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

Skylark April 16 - 20 21 - 25 26 - 30 May 01 - 05 06 - 10 11 - 15 16 - 20 21 - 25 26 - 30 31 - 04 June 05 - 09 10 - 14 15 - 19 20 - 24 25 - 29 30 - 04 July 05 - 09 10 - 14 15 - 19 20 - 24 25 - 29 30 - 03 August 04 - 08 09 - 13 14 - 18 19 - 23 Total 302 1 (0.3) 3 (1.0) 12 (4.0) 18 (6.0) 52 (17.2) 31 (10.3) 23 (7.6) 20 (6.6) 38 (12.6) 24 (8.0) 28 (9.3) 16 (5.3) 14 (4.6) 6 (2.0) 8 (2.7) 4 (1.3) 3 (1.0) 1 (0.3)


Meadow Pipit


Reed Bunting

Willow Warbler


Sedge Warbler

1 (8.3) 2 16.7) 3 (25.0) 2 (16.7) 2 (16.7) 1 (1.5) 2 (3.0) 11 (16.7) 11 (16.7) 16 (24.2) 3 (4.5) 5 (7.6) 1 (8.3) 2 (3.0) 5 (7.6) 1 (1.5) 4 (6.1) 1 (8.3) 3 (4.5) 1 (1.5) 1 (1.4) 3 (4.2) 10 (14.1) 14 (19.7) 15 (21.1) 9 (12.7) 4 (5.6) 3 (4.2) 2 (2.8) 2 (2.8) 1 (1.4) 2 (2.8) 2 (2.8) 1 (1.2) 2 (2.3) 12 (14.0) 14 (16.3) 23 (26.7) 18 (20.9) 3 (3.5) 2 (2.3) 2 (2.3) 2 (2.3) 1 (1.2) 2 (2.3) 1 (1.2) 1 (1.2) 1 (1.5) 1 (1.4) 1 (1.4) 1 (3.2) 2 (2.3) 1 (3.2) 1 (3.6) 3 (10.7) 2 (6.5) 1 (3.6) 10 (35.7) 5 (17.9) 6 (21.4) 2 (7.1) 13 (41.9) 8 (25.8) 3 (9.7) 1 (3.2) 2 (6.5) 2 1 2 1 1 1 1

1 (1.4) 12 * 66 71 86 28* 31* 9

* data from 1986 included

Table 3 – Skylark brood ringing dates 1977-1985
1977 April 16 - 20 21 - 25 26 - 30 01 - 05 06 - 10 11 - 15 16 - 20 21 - 25 26 - 30 31 - 04 05 - 09 10 - 14 15 - 19 20 - 24 25 - 29 30 - 04 05 - 09 10 - 14 15 - 19 20 - 24 25 - 29 1978 1979 1 (2.5) 3 (7.5) 1 (2.5) 7 (17.5) 1 (2.5) 1 (2.5) 3 (7.5) 7 (17.5) 6 (15.0) 1 (2.5) 2 (5.0) 1 (2.5) 2 (5.0) 3 (7.5) 1980 1 (2.1) 2 (4.2) 8 (16.7) 6 (12.5) 4 (8.3) 4 (8.3) 3 (6.3) 5 (10.4) 5 (10.4) 2 (4.2) 5 (10.4) 3 (6.3) 1981 1982 1 (3.0) 2 (6.1) 2 (6.1) 8 (24.2) 1 (3.0) 1 (3.0) 2 (6.1) 5 (15.2) 6 (18.2) 2 (6.1) 1 (3.0) 2 (6.1) 1 (3.4) 1 (2.5) 1 (2.5) 1983 1984 1985

4 (12.1) 1 (5.6) 4 (22.2) 3 (16.7) 4 (22.2) 2 (11.1) 4 (12.1) 3 (9.1) 1 (3.0) 1 (3.0) 12 (36.4) 2 (6.1) 4 (12.1)



1 (3.2) 4 (12.9) 4 (12.9) 3 (9.7) 1 (3.2) 7 (22.6) 3 (9.7) 2 (6.5) 1 (3.2) 3 (9.7) 2 (6.5)

1 (3.3) 2 (6.7) 9 (30.0) 2 (6.7) 7 (23.3) 1 (3.3) 3 (10.0) 1 (3.3) 3 (10.0) 1 (3.3)

1 (2.5) 10 (25.0) 14 (35.0) 3 (7.5) 3 (7.5) 1 (2.5) 2 (5.0) 3 (7.5) 2 (5.0)

1 (3.4) 12 (41.4) 1 (3.4) 1 (3.4) 6 (20.7) 1 (3.4) 5 (17.2) 1 (3.4)

3 (16.7) 1 (5.6)

2 (6.1)


Table 4 – Meadow Pipit brood ringing dates 1980-1985
1980 April May 26 - 30 01 - 05 06 - 10 11 - 15 16 - 20 21 - 25 26 - 30 31 - 04 05 - 09 10 - 14 15 - 19 20 - 24 25 - 29 30 - 04 05-09 10-14 15 - 19 20 - 24 25 - 29 1981 1 (12.5) 1 (12.5) 3 (37.5) 1 (12.5) 1982 1983 1984 1985

1 (11.1) 1 (11.1) 1 (11.1) 1 (11.1) 2 (22.1)


1 (12.5) 2 (22.1) 1 (11.1) 1 (12.5)

1 (12.5) 1 (12.5) 1 (12.5) 2 (25.0) 1 (12.5) 1 (12.5)

3 (23.1) 1 (7.7) 7 (53.8) 1 (7.7) 1 (7.7)

3 (27.3) 3 (27.3)

1 (6.3) 4 (25.0) 3 (18.8) 4 (25.0) 1 (6.3)

1 (9.1) 1 (9.1) 1 (9.1)

1 (6.3) 1 (6.3) 1 (6.3)

1 (12.5) 1 (9.1)


1 (9.1)

Table 5 – Linnet brood ringing dates 1977-1985
1977 April 16 - 20 21 - 25 26 - 30 01 - 05 06 - 10 11 - 15 16 - 20 21 - 25 26 - 30 31 - 04 05 - 09 10 - 14 15 - 19 20 - 24 25 - 29 30 - 04 05 - 09 10 - 14 15 - 19 20 - 24 25 - 29 20 1978 1979 1980 1981 1982 1983 1984 1985


1 (4.8) 1 2 1 1 2 (50.0) 1 (25.00 1 (25.00 1 (4.3) 2 (8.7) 7 (30.4) 7 (30.4) 2 (8.7) 1 (4.3) 1 (4.3) 1 (4.3) 1 1 1 (4.3) 4 (19.0) 2 (9.5) 3 (14.3) 1 (4.8) 2 (9.5) 1 (4.8) 1 (4.8) 1 (4.8) 1 (4.8) 1 (4.8)

1 2 2

1 2


1 (12.5) 3 (37.5) 1 (12.5) 1 (12.5) 1 (12.5) 1 (12.5)



1 (4.8) 1 (4.8) 1 (4.8)

Table 6 – Reed Bunting brood ringing dates 1977-1985
1977 April 16 - 20 21 - 25 26 - 30 01 - 05 06 - 10 11 - 15 16 - 20 21 - 25 26 - 30 31 - 04 05 - 09 10 - 14 15 - 19 20 - 24 25 - 29 30 - 04 05 - 09 10 - 14 15 - 19 20 - 24 25 - 29 1978 1979 1980 1981 1982 1983 1984 1985


1 (12.5) 2 1 1 (7.1) 1 (7.1) 6 (42.9) 5 (35.7) 2 (15.4) 2 (15.4) 3 (23.1) 2 (15.4) 1 (7.7) 2 (25.0) 1 (12.5) 3 (37.5) 2 (28.6) 1 (14.3) 2 (28.6) 1 (14.3) 1 1 (14.3) 1 (7.7) 1 (7.7) 1 (7.1) 1 (7.7) 1 (11.1) 1 (12.5) 1 (11.1) 1 (7.1) 1 (7.1) 1 (11.1) 2 (22.2) 2 (22.2) 2 (22.2) 2 (16.7) 3 (25.0) 2 (16.7) 2 (16.7) 3 (25.0)


2 1 2

1 (7.1) 6 (42.9) 4 (28.6) 1 (7.1)


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 45 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 1020 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 (MayerGross 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.


Table 7 – Summary of breeding data collected from eight passerine species i. laying dates for the first eggs of clutch * = insufficient data

Skylark Stonechat Meadow Pipit Linnet Reed Bunting Willow Warbler Whitethroat Sedge Warbler

Earliest 06 April 05 April 11 April 16 April 21 April 08 May 13 May 22 May

Peak 18-22 April 12-16 April 28 Apr– 2 May 04-08 May 08-12 May 11-15 May 13-17 May *

Latest 04 July 31 May 28 June 29 July 02 July 16 June 15 July 20 June

Duration, days 90 57 79 105 73 40 64 35*

ii. ringing dates * = insufficient data

Skylark Stonechat Meadow Pipit Linnet Reed Bunting Willow Warbler Whitethroat Sedge Warbler

Earliest 24 April 29 April 03 May 09 May 13 May 01 June 06 June 13 June

Peak 06-10 May 06-10 May 21-25 May 26-30 May 31 May- 4 June 05-09 June 05-09 June *

Latest 22 July 25 June 21 July 20 August 24 July 09 July 07 August 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-transsaharan 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 transsaharan 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.