A SUMMARY OF NEST RECORDING AND RINGING PULLI IN SOUTH-

WEST LANCASHIRE
IAN H WOLFENDEN

Since obtaining a “C” permit in 1974, I have concentrated my ringing effort on

studying the birds of the Crosby and Hightown dunes. This area had been my
favourite bird watching haunt and a permit allowed me to study the birds that I
had watched for many years. When I was granted a pullus endorsement a few
years later, I was enabled, theoretically, to investigate the birds from egg until
death. By mist-netting fully grown birds throughout the year and by ringing
nestlings during the breeding season it is possible to obtain a complete picture of
the populations of passerines inhabiting the dunes and associated meadows.
Ringing passerine pulli in a duneland habitat is a time-consuming pastime, as the
majority of nests have to be located by watching adult birds that are feeding
young. Only a small number of nests are “stumbled across” whilst walking in the
dunes. Active searching in dense low vegetation is not to be recommended as it
endangers the nest and its contents in a number of ways. First many birds leave
the nest at the last second and very quietly. Therefore it would be easy to
damage supporting vegetation or even the nest itself with disastrous
consequences. Secondly many of the duneland plants are damaged by walking
through them and therefore it is easy to leave a trail to the nest which might
attract predators. The reason for the nest being built in dense vegetation in the
first place is to afford it protection. Having watched a bird back to its nest from a
distance that does not disturb it, the nest is pin-pointed with reference to a
distinct characteristic such a clump of 3 flower heads or a dead flower lying at a
strange angle. The nest is then approached directly keeping in a straight line from
observation point to assumed nest site, stopping occasionally to check with
binoculars. In the final approach to the nest it is preferable to step over and
between plants, thus causing as little damage as possible and also breaking the
trail. Having located the nest, ringed any young and taken details for BTO record
card, the vicinity is left using the same footprints and carefully rearranging
vegetation. Nests in sensitive sites are only revisited if the young are too small to
ring on the first visit. Many Linnets lay in the clumps of marram and sea lyme
grass around the main dune ridge and therefore are often close to paths. It is
possible to find these nests without leaving an obvious trail, and this is how most
Linnet nests are located; the fact that many are close to paths must contribute to
predation by humans and other mammals.

Pulli have been ringed on these dunes since the early 1960’s but not on a
regular basis. Since commencing this study I have tried to visit the area every 2 or
3 days between the third week in April and mid July. By regular visiting, nests can
be located before they contain young that are too big to ring or have even
fledged. By dividing an area into ‘walks’ the observer becomes aware of the birds
in the area and by watching them closely, notices when they are carrying food.
Often the first hint that a bird has young is the constant alarming when the
observer enters the territory. After a few years watching it becomes apparent
that certain areas are favoured. Often the same clump of willow scrub or patch of
grass will hold a pair of birds of a particular species every year. Nests are
sometimes situated close to nests of previous seasons. With experience the
observer gains an eye for where a nest is likely to be, and further indications
come from the alarmed bird. Locating nests is best done alone otherwise
conversation occurs and bird calls are missed. Once a feeding bird has been
located it is usually more reluctant to return to its nest when there are more than
one person watching. The observer may also have to move further away for the
bird to visit its nest. As a consequence the average time taken to find a nest is
longer. Also if more than one person is searching there is a greater change of
damage.

As experience increases more nests are located and more pulli ringed as can be
seen from the annual totals for pulli ringed in this study. It becomes apparent
that most clutches of a particular species hatch around the same time every year.
Consequently it is possible to concentrate on each species at a time when their
young are at a ringable age. Locating nests and ringing pulli is an enjoyable and
worthwhile pastime. Besides the satisfaction of outwitting the parent birds there
is also the opportunity to marvel at the variation and structure of both nest and
nestling. The transformation that takes place between newly hatched chick and
fledgling in 14 days is remarkable. The information collected on nest record cards
and by ringing pulli is of value ornithologically. The former provides breeding
biology data to be included in national files and the latter provides ringed birds of
known age and place of hatching which can be important after recovery.

This article deals with a small part of the information collected on the biology
of the duneland birds. Later group reports will cover ringing recoveries and
retraps of birds ringed as pulli.

The Crosby and Hightown dune system has a surprising variety of habitats for
its size (about 188 hectares). These include frontal dunes, damp slacks, damp
meadows, reed-fringed pools, dry grassy areas, willow scrub, sea buckthorn
thickets, a small wood, copses, a reed bed, and some saltmarsh. The adjacent golf
course has large areas of ‘fairway’ and ‘green’. A more comprehensive
description of the area is to be found in the SWLRG report for 1980.

Some of the information here may be useful to other ringers planning a season
of passerine pulli ringing in similar habitats. It may also be of use to those training
for a pullus endorsement, as it gives them some guidance as to when to persuade
their trainer to go out.

Table 1 – Number of pulli ringed annually 1977 to 1985

YEAR 1977 1978 1979 1980 1981 1982 1983 1984 1985
TOTAL 117 198 226 311 235 268 258 331 372

Table 2 shows the dates of ringing broods of the most ringed species during the
years 1977 to 1985. The breeding season has been divided into 5-day periods.
The number of broods ringed in each 5-day period is expressed in parentheses as
a percentage of the total. For a small number of broods that were too old to ring,
the dates have been calculated for when they might have been ringed. Data from
1986 have been included in the analyses for Stonechat and all warbler species
because of the relative paucity of information.

A small number of broods of Skylark, Meadow Pipit and Reed Bunting ringed at
Seaforth Dock Pools have been included in the data also as these species breed in
habitat similar to a part of the dunes, and the site is not far away. Table 2 shows
the period of time during which species have young of a ringable age and also
demonstrates that all species have at least one peak during this period,
suggesting that a sizeable proportion of the birds start laying simultaneously. It
also shows that all species have a discrete breeding season with earliest pairs
starting to breed at different times. It shows that the first nesting species is the
Skylark followed sequentially by Stonechat, Meadow Pipit, Linnet, Reed Bunting,
Willow Warbler, Whitethroat and Sedge Warbler. Those that are not trans-
saharan migrants commence breeding at roughly 5-day intervals. There is then a
lapse of approximately 20 days before the first Willow Warblers start laying
followed 5 and 10 days later by Whitethroat and Sedge Warbler respectively.

The timing of pulli ringing can be directly related to egg-laying and to the timing
of breeding of every pair. Tables 3 to 6 show the ringing dates, by year, for each
species for which sufficient data are collected.
 Table 2 – Brood ringing dates 1977-1985

Meadow Reed Willow Sedge

Skylark Stonechat Pipit Linnet Bunting Warbler Whitethroat Warbler
April 16 - 20
21 - 25 3 (1.0)
26 - 30 12 (4.0) 1 (8.3)
May 01 - 05 18 (6.0) 2 16.7) 1 (1.5)
06 - 10 52 (17.2) 3 (25.0) 2 (3.0) 1 (1.4)
11 - 15 31 (10.3) 2 (16.7) 11 (16.7) 3 (4.2) 1 (1.2)
16 - 20 23 (7.6) 11 (16.7) 10 (14.1) 2 (2.3)
21 - 25 20 (6.6) 2 (16.7) 16 (24.2) 14 (19.7) 12 (14.0)
26 - 30 38 (12.6) 3 (4.5) 15 (21.1) 14 (16.3)
31 - 04 24 (8.0) 5 (7.6) 9 (12.7) 23 (26.7) 1 (3.6)
June 05 - 09 28 (9.3) 1 (8.3) 2 (3.0) 4 (5.6) 18 (20.9) 10 (35.7) 13 (41.9)
10 - 14 16 (5.3) 5 (7.6) 3 (4.2) 3 (3.5) 5 (17.9) 8 (25.8) 2
15 - 19 14 (4.6) 1 (1.5) 2 (2.8) 2 (2.3) 6 (21.4) 3 (9.7) 1
20 - 24 6 (2.0) 4 (6.1) 2 (2.8) 2 (2.3) 2 (7.1) 1 (3.2) 2
25 - 29 8 (2.7) 1 (8.3) 3 (4.5) 1 (1.4) 2 (2.3) 2 (6.5) 1
30 - 04 4 (1.3) 1 (1.5) 2 (2.8) 1 (1.2) 1 (3.6) 1
July 05 - 09 3 (1.0) 2 (2.8) 2 (2.3) 3 (10.7) 1
10 - 14 1 (0.3) 1 (1.2) 2 (6.5) 1
15 - 19 1 (1.2)
20 - 24 1 (0.3) 1 (1.5) 1 (1.4) 2 (2.3) 1 (3.2)
25 - 29 1 (1.4)
30 - 03
August 04 - 08 1 (3.2)
09 - 13
14 - 18
19 - 23 1 (1.4)
Total 302 12 * 66 71 86 28* 31* 9

* data from 1986 included

 Table 3 – Skylark brood ringing dates 1977-1985

1977 1978 1979 1980 1981 1982 1983 1984 1985

April 16 - 20
21 - 25 1 (2.5) 1 (2.1) 1 (3.0)
26 - 30 4 (12.1) 3 (7.5) 2 (4.2) 1 (3.2) 2 (6.1)
May 01 - 05 1 (2.5) 8 (16.7) 4 (12.9) 2 (6.1) 1 (3.3) 1 (2.5) 1 (3.4)
06 - 10 1 (5.6) 4 (12.1) 7 (17.5) 6 (12.5) 4 (12.9) 8 (24.2) 10 (25.0) 12 (41.4)
11 - 15 4 (22.2) 3 (9.1) 4 (8.3) 3 (9.7) 1 (3.0) 2 (6.7) 14 (35.0)
16 - 20 3 (16.7) 1 (3.0) 1 (2.5) 4 (8.3) 1 (3.2) 9 (30.0) 3 (7.5) 1 (3.4)
21 - 25 4 (22.2) 1 (3.0) 1 (2.5) 7 (22.6) 1 (3.0) 2 (6.7) 3 (7.5) 1 (3.4)
26 - 30 2 (11.1) 12 (36.4) 3 (7.5) 3 (6.3) 3 (9.7) 2 (6.1) 7 (23.3) 6 (20.7)
31 - 04 2 (6.1) 7 (17.5) 5 (10.4) 2 (6.5) 5 (15.2) 1 (3.3) 1 (2.5) 1 (3.4)
June 05 - 09 6 (15.0) 5 (10.4) 1 (3.2) 6 (18.2) 3 (10.0) 2 (5.0) 5 (17.2)
10 - 14 4 (12.1) 1 (2.5) 2 (4.2) 3 (9.7) 2 (6.1) 1 (3.3) 3 (7.5)
15 - 19 2 (5.0) 5 (10.4) 1 (3.0) 3 (10.0) 2 (5.0) 1 (3.4)
20 - 24 3 (6.3) 2 (6.5) 1 (3.3)
25 - 29 3 (16.7) 2 (6.1) 1 (2.5) 2 (6.1)
30 - 04 1 (5.6) 2 (5.0) 1 (3.4)
July 05 - 09 3 (7.5)
10 - 14 1 (2.5)
15 - 19
20 - 24 1 (2.5)
25 - 29

Table 4 – Meadow Pipit brood ringing dates 1980-1985

1980 1981 1982 1983 1984 1985

April 26 - 30
May 01 - 05 1 (12.5)
06 - 10 1 (12.5) 1 (6.3)
11 - 15 1 (11.1) 3 (37.5) 3 (23.1) 4 (25.0)
16 - 20 1 (11.1) 1 (12.5) 1 (12.5) 1 (7.7) 3 (27.3) 3 (18.8)
21 - 25 1 (11.1) 1 (12.5) 7 (53.8) 3 (27.3) 4 (25.0)
26 - 30 1 (11.1) 1 (12.5) 1 (6.3)
31 - 04 2 (22.1) 2 (25.0) 1 (7.7)
June 05 - 09 1 (12.5) 1 (9.1)
10 - 14 1 (12.5) 1 (12.5) 1 (7.7) 1 (9.1) 1 (6.3)
15 - 19 1 (6.3)
20 - 24 2 (22.1) 1 (12.5) 1 (9.1)
25 - 29 1 (11.1) 1 (12.5) 1 (6.3)
30 - 04 1 (9.1)
05-09
10-14
July 15 - 19
20 - 24 1 (9.1)
25 - 29
 Table 5 – Linnet brood ringing dates 1977-1985

1977 1978 1979 1980 1981 1982 1983 1984 1985

April 16 - 20
21 - 25
26 - 30
May 01 - 05
06 - 10 1 (4.8)
11 - 15 2 1 (4.3)
16 - 20 1 1 2 (50.0) 2 (8.7) 4 (19.0)
21 - 25 1 1 1 (25.00 1 1 (12.5) 7 (30.4) 2 (9.5)
26 - 30 2 3 (37.5) 7 (30.4) 3 (14.3)
31 - 04 2 1 (25.00 2 1 (12.5) 2 (8.7) 1 (4.8)
June 05 - 09 1 (12.5) 1 (4.3) 2 (9.5)
10 - 14 1 (12.5) 1 (4.3) 1 (4.8)
15 - 19 1 (12.5) 1 (4.8)
20 - 24 1 (4.3) 1 (4.8)
25 - 29 1 (4.8)
30 - 04 1 1 (4.8)
July 05 - 09 1 1 (4.3)
10 - 14
15 - 19
20 - 24 1 (4.8)
25 - 29 1 (4.8)
Aug 20 1 (4.8)
Table 6 – Reed Bunting brood ringing dates 1977-1985

1977 1978 1979 1980 1981 1982 1983 1984 1985

April 16 - 20
21 - 25
26 - 30
May 01 - 05
06 - 10
11 - 15 1 (12.5)
16 - 20 2 (15.4)
21 - 25 2 1 (7.1) 2 (15.4) 2 (25.0) 2 (28.6) 1 (11.1) 2 (16.7)
26 - 30 1 2 1 (7.1) 3 (23.1) 1 (12.5) 2 (22.2) 3 (25.0) 1 (7.1)
31 - 04 1 6 (42.9) 2 (15.4) 3 (37.5) 1 (14.3) 2 (22.2) 2 (16.7) 6 (42.9)
June 05 - 09 2 5 (35.7) 1 (7.7) 2 (28.6) 2 (22.2) 2 (16.7) 4 (28.6)
10 - 14 3 (25.0)
15 - 19 1 (14.3) 1 (7.1)
20 - 24 1 1 (11.1)
25 - 29 1 (14.3) 1 (7.1)
30 - 04 1 (7.1)
July 05 - 09 1 (7.7) 1 (12.5)
10 - 14 1 (7.7)
15 - 19 1 (7.1)
20 - 24 1 (7.7) 1 (11.1)
25 - 29

Skylark

For this species ringing occurs mainly between 6-15 May and 26 May – 9 June
(first egg dates approximately 18-27 April and 7-21 May). The period 6-10 May
shows the only pronounced peak; the timing of any other peak is variable from
year to year (Table 3). Table 2 shows that Skylarks have the second longest
breeding season, laying their first eggs over, approximately, a 90-day period. It
also shows that for about half the period breeding numbers are fairly level. The
Skylark colour-ringing study has found little evidence of double broods in this
population and so it must not be assumed that most broods ringed during the
season are the result of first nesting attempts. It would be expected that most
pairs start breeding about the same time when food availability and climatic
conditions allow. There are several possible reasons for this leisurely breeding
season. The colour-ringing study has shown that many Skylarks winter close to
the breeding site on nearby farmland and should be able to commence breeding
without the constraints imposed by moult and the preparation for migration. The
density of the breeding population is very high and hence the birds are highly
territorial. This could result in only a limited number of pairs managing to obtain
territories large enough in which to breed. As the season progresses the birds
become less territorial and this may allow non-breeding birds in to start
breeding. When birds are feeding young there are few disputes and birds
regularly cross boundaries. Further, nests are often found fairly close together,
although these cannot belong to the same pair. The colour-ringing study has also
indicated that many breeding birds are first-year and it is possible that these tend
to lay later as territories become available. Another reason for the long breeding
season may be that the duneland consists of a variety of habitats and birds in
different habitats may be governed by differing availabilities. The data collected
so far could be re-analysed with this in mind, but many territories include a
mixture of habitats. The small number of breeding attempts in the last third of
the season may be due to birds re-laying and a few second broods. Table 3 shows
that for the years 1983 – 1985, Skylarks have commences breeding approximately
10 days later than in previous years possibly due to the effects of harsh winters
and cold springs. In these years the ringing peaks are noticeably greater, as
though a number of pairs have been delayed by the same factor. The earliest and
latest dates recorded for the laying of the first egg of a clutch were 6 April and 4
July. These were calculated using an incubation period of 11 days (Mayer-Gross
1970, The Nest Record Scheme), age at ringing and a clutch size taken to be the
same as the brood size plus the number of unhatched eggs.

Stonechat

This is the least common of the duneland birds and in recent years has
declined from the 4-5 pairs present in the late 1970’s. This may reflect the
harsher winters and poorer springs of recent years, although it may be a cyclical
variation returning to the same low level that occurred prior to 1973. However
further evidence linking the decline to climatic change is the fact that no broods
have been ringed prior to 6 May since 1981. This shows delayed breeding in
recent years as in the case of the Skylark. Stonechats breeding on the dunes
recently have nested close to the houses at Hightown and the car park at Hall
Road, and consequently have suffered from disturbance. This seems a strange
habit considering that there are only 1 or 2 pairs and plenty of apparently suitable
habitat. Nevertheless they rear young successfully every year, sometimes
producing two broods at the Hightown end. Brood sizes are usually 5-6. The
habitat at the Hall Road end in mainly grass, marram and sea lyme grass with
scattered bushes and some bramble. At the Hightown end willow scrub
predominates with some grass and scattered bushes. Table 2 shows that 75% of
the Stonechat pulli are ringed in May and that ringing takes place mostly between
6-10 May at the same time as most Skylark ringing occurs. It also shows that
almost 60% of broods were ringed during the first 15 days of May. The single
brood ringed on 6 June is a known re-lay after predation and the birds ringed on
25 June were from the only second brood. These were ringed 42 days after the
first brood, and belonged to a male almost certainly mated with a lone female
whose brood was ringed on 21 May. Second broods are more common that these
data suggest. Locating Stonechat nests is difficult owing to their well hidden
positions and to the fact that some birds show little alarm until the young have
fledged. This observation refers particularly to second broods. Stonechat pulli
are ringed at an age of about 6-7 days. The earliest and latest dates estimated for
the laying of the first egg are 5 April and 31 May. These were calculated using an
incubation period of 14 days (Mayer-Gross 1970), and the age and brood size at
ringing. Clutch size was taken to be the same as the brood size plus unhatched
eggs.

Meadow Pipit

This species only colonised the dunes in recent years possibly associated with
the drying out of the habitat and the increase of grassy areas. It was first proved
to be breeding at Seaforth Docks in 1978 and on the dunes in 1980. Around 30
pairs now nest of the dunes and leave the area during the winter. Table 4 shows
annual ringing dates and Table 2 the overall summary. A further 7 Meadow Pipit
nests were located in 1984 but these held young Cuckoos. Due to lack of
experience of ageing Cuckoos and therefore a risk of bias, these nests are not
included in the data. Table 2 shows that Meadow Pipits have been ringed over an
80 day period, 58% of them between 11-25 May, about 15 days later than
Skylarks and Stonechats. After the peak, ringing continues at a lower rate,
presumably including re-lays, late breeding and a few second broods. As with the
Skylark, breeding density is high in some areas and some birds may delay
breeding due to competition. The habitats occupies by Meadow Pipits are similar
to those already described for Stonechats, with some birds breeding also in areas
of mainly grass. Brood sizes are usually 4-5 but 3 and 6 have been recorded. The
earliest and latest dates for the laying of the first egg are estimated to be 11 April
and 28 June. These were calculated using an incubation period of 13 days
(Mayer-Gross 1970), and the age and brood size at the time of ringing. Clutch size
was taken to be the same as the brood size at the time of ringing plus unhatched
eggs. Most first eggs are laid between 28 April and 2 May. Table 4 shows that the
time of peak ringing activity varies from year to year, although the small amount
of data may influence this finding.

Linnet

This species is a common breeding finch throughout the dunes with 30-50 pairs
occurring in most years. It is difficult to estimate the size of the breeding
population as birds feed well away from the nest, often in small parties. Also
males frequently sing away from the breeding area. Very little territorial
behaviour is shown and nests are often found close together in the best habitats.
Favoured sites are clumps of sea lyme grass and marram, often in the frontal
dunes. Other common nest sites are in willow scrub and in steep banks and
exposed roots on the sides of eroding dunes. Very few nests are found in bushes.
This is due possibly to the exposed nature of the site where nests could be
damaged by strong winds. Most nests are close to the ground and there is much
predation by a variety of mammals. The choice of the frontal dunes leads to
further trouble from sun-bathers and children. A few nests have been found in
grass clumps less than 6 inches from the ground. In 1986 strong to gale-force
winds in late May caused the failure of most of the nests in the exposed roots of
eroded dunes because blown sand filled the nest and the birds deserted eggs and
young. In one nest, all that remained was a dead chick lying on the rim of a sand
filled nest cup. Linnet nests are the only ones that I actively search for. By
walking along the paths between the marram and sea lyme grass clumps it is easy
to locate nests without causing harm to them. Linnets are the only common
species that feed their young on seeds and it would appear that they time their
breeding to coincide with the seeding of coltsfoot and early dandelions which
abound in this dune system. The Linnet is the fourth of the studies species to
start breeding. The earliest brood was ringed on 9 May. Table 2 shows that the
peak time for ringing Linnet pulli is 21-30 May (first egg dates approximately on
29 April to 8 May). It also shows that 68% of pulli are ringed during the 20-day
period of 16 May to 4 June. After this date, breeding and therefore pulli ringing
continues at a lower level until late August with the latest ringing occurring on 20
August. Many of these later broods must be re-lays after predation, but some are
second and possibly third broods, although it is obvious from these data that
second attempts are uncommon. Nests are heavily predated at both the egg and
chick stage. The high failure rate can be seen if the data for 1984 are examined.
This was a particularly good year, with 38 nests located. Of these, 11 (29%) were
predated, 5 (13%) were almost certainly successful, 4 (11%) were deserted at the
egg stage, 2 (5%) were definitely successful (pulli mist-netted when fully fledged)
and 1 (3%) clutch was infertile. The outcome of 15 nests (39%) was unknown.
Linnets are the only species which I have recorded incubating infertile or addled
eggs beyond the normal incubation time (2 records). The birds incubated eggs for
at least 17 days compared with the normal period of 12 days. They also regularly
have unhatched, presumed infertile eggs, with up to 4 recorded in a nest. A
brood of 1 with 4 unhatched eggs was recorded. Possible reasons include
weather, poor breeding conditions of the birds and residues from week killers
used on nearby farmland or in the wintering area. Further study with analysis of
eggs may be of value. Linnet flocks were a regular sight on the dunes from late
summer through the autumn until the mid 1970’s. Since then small parties have
been the norm. As the population has shown no serious decline, it must be
assumed that changes in habitat have reduced the summer and autumn feeding
possibilities. Linnets are shown, in Table 2, to have the longest breeding season
of any of the species studies, laying their eggs over a period of approximately 105
days. This is assumed to be due to a constant supply of seeding plants on which
to feed their young. The other species studies are all insectivorous, feeding the
young on a variety of grubs, larvae and other insects which hatch over a shorter
season. The earliest and latest dates for laying of the first egg were calculated to
be 16 April and 29 July. These were based on age and brood size at the time of
ringing and on an incubation period of 12 days (Mayer-Gross 1970). Clutch size
was taken to be the same as the brood size plus unhatched eggs.

Reed Bunting

This is the last of the ‘non-trans-saharan migrants’ to start laying. They appear
to feed their young on small green caterpillars, mainly collected from the willow
scrub. As the other insectivorous species feed their young a more varied diet, it is
probable that the availability of these caterpillars has some effect on the timing of
the Bunting’s breeding season. They appear to start laying about 10 days after
the Meadow Pipits. The peak occurs 10 days later also, with peak ringing taking
place between 31 May and 4 June. Most pairs start breeding within a 20 day
period. This is shown by the fact that 78% of broods are ringed between 21 May
and 9 June. After that, breeding and therefore ringing, continues at a slow but
steady rate. This indicates that few if any second and third broods are reared.
Peak time for laying of the first egg of a clutch is between 8-12 May, while 78% of
pairs have their first egg between 28 April and 17 May. The fact that such a high
proportion start breeding during such a short time span suggests a powerful
limiting factor that could be the food supply. There can be little delayed
breeding, due to competition for territories as suggested for Skylarks and possibly
Meadow Pipit. Of the non-trans-saharan migrants, this species shows the most
synchronized breeding pattern. Reed Buntings nest close to the ground,
occasionally on it. Nests are usually amongst grass or willow scrub and, when
built amongst the latter, are usually hidden in a grass clump. Nests in the open
dry dunes are built in marram, sea lyme grass and brambles. Clutch sizes are of 4-
5 eggs but 6 has been recorded once.

The earliest and latest dates for laying of the first egg of a clutch were calculated
to be 21 April and 2 July. This was based on brood size and age on the date of
ringing and an incubation period of 13 days (Mayer-Gross 1970). Clutch size was
taken to be the same as brood size plus unhatched eggs. There is some slight
evidence that the first breeding attempts by Reed Buntings have been 5-10 days
later in the last few years.

Willow Warbler

This is the first trans-saharan breeding migrant to arrive on the dunes, with the
first males arriving about 20 April. Most of the females arrive during the first
week of May. They pair and start nest building soon after because the peak time
for ringing is 5-9 June, therefore suggesting a first egg date of about 11-15 May
allowing for a brood size of 6, an age of 7days at ringing and an incubation period
of 13 days. The building of the nest must take 2-3 days, since it has an intricate
domed roof and a well feathered lining. This would allow the earliest female
about 3-5 days to choose a mate. Table 2 shows that the majority of Willow
Warbler pulli (75%) are ringed over a 15 day period 5-19 June. These birds will
have their first egg about 11-25 May. Table 2 also shows that the peak time of
ringing and therefore of breeding, is at the very start of the species breeding
season. This contrasts with the non-trans-saharan migrants which have peaks 10-
20 days after the dates of first attempts. The reason for this “tight” breeding
season must be that the birds need to breed as soon as possible to allow time to
moult before migration. The information in Table 2 suggests that second broods
are unlikely as the time between the earliest and latest broods is too short, unless
the female were to build a second nest shortly after the first brood has fledged.
The 4 later broods must be re-lays or exceptionally late first broods.

All nests found at Hightown except for one were built on the ground amongst
grass, willow scrub, bramble or a mixture of these plants. One was found about a
foot off the ground amongst grass and willow scrub. The entrances to the well
hidden domed nests are often hard to locate. Nests are often at the base of small
saplings or bushes but sometimes they are well away from the nearest bush in
willow scrub. Several have been located close to public footpaths used frequently
by dog-walkers. The fact that these were not predated shows how well hidden
they are. Brood sizes are usually 6-7 and when smaller are invariably
accompanied by unhatched eggs. Normal clutch would appear to be 6-7. The
earliest and latest dates for laying of the first egg of a clutch were calculated to be
8 May and 16 June. These were based on brood size, age at date of ringing, and
an incubation period of 13 days (Mayer-Gross 1970). Clutch size was taken to be
brood size plus unhatched eggs. Willow Warbler pulli were ringed over a 40-day
period.

Whitethroat

This is the second of the trans-saharan migrants to start breeding. The first
males arrive in the last few days of April or the first few days of May,
approximately 7-10 days after the Willow Warblers. From Table 2 it can be seen
that, like the Willow Warbler, peak ringing activity and therefore breeding takes
place at the start of this species breeding season. In fact it is more pronounced
than the Willow Warbler with 68% of broods being ringed over a 10-day period.
Peak ringing of pulli occurs between 5-9 June. It is interesting to note that,
despite arriving later than the Willow Warblers, the species achieves more young
of a comparable age at an earlier stage. There are several possible factors
contributing to this observation. First, female Whitethroats probably arrive
shortly after the males. Secondly, the nest of the Whitethroat is less bulky and
robust than that of the Willow Warbler and, therefore, takes less time to build.
Thirdly the clutch is smaller by 1-2 eggs. All these factors could allow the
Whitethroat to “catch up” with the Willow Warblers after its late arrival. The fact
that more pairs of Whitethroats than Willow Warblers start breeding in the first
10 days of the species breeding season indicates that either the females’ arrival is
more coordinated or that they take a shorter time to choose a mate and start
building. Perhaps female Whitethroats are in better condition on arrival. Table 2
shows that few pairs rear second broods. The very late brood ringed on 7 August
is almost certainly a second brood, even possibly a third. It is the latest date
recorded for ringing pulli of a species feeding its young on insectivorous food.
Only Linnets, feeding their young on seeds, have been recorded on the dunes
with young in the nest later than this. While this brood was still in the nest, the
majority of local Whitethroats would have migrated. The adults would have little
time to moult after rearing the brood unless they were going to stay later than
usual. Several of this brood were mist-netted on 27 August and were already
undergoing juvenile moult of wing coverts and body feathers, even though their
primaries wand retrices were still growing. From Table 2 it can be seen that 87%
of Whitethroat broods are ringed (and therefore 87% of breeding attempts
started) over a 25-day period, very similar to the Willow Warbler at 86%. The fact
that most pairs breed at about the same time suggests an adaption to migratory
needs, allowing them to moult before moving south. Whitethroat pulli have been
ringed over a 63 day period and breeding commenced over a calculated 64 day
period. Whitethroat nests are usually located close to the ground, either
suspended between vertical plant stalks or supported by willow scrub, bramble or
meadow sweet. The commonest brood size is 5 but 3-6 have been recorded. The
earliest and latest dates calculated for the laying of the first egg of a clutch are 13
May and 15 July. This is based on the age of young and brood size at the time of
ringing. The clutch size was taken to be the same as the brood size plus
unhatched eggs, while the incubation period was taken to be 13 days (Mayer-
Gross 1970).

Sedge Warbler

Probably the least common of the 3 species of breeding warblers studied.

Certainly its nest is the most difficult to locate, often being well hidden in dense
vegetation. The adults often disappear into the vegetation, well before they have
reached the nest, which is typically built low down, sometimes even on the
ground. Nests are sometimes supported between the vertical stems of plants,
like the nests of the Reed Warbler. Typical associated vegetation includes
grasses, nettles, reed mace, willow scrub and loosestrife. Brood sizes of 4-6 are
the norm.

The first males return towards the end of April or the beginning of May,
arriving after the Willow Warbler but before the Whitethroat. It is therefore
interesting to note from Table 2 that it is the last of the warbler species to breed.
The first Sedge Warblers to breed lay their eggs approximately 5 days after the
first Whitethroats. There are several possible reasons for this delay, such as:
 i. females arriving a lot later than the males;
ii. females taking longer to choose a mate and/or nest site;
iii. the birds are not in breeding condition when they arrive. This could
be due to long distance non-stop flight depleting their reserves.
Other species could migrate more leisurely with rests to feed and
so be in better condition when they arrive;
iv. the vegetation in which they breed may not have grown enough for
them to build their nests.

I think that the lack of vegetation is the most likely explanation. Many of the
nests have been found amongst plants which grow from ground level every year
and do not reach full growth until July or August. The birds would therefore have
to wait for sufficient growth to allow nest support and concealment. The fairly
even spread of breeding attempts throughout the breeding season must also
reflect the fact that males and presumably females take up territories as late as
the end of May or even early June. There are two possible reasons for this. First,
birds may have just arrived from Africa. Secondly, birds may be moving around
looking for suitable breeding habitat and as vegetation grows, areas which at first
were unsuitable become viable nesting sites. These presumably are secondary
habitats. A recent study showed that singing males in late April to early May had
territories in wet areas with plenty of undergrowth whilst later occupied
territories were in drier areas with less dense undergrowth and more grassland.
Presumably these territories become more suitable as fresh growth provides
better cover. Table 2 shows no peak time for ringing pulli (and therefore for the
start of breeding) although the data are rather limited reflecting a difficulty in
nest location. It does suggest that more breeding attempts are made in the first
half of the breeding season. First eggs are laid over a period of approximately 35
days, the shortest span recorded for any of the study species. This is probably
shorter than the true time period as mist-netting of young birds with ‘wings in
pin’ can take place will into August. This generally late breeding season may be
the reason why the adults do not moult before migrating south and therefore
allows them to depart earlier than the other warblers.

The earliest and latest dates for the laying of the first egg of a clutch were
calculated to be 22 May and 20 June. This was based on age and brood size at
time of ringing and an incubation period of 13 days (Mayer-Gross 1970). Clutch
size was taken to be brood size plus any unhatched eggs at time of ringing.
 SUMMARY AND DISCUSSION

Table 7 – Summary of breeding data collected from eight passerine species

i. laying dates for the first eggs of clutch * = insufficient data

Earliest Peak Latest Duration, days

Skylark 06 April 18-22 April 04 July 90
Stonechat 05 April 12-16 April 31 May 57
Meadow Pipit 11 April 28 Apr– 2 May 28 June 79
Linnet 16 April 04-08 May 29 July 105
Reed Bunting 21 April 08-12 May 02 July 73
Willow Warbler 08 May 11-15 May 16 June 40
Whitethroat 13 May 13-17 May 15 July 64
Sedge Warbler 22 May * 20 June 35*

ii. ringing dates * = insufficient data

Earliest Peak Latest

Skylark 24 April 06-10 May 22 July
Stonechat 29 April 06-10 May 25 June
Meadow Pipit 03 May 21-25 May 21 July
Linnet 09 May 26-30 May 20 August
Reed Bunting 13 May 31 May- 4 June 24 July
Willow Warbler 01 June 05-09 June 09 July
Whitethroat 06 June 05-09 June 07 August
Sedge Warbler 13 June * 14 July

Although Tables 3 – 6 show that there is variation from year to year in the
timing of the breeding season for each species, Table 2 shows that overall, each
species shows fairly constant trends. In the case of non-trans-saharan migrants,
there are a few early breeding pairs followed by a period during which the largest
proportion of the birds start breeding simultaneously. This is then followed by a
decrease in numbers starting to nest and breeding then continues at a low level
until the end of the season. Only the Skylark departs from this pattern. It shows
the lowest peak, with breeding commencing at a more even rate throughout the
season. The trans-saharan migrants show a different breeding strategy with a
high proportion of pairs starting to breed simultaneously at the beginning of the
season, presumably related to the need to breed as soon as possible and to the
timing of the females arrival. They also have shorter breeding seasons than the
other species, except the Stonechat, presumably because of the need to migrate.
Willow Warbler and Whitethroat start breeding as soon as the females arrive, but
Sedge Warblers appear to delay for a time possibly because of slow vegetation
growth at the nesting sites or the later arrival of the females, who may be in poor
condition after a longer non-stop flight. It is interesting to note that Sedge
Warblers, at least the first males, arrive before the first Whitethroats. Willow
Warblers, Whitethroats and probably Sedge Warblers show much less variation
from year to year in the timing of their breeding. No doubt this is the result of the
tight schedule of two migrations, a breeding season and a complete moult (except
for Sedge Warblers) in 5-6 months. Table 7 is a summary of the information
gathered and shows earliest, latest and peak ringing dates for each species. It
also shows the calculated earliest, latest and peak laying dates for the first eggs of
a clutch. From Table 7 it can be seen that the order of ringing the first brood of
each species is Skylark, Stonechat, Meadow Pipit, Linnet, Reed Bunting, Willow
Warbler, Whitethroat and Sedge Warbler. A period of 4 to 19 days passes
between the ringing of the first brood of consecutive species. The non-trans-
saharan migrants start breeding first, with periods of between 4-6 days separating
the ringing of first broods of consecutive species. There is then a period of 19
days before the ringing of the first broods of trans-saharan migrants (Willow
Warblers) followed by 5-7 days separating the first broods of the other trans-
saharan migrants. Peak ringing times are also staggered although Stonechat and
Skylark share similar dates, as do Willow Warbler and Whitethroat. When the
timing of the first egg dates is examined it is found that Stonechats are the first to
lay with Skylarks second. This change in order is due to the fact that Skylarks lay
smaller clutches, incubate for a shorter time, and are ringed at a younger age.
Apart from this, the order of laying is the same as for ringing. When the “Peak
laying” only is analysed we find that Stonechats lay almost a week earlier than
Skylarks, otherwise the order is the same as for ringing. The species order for
laying of the latest clutches is, Stonechat, Willow Warbler, Sedge Warbler,
Meadow Pipit, Reed Bunting, Skylark, Whitethroat and Linnet. The order for
ringing of the latest broods is the same except that the Skylark and Reed Bunting
change positions. Linnets, the only species feeding seeds to its young breed the
latest, presumably due to the abundant summer seeding plants and grasses. It is
interesting to note that the Whitethroat, a trans-saharan migrant that moults
before migrating south, is recorded as the latest breeding insectivorous species.
The species order for length of breeding season (defined as the period of time
during which first eggs are laid) is Linnet, Skylark, Meadow Pipit, Reed Bunting,
Whitethroat, Stonechat, Willow Warbler and Sedge Warbler. Linnet shows the
longest breeding season while the trans-saharan migrants are among the species
with the shortest. There appears to be no correlation between the date that a
species starts and the length of its breeding season apart from the observation
that species starting to breed earlier, have on average, longer breeding seasons,
as is to be expected. It is also interesting to note that Willow Warbler and
Whitethroat have very similar peak breeding periods despite the difference in
arrival times. The non-trans-saharan migrants arrive on the dunes at around the
same time. Why then, do they not start breeding at the same time? Presumably
because of differing diets. If we look at feeding habits it becomes apparent that
the species collect their food in different ways and at different levels in the
vegetation. This leads to differing diets and also minimises inter-specific
competition. Skylarks collect their food at ground level often along the edges of
paths. Stonechats collect food by dropping from a perch onto prey, either on
vegetation or the ground. Alternatively, they catch insects like a flycatcher.
Meadow Pipits mostly pick food from vegetation close to the ground, often while
walking along the edges of paths. Linnets collect their food from flower or grass
heads. Reed Buntings eat mainly caterpillars found in willow scrub and other low
vegetation. They differ from Meadow Pipits in that they climb around in the
foliage usually at a higher level. Willow Warblers collect most of their food from
amongst the willow bushes whilst Whitethroats appear to use similar habitat to
both Willow Warbler and Reed Bunting. Sedge Warblers search willow bushes
and low vegetation in damp areas and so are only likely to come into conflict with
Reed Buntings. This variation in feeding ecology between the species will reduce
inter-specific competition and affect the timing of the breeding season. From
Tables 2 and 7 it can be seen that ringing of pulli takes place during a period of
time that varies from species to species. This is invaluable to the ringer as he
need not waste time watching a species when it is “out of season” and can
concentrate when it should have young in the nest. Consequently Tables 2 and 7
should be of use to ringers who intend to ring pulli of the species studied.
Although the dates for each species may not be directly comparable to other
study areas, due to differences in latitude and height above sea level, the order of
first breeding attempts and pulli ringing may remain the same. It is hoped that
further articles on the results of pulli ringing on the dunes will appear in future
reports. Among intended subjects are:

1. Recoveries (very few so far!)

2. Retraps in future years.
3. Sex of retraps in future years – do more males than females
return to the area?
4. Longevity.
5. Variation of brood sizes in a season and from year to year.