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This thesis has been submitted in fulfilment of the requirements for a postgraduate degree (e.g. PhD, MPhil, DClinPsychol) at the University of Edinburgh. Please note the following terms and conditions of use: • This work is protected by copyright and other intellectual property rights, which are retained by the thesis author, unless otherwise stated. • A copy can be downloaded for personal non-commercial research or study, without prior permission or charge. • This thesis cannot be reproduced or quoted extensively from without first obtaining permission in writing from the author. • The content must not be changed in any way or sold commercially in any format or medium without the formal permission of the author. • When referring to this work, full bibliographic details including the author, title, awarding institution and date of the thesis must be given. Causes of decline and conservation solutions for Corn Buntings Emberiza calandra in eastern Scotland Allan J. Perkins PhD The University of Edinburgh 2012 i DECLARATION I hereby declare that I alone composed this thesis, that the work is my own, except for contributions from colleagues clearly stated at the start of each chapter, and that this work has not been submitted for any other degree or professional qualification. Date: Allan J. Perkins ii ACKNOWLEDGEMENTS I start by thanking my employers, the Royal Society for the Protection of Birds (RSPB), for funding this PhD, and Jeremy Wilson for proposing the project. As my supervisor, Jeremy, along with John Deag (University of Edinburgh), provided encouragement, support and constructive criticism thoughout. I also thank my colleagues and friends Hywel Maggs and Adam Watson, without whom this project would not have been possible. Adam gave complete access to his 20+ year dataset on Corn Buntings, and has been tremendously helpful and supportive. Hywel ran the RSPB’s Farmland Bird Lifeline (FBL) project, and we shared equally almost all aspects of fieldwork. Field trips with Adam and Hywel to look for and discuss Corn Buntings are always a pleasure, and will forever be remembered. My thanks also go to the RSPB field assistants – Amanda Biggins, Alan Bull, Steven Coyne, Richard Firmin, Clive McKay and John McMahon – for collecting much of the data that form the basis of this thesis, and other RSPB staff who have helped out with fieldwork – Karen Cunningham, Andy Wight, Stuart Benn and Vicky Anderson. Volunteer Ken Bruce also generously gave up his time to help with fieldwork, whilst Chris Smout provided data from his annual Corn Bunting surveys in Fife. Thanks to the RSPB team at Aberdeen for their support and friendship over the years, and to various colleagues for comments on manuscripts or general advice and discussion – Martin Auld, Ian Francis, Graham Rebecca, Mark Hancock, James Pearce-Higgins, Steven Ewing, David Douglas, Guy Anderson, Richard Bradbury, Paul Donald, David Gibbons and others. Several referees also provided constructive comments to improve manuscripts, including Nick Brickle, Rhys Green, David Harper, Dan Chamberlain, Stephen Browne and Mark Whittingham, and those of examiners Ian Hartley and Aubrey Manning helped to improve the final version of the thesis. Thanks to the many farmers and landowners for their involvement in the project, allowing access and testing management solutions for Corn Buntings on their farms. Thankyou to Scottish Natural Heritage (in particular Ewan Lawrie) and the Biodiversity Action Grant Scheme (Scottish Government) for their financial support of FBL, and Steff Ferguson, Nicky Penford and Flora Grigor-Taylor for drawing up many of the FBL management plans. Thanks to my family for their support, even though I’m not entirely sure they knew what this was all about! Finally, a special thankyou to Amanda for her tolerance and patience with me, especially during these final few months of writing up, and of course for her fieldwork contributions on those “poxy sparras” as she so affectionately calls them! iii ABSTRACT The Corn Bunting Emberiza calandra is one of the most severely declining farmland birds across Europe. In the UK, numbers fell by 86% between 1967 and 2008. Corn Buntings favour open landscapes, nest on or close to the ground, are often polygynous, double- brooded, and have a seed-based diet supplemented in summer by invertebrates. This study investigated the recent causes of decline in arable and mixed farmland in eastern Scotland, and sought to identify potential conservation solutions that could be delivered through agri- environment schemes (AES). Combining new data with analyses of existing long-term datasets, I investigated habitat associations during summer and winter, the timing and success of nesting attempts, and measured reproductive and population responses to AES. Corn Buntings declined almost to extinction in one study area where, over 20 years, the main recorded intensifications of farming were reduced weed abundance within crops and removal of boundaries to make bigger fields. Territory locations, late-summer occupancy and polygyny were all strongly associated with weedy fields. There were also positive associations with overhead wires and in early summer with winter barley and forage grasses. Late-summer occupancy was associated with spring-sown cereals, crops that are amongst the last to be harvested. Changes in habitat associations and to aspects of the mating system as the population declined and agriculture intensified are discussed. Intensive monitoring showed that Corn Buntings laid clutches from mid-May to mid-August, mostly in fields of forage grasses and autumn-sown cereals in early summer, and spring- sown cereals in late summer. A preference for nesting in dense swards explained this seasonal variation. Breeding success in forage grasses was poor, due to high rates of nest loss during mowing. However, in experimental trials, nest success in fields with delayed mowing was fivefold that of control fields. With sufficient uptake through AES, delayed mowing could raise productivity to levels required to reverse population declines. In winter, cereal stubbles and AES unharvested crop patches were the main foraging habitats used. Unharvested crops with abundant cereal grain in their first winter of establishment were favoured. Population monitoring over seven years and 71 farms revealed increases on farms with AES targeted at Corn Buntings, no significant change on farms with general AES, and declines on control farms. In arable-dominated farmland, management that increased food availability reversed declines, but on mixed farmland where Corn Buntings nested in forage grasses, delayed mowing was essential for population increase. This study has already influenced the design of AES targeted at Corn Buntings in Scotland, and I make further recommendations for the species’ conservation and design of AES that are applicable to farmland throughout Britain and Europe. iv CONTENTS Declaration ii Acknowledgements iii Abstract iv Contents v List of tables xi List of figures xii Photographs xiv 1 Introduction – changes in agriculture & farmland bird population declines 1 1.1 Agricultural Intensification 1 1.1.1 Changes to farming in the UK 2 1.1.2 Changes to farming in other parts of Europe 5 1.1.3 The cost to biodiversity 7 1.2 Changes in farmland bird populations 7 1.2.1 UK trends 7 1.2.2 European trends 9 1.3 Ecological and life-history traits of common European farmland birds 14 1.4 Agricultural changes driving population trends of farmland birds 21 1.4.1 Arable management intensity 21 1.4.2 Grassland management intensity 23 1.4.3 Crop diversity 24 1.4.4 Area of fallow or semi-natural habitat 25 1.4.5 Field wetness 33 1.4.6 Area of scrub/woodland 34 1.4.7 Field size 35 1.4.8 New crops 36 1.4.9 Summary 36 1.5 Measures to restore farmland biodiversity – agri-environment schemes 37 1.5.1 A brief history of UK agri-environment schemes 37 1.5.2 Effectiveness of agri-environment schemes 39 1.6 Corn Buntings in eastern Scotland – a case study 40 v 2 Study aims, species and area – Corn Buntings in eastern Scotland 41 2.1 Background 41 2.2 Aims of project 41 2.3 Outline of chapters 41 2.4 The study species – Corn Bunting 44 2.4.1 Size, plumage and song 44 2.4.2 Distribution, habitats and movements 44 2.4.3 Diet, foraging, breeding and habitat associations 45 2.4.4 Some unusual traits 46 2.5 Corn Bunting population trends 47 2.5.1 Europe 47 2.5.2 UK 47 2.5.3 Eastern Scotland 48 2.5.3.1 Data analysis 49 2.5.3.2 Results 49 2.5.3.3 Discussion 51 3 Habitat associations with territory distribution and mating system 59 3.1 Introduction 59 3.2 Methods 61 3.2.1 Study area 61 3.2.2 Field methods 61 3.2.3 Characterising Corn Bunting territories 62 3.2.4 Statistical modelling 63 3.2.5 Predictors and sets of models 64 3.3 Results 65 3.3.1 Trends in Corn Buntings and habitats 65 3.3.2 Habitat associations of territory locations, late-summer occupancy & polygyny 66 3.3.2.1 Preliminary model selection 66 3.3.2.2 Habitat associations and year effects – territory location 66 3.3.2.3 Habitat associations and year effects – late-summer occupancy 67 3.3.2.4 Habitat associations and year effects – polygyny 67 3.3.3 Comparison between 20-year and 3-year analyses 68 3.4 Discussion 68 3.4.1 Habitat associations of territory locations 69 3.4.2 Seasonal effects 70 vi 3.4.3 Polygyny 71 3.4.4 Temporal trends in habitat associations 72 3.4.5 Conservation implications 72 3.4.6 Value of long-term studies 73 4 Habitat selection by females for nesting 93 4.1 Introduction 93 4.2 Methods 94