This article was downloaded by: [Ove Arup Partnership] On: 16 November 2011, At: 04:26 Publisher: Taylor

& Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK

Bird Study
Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/tbis20

Breeding ecology of Turtle Doves Streptopelia turtur in Britain during the period 19412000: an analysis of BTO nest record cards: Capsule No trends over time were detected in any aspect of Turtle Dove breeding ecology and only slight regional variation, based on individual nesting attempts recorded.
Stephen J. Browne, Nicholas J. Aebischer & Humphrey Q.P. Crick Available online: 29 Mar 2010

To cite this article: Stephen J. Browne, Nicholas J. Aebischer & Humphrey Q.P. Crick (2005): Breeding ecology of Turtle Doves Streptopelia turtur in Britain during the period 19412000: an analysis of BTO nest record cards: Capsule No trends over time were detected in any aspect of Turtle Dove breeding ecology and only slight regional variation, based on individual nesting attempts recorded., Bird Study, 52:1, 1-9 To link to this article: http://dx.doi.org/10.1080/00063650509461368

PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: http://www.tandfonline.com/page/terms-and-conditions This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae, and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand, or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material.

Bird Study (2005) 52, 19

Breeding ecology of Turtle Doves Streptopelia turtur in Britain during the period 19412000: an analysis of BTO nest record cards
STEPHEN J. BROWNE1*, NICHOLAS J. AEBISCHER1 and HUMPHREY Q.P. CRICK2
1The

Game Conservancy Trust, Fordingbridge, Hampshire SP6 1EF, UK and 2British Trust for Ornithology, The Nunnery, Thetford, Norfolk IP24 2PU, UK

Downloaded by [Ove Arup Partnership] at 04:26 16 November 2011

Capsule No trends over time were detected in any aspect of Turtle Dove breeding ecology and only slight regional variation, based on individual nesting attempts recorded. Aim To present information on the breeding ecology of Turtle Doves and identify any temporal or regional trends that may have contributed to the decline of the species. Methods Information on nesting habitat, type of bush used, nest height, clutch size, brood size and nest outcome was extracted from 1925 Turtle Dove nest record cards from 1941 to 2000, and examined for temporal and regional trends. Results The majority of Turtle Doves nest in thorny trees within scrub habitats, where mean nest height was 2.27 0.02 m. Mean first-egg date was 18 May 1 day (annual range 28 April to 26 May). Mean clutch size was 1.84 0.01 (annual range 1.651.93), producing a mean brood size of 1.82 0.01 (range 1.502.00). Based on the recorded fate of individual nests, 41.3 1.4% were successful, 44.9 1.7% were predated and the rest (13.9 1.4%) were lost to other causes. Nest survival rate averaged 0.577 0.019 during the 14-day incubation period and 0.771 0.019 during the 15-day nestling period giving an overall rate during the entire nesting period of 0.445 0.018. Although there was some significant variation in these parameters between time periods and regions, there were no significant linear trends during the period 19412000. Conclusion The population decline experienced by Turtle Doves breeding in Britain is not due to lower success of individual nesting attempts. If breeding productivity has played a role in the decline, it must be through a reduction in the average number of nesting attempts per pair.

The Turtle Dove Streptopelia turtur is one of a number of British and European bird species that inhabit farmland and have undergone sustained declines in abundance and contractions in range (Gibbons et al. 1993, Fuller et al. 1995, Heath et al. 2000, Baillie et al. 2002). In Britain, Turtle Doves have declined in abundance by 70% and in range by 25% over the last 40 years (Gibbons et al. 1993, Baillie et al. 2002). In view of these declines the Turtle Dove is recognized as a Red List species of conservation concern in the UK (Gregory et al. 2002), is one of the UKs Biodiversity Action Plan species (Anon 1995) and is a Category 3 species of European Conservation Concern (Heath et al. 2000). Following one of the recommendations of the Biodiversity Action Plan, an intensive research project
*Correspondence author. Email: sbrowne@gct.org.uk

was undertaken in Britain between 1996 and 2001 to identify causes of the species decline and to produce recommendations for its conservation (Calladine et al. 1997, Browne & Aebischer 2001, 2003b, 2004). Browne & Aebischer (2004) found that general aspects of Turtle Dove breeding ecology had not changed since an early study by Murton (1968), but that the average number of nesting attempts undertaken per pair each year had almost halved. As independent corroboration of this result, the timing of Turtle Dove autumn migration has become earlier and matches the earlier termination of the species breeding season (Browne & Aebischer 2003a). The breeding productivity of British Turtle Doves during 1998-2000, as measured by Browne & Aebischer (2004), would result in an annual population decrease of 17%, which easily accounts for the observed rate of decline. The underlying cause of this reduction in breeding performance remains

2005 British Trust for Ornithology

S.J. Browne, N.J. Aebischer and H.Q.P. Crick

Downloaded by [Ove Arup Partnership] at 04:26 16 November 2011

unknown, but is thought to result from the removal and degradation of nesting habitats (Browne & Aebischer 2004) and a possible switch in diet brought about by reduced food availability (Browne & Aebischer 2003b). Historical information held by the British Trust for Ornithologys Nest Record Scheme (Crick & Baillie 1996), a data set of nesting attempts undertaken by UK breeding birds, offers the opportunity to examine Turtle Dove nesting ecology at a much larger geographical scale than allowed by the intensive study of Browne & Aebischer (2004). A number of studies using Nest Record Scheme data have successfully related demographic parameters to population and environmental changes (Crick et al. 1994, Thomson et al. 1997, Peach et al. 1999, Siriwardena et al. 2000, 2001). We investigate whether Turtle Dove nesting ecology, as measured by Nest Record Scheme data, has changed over the last 60 years, to seek further insight into the possible reasons for the species decline. Potential changes that the Nest Record Scheme data have enabled us to consider include: (1) changes in the type of habitat or tree used for nesting, following alterations to habitat availability and quality (OConnor & Shrubb 1986, Macdonald & Johnson 2000); (2) these may in turn have altered nest height or increased predation risk; (3) first-egg date may change, as birds may need to spend more time at the start of the breeding season finding suitable habitat or attaining suitable body condition; (4) this may also have affected clutch and brood size; (5) increased predator abundance (Gregory & Marchant 1996) may have affected nest outcome.
METHODS

The regions were established by eye after visual inspection of information provided by Gibbons et al. (1993) on Turtle Dove distribution, abundance and range contraction. The regionsa were based on county boundaries and were defined as (1) counties supporting the highest abundance of Turtle Doves; (2) counties with lowest abundance; (3) counties where Turtle Doves were present during 196872, but absent during 198891 (Fig. 1). Six cards, for which the regions were either not recorded or were well outside the defined regions (i.e. in Scotland), were excluded from the analysis. When investigating temporal trends and to overcome the problem of small sample sizes, particularly in the early and late years, data were grouped into five-year periods, from 194145 through to 19962000 (Table 1). In order to investigate breeding success amongst periods with different rates of population change we defined three phases in the population trend of British Turtle Doves, based on information provided by Marchant et al. (1990), Siriwardena et al. (1998) and Browne & Aebischer (2003c). These phases corresponded to a period of population increase (196175), stability (197685) and decrease (19862000).
Nest site

In order to consider the habitat where the nest was located, information on the cards was grouped into five broad categories based on the descriptions of general

In total, 1931 Turtle Dove nest record cards (Crick & Baillie 1996) were submitted to the British Trust for Ornithology for the period 19412000 and the information they contain partially computerized. However, not all of the cards contain all the potential information and many are based on only one visit to the nest, whereas several aspects of nesting ecology require at least two visits for evaluation. Information on the habitat where the nest was located, type of woody vegetation supporting the nest, nest height, clutch size, brood size and nest outcome was extracted from the nest record cards. Not all of the information for these parameters was available for the entire period 19412000, so some were considered over a shorter period. This information was assigned to three geographical regions, 12 five-year periods and three phases in the population trend of British Turtle Doves.
2005 British Trust for Ornithology, Bird Study,
52, 19

Figure 1. Map of regions used in the analyses. Region 1 comprises counties supporting the highest abundance of Turtle Doves, Region 2 those with lowest abundance and Region 3 those where Turtle Doves were present during 196872 but absent during 198891 (Gibbons et al. 1993).

Temporal changes in Turtle Dove breeding ecology

Table 1. Total number of Turtle Dove nest records that have been submitted to the BTO during each five-year period between 1950 and 2000 and the number that had more than one visit (see Fig. 1 for regions). Region Five-year period 194145 194650 195155 195660 196165 196670 197175 197680 198185 198690 199195 19962000 Total 1 51 4 46 43 79 191 158 162 62 55 32 71 954 2 1 10 52 38 145 142 119 137 96 69 32 25 866 3 0 2 9 5 35 12 7 11 10 10 3 1 105 Total no. of cards 52 16 107 86 259 345 284 310 168 134 67 97 1925 No. of cards with multiple visits 10 9 49 38 110 150 131 153 84 76 40 54 904

Downloaded by [Ove Arup Partnership] at 04:26 16 November 2011

nesting habitats (Crick 1992). These were hedgerow, plantation, scrub, woodland and isolated bushes (which included bushes in gardens). The woody vegetation supporting the nest was categorized into a number of broad types based on the main tree species identified by Browne & Aebischer (2004) as being important for Turtle Doves. The categories were: Elder Sambucus nigra, fruit trees, thorn (Hawthorn Crataegus monogyna and Blackthorn Prunus spinosa), broadleaved (excluding Elder, thorn and fruit trees) and coniferous. For each card, nest outcome was categorized as: successful, predated, lost from causes other than predation, and unknown. Multinomial logistic regressionb was used to compare the number of cards in each of the habitat, woody vegetation and outcome categories, based on counts of cards within each region, in each of the five-year periods and during the three different population phases. An analysis of variance was used to compare mean nest height (log-transformed) between regions, fiveyear periods and the three different population phases. Before the mid-1980s nest height was usually given in feet and this was converted to metres before analysis.
Timing of breeding

although other authors (Siriwardena et al. 2000) have used 20 days as the length of the nestling period. For some cards it was possible to calculate only minimum and maximum first-egg dates; in this case a mean of the two values was used, provided that the difference between these two values was less than 14 days, otherwise the data were excluded. Days were assigned a numerical value where 1 March = 60. Differences in mean first-egg dates between the regions, the five-year periods and the three different population phases were tested with an analysis of variance. In each of the fiveyear periods 194145 and 19962000 it was only possible to calculate first-egg date for one card in each period in region 3, so the analysis was restricted to regions 1 and 2 during these two periods.
Clutch and brood size

First-egg date was calculated for all nest record cards where there were sufficient visits to enable an accurate estimate to be made. First-egg date estimates took into account laying period, the timing of hatching and fledging. They were based on an incubation period of 14 days and nestling periods of 15 days based on the intensive study of Browne & Aebischer (2004),

Multinomial logistic regressionb was used to compare the clutch and brood size between the regions, the fiveyear periods and the three different population phases, based on counts of cards having either one egg or nestling and two or more eggs and nestlings. It was not possible to identify precisely the clutch size at onset of incubation, the brood size at hatching or the brood size at fledging, because nest contents were reported only when each nest was visited. Instead the maximum clutch and brood size recorded for each nest was used.
Nest success

The survival of nests recorded on the nest record cards was analysed using the Mayfield method (Mayfield
2005 British Trust for Ornithology, Bird Study,
52, 19

S.J. Browne, N.J. Aebischer and H.Q.P. Crick

Downloaded by [Ove Arup Partnership] at 04:26 16 November 2011

1961, 1975, Hensler & Nichols 1981), which was used to calculate the daily survival rate for Turtle Dove nests during the incubation and nestling periods separately. The probabilities of nest survival over the whole of the incubation and of the nestling period were estimated by raising the corresponding daily survival rates to the power 14 and 15 respectively; their product gave an estimate of overall success. Standard errors on these estimates were calculated following Hensler (1985). Only nests visited more than once were included in the analysis. The analysis was based on nest outcome and partial losses were not considered. If the precise day of failure or hatching was not known the mid-point between the two visits either side of the event was used. An extension to the Mayfield method using logistic regression (Aebischer 1999) was used to investigate variations in nest survival rates during the incubation and nestling periods, in relation to region, the five-year periods and the three different population phases. Linear trends over time in median nest height, mean first-egg date, mean clutch and brood size and overall nest survival rates during the incubation and nestling stages were tested using regression against the midpoint of each five-year period, weighting by 1/se2 to take into account the accuracy of each estimate.

It is important to consider that there are potential biases in looking at the frequencies of habitat use, nest location and nest heights recorded on nest record cards because these depend, to a certain extent, on searching preferences of the observers. However, it was assumed that this information is broadly representative and that when considering differences between regions and years any biases were assumed to be constant between regions and years. Where appropriate the interactions between the factors were considered in all the analyses. The analyses were carried out using the statistical packages SYSTAT 10 (SPSS Inc., Chicago) and GENSTAT 6 (Genstat Committee 2002).
RESULTS Nest site

The nesting habitat recorded most often was scrub, followed by woodland and then hedges, with coniferous plantations being recorded least often (Table 2). There was no significant interaction between time period and region (288 = 65.37, P > 0.05). Although there was a significant difference in the proportion of nests

Table 2. Habitats used by Turtle Doves (%) for nesting during the period 19412000, in three geographical regions and during the three population phases as recorded on nest record cards (see Fig. 1 for regions). Classification Five-year period 194145 194650 195155 195660 196165 196670 197175 197680 198185 198690 199195 19962000 Region Region 1 Region 2 Region 3 Population phase Increase Stable Decrease Overall No. of cards Hedge Plantation Scrub Woodland Isolated bush

46 17 99 86 245 331 270 291 146 129 61 97

60.8 33.3 7.1 13.9 10.1 19.1 11.8 17.5 19.2 19.4 22.7 25.7

0.0 6.7 7.1 4.6 10.9 8.2 7.4 5.8 5.5 3.1 15.2 0.0

32.6 26.7 61.6 52.3 50.0 41.9 50.0 47.7 50.7 51.2 37.8 39.2

0.0 13.3 13.2 18.6 22.9 25.4 26.3 22.7 21.9 13.2 13.6 5.2

6.5 20.0 11.1 10.5 6.1 5.4 4.4 6.2 2.7 13.2 10.6 29.9

904 817 97

22.2 12.6 12.4

5.1 9.1 5.2

42.0 52.6 55.7

20.6 20.4 18.6

10.1 5.3 8.2

437 661 560 1818

21.3 15.1 14.8 17.4

5.0 8.8 6.6 6.9

46.5 46.4 48.9 47.5

14.4 23.8 24.3 20.4

12.8 5.9 5.4 7.8

2005 British Trust for Ornithology, Bird Study,

52, 19

Temporal changes in Turtle Dove breeding ecology

Downloaded by [Ove Arup Partnership] at 04:26 16 November 2011

recorded in each of the habitat categories in each of the time periods between 1941 and 2000 (244 = 248.14, P < 0.001) and in each of the population phases (28 = 67.76, P < 0.001), no clear pattern over time was apparent (Table 2). Nesting habitat differed significantly among regions (28 = 40.7, P < 0.001), with birds nesting in region 1 making more use of hedges and isolated bushes whereas birds in the other regions used scrub more. Considering the type of woody vegetation supporting the nest, thorny trees (principally Hawthorn and Blackthorn) were the type of tree most often recorded (Table 3). There was no significant interaction between region and time period (272 = 41.42, P > 0.05). Although the number of nests in each of the tree type categories varied significantly between the time periods (236 = 70.91, P < 0.001), there was no discernible trend over time or between population phases (28 = 14.01, P > 0.05). The type of woody vegetation supporting the nest differed significantly between regions (28 = 73.11, P < 0.001). Birds in region 3 sited their nests more frequently in coniferous and broadleaved vegetation and less frequently in thorn and elder than in the other regions.
Nest height

significantly between the five-year periods from 1941 to 2000 (F11,1837 = 1.56, P > 0.05) or between regions (F2,1837 = 1.07, P > 0.05). There was no significant linear trend in the means during 1941 to 2000 (r10 = 0.104, P > 0.05) or during the population phases (F2,1702 = 2.02, P > 0.05). The majority of nests were found at between 1 and 3 m and overall mean nest height during 1941 to 2000 was 2.27 0.02 m (n = 1854, range = 0.2012.22 m).
First-egg date

First-egg date was calculated for 978 nests found during the period 19512000. There was no significant interaction between five-year period and region when considering first-egg date (F17,943 = 1.59, P > 0.05). First-egg date was significantly different between fiveyear periods (F9,959 = 2.55, P = 0.006), but not region (F2,959 = 0.76, P > 0.05). There was no significant linear trend over the period 19502000 (r8 = 0.058, P > 0.05) in mean first-egg date or during the population phases (F2,892 = 0.89, P > 0.05). Annual mean first-egg date during the period 19512000 ranged from 28 April to 26 May (Table 4).
Clutch and brood size

There was no significant interaction between five-year period and region when considering nest height (F21,1817 = 1.39, P > 0.05). Nest height did not vary

Clutch size was recorded for 1506 nests during the period 19412000. Sixteen per cent of nests contained one egg, 83% contained two eggs and only 1%

Table 3. Use (%) of different types of tree by nesting Turtle Doves during the period 19412000, in three geographical regions as recorded on nest record cards (see Fig. 1 for regions). Classification Five-year period 194650 195155 195660 196165 196670 197175 197680 198185 198690 199195 19962000 Region Region 1 Region 2 Region 3 Overall No. of cards Elder Fruit Thorn Broadleaved Coniferous

14 94 73 237 319 266 288 153 128 60 96

7.1 14.9 17.8 22.4 18.2 18.0 7.6 14.4 14.1 6.7 10.4

0.0 1.1 2.7 8.0 2.8 2.6 1.0 1.3 1.6 1.7 0.0

64.3 52.1 45.2 43.0 53.6 53.8 59.4 51.6 63.3 53.3 71.9

21.4 23.4 27.4 19.8 18.5 18.4 24.0 21.6 15.6 21.7 15.6

7.1 8.5 6.8 6.8 6.9 7.1 8.0 11.1 5.5 16.7 2.1

835 799 94 1728

17.5 13.8 7.5 15.2

4.0 1.4 2.2 2.7

56.8 53.2 43.0 54.3

17.5 21.7 33.3 20.3

4.3 10.0 15.1 7.5

2005 British Trust for Ornithology, Bird Study,

52, 19

S.J. Browne, N.J. Aebischer and H.Q.P. Crick

Table 4. Mean ( se) first-egg date and clutch size of Turtle Dove nests recorded on nest record cards. First-egg dates are expressed as numeric values, where 1 March = 60. It was not possible to calculate first-egg date during 194150. First-egg date Five-year period 194145 194650 195155 195660 196165 196670 197175 197680 198185 198690 199195 199600 No. of cards Mean s.e Clutch size No. of cards Mean se 40 12 87 71 197 266 223 260 122 102 52 74 1.65 1.75 1.83 1.85 1.87 1.87 1.85 1.83 1.77 1.81 1.81 1.93 0.08 0.18 0.04 0.04 0.03 0.02 0.02 0.02 0.04 0.04 0.05 0.03

differences were detected between the population phases (26 = 5.25, P > 0.05). Brood size did not vary significantly with region (24 = 0.69, P > 0.05).
Nest fate

Downloaded by [Ove Arup Partnership] at 04:26 16 November 2011

46 44 133 159 124 156 102 80 41 93

127.4 133.0 144.4 134.3 138.1 140.7 136.1 133.7 139.6 136.2

4.2 3.2 2.6 1.9 2.0 1.9 2.1 2.6 3.3 2.1

The fate of 1263 nests (65.5%) was recorded on nest record cards during the period 19412000 (Table 5). For the nests where nest fate was known, there was no significant interaction between time period and region (216 = 49.25, P > 0.05). Nest fate varied significantly with region (24 = 18.40, P = 0.002) but not time period (222 = 29.61, P > 0.05) or population phase (24 = 2.35, P > 0.05). Overall 41.3 1.5% of nests successfully produced young, 44.9 1.5% were predated, with the remainder (13.9 1.4%) being lost to other causes (Table 5). More nests were successful and fewer nests were predated in region 1 than in regions 2 and 3 (Table 5).
Nest success

contained three eggs. Mean clutch size in each of the five-year periods ranged from 1.65 0.07 to 1.93 0.03. There was no significant interaction between region and five-year period when considering clutch size (244 = 2.94, P > 0.05). There was a significant difference in clutch size between the five-year periods (222 = 49.49, P < 0.001), but there was no significant linear trend in mean clutch size (r10 = 0.297, P > 0.05; Table 4) or differences between the population phases (26 = 5.71, P > 0.05). Clutch size did not vary significantly with region (24 = 4.25, P > 0.05). Brood size was known for 921 nests during the period 19412000. Eighteen per cent of nests contained one nestling, 82% contained two nestlings and less than 1% contained three young. Mean brood size in each of the five-year periods ranged from 1.50 0.11 to 2.00 0.18. There was no significant interaction between five-year period and region (244 = 12.83, P > 0.05). Mean brood size did not vary significantly between the five-year periods (222 = 16.91, P > 0.05), there was no significant linear trend in mean brood size (r10 = 0.512, P > 0.05) and no

Incubation

The daily survival rate of individual Turtle Dove nests during the incubation period over the five-year periods between 1951 and 2000 ranged from 0.948 to 0.972, and averaged 0.961 0.002. This meant that overall nest survival during the 14-day incubation period ranged from 0.479 to 0.674 (average 0.577 0.019); there was no evidence of a linear trend over time (r8 = 0.368, P > 0.05). The logistic regression analysis of nest success during the incubation period included the terms five-year period and region in the model. The interaction between five-year period and region was not significant (220 = 19.15, P > 0.05). After dropping the nonsignificant interaction term from the model, nesting success did not vary with five-year period (211 = 12.01, P > 0.05) or region (22 = 0.64, P > 0.05) or population phase (22 = 2.18, P > 0.05).

Table 5. Nest fate (%) for Turtle Dove nests during the period 19412000, in three geographical regions, as recorded on nest record cards (see Fig. 1 for regions). No. of cards Region Region 1 Region 2 Region 3 Overall Fate known 590 606 67 1263 Fate unknown 364 260 38 662 Lost other than predation 14.6 13.0 14.9 13.9 1.4 Predated 39.7 49.5 49.3 44.9 1.5 Nest fate (%) Successful 45.8 37.5 35.8 41.3 1.5

2005 British Trust for Ornithology, Bird Study,

52, 19

Temporal changes in Turtle Dove breeding ecology

Nestling period

Downloaded by [Ove Arup Partnership] at 04:26 16 November 2011

The daily survival rate of Turtle Dove nests during the five-year periods between 1951 and 2000 ranged from 0.977 to 0.988 and averaged 0.982 0.002. This corresponded to overall nest survival during the 15-day nestling period ranging from 0.707 to 0.853 (average 0.771 0.019). There was no evidence of a linear trend over time in overall nest survival during the nestling period (r8 = 0.352, P > 0.05). The interaction between five-year period and region was not significant (219 = 22.07, P > 0.05). After dropping the non-significant interaction term from the model, nest success did not vary with five-year period (211 = 11.36, P > 0.05) or region (22 = 0.32, P > 0.05) or population phase (22 = 0.41, P > 0.05).
Overall nesting period

Overall breeding success during the entire nesting period ranged from 0.389 to 0.514 (average 0.445 0.018). As expected from the lack of temporal trend during incubation or brood-rearing, there was no evidence of a linear trend over time in overall breeding success (r8 = 0.214, P > 0.05) or of a difference between population phases (22 = 2.99, P > 0.05).
DISCUSSION

The results presented here show that there have not been any detectable trends over time in any aspect of Turtle Dove nesting ecology, based on individual nesting attempts during the period 19412000 and recorded on nest record cards, but some differences between regions. The habitat and type of tree used for nesting varied between regions and this may reflect habitat availability. Scrub is the preferred nesting habitat for Turtle Doves (Mason & Macdonald 2000, Browne & Aebischer 2004) and because this type of habitat is relatively uncommon, especially when compared to the availability of hedgerows, it is likely to be used more frequently when Turtle Dove density is low, as in Regions 2 and 3. However in Region 1, where Turtle Dove density is highest, more birds are likely to overspill into hedgerows and a higher frequency of nests would be recorded within them. Nests in region 1 were more successful and were predated less than nests in the other regions. This may be due to the lower numbers of Magpies Pica pica, one of the Turtle Doves main nest predators, in region 1 compared to the other regions (Tapper 1992). There is therefore little evidence that the population

decline experienced by Turtle Doves breeding in Britain is due to the lowered success of individual attempts. If productivity has played a role in the decline, it must be through a reduction in the number of nesting attempts per pair during the breeding season. In an analysis of Nest Record Scheme data that considered Turtle Dove amongst other species, Siriwardena et al. (2001) concluded that although a number of measures of breeding performance fluctuated with population trends, the precise causes of population decline could not be identified. It was thought likely that post-fledging survival and the number of nesting attempts made within a season were important (Siriwardena et al. 2001). The latter view is supported by an intensive study of Turtle Dove breeding ecology undertaken by Browne & Aebischer (2004) which showed that the number of nesting attempts undertaken by each pair per breeding season was significantly lower in the late 1990s compared with the early 1960s (Murton 1968, Browne & Aebischer 2004). Additionally, Turtle Doves now have an earlier conclusion of the breeding season and undertake autumn migration earlier than in the 1960s (Browne & Aebischer 2003a). Other studies of the breeding ecology of declining farmland birds have also shown that it is the number of attempts undertaken per year, rather than the success of each individual attempt, that has reduced breeding performance and contributed to the decline of a range of species, including Corn Buntings Miliaria calandra (Brickle & Harper 2002), Skylarks Alauda arvensis (Donald et al. 2002), Barn Swallows Hirundo rustica (Mller 2001), Song Thrush Turdus philomelos (Thomson & Cotton 2000) and Linnet Carduelis cannabina (Moorcroft & Wilson 2000). The precise cause for the reduction in the number of nesting attempts undertaken by Turtle Doves is unknown, but it is likely that the underlying causes are reduced food availability affecting diet (Browne & Aebischer 2003b) and reduced nesting habitat availability (Browne & Aebischer 2004), brought about by agricultural intensification.
ACKNOWLEDGEMENTS We thank all Nest Record Scheme surveyors for their efforts over the last 60 years in collecting the data used here. The Nest Record Scheme is part of the BTOs Integrated Population Monitoring Programme carried out under partnership with the Joint Nature Conservation Committee (on behalf of English Nature, Scottish Natural Heritage, the

2005 British Trust for Ornithology, Bird Study,

52, 19

S.J. Browne, N.J. Aebischer and H.Q.P. Crick

Countryside Council for Wales and the Environment & Heritage Service in Northern Ireland). English Nature funded the analysis and preparation, and we thank Phil Grice of English Nature for his support and advice. Professor Chris Mason, Dr John OHalloran, Dr Rhys Green and an anonymous referee made useful comments.

ENDNOTES a. Region 1 (n = 954, 50%) comprised Cambridgeshire, Essex, Kent, Lincolnshire (North and Humberside), Norfolk, Suffolk, Sussex (East and West) and Yorkshire (East). Region 2 (n = 867, 45%) comprised Bedfordshire, Buckinghamshire, Derbyshire, Hampshire, Hertfordshire, Leicestershire, London (greater and inner), Northamptonshire, Nottinghamshire, Oxfordshire, Rutland, Surrey, Warwickshire and Yorkshire (South and West). Region 3 (n = 105, 5%) comprised Avon, Cheshire, the old county of Denbighshire, Devon, Dorset, the old county of Flintshire, Gloucestershire, Herefordshire, Lancashire, Merseyside, Shropshire, Somerset, Staffordshire, West Midlands, Wiltshire, and Worcestershire. b. Multinomial logistic regression models have multi-level dependent variables and are essentially a combination of binary logits estimated simultaneously (Agresti 1990). Hypotheses were tested using Wald statistics, which are distributed as chi-square.

REFERENCES
Aebischer, N.J. 1999. Multi-way comparisons and generalised linear models of nest success: extensions of the Mayfield method. Bird Study 46(suppl.): 2231. Agresti, A. 1990. Categorical Data Analysis. JohnWiley & Sons, Inc., New York. Anon. 1995. Biodiversity: The UK Steering Group Report, Vol. 2: Action Plans. HMSO, London. Baillie, S.R., Crick, H.Q.P., Balmer, D.E., Beaven, L.P., Downie, I.S., Freeman, S.N., Leech, D.I., Marchant, J.H., Noble, D.G., Raven, M.J., Simkin, A.P., Thewlis, R.M. & Wernham, C.V. 2002. Breeding Birds in the Wider Countryside: Their Conservation Status 2001. BTO Research Report No. 278. British Trust for Ornithology, Thetford. Brickle, N.W. & Harper, D.G.C. 2002. Agricultural intensification and the timing of breeding of Corn Buntings Miliaria calandra. Bird Study 49: 219228. Browne, S.J. & Aebischer, N.J. 2001. The Role of Agricultural Intensification in the Decline of the Turtle Dove Streptopelia turtur. English Nature Research Report No. 421. English Nature, Peterborough. Browne, S.J. & Aebischer, N.J. 2003a. Temporal changes in the migration phenology of Turtle Doves Streptopelia turtur in Britain, based on sightings from coastal bird observatories. J. Avian Biol. 34: 6571. Browne, S.J. & Aebischer, N.J. 2003b. Habitat use, foraging ecology and diet of Turtle Doves Streptopelia turtur in Britain. Ibis 145: 572582.

Browne, S.J. & Aebischer, N.J. 2003c. Temporal variation in the biometrics of Turtle Doves Streptopelia turtur caught in Britain between 1956 and 2000. Ringing Migr. 21: 203208. Browne, S.J. & Aebischer, N.J. 2004. Temporal changes in the breeding ecology of Turtle Doves Streptopelia turtur in Britain, and implications for conservation. Ibis 146: 125137. Calladine, J.R., Buner, F. & Aebischer, N.J. 1997. The Summer Ecology and Habitat Use of the Turtle Dove Streptopelia turtur: a Pilot Study. English Nature Research Report No. 219. English Nature, Peterborough. Crick, H.Q.P. 1992. A bird-habitat coding system for use in Britain and Ireland incorporating aspects of land management and human activity. Bird Study 39: 112. Crick, H.Q.P., Dudley, C., Evans, A.D. & Smith, K.W. 1994. Causes of nest failure among buntings in the UK. Bird Study 41: 8894. Crick, H.Q.P. & Leech, D.I. 2003. The UK Nest Record Scheme: its value for science and conservation. Bird Study 50: 254270. Donald, P.F., Evans, A.D., Muirhead, L.B., Buckingham, D.L., Kirby, W.B. & Schmitt, S.I.A. 2002. Survival rates, causes of failure and productivity of Skylark Alauda arvensis nests on lowland farmland. Ibis 144: 652664. Fuller, R.J., Gregory, R.D., Gibbons, D.W., Marchant, J.H., Wilson, J.D., Baillie, S.R. & Carter, N. 1995. Population declines and range contractions among lowland farmland birds in Britain. Conserv. Biol. 9: 14251441. Genstat Committee 2000. The Guide to Genstat. Lawes Agricultural Trust, Rothamsted. Gibbons, D.W., Reid, J.B. & Chapman, R.A. 1993. The New Atlas of Breeding Birds in Britain and Ireland: 19881991. T. & A.D. Poyser, London. Gregory, R.D. & Marchant, J.H. 1996. Population trends of Jays, Magpies, Jackdaws and Carrion Crows in the UK. Bird Study 43: 2837. Gregory, R.D., Wilkinson, N.I., Noble, D.G., Robinson, J.A., Brown, A.F., Hughes, J., Procter, D.A., Gibbons, D.W. & Galbraith, C.A. 2002. The population status of birds in the United Kingdom, Channel Islands and Isle of Man: an analysis of conservation concern 20022007. Br. Birds 95: 410450. Heath, M., Borggreve, C. & Peet, N. (eds) 2000. European Bird Populations: Estimates and Trends. Birdlife Conservation Series No. 10. Birdlife International, Cambridge. Hensler, G.L. 1985. Estimation and comparison of functions of daily nest survival probabilities using the Mayfield method. In Morgan, B.J.T. & North, P.M. (eds) Statistics in Ornithology: 289301. Springer-Verlag, Berlin. Hensler, G.L. & Nichols, J.D. 1981. The Mayfield method of estimating nesting success: a model, estimators and simulation results. Wilson Bull. 93: 4253. Macdonald, D.W. & Johnson, P.J. 2000. Farmers and the custody of the countryside: trends in loss and conservation of non-productive habitats 19811998. Biol. Conserv. 94: 221234. Marchant, J.H., Hudson, R., Carter, S.P. & Whittington, P. 1990. Population Trends in British Breeding Birds. British Trust for Ornithology, Tring. Mason, C.F. & Macdonald, S.M. 2000. Influence of landscape and land-use on the distribution of breeding birds in farmland in eastern England. J. Zool (Lond.) 251: 339348. Mayfield, H. 1961. Nesting success calculated from exposure. Wilson Bull. 73: 255261. Mayfield, H. 1975. Suggestions for calculating nest success. Wilson Bull. 87: 456466. Moorcroft, D. & Wilson, J.D. 2000. The ecology of Linnets

Downloaded by [Ove Arup Partnership] at 04:26 16 November 2011

2005 British Trust for Ornithology, Bird Study,

52, 19

Temporal changes in Turtle Dove breeding ecology

Downloaded by [Ove Arup Partnership] at 04:26 16 November 2011

Carduelis cannabina on lowland farmland. In Aebischer, N.J., Evans, A.D., Grice P.V. & Vickery, J.A. (eds) Ecology and Conservation of Lowland Farmland Birds: 173181. British Ornithologists Union, Tring. Mller, A.P. 2001. The effect of dairy farming on barn swallow Hirundo rustica abundance, distribution and reproduction. J Appl. Ecol. 38: 378389. Murton, R.K. 1968. Breeding, migration and survival of Turtle Doves. Br. Birds 61: 193212. OConnor, R.J. & Shrubb, M. 1986. Farming and Birds. Cambridge University Press, Cambridge. Peach, W.J., Siriwardena, G.M. & Gregory, R.D. 1999. Longterm changes in the abundance and demography of British reed buntings Emberiza schoeniclus. J. Appl. Ecol. 36: 798811. Siriwardena, G.M., Baillie, S.R., Buckland, S.T., Fewster, R.M., Marchant, J.H. & Wilson, J.D. 1998. Trends in the abundance of farmland birds: a quantitive comparision of smoothed CBC indices. J. Appl. Ecol. 35: 2443. Siriwardena, G.M., Baillie, S.R., Crick, H.Q.P. & Wilson, J.D.

2000. The importance of variation in the breeding performance of seed-eating birds in determining their population trends on farmland. J. Appl. Ecol. 37: 128148. Siriwardena, G.M., Baillie, S.R., Crick, H.Q.P. & Wilson, J.D. 2001. Changes in agricultural land-use and breeding performance of some granivorous farmland passerines in Britain. Agric. Ecosyst. Environ. 84: 191206. Tapper, S. 1992. Game Heritage: an Ecological Review from Shooting and Gamekeeping Records. Game Conservancy Ltd, Fordingbridge. Thomson, D.L. & Cotton, P.A. 2000. Understanding the decline of the British population of Song Thrushes Turdus philomelos. In Aebischer, N.J., Evans, A.D., Grice, P.V. & Vickery, J.A. (eds) Ecology and Conservation of Lowland Farmland Birds: 151155. British Ornithologists Union, Tring. Thomson, D.L., Baillie, S.R. & Peach, W.J. 1997. The demography and age-specific annual survival of British song thrushes Turdus philomelos during periods of population stability and decline. J. Anim. Ecol. 66: 414424.

(MS received 24 April 2003; revised MS accepted 16 January 2004)

2005 British Trust for Ornithology, Bird Study,

52, 19