not result in an increased goodness of
squares containing woodland. Climate data were expressed as 30fit and are not discussed further.

year mean values interpolated over a 10-km grid. The variables of

January minimum temperature, July maximum temperature, annual

precipitation and total number of frost days at mean altitude were
Temporal changes in distribution

used, following Carey et al. (1995). Not surprisingly, some of the

climate variables were correlated (Table 2). Also, broadleaved
Changes in the breeding distribution of the nuthatch, both in the
woodland cover tends to be higher in the south and west of Britain,
study
the areas of the country which also have a milder climate, leading
to area and nationally, were deduced from published sources.
Information on the past distribution of nuthatches was taken from
some correlation between woodland and climate variables.
five bird atlases covering the area: Holloway (1996), relative
The approach used was to model the probability of nuthatch
occurrence as a function of some, or all, of these variables and thenabundance of species on a county basis for 1875-1900; Sharrock

(1977), breeding distributions on a 10-km-square basis for 1968to use the model to interpolate the probabilities of nuthatch pres1972; Limentani et al. (1988), breeding distributions on a 2 x 2 km
ence in the sample squares in the Monks Wood study area. Mean
square
basis for 1979-1983; Lack (1986), winter distributions on a
woodland size was log-transformed, and percentage woodland
cover angularly transformed, before model fitting, to correct for 10-km-square basis for 1981-1984; Gibbons et al. (1993), breeding
distributions on a 10-km-square basis for 1988-1991. Details of
non-normal distributions. The modelling involved a series of
logistic regressions, without interactions, of the general form sites at which nuthatches have been recorded were also extracted
from the Huntingdonshire Flora and Fauna Society annual reports
1949-1995 and from Clark (1996). Huntingdonshire is now iny = a + b\xl + b2x2... bnXn (3)
cluded within Cambridgeshire. Further records of nuthatches in the
where
study area from 1990 to 1994 were obtained from the county bird
recorder (G. Elliott, personal communication) and from field surp = exp(y)/[l + exp(y)] (4)

veys by SAH and PEB. The results are necessarily subjective and
incomplete as the
sourcesp
vary=
greatly
in accuracy,
detail,
and xi = an environmental often
variable
and
the
probability

occurrence (Hosmer and Lemeshow 1989). The grid squares con-

scale and completeness of recording.

Table 2 Variables and abbreviations used in the analysis of

Wood study area) to aid interpretation of results from logistic regressions (total number of squares = 2793)

nuthatch distribution within Britain at a 10-km square scale, and

the intercorrelations between the variables (excluding the Monks

Variable

name

Unit

Abbreviation

January mean minimum temperature, 1960-1990 C Janmin

July mean maximum temperature, 1960-1990 C Julmax
Mean annual precipitation, 1960-1990 mm Annprep
Mean frost days per year, 1960-1990 days Frost

Total area of deciduous woodland ha Totarea

Mean deciduous woodland size ha Meanarea

Total area of deciduous woodland in the eight surrounding 10-km squares ha Surrarea
The number of surrounding squares containing deciduous woodland 0-8 Surrsq
Correlations between variables

Janmin Julmax Annprep Frost Totarea Meanarea Surrarea

Julmax 0.29

Annprep -0.27 -0.67

Frost -0.90 -0.21 0.29

Totarea 0.07 0.36 -0.07

-0.02

Meanarea 0.06 0.21 -0.07 -0.03 0.30

Surrarea -0.62 -0.36 0.01 0.54 -0.15

-0.13

Surrsq -0.41 0.19 0.09 0.42 0.29 0.08 0.15

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131

Nuthatches were recorded in 4 of the 7 woods for which

Results

the estimated probability of breeding was > 0.5, whereas

they were seen (without evidence for breeding) in only

2 of the 21 woods for which the estimated probability

Habitat quality

of breeding was < 0.5 (Table 4).

The smallest of the three woods (6.0 ha) visited by the

expert group also had the least suitable habitat, with a
score of 2. There were few oaks, Quercus sp., with elm,
Large-scale landscape structure and climate
Ulmus sp. being the dominant tree. Trees were also
younger in this wood than in the other two. The next
All models at the national scale gave a reasonable dewood (71.4 ha) was of much higher quality for nutscription of nuthatch distribution. The model that gave
hatches, with large mature broadleaved trees, many of
which were oaks. Hazel, Corylus avellana, an important

the best fit to the data not used in the model development
was the one that included all four landscape variables,
autumn-winter food supply, was also present in the
but no climate variables. However, this model (model 2
shrub layer. There were several large areas which had
in Table 5) should be considered with caution, because

been clear felled and replanted, so only a proportion of

the effect of the area of woodland in the surrounding
the wood was suitable for nuthatch breeding. However,
squares was negative. This variable was also negatively
habitat quality could not explain the lack of nuthatches
correlated with woodland area within the squares
in this wood. From the experience of studies on nut(Table 2), which suggested that its influence may have
hatches in Sweden and The Netherlands, this wood was
been due more to the pattern of woodland in the landexpected to hold about 20 pairs. The largest wood visscape than to a direct effect of each variable on the birds.
ited by the expert group was Monks Wood (157.5 ha)
The other models tended to underestimate probabilwhich was very variable in habitat quality for nutities of occurrence at medium probabilities. The models
hatches. Within the wood there were some grassy fields,
with climate variables (e.g. model 1) described well the
and areas of coppice, young trees and scrub. There were,
observed lack of nuthatches in Scotland, but gave a poor
however, some areas with enough large oaks to support
an estimated 10-20 pairs of nuthatches.

Although habitat within the woods, especially the

Table 4 A comparison of the estimated probabilities of the
two largest, seemed suitable, the agricultural landscape presence of breeding nuthatches and the predicted numbers of
breeding pairs [from regression models, Eqs. 1 and 2 see text] with
surrounding them was noted as being very poor for
nuthatch movement. There were few individual mature

the observed numbers of nuthatches recorded between 1990-1996.

A value of 0.5 recorded pairs indicates a single bird recorded with

trees or rows of trees to assist in the dispersal of nutno evidence of breeding

hatches between woods.

Woodland Probability Predicted Maximum

Of the 64 woods which were surveyed, 28 contained
area (ha) of breeding number number of
good-quality habitat (categories 3-5), but 16 of these
of pairs pairs recorded
were too small (< 2 ha) to support a pair of nuthatches
(Table 3). Larger woods were more likely to contain
0.1
0.00
0.07
0
0.1
0.00
0.09
0
good nuthatch habitat (Table 3).
0.3 0.00 0.09 0
0.3 0.01 0.14 0
0.5
0.01
0.11
0
Occupancy of individual woods
0.7
0.00
0.03
0
0.7
0.00
0.00
0
0.8
0.00
0.00
0
The numbers of breeding pairs predicted from the model
1.0
0.09
0.34
0
using local landscape structure (Eq. 2) suggested that the 1.2 0.01 0.14 0
largest woods in the study area would be expected to
1.5
0.08
0.35
0
hold more than 10 pairs - around half the numbers ex- 1.6 0.01 0.16 0.5
1.7
0.02
0.21
0
pected from area alone. The field observations revealed
actual numbers to be lower than predicted by the model. 1.8 0.00 0.08 0
1.8
0.04
0.26
0
2.0 0.02 0.19 0
2.9 0.01 0.15 0
2.9 0.17 0.54 0
Table 3 Habitat suitability for breeding nuthatches (see Table 1)
3.9
0.36
0.84
0
recorded in different size categories of wood in the study area
4.0 0.17 0.57 0.5
4.2 0.23 0.66 0
Wood Habitat suitability Total
size (ha)
4.2
0.56
1.17
0
0
1
2
3
4
5
7.5
0.62
1.45
0
8.0
0.87
2.5
1
<2 4 8 19 6 7 3 47
8.8
0.77
1.98
0
2-10 - 1 5 8 1 - 15
19.4 0.91 3.52 0.5
>10
1
2
3
71.4 1.0 22.10 0.5
Total 4 9 24 15 10 3 65
157.5 1.0 63.03 2

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132
r

Fig. 3 Observed and modelled breeding distribution of nut

in Britain at the 10-km-square scale. BTO breeding was re
during 1988-1991 (from Gibbons et al. 1993), presence of
nuthatches indicated by a probability of 1.0, and absence by a
probability of 0, and the probabilities of occurrence according to
models 1-5. Details of models are given in Table 5 and in the text

L.
0 . _ .-

O/3
(N *_

3 - D0
~ <.,0- 0

description of the distribution in southern England and

"o

>.E

rn'- 0 E

'5 0

a) .
. C).

40
a)

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Q
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0

Cf

.^ C,

a _O

Ul

-^^S^ - c^O
1O

D1c
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Wales, in that they suggested a distribution centred

on the south-east, rather than on the south-west, of
England (Fig. 3).
The models without climate variables (e.g. models 2
and 3) suggested that the study area was indeed unlikely
to have breeding nuthatches, implying that landscape
structure was unsuitable. Of the 12 10-km squares
around the study area, only 1 according to model 3 and
4 according to model 2 were predicted to have a greater
than 50% probability of occupation (Table 6). This
likelihood was far greater if the models which include
climate variables are used (Fig. 3).

Ct

C)
-u

oZ

o-6
,J: .~
.! .

Holloway (1996) considered the nuthatch to be a com-

CV mC

mon breeding bird in Huntingdonshire between 1875

and 1900. However, Peake (1926) also used published
and unpublished information from the same era and
stated that the nuthatch occurred "only in limited

-.o

^^CX

*O Y
03 C.Ce

C)

_ ,,

Temporal changes in distribution

0 -

._

c X

numbers and unknown in the north and east of the

0

m >

- O

rS

- e)

0a)

U I

:X

c ^

[-

L..,

county". He mentioned only a single site where a few

pairs bred. Tebbut (1967) listed nine regular breeding

<

..

00
ct

>

e-t+

<

0
- .2

'- U,

square during 1968-1972 to five 10-km squares during

1988-1991. Moreover, increasing numbers of nuthatches
have been recorded from woods in this area since they

> '

0 0
0o r

IUs

r-

were first recorded there in the 1970s, with several woods

" I
.-0

C) t

0 ,9
0
X

-eo
o 0-

0n'2 m

't

au

cLt

b1 D

._

-0

>^

Discussion

"s 0

._)^ >

0.

;Y o X2

0 0.
E
r0 0)

) U,.
CZ 4.
CZ

Cu C-. Al
* c/l V ,
U)

holding up to five pairs by 1995 (Clarke 1996). The

nuthatch breeding range in south Cambridgeshire,
approximately 20 km south-east of Monks Wood, has
also expanded in recent years, from two to four 10-km
squares (Bircham et al. 1994).

._

in.

U C) o

area. Over recent years, the few sites in the study area
which previously had nuthatches appear to have lost
them as a breeding species. However, there has been a
spread in the distribution in an area approximately
18 km north-west of Monks Wood, from one 10-km

Ct +

c) 0

au^

sites in the county, mostly to the south-west of the study

V5

CZ

.L

- u 1

Li

'A

0
0

-0

ia)
0. C

E -X

V)

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&
OO
0+

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; <t Y

re)

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W)

Comparison of individual woods with the criteria for

habitat quality strongly suggested that the study area did
include suitable habitat for nuthatches in a number of

woods. Some woods were large enough to support core

populations of more than ten pairs with a low probability of extinction (Verboom et al. 1993). Therefore,
habitat quality did not appear to limit the presence of
nuthatches, and yet, no more than two breeding pairs

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133

Probability of finding nuthatch

km

Ei 0 I 0.2 i 0.4 U 0.6 * 0.8 U 1.0

0 I(00) 200 300

IT

Probability of finding nuthatch

El 0 g 0.2 M 0.4 U 0.6 * 0.8 * 1.0

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All use subject to http://about.jstor.org/terms

ki

0 100 200 300

134

Table 6 Estimated probability of occurrence of nuthatches within

10-km grid squares comprising the Monks Wood study area and
surroundings, on the basis of all regression models (Table 5).
Squares 51E 28N and 52E 28N contain most of the local study

area. Actual occurrence: presence (1) or absence (0) of breeding

nuthatches within the 10-km square (Gibbons et al. 1993). See text
for details, see also Fig. 4

Grid refs. of squares Woodland Mean woodland Probability of breeding from regression models Actual

cover

Easting
50
51
52
53
50
51
52

53
50
51
52
53

27
27
27
27
28
28
28

28
29
29
29
29

0.56
1.72
0.67
0.36
3.79
2.61
3.60

0.63
5.86
1.58
0.44
0.21

(%)

area

Northing

0.02
0.39
0.02
0.01
0.44
1.31
0.24

0.04
0.05
0.08
0.01
0.02

0.82
0.84
0.84
0.83
0.83
0.83
0.84

0.41
0.43
0.28
0.20
0.54
0.85
0.51

(km2)

0.44
0.32
0.13
0.09
0.50
0.92
0.34

0.66
0.72
0.67
0.57
0.82
0.86
0.81

occurrence

0.67 0.58 0
0.72 0.69 0
0.67 0.70 0
0.56 0.59 0
0.83 0.80 1
0.87 0.74 0
0.81 0.80 0

0.84 0.23 0.10 0.61 0.61 0.63 0

0.81 0.51 0.36 0.85 0.85 0.87 1
0.84 0.28 0.14 0.66 0.66 0.68 1
0.84 0.20 0.09 0.58 0.58 0.61 0
0.84 0.20 0.09 0.59 0.58 0.61 0

nilesof
were more
likelywoods
to leave their natal
area inno
frag- w
were observed in any
the
and
mented
habitat, even year.
though there were vacant
breeding nuthatches in
every
territories. This resulted in vacant territories in fragAlthough there was suitable habitat available, it
amounted to only 0.7% of the total cover, suggesting mented woodland being occupied by single birds or
that landscape effects, particularly isolation, could be aremaining empty more frequently than those in more
problem. Certainly, predictions of numbers of breedingwooded areas (Matthysen and Currie 1996). This dispairs, taking isolation into account, were much lower parity in dispersal patterns between fragmented and
than expected from habitat area alone, implying that more wooded landscapes could mean that birds were
isolation of woodland in the landscape was reducing the more likely to move out of a fragmented area than move
potential of these woods to support nuthatch popula- into it. While Matthysen et al. (1995) did not find a net
loss from their fragmented study area, they did not look
tions. Most woods with suitable habitat were small and
unlikely to support breeding nuthatches, so only theat the landscape surrounding it. In our study area, the
fact that the suitable woods were separated from the
suitable habitat occurring in woods of at least 5 ha
nearest nuthatch populations by about 18 km, with very
contributed to the ability of nuthatches to breed in this
little suitable habitat in the intervening landscape, could
area. However, even the predicted numbers in these
mean
that new populations were unlikely to be establarger woods were higher than the actual numbers
lished, and those that were present would be prone to
observed on the ground (Table 4).
The interpretation of the distribution at the 10-km local extinction.
The nuthatch has been a scarce breeding bird in the
grid scale across the whole country depended to some
study area throughout the 20th century. During this
extent on which model was chosen. Models including
time, potential habitat has undergone many changes
climate suggested that the study area was suitable for the
management of woods as well as losses and
nuthatch, but these models tended to overemphasise through
the
gains in woodland cover. There was an estimated loss of
species occurrence in south-eastern England. The models
20% of broadleaved woodland between 1945-1985 due
involving just landscape parameters were therefore more
to conversion to arable land. This trend may have halted
appropriate, even though they suggested that nuthatches
and even reversed in recent years with incentives for
may be found further north into Scotland than climate
woodland planting (Peterken and Allison 1989).
limits may allow. Both the model including the effects farm
of
Even
areas
which remained wooded over long periods
neighbouring squares and the model excluding them
suggested that the landscape of the study area was vary
far considerably in quality with time due to changes in
woodland management. Traditional management until
from ideal for nuthatches. Overall, the results suggested
that, while there were areas of suitable habitat, somethe
of early 20th century maintained woodland in early
successional stages with a low density of mature trees
which should support viable populations of nuthatches,
1980). Although this practice has now largely
there was simply too little broadleaved woodland in (Rackham
the
ceased,
there
has been a loss of mature trees, largely
region in and around the study area to make the esoaks, due to timber removal during the two world wars,
tablishment and persistence of such populations likely.
conversion of broadleaved woodland to conifer plantaThe results from a study by Matthysen et al. (1995) in
tions, and selective felling of timber trees (Hooper 1973;
Belgium supported this view. They found that nuthatch
Peterken and Allison 1989). As well as the loss of mature
dispersal distances in fragmented habitat (median
trees from woodland, there has also been a loss of trees
> 3 km) were greater than those found in continuous
habitat (median ca. I km). They also found that juve-in the surrounding farmland. For example, 90% of non-

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All use subject to http://about.jstor.org/terms

135

woodland trees in two areas of Cambridgeshire were lost

between 1947 and 1983 (Peterken and Allison 1989).

Clarke JS (1996) The birds of Huntingdon and Peterborough.

Clarke, Hilton, UK

Cramp S, Perrins CM (eds) (1993) The birds of the Western Palearctic, vol 7. Oxford University Press, Oxford, pp 298-313
woodland trees, with fewer now occurring in linear
Enoksson B (1987) Local movements in the nuthatch Sitta eurofeatures and most occurring in isolated clumps.
paea. Acta Regiae Soc Sci Litt Gothob Zool 14: 36-47
It appears from current woodland management that Enoksson B (1988) Prospective resource defense and its consequences in the nuthatch Sitta europaea L. PhD thesis, Upssala
the quality of present woods will increase for nuthatches,

There has also been a change in the distribution of non-

University

as coppice matures, and new plantings mature into high

Enoksson B (1990) Autumn territories and population regulation

forest. However, there is no indication of any increase in

in the nuthatch Sitta europaea: an experimental study. J Anim

the numbers of trees in the surrounding landscape ma- Ecol 59: 1047-1062
trix. The result may well be an overall increase in the Enoksson B, Nilsson SG (1983) Territory size and population
density in relation to food supply in the nuthatch Sitta europaea
number of woods which are of suitable habitat quality, (Aves). J Anim Ecol 52: 927-935
which would have the effect of increasing the probable
Enoksson B, Angelstam P, Larsson K (1995) Deciduous forest and

numbers of breeding nuthatches, assuming that they can

resident birds: the problem of fragmentation in a coniferous

become established. The probability of this happening is forest landscape. Landscape Ecol 10: 267-275
increasing, as the U.K. national distribution of the

Fuller RM, Groom GR, Jones AR (1994) The landcover map of

Great Britain: an automated classification of Landsat thematic

nuthatch has been expanding northwards and eastwards

mapper data. Photogramm Eng Remote Sensing 40: 553-562
over the last 20 years, part of a Europe-wide expansion
Gibbons DW, Reid JB, Chapman RA (1993) The new atlas of
breeding birds in Britain and Ireland: 1988-1991. Poyser,
of range (Marchant et al. 1990; Svensson 1992; Cramp
London
and Perrins 1993; Gibbons et al. 1993). Establishment in
Glutz von Blotzeim UN, Bauer KN (1993) Handbuch der Vogel
the study area may well arise due to immigration from
Mitteleuropas, band 13/11. Aula, Weisbaden
one of two increasing populations of nuthatch, each
Hinsley SA, Bellamy PE, Newton I (1995a) Bird species turnover
approximately 18 km from suitable woods in the study and stochastic extinction in woodland fragments. Ecography
18: 41-50

area.

Hinsley SA, Bellamy PE, Newton I, Sparks TH (1995b) Habitat and

landscape factors influencing the presence of individual breeding

Acknowledgements We would like to thank the many people

who
bird
species in woodland fragments. J Avian Biol 26: 94-104
kindly gave us access to their woods to collect field data. We also
Hinsley SA, Bellamy PE, Newton I, Sparks TH (1996a) Influences
thank the many birdwatchers who undertook the fieldworkofupon
population size and woodland area on bird species distriwhich the bird atlases were based. The woodland map of thebutions
local in small woods. Oecologia 105: 100-106
study area was digitised by Jackie Ulyett, while Ruth Swetnam
Hinsley SA, Pakeman RJ, Bellamy PE, Newton I (1996b) Influedited and extracted data from it. The climate data were provided,
ences of habitat fragmentation on bird species distributions and
through the LINK project, by the Climatic Research Unit of the
regional population size. Proc R Soc Lond B 263: 307-313
University of East Anglia. The British Trust for Ornithology supHolloway S (1996) The historical atlas of breeding birds in Britain
plied the data from the New atlas of breeding birds. Dave Howard
and Ireland 1875-1900. Poyser, London
generated the maps of the national models. Tim Sparks, Ian
Hooper MD (1973) History. In: Steele RC, Welch RC (eds) Monks
Newton and Lars Gabrielson provided valuable comments on
Wood, a nature reserve record. The Nature Conservancy,
earlier drafts. B.E. was supported by the Swedish Environmental
Huntingdon, pp 22-35
Protection Agency. This work contributes both to LANDECOHosmer DW, Lemeshow S (1989) Applied logistic regression.
NET, a research programme part funded by the European Com- Wiley, New York
mission, and to the NERC TIGER IV.2a programme on the
Krebs CJ (1978) Ecology: the experimental analysis of distribution
implications of climate change on populations. Finally, we would
and abundance, 2nd edn. Harper & Row, New York
like to thank the participants of a LANDECONET workshopLack
at P (1986) The atlas of wintering birds in Britain and Ireland.
Monks Wood, November 1994, whose ideas and enthusiasm pro- Poyser, Calton
vided the stimulus for this work; details of the workshop and of the

Limentani J, Elliott G, Everett M (1988) The breeding birds of

LANDECONET programme as a whole can be found on http://Huntingdon and Peterborough, 1979-83. Elliott & Everett

www.nmw.ac.uk/ite/econet/index.html.

Enterprises, Buckden

Lui J (1993) Discounting initial population sizes for prediction of

extinction probabilities in patchy environments. Ecol Modell
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