Atlas of the Mammals of Great Britain and Northern Ireland 1784272043, 9781784272043

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ATLAS OF THE

MAMMALS OF GREAT BRITAIN AND NORTHERN IRELAND

ATLAS OF THE

MAMMALS OF GREAT BRITAIN AND NORTHERN IRELAND

DEREK CRAWLEY, FRAZER COOMBER, LAURA KUBASIEWICZ, COLIN HARROWER, PETER EVANS, JAMES WAGGITT, BETHANY SMITH & FIONA MATHEWS

PELAGIC PUBLISHING

Compiled and edited by: Derek Crawley1*, Frazer Coomber1,2*, Laura Kubasiewicz1, Colin Harrower3, Peter Evans4,5, James Waggitt4,5, Bethany Smith1 and Fiona Mathews1,2. * Made equal contributions 1. 2. 3. 4. 5.

The Mammal Society, 8 St John’s Church Road, London E9 6EJ. The University of Sussex, John Maynard Smith Building, Falmer BN1 9QG. NERC Centre for Ecology and Hydrology, Crowmarsh Gifford, Wallingford OX19 8BB. Sea Watch Foundation, Ewyn y Don, Bull Bay, Amlwch, Anglesey LL68 9SD. School of Ocean Sciences, Bangor University, Menai Bridge, Anglesey LL59 5AB.

Species accounts written by: Diana Bell, Sam Berry, Johnny Birks, Jenny Bryce, Simone Bullion, Ruairidh Campbell, Roisin Campbell-Palmer, Anna Champneys, Paul Chanin, Arnold Cooke, Frazer Coomber, Derek Crawley, Elizabeth Croose, Mike Dean, John Dutton, Peter Evans, John Flowerdew, Anita Glover, Martyn Gorman, John Gurnell, John Haddow, Daniel Hargreaves, Jochen Langbein, Penny Lewns, Fiona Mathews, Robbie McDonald, Tony Mitchell-Jones, Marion O’Neil, Josephine Pemberton, Rory Putman, Nigel Reeve, Hugh Rose, Henry Schofield, Dawn Scott, Jo Sharplin, Craig Shuttleworth, Peter Smith, Charles Smith-Jones, David Thompson, Roger Trout, Alastair Ward, Daniel Whitby and Ian White.

Published by Pelagic Publishing PO Box 874 Exeter EX3 9BR UK www.pelagicpublishing.com

Atlas of the Mammals of Great Britain and Northern Ireland ISBN 978-1-78427-204-3 ISBN 978-1-78427-205-0 ISBN 978-1-78427-206-7

Hardback ePub PDF

© The Mammal Society 2020 The moral rights of the authors have been asserted. All rights reserved. Apart from short excerpts for use in research or for reviews, no part of this document may be printed or reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, recording, now known or hereafter invented or otherwise without prior permission from the publisher. A CIP record for this book is available from the British Library Cover photograph: Dan Rushton

CONTENTS

FOREWORDviii ACKNOWLEDGEMENTSx INTRODUCTION1 METHODS3 Data collection 3 Distribution maps 5 Record coverage across the UK 5 SPECIES ACCOUNTS13 Hedgehog (Erinaceus europaeus; Linnaeus, 1758) 14 European mole (Talpa europaea; Linnaeus, 1758) 16 Common shrew (Sorex araneus; Linnaeus, 1758) 18 Pygmy shrew (Sorex minutus; Linnaeus, 1766) 20 Water shrew (Neomys fodiens; Pennant, 1771) 22 Lesser white-toothed shrew (Crocidura suaveolens; Pallas, 1811) 24 European rabbit (Oryctolagus cuniculus; Linnaeus, 1758) 26 Brown hare (Lepus europaeus; Pallas, 1778) 28 Mountain hare (Lepus timidus; Linnaeus, 1758) 30 Red squirrel (Sciurus vulgaris; Linnaeus, 1758) 32 Grey squirrel (Sciurus carolinensis; Gmelin, 1788) 34 Eurasian beaver (Castor fiber; Linnaeus, 1758) 36 Hazel dormouse (Muscardinus avellanarius; Linnaeus, 1758) 38 Edible dormouse (Glis glis; Brisson, 1762) 40 Bank vole (Myodes glareolus; Schreber, 1780) 42 Field vole (Microtus agrestis; Linnaeus, 1761) 44 Common vole (Microtus arvalis; Pallas, 1778) 46 Water vole (Arvicola amphibius; Linnaeus, 1758) 48 Harvest mouse (Micromys minutus; Pallas, 1771) 50 Wood mouse (Apodemus sylvaticus; Linnaeus, 1758) 52 Yellow-necked mouse (Apodemus flavicollis; Melchior, 1834) 54 House mouse (Mus musculus; Linnaeus, 1758) 56

vi  Atlas of the Mammals of Great Britain and Northern Ireland Brown rat (Rattus norvegicus; Berkenhout, 1769) Black rat (Rattus rattus; Linnaeus, 1758) Wildcat (Felis silvestris; Miller, 1907) Red fox (Vulpes vulpes; Linnaeus, 1758) Badger (Meles meles; Linnaeus, 1758) Otter (Lutra lutra; Linnaeus, 1758) Pine marten (Martes martes; Linnaeus, 1758) Stoat (Mustela erminea; Linnaeus, 1758) Weasel (Mustela nivalis; Linnaeus, 1758) Polecat (Mustela putorius; Linnaeus, 1758) American mink (Neovison vison; Schreber, 1777) Wild boar (Sus scrofa; Linnaeus, 1758) Red deer (Cervus elaphus; Linnaeus, 1758) Sika deer (Cervus nippon; Temminck, 1838) Fallow deer (Dama dama; Linnaeus, 1758) Roe deer (Capreolus capreolus; Linnaeus, 1758) Chinese water deer (Hydropotes inermis; Swinhoe, 1870) Reeves’ muntjac deer (Muntiacus reevesi; Ogilby, 1839) Greater horseshoe bat (Rhinolophus ferrumequinum; Schreber, 1774) Lesser horseshoe bat (Rhinolophus hipposideros; Bechstein, 1800) Alcathoe bat (Myotis alcathoe; von Helversen & Heller, 2001) Whiskered bat (Myotis mystacinus; Kuhl, 1817) Brandt’s bat (Myotis brandtii; Eversmann, 1845) Bechstein’s bat (Myotis bechsteinii; Kuhl, 1817) Daubenton’s bat (Myotis daubentonii; Kuhl, 1817) Greater mouse-eared bat (Myotis myotis; Borkhausen, 1797) Natterer’s bat (Myotis nattereri; Kuhl, 1817) Serotine bat (Eptesicus serotinus; Schreber, 1774) Leisler’s bat (Nyctalus leisleri; Kuhl, 1817) Noctule bat (Nyctalus noctula; Schreber, 1774) Common pipistrelle bat (Pipistrellus pipistrellus; Schreber, 1774) Soprano pipistrelle bat (Pipistrellus pygmaeus; Leach, 1825) Nathusius’ pipistrelle bat (Pipistrellus nathusii; Keyserling & Blasius, 1839) Barbastelle bat (Barbastella barbastellus; Schreber, 1774) Brown long-eared bat (Plecotus auritus; Linnaeus, 1758) Grey long-eared bat (Plecotus austriacus; Fischer, 1829) Grey seal (Halichoerus grypus; Fabricius, 1791) Harbour seal (Phoca vitulina; Linnaeus, 1758) North Atlantic right whale (Eubalaena glacialis; Müller, 1776) Bowhead whale (Balaena mysticetus; Linnaeus, 1758) Humpback whale (Megaptera novaeangliae; Borowski, 1781) Blue whale (Balaenoptera musculus; Linnaeus, 1758) Fin whale (Balaenoptera physalus; Linnaeus, 1758) Sei whale (Balaenoptera borealis; Lesson, 1828) Minke whale (Balaenoptera acutorostrata; Lacepede, 1804)

58 60 62 64 66 68 70 72 74 76 78 80 82 84 86 88 90 92 94 96 98 100 102 104 106 108 110 112 114 116 118 120 122 124 126 128 130 132 134 136 138 140 142 144 146

Contents vii Northern bottlenose whale (Hyperoodon ampullatus; Forster, 1770) Cuvier’s beaked whale (Ziphius cavirostris; Cuvier, 1823) Sowerby’s beaked whale (Mesoplodon bidens; Sowerby, 1804) Pygmy sperm whale (Kogia breviceps; Blainville, 1838) Dwarf sperm whale (Kogia sima; Owen, 1866) Sperm whale (Physeter macrocephalus; Linnaeus, 1758) Beluga (Delphinapterus leucas; Pallas, 1776) Killer whale or Orca (Orcinus orca; Linnaeus, 1758) False killer whale (Pseudorca crassidens; Owen, 1846) Long-finned pilot whale (Globicephala melas; Traill, 1809) Risso’s dolphin (Grampus griseus; Cuvier, 1812) Atlantic white-sided dolphin (Lagenorhynchus acutus; Gray, 1828) White-beaked dolphin (Lagenorhynchus albirostris; Gray, 1846) Common dolphin (Delphinus delphis; Linnaeus, 1758) Striped dolphin (Stenella coeruleoalba; Meyen, 1833) Bottlenose dolphin (Tursiops truncatus; Montagu, 1821) Harbour porpoise (Phocoena phocoena; Linnaeus, 1758)

148 150 152 154 156 158 160 162 164 166 168 170 172 174 176 178 180

CETACEANS KNOWN IN BRITAIN AND IRELAND ONLY FROM STRANDINGS  182 Blainville’s beaked whale (Mesoplodon densirostris; Blainville, 1817) 182 Gervais’ beaked whale (Mesoplodon europaeus; Gervais, 1855) 182 True’s beaked whale (Mesoplodon mirus; True, 1913) 183 Narwhal (Monodon monoceros; Linnaeus, 1758) 183 Melon-headed whale (Peponocephala electra; Gray, 1846) 184 Fraser’s dolphin (Lagenodelphis hosei; Fraser, 1956) 184 VAGRANT SPECIES AND THOSE WITHOUT ESTABLISHED POPULATIONS IN THE UK  185 Raccoon (Procyon lotor; Linnaeus, 1758) 185 Red-necked wallaby (Macropus rufogriseus; Desmarest, 1817) 185 Reindeer (Rangifer tarandus; Linnaeus, 1758) 186 Pond bat (Myotis dasycneme; Boie, 1825) 186 Geoffroy’s bat (Myotis emarginatus; Geoffroy, 1806) 186 Northern bat (Eptesicus nilssonii; Keyserling and Blasius, 1839) 187 Parti-coloured bat (Vespertilio murinus; Linnaeus, 1758) 187 Kuhl’s pipistrelle bat (Pipistrellus kuhlii; Kuhl, 1817) 188 Savi’s pipistrelle bat (Hypsugo savii; Bonaparte, 1837) 188 Bearded seal (Erignathus barbatus; Erxleben, 1777) 189 Harp seal (Pagophilus groenlandicus; Erxleben, 1777) 189 Hooded seal (Cystophora cristata; Erxleben, 1777) 189 Ringed seal (Pusa hispida; Schreber, 1775) 190 Walrus (Odobenus rosmarus; Linnaeus, 1758) 190 FERAL COLONIES AND POPULATIONS191

FOREWORD

While the maps in this Atlas are fascinating in their own right, they are also an important resource for the conservation and management of mammal populations. Fulfilling the Society’s role in advocating science-led mammal conservation, they underlie the generalised distribution maps provided in the Guide to the Population and Conservation Status of Britain’s Mammals (Mathews et al., 2018). In this Atlas, information on reported occupancy is shown in higher spatial resolution, and the maps also document changes in recorded distribution since 1960. They also provide an essential reference for ecological consultants wishing to know which species of protected mammals could potentially be affected by a development in a particular area. This is the latest in a series of publications based on the systematic collection of distribution records by the Mammal Society and its members extending back to the 1960s. Provisional Atlases were published in 1971, 1978 and 1984, while the first Atlas of Mammals in Britain, compiled and edited by Henry Arnold, was published by the Institute of Terrestrial Ecology and the Joint Nature Conservation Committee in 1993. In addition to Henry, who also edited the second and third provisional Atlases, and to Gordon Corbet, editor of the first of these, a small army of people must be thanked for their contributions. This includes you, the reader; if you have ever submitted a mammal record, whether directly, through one of our partners, or via the Society’s apps – Mammal Tracker or Mammal Mapper – I would like to thank you on behalf of the Society. Over the past five years a small group of people have co-ordinated collectors of records, cajoled contributors and tried to coerce computers into providing suitable maps for this Atlas. Derek Crawley deserves a special mention and my undying gratitude for taking on the co-ordinating role, following the untimely death of Derek Yalden, my predecessor as President. Fiona Mathews, Laura Kubasiewicz, Frazer Coomber, Peter Evans, James Waggitt, Colin Harrower, Bethany Smith, Richard Shore, David Roy and Martin Harvey worked with Derek to turn a collection of distribution records into an informative atlas for which we are very grateful. The Society is also deeply indebted to the mammal groups, mammal recorders, record verifiers and authors of the species accounts who all played their part in creating this volume. There is one person, however, whose contribution overshadows all of these: Derek Yalden was an inspirational mammalogist whose boundless enthusiasm and extensive know­ledge infected generations of naturalists with the mammal-recording bug. Derek’s unexpected death in 2013 was a great blow to the Society. In an obituary for Derek (Mammal Notes, no. 166, 2013) Pat Morris recalls that he and Derek, who

Foreword ix met while still at school, ‘took up the challenge to fill in blank squares on [the] distribution maps’ produced for the London Natural History Society’s mammal recorder. This habit persisted throughout Derek’s life and was particularly, but by no means exclusively, focused on the Peak District where he studied the distribution and ecology of mountain hares, red-necked wallabies and even birds such as the golden plover and common sandpiper. Derek was a member of the Society for 50 years and filled many significant roles. He was editor of Mammal Review for 32 years; wrote or contributed many technical publications, including all four editions of the Handbook of British Mammals (co-editing the last of these, an 800-page blockbuster published by the Society in 2008); and was President for 16 years, a period equalled only by the first Society President, the fourth Earl of Cranbrook, John Gathorne-Hardy. During his presidency, Derek proposed that the Society should produce a Mammal Atlas for the twenty-first century and was the driving force behind its development. This volume is a testament to his vision, his enthusiasm and his great talent for involving other people in studying mammals. We owe him a great deal and we still miss him. Paul Chanin, June 2018

ACKNOWLEDGEMENTS

We would like to thank the many volunteer surveyors and ecological consultants who contributed the data used to produce the Atlas distribution maps. We are also grateful for the support of local Biological Records Centres and other organisations in sharing their datasets: our task would have been much harder without the tremendous contribution of county recorders and verifiers. Tom Hunt at the Association of Local Records Centres provided valuable assistance in ensuring the smooth transition of data, and we are also grateful to the National Biodiversity Network Trust. Specific records were supplied by Baseline Ecology, the Bat Conservation Trust, Bedfordshire and Luton Biodiversity Recording and Monitoring Centre, Biodiversity Gatwick Project, Biodiversity Information Service for Powys and Brecon Beacons National Park, Biological Records Centre, Boat of Garten Wildlife Group, Bristol Regional Environmental Records Centre, British Deer Society, British Trust for Ornithology, Buckinghamshire and Milton Keynes Environmental Records Centre, Cambridgeshire & Peterborough Environmental Records Centre, Canals and Rivers Trust, Central Scotland Green Network Trust, Cofnod (North Wales Environmental Information Service), Cumbria Biodiversity Data Centre, Derbyshire Biological Records Centre, Derbyshire Mammal Atlas, Derbyshire Mammal Group, Devon Biodiversity Records Centre, Doncaster MBC Biological Records Centre, Dorset Environmental Records Centre, Dorset Mammal Group, Dr Francis Rose Field Notebook Project, Dumfries and Galloway Environmental Resources Centre, East Ayrshire Countryside Ranger Service, EcoRecord, Environment Agency, Environmental Records Centre for Cornwall and the Isles of Scilly, Environmental Records Information Centre North East, Essex Wildlife Trust, Fife Nature Records Centre, Focus Ecology Ltd, Furesfen Ecological Consultancy, Glasgow Museums BRC, Gloucestershire Centre for Environmental Records, Greater Lincolnshire Nature Partnership, Greater Manchester Ecology Unit, Greenspace Information for Greater London (GiGL), Hampshire Biodiversity Information Centre, Herefordshire Biological Records Centre, Hertfordshire Natural History Society, Herts Environmental Records Centre, Highland Biological Recording Group, Humber Environmental Data Centre, Inner Forth Nature Counts, iRecord, Isle of Wight Local Records Centre, iSpot, IW Council Parks and Countryside Section, John Muir Trust, Kent & Medway Biological Records Centre, Kent Biological Records Centre, Lancashire Environment Record Network, Leicestershire and Rutland Environmental Records Centre, Lorn Natural History Group, Lothian and Borders Mammal Group, Lymington Naturalists, Mammals of Suffolk, MaWSE project, Merseyside BioBank, Ministry of Justice, MKA Ecology Ltd, National Trust, National Trust for Scotland, Natural Resources Wales, Nonsuch Watch, Norfolk

Acknowledgements xi Biodiversity Information Service, North & East Yorkshire Ecological Data Centre, North Ayrshire Countryside Ranger Service, North East Scotland Biological Records Centre, Northamptonshire Biodiversity Records Centre, Nottinghamshire Mammal Database, Outer Hebrides Biological Recording Project, People’s Trust for Endangered Species, Phlorum Consultancy, PJC Consultancy, Powys and BBNP Biodiversity Information Service, Preston Montford Field Studies Council Centre, Riverbank Wildlife Area, Rotherham Biological Records Centre, Royal Horticultural Society, RPS, RSPCA, SCC Open Space, Scottish Natural Heritage, Scottish Wildlife Trust, Sheffield Biological Records Centre, Shetland Biological Records Centre, Shire Group of Internal Drainage Boards, Shropshire Ecological Data Network, Somerset Environmental Records Centre, Sorby Mammal Group, Sorby Natural History Society, South East Wales Biodiversity Records Centre, St Helens Wildlife Recording Group, Staffordshire Ecological Record, Suffolk Biological Records Centre, Surrey Biodiversity Information Centre, Surrey Biological Records Centre, Surrey Dormouse Group, Surrey Mammal Group, Sussex Biodiversity Record Centre, Sustrans, Tawny Croft Wildlife Consultants, Thames Valley Environmental Records Centre, The Biodiversity Information System for Cheshire, Halton, Warrington and the Wirral, The Ecology Consultancy, The Magnificent Science Company Limited, Unsted Wildlife Monitoring, Vincent Wildlife Trust, Warwickshire Biological Records Centre, West Wales Biodiversity Information Centre, West Yorkshire Ecology Service, Wildlife Information Centre, WildWatch Project Records, Wiltshire and Swindon Biological Records Centre, Worcestershire Biological Records Centre, Yorkshire Naturalists’ Union, Yorkshire Wildlife Trust, Declan Barraclough, Johnny Birks, A. Blunden, Ian Bond, I. Boyd, Helen Butler, Paul Chanin, Robert and Val Clinging, Derek Crawley, John Dobson, John Durkin, Dave and Joyce Earl, Peter Follett, Liz Halliwell, Annie Haycock, Zoe Haysted, Harriet James, David Jardine, Jenny Jones, Rosy Jones, G. Knass, Robert Lamb, Penny Lewns, Steve Lonsdale, Anne Marston, Chris Matcham, Brenda Mayles, Eve Mulholland, Ed Pooley, C.R. Pope, Brian Ribbands, Phil Richardson, Kirstie and Calum Ross, Graham Scholey, Paul Seligman, Rob Spencer, Stephanie Tyler, Tracy Underwood, Michael Walker, Debbie Wallace, Ken Walton and Antony Witts. The data sources for cetacean records are shown in Table 1. These records were collated as part of the NERC/DEFRA funded Marine Ecosystems Research Programme (NE/L003279/1). We would especially like to thank: Joana Andrade, Mick Baines, Oliver Boisseau, Gareth Bradbury, Tom Brereton, Kees Camphuysen, Jose Martinez-Cedeira, Carol Cooper, Jan Durinck, Tom Felce, Isabel Garcia-Baron, Stefan Garthe, Steve Geelhoed, Anita Gilles, Martin Goodall, John Goold, Jan Haelters, Sally Hamilton, Phil Hammond, Lauren Hartny-Mills, Nicola Hodgins, John Houghton, Kathy James, Mark Jessopp, Nia Jones, Ailbhe Kavanagh, Mardik Leopold, Katrin Lohrengel, Maite Louzao, Colin MacLeod, Oliver O’Cadhla, Sarah Perry, Graham Pierce, Vincent Ridoux, Kevin Robinson, Camilo Saavedra, Begoña Santos, Richard Shucksmith, Henrik Skøv, Eric Stienen, Signe Sveegaard, Paul Thompson, Nicolas Vanermen, Dave Wall, Andy Webb and Jared Wilson. We would also like to thank all the mammal record verifiers who have helped to ensure that the data are of the highest quality: Lorcan Adrain, Debbie Alston, Simone Bullion, Helen Butler, Rebecca Collins, Nathalie Cossa, Derek Crawley, John Ellis, Dan Forman, Claudia Gebhardt, Penny Green, Richard Grogan, Gareth Harris, Annie Haycock, Gary Hedges, Mark Hows, Barry Ingram, Jenny Jones, Jim Jones, Dave Kilbey, Ellie Knott, Emma Koblizek, Richard Lawrence, Ellen Lee, Sorcha Lewis, Declan Looney, John Mackintosh, Chris Manning, John Martin, Jacqueline

xii  Atlas of the Mammals of Great Britain and Northern Ireland Merrick, Tony Mitchell-Jones, Phillip Morgan, Helen O’Brien, Gareth Parry, Jools Partridge, Katherine Pinnock, Jonathan Pounder, John Ray, Brian Ribbands, Andy Riches, Angela Ross, Andy Rothwell, David Roy, Ro Scott, Graham Smith, Tamasine Stretton, Sarah Underwood, Lisa Wade, Michael Walker, Richard Webb, David Williams, Graeme Wilson and John Young. Liam Lysaght has been helpful with ensuring co-ordination with the Atlas of Mammals of Ireland, and Johnny Birks, as then chair of the Mammal Society, was integral to setting up the current Atlas project. Thanks are owed to the mammal experts who have written the species accounts, and their names are presented with each individual account. In addition, we would like to thank all the photographers who have submitted photographs to the Mammal Society, including entrants to our amateur Mammal Photographer of the Year competition. The editors would like to apologise for any oversight in our thanks. A huge number of people have been involved with this project, submitting, collecting, preparing and verifying records; writing text; and by giving general support. Thank you all.

INTRODUCTION

The aim of the Atlas of Mammals of Great Britain and Northern Ireland is to increase understanding of the distribution of mammals, and to document changes over time since the last Atlas (1960–92; Arnold, 1993). The current project began as the Mammal Watch South East (MaWSE). Part-funded by the Heritage Lottery Fund (HLF), MaWSE created the Mammal Tracker app, which provided an easy interface for the public to submit records from across the UK. This local project then expanded to encompass the whole of the UK, the Channel Islands and the Isle of Man, and, with the assistance of the Sea Watch Foundation, was extended to include cetaceans. The species accounts are ordered as terrestrial mammals (including seals), cetaceans, cetacean species known only from strandings, vagrant species and feral populations. The terrestrial mammals are arranged in the same order as the Review of the Population and Conservation Status of British Mammals (Mathews et al., 2018) followed by the seal species, which were not included in that review. The cetacean species are organised according to taxonomic similarities. The common names used in this Atlas are those used in the Review of the Population and Conservation Status of British Mammals, or, for seals and cetaceans, those in the Mammal Society’s online species hub. The Atlas, unlike the Review, includes the Channel Islands. The Orkney and Guernsey vole are therefore presented in a single species report as they are both the common vole Microtus arvalis. Island subspecies, such as the Skomer vole (a subspecies of the bank vole Myodes glareolus) and the Irish hare (a subspecies of the mountain hare Lepus timidus), are not presented individually. Each species account, with the exception of the vagrant species, contains information on the species distribution, ecology and identification. This is accompanied by a distribution map, an illustration of the seasonal distribution of occurrence records, and a photograph intended to aid identification. For further information on the status of terrestrial mammals in Britain, readers are referred to the Review of the Population and Conservation Status of British Mammals and the Regional Red List for mammals that has been compiled by the Society for the Statutory Nature Conservation Organisations according to International Union for the Conservation of Nature (IUCN) standards. For cetaceans, the species accounts include sections on distribution, ecology and identification. These have been adapted for this Atlas from larger accounts written for the book European Whales, Dolphins and Porpoises to commemorate the 25th anniversary of the United Nations Environment Programme’s regional agreement, ASCOBANS (Evans, 2019). The photographs were sourced from the Mammal Society’s catalogue of photographs, species experts and various appeals. Many were submitted by members

2  Atlas of the Mammals of Great Britain and Northern Ireland of the public as part of the Mammal Society’s Mammal Photographer of the Year competition. Wherever possible, the Atlas uses photographs taken of wild animals within the UK and its Crown dependencies. Selection of photographs was based on the display of key identification features rather than aesthetic appeal. The Mammal Society is extremely grateful to all the photographers, and their names appear adjacent to the relevant image.

BIBLIOGRAPHY ARNOLD, H.R. 1993. Atlas of Mammals in Britain. ITE Research Publication No. 6. London: Joint Nature Conservation Committee and Institute of Terrestrial Ecology, HMSO. EVANS, P.G.H. 2019. European Whales, Dolphins and Porpoises. Marine Mammal Conservation in Practice. London and New York: Elsevier/Academic Press. MAMMAL SOCIETY. 2018. Full species list. Accessed at http://www.mammal.org.uk/ species-hub/uk-mammal-list/ (30 July 2019). MATHEWS, F., KUBASIEWICZ, L.M., GURNELL, J., HARROWER, C.A., McDONALD, R.A. & SHORE, R.F. 2018. A Review of the Population and Conservation Status of British Mammals. A Report by the Mammal Society Under Contract to Natural England, Natural Resources Wales and Scottish Natural Heritage. Peterborough: Natural England.

METHODS

DATA COLLECTION British mammal occurrence data at 10 km resolution or higher were gathered from the following sources: National Biodiversity Network (NBN) gateway (NBN Atlas), local biological record centres, local and national monitoring schemes, iRecord, Natural England, the Mammal Society, Sea Watch Foundation, the Mammal Tracker app, and records provided by individuals. A full list of data providers is supplied in the acknowledgements. Validation and verification was conducted by the Mammal Society’s appointed verifiers, using protocols drawn up by Derek Crawley with help from the Biological Records Centre (BRC). Unverified mammal records derived from organisations that did not have mammal surveying as their primary objective were only used for those species that were unlikely to be misidentified, namely moles Talpa europaea, rabbits Oryctolagus cuniculus, badgers Meles meles, foxes Vulpes vulpes and hedgehogs Erinaceus europaeus. Most of these unverified records were supplied by the British Trust for Ornithology. Thirty-six species of cetaceans have been recorded in European seas, 30 of which have been recorded at one time or another in the waters around Britain and Ireland. Species accounts are given for all 30 species, although distribution maps and fullpage accounts are included only for those species recorded from live sightings. Six species have only been recorded from strandings. In Britain, these include Gervais’ beaked whale Mesoplodon europaeus, Blainville’s beaked whale Mesoplodon densirostris, narwhal Monodon monoceros, melon-headed whale Peponocephala electra and Fraser’s dolphin Lagenodelphis hosei; in Ireland, True’s beaked whale Mesoplodon mirus and Gervais’ beaked whale are both known only from strandings. True’s beaked whale is the only species recorded in Ireland but not in Britain. The distribution maps presented here are compiled from verified sightings between 1980 and the present, held within the Sea Watch Foundation’s national database and the NERC/Defra-funded Marine Ecosystem Research Programme (see Table 1 for details). These records are derived from observations made from the coast, from vessels, and from aerial surveys (both dedicated and opportunistic). Together, they total more than 200,000 records of live sightings. Strandings have not been included on the maps, although some live sightings near the coast were followed by the animal stranding. Occurrence data are shown in this Atlas for two specific time periods: a historic period (1960–92; for cetaceans, 1980–99) and the current time period (2000–16). The historic period was chosen to coincide with the time frame of the 1993 mammal atlas (Arnold, 1993; hereafter referred to as the Arnold Atlas), to facilitate comparisons

4  Atlas of the Mammals of Great Britain and Northern Ireland TABLE 1  Sources of cetacean sightings data Data source

No. of records

Data source

No. of records

Sea Watch Foundation

122,139

SAMM Survey

835

JNCC European Seabirds at Sea

14,527

Manx Whale & Dolphin Group

795

Dutch Seabird Group

10,447

Irish National Parks & Wildlife Service

701

The University of Veterinary Medicine Hannover

8,871

NORCET Surveys

669

WWT Consulting

6,796

MARINELife

616

JNCC MMO data

5,310

HiDef

449

Hebridean Whale & Dolphin Trust

4,988

CODA Survey

321

Cardigan Bay Marine Wildlife Centre

3,722

Aarhus University

227

Irish Whale & Dolphin Group

3,184

Marine Awareness North Wales

203

Institute for Marine Resources and Ecosystem Studies

3,015

Mardik Leopold

102

ORCA

2,194

Cornwall Wildlife Trust

85

Cetacean Research & Rescue Unit

2,192

SIAR Survey

80

SCANS-2 Survey

2,031

Coordinadora para o Estudo dos Mamíferos Mariños

65

Royal Belgium Institute for Natural Sciences

1,371

University of Swansea

58

Whale & Dolphin Conservation

1,240

RWE nPower

52

SCANS-1 Survey

1,198

PELGAS Survey

26

Bundesamt für Naturschutz

1,191

APEM

19

Instituut voor Natuur- en Bosonderzoek

928

ATLANCET Survey

1

Marine Conservation Research

911

JUVENA Survey

1

University of Aberdeen

911

TOTAL

between the two atlases. It should be noted that the information displayed in the current publication does not exactly replicate that shown for the same period in the Arnold Atlas. Ongoing updates to, and cleaning of, the dataset mean that there are additional data points that were missing from the Arnold Atlas; and conversely, some of the records shown in the Arnold Atlas have been removed or relocated. Cetaceans were only sparsely and sporadically recorded before 1980, though records have become relatively common subsequently. Given the scarcity of data prior to the 1980s, the relative abundance of data in the 1990s, and the fact that the Arnold Atlas excluded cetaceans, there was little benefit from aligning the historical time period with the Arnold Atlas. Therefore, for cetaceans, the historical time period begins when recording became more common (1980) and ends at 1999, i.e. immediately before the current time period. The current period for the Atlas is considered to be from 2000 to 2016 for both terrestrial mammals and cetaceans. The dates were set so that available data from published local county atlases could be used. However, for a few species that have experienced rapid changes in distribution – red squirrel Sciurus vulgaris, grey squirrel Sciurus carolinensis and water vole Arvicola amphibius – shorter time periods have been used. The current period is set at 2005–16 for the water vole, and 2010–16 for both squirrel species.

202,471

Methods 5

DISTRIBUTION MAPS The distribution maps were produced from the collated occurrence data by Colin Harrower (for terrestrial species) and James Waggitt (cetaceans). Terrestrial mammal and seal maps were created using the ‘BRCmap’ package and the cetacean maps were produced using ‘raster’ (Hijmans, 2013) and ‘maptools’ (Bivand & Lewin-Koh, 2015) packages, within the analytical framework R (R Core Team, 2017). The data for both terrestrial mammals and cetaceans are mapped at a 10 km square resolution. The terrestrial species used the Ordnance Survey of Great Britain and Ordnance Survey of Ireland 10 km resolution grid squares. From here on, both marine and terrestrial grids will be referred to as hectads. For each hectad, a symbol is plotted to indicate whether a species was present in that hectad during the historical period, the current period, or both periods. There may be some discrepancies between the Atlas maps and the known distributions of some species. This is because the maps were created using the available records for each species, and these records are not always complete. In some areas, for example the Scottish Highlands and Islands, the numbers of records are low (Figure 1). Lower recording effort may create artefactual gaps in distributions, and means that populations may not be well mapped on islands. In addition, data-sharing agreements mean that some records might not have been accessible. There is also the possibility of false presence records if the species can easily be misidentified, for instance stoat Mustela erminea and weasel Mustela nivalis, or bank and field voles Microtus agrestis. Although most errors of this kind have been removed during the data-cleaning process, it is very difficult to identify false positives where they lie within the known geographical range of the species and occur at an appropriate time of year. It must also be stressed that a positive hectad represents a record, not necessarily an established breeding population. Therefore, the distribution maps display the true distribution from all verified records. As with all distribution atlases, the maps will only be as good as the available records from which they were created. So, many will represent the actual species distribution as well as, to some extent, the distribution of the observers. Since cetaceans are very mobile and do not observe political boundaries, we have taken more natural areas and included records from across the North Sea and within the shelf seas surrounding Britain and Ireland. Figure 2 shows the boundaries of the area for which cetacean records have been presented and also includes the UK’s Exclusive Economic Zone to represent the extent of British waters. The seasonal distribution, or phenology, of records for each species is presented alongside the distribution maps. These phenology plots show a breakdown of the number of records per month to highlight the times of the year in which recordings for each species tend to be made. The phenology plots only include data from the current atlas period, 2000–16, and are only displayed for species with ten or more records.

RECORD COVERAGE ACROSS THE UK Terrestrial mammal records from the current atlas period (2000–16) were collected from across the UK, the Channel Islands and the Isle of Man (Figure 1). In this Atlas, the terms ‘UK’ and ‘Great Britain’ are used as geographical terms that encompass the Crown dependencies, rather than political boundaries, and ‘Ireland’ refers to the island of Ireland. There was at least one record for most hectads including many islands, such as the Northern Isles, the Hebrides, the Isles of Scilly and the Isle of Wight. The number of records for each hectad ranged dramatically from one to over 2,000, and the distribution of records per hectad revealed distinct areas of low and

6  Atlas of the Mammals of Great Britain and Northern Ireland

Number of Records

FIGURE 1 ​The number of terrestrial mammal records per hectad from the 2000–16 atlas period across Great Britain and Northern Ireland.

Methods 7 FIGURE 2  Map of the area for which cetacean records have been reported. The UK’s Exclusive Economic Zone has also been included to represent the extent of British waters.

high numbers of records. Areas with high numbers of records per hectad are located in the following regions: south of the line from the Bristol Channel to the Wash; Nottinghamshire; Derbyshire; Cumbria; and Northumberland. Areas of low numbers of records include the following: Pembrokeshire and Glamorgan; East Riding of Yorkshire; Northern Ireland; southern Scotland; and north-west Scotland (Figure 1). Every species of terrestrial mammal covered in this Atlas, with the exception of the raccoon Procyon lotor, soprano pipistrelle bat Pipistrellus pygmaeus and the Alcathoe bat Myotis alcathoe (the latter two only being identified recently), has records from 2000 to 2016 and was present in at least one hectad in both the 2000–16 and the 1960–92 time periods. There were nearly 300,000 mammal occurrence records from the historical time period. This increased dramatically (by 435%) to just over 1.3 million records for the current period, despite the time period under consideration being shorter (17 years compared with 33 years), highlighting the increased recording of mammals between the two periods. All species except the common vole, wildcat Felis silvestris and house mouse Mus musculus had more records in the current than the historical time period (Table 2). Owing to very small numbers of records, black rat Rattus rattus and greater mouse-eared bat Myotis myotis are excluded from time-trend assessments. There have been some noticeable changes in the distribution of some species: of the 60 resident terrestrial mammals, 13 have shown a decline in the number of occupied hectads. Declines of >30% were seen for the wildcat, red squirrel, harvest mouse Micromys minutus and house mouse. It is also apparent that some mustelids,

8  Atlas of the Mammals of Great Britain and Northern Ireland TABLE 2 ​The number of terrestrial mammal occurrence records and the number of occupied hectads. The final column shows the change in number of occupied hectads between the two time periods indicating distributional change. Symbols next to the species name highlight cases where the current time period differs from the standard definition of 2000–16 (+ indicates 2005–16 and * indicates 2010–16). Where species are non-native (NN), are reintroduced, or are naturalised (nat), this is shown in parentheses next to the common name. Naturalised species are those that are present in the UK owing to human activities, but have been present since at least Roman times. Records 1960–92

Records 2000–16

Hectads 1960–92

Hectads 2000–16

Hectad change

Hedgehog

17,618

134,753

2,191

2,507

316

European mole

21,807

65,493

2,259

2,325

66

Common shrew

7,449

8,514

1,630

1,445

–185

Pygmy shrew

3,374

4,218

1,195

1,008

–187

Water shrew

1,946

3,318

834

972

138

Common name

Lesser white-toothed shrew (nat)

38

64

4

4

0

European rabbit (nat)

21,251

106,569

2,491

2,540

49

Brown hare (nat)

13,948

49,378

1,847

1,967

120

Mountain hare

2,186

5,945

475

607

132

Red squirrel (*)

9,329

27,565

1,193

806

–387

14,833

42,575

1,511

1,772

261

0

164

0

53

53

2,717

30,898

453

572

119

Grey squirrel (*, NN) Eurasian beaver (reintroduced) Hazel dormouse Edible dormouse (NN)

128

154

13

23

10

Bank vole

4,736

9,156

1,258

1,284

26

Field vole

6,036

12,340

1,482

1,624

142

46

45

17

9

–8

Water vole (+)

9,879

32,764

1,326

1,224

–102

Harvest mouse

2,566

2,926

788

649

–139

Wood mouse

6,807

20,296

1,627

1,641

14

693

3,467

271

371

100

Common vole (nat)

Yellow-necked mouse House mouse (nat)

3,242

2,542

1,074

686

–388

Brown rat (NN)

6,933

16,824

1,621

1,561

–60

159

29

82

16

–66

Black rat (nat) Wildcat

602

371

351

111

–240

Red fox

13,686

91,616

2,035

2,452

417

Badger

19,825

80,110

2,048

2,292

244

Otter

24,760

82,866

2,051

2,394

343

Methods 9 Records 1960–92

Records 2000–16

Hectads 1960–92

Hectads 2000–16

Hectad change

767

3,268

329

559

230

Stoat

7,645

13,675

1,791

1,771

–20

Weasel

7,565

8,735

1,664

1,527

–137

Polecat

1,915

4,210

332

775

443

American mink (NN)

5,311

12,213

1,402

1,617

215

5

468

3

67

64

4,827

41,262

970

1,222

252

850

8,488

266

411

145

Fallow deer (nat)

2,701

19,507

636

1,028

392

Roe deer

8,234

87,385

1,470

2,091

621

400

3,251

71

120

49

Reeves’ muntjac deer (NN)

2,998

20,622

680

966

286

Greater horseshoe bat

4,121

5,451

192

293

101

Lesser horseshoe bat

8,196

17,856

334

460

126

Common name Pine marten

Wild boar (reintroduced) Red deer Sika deer (NN)

Chinese water deer (NN)

Alcathoe bat Whiskered bat Brandt’s bat Bechstein’s bat

0

35

0

19

19

1,357

3,018

446

725

279

337

798

112

241

129

174

1,468

33

134

101

3,506

24,521

655

1,479

824

95

78

5

1

–4

Natterer’s bat

3,389

13,759

679

1,190

511

Serotine bat

1,420

9,304

300

708

408

Daubenton’s bat Greater mouse-eared bat

Leisler’s bat

463

1,419

124

453

329

Noctule bat

1,834

16,199

531

1,362

831

Common pipistrelle bat

3,037

63,549

527

1,999

1,472

Soprano pipistrelle bat Nathusius’ pipistrelle bat Barbastelle bat Brown long-eared bat Grey long-eared bat Grey seal Harbour seal

0

37,920

0

1,848

1,848

12

1,363

11

432

421

144

2,815

82

436

354

8,853

29,375

1,378

1,768

390

72

246

33

45

12

1,920

11,058

643

1,007

364

995

6,245

450

725

275

10  Atlas of the Mammals of Great Britain and Northern Ireland

Number of Species

FIGURE 3 Species richness, defined as the number of species recorded per hectad, for terrestrial mammals from the current atlas period (2000–16).

Methods 11

FIGURE 4  Species richness of cetaceans, calculated as the number of species recorded per hectad from the current period (2000–2016).

deer and bat species have expanded their distribution. However, it can be difficult to determine whether the observed change of distribution reflects changes in observer effort (all), increases in recording technology (bats) and better species identification (bats), or an actual biological change in distribution (mustelids). The terrestrial species richness across the UK, defined here as the number of different species within each hectad, ranges from one to over 35 in the current time period (Figure 3). There is a distinct pattern, with the hectads in the south of Britain having a higher species richness than those in northern England, north and west Scotland, and Northern Ireland. For cetaceans, the number of species recorded per cell is greatest along Atlantic coasts (Figure 4): the Hebrides, southern Ireland, south-west England, and in the Irish Sea around Pembrokeshire, the Lly^ n Peninsula, Anglesey and the Isle of Man. In the North Sea, the greatest number of species occurs in the north-west

12  Atlas of the Mammals of Great Britain and Northern Ireland

FIGURE 5  The number of cetacean records per hectad from the current atlas period from across the North Sea and within the shelf seas surrounding Britain and Ireland.

of the region, around north and east Scotland. These are also the areas where the greatest number of records occur (Figure 5), influenced to some extent by variation in observer effort. However, sightings rates and species diversity from effort-corrected surveys show a similar pattern.

BIBLIOGRAPHY ARNOLD, H.R. 1993. Atlas of Mammals in Britain. ITE Research Publication No. 6. London: Joint Nature Conservation Committee and Institute of Terrestrial Ecology, HMSO. BIVAND, R. and LEWIN-KOH, N. 2015. Maptools: Tools for Reading and Handling Spatial Objects. R package version 0.8–36. HIJMANS, R.J. 2013. Raster: Geographic data analysis and modelling. R package version 2.1–66. R CORE TEAM. 2017. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. Accessed at https://www.R-project.org/ (30 July 2019).

SPECIES ACCOUNTS

14  Atlas of the Mammals of Great Britain and Northern Ireland

Hedgehog

Erinaceus europaeus (LINNAEUS, 1758) They were probably introduced to mainland Ireland and Beginish Island (Kerry).

ECOLOGY The hedgehog is a nocturnal insectivore found in a wide range of rural and urban environments. It eats a variety of macroinvertebrates including worms, beetles, insect larvae and gastropod molluscs, but it may also take carrion and small vertebrates. The species spends the day in a nest in undergrowth which is typically constructed of broad leaves and/or grasses. The hedgehog hibernates in winter.

IDENTIFICATION The hedgehog is the UK’s only spiny mammal. Sharp spines completely replace hair on the back and crown of the head. It ‘rolls up’ in defence leaving only spiny skin exposed. The pelage is typically mid-brown. It has a short tail, small eyes and a tapering snout.

BIBLIOGRAPHY

MARK BALDWIN

DISTRIBUTION The hedgehog remains widespread in suitable habitats across the UK, but there is good evidence of a serious and continuing decline. Many possible factors can be potentially associated with this change in abundance, including agricultural intensification (particularly the use of biocides and a reduction in the availability of prey, nest sites and refuges), habitat loss and degradation, landscape fragmentation, and mortality from road traffic and predation. This species is generally absent from wetlands, moors, uplands, coniferous forests and city centres. Hedgehogs have been introduced to many archipelagos and islands: Shetland, Orkney, the Western Isles (Lewis, Harris, Benbecula, North and South Uist), Skye, Soay, Canna, Coll, Mull, Ulva, Luing, Islay, Arran, Bute, the Isle of Man, Anglesey, the Isle of Wight, St Mary’s (Isles of Scilly), Alderney, Guernsey, Jersey and Sark.

HOF, A.R. 2009. A study of the current status of the hedgehog (Erinaceus europaeus), and its decline in Great Britain since 1960. Unpublished PhD thesis, Royal Holloway University of London. MORRIS, P.A. & REEVE, N.J. 2008. Hedgehog Erinaceus europaeus. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 241–249. PTES & BHPS. 2015. The State of Britain’s Hedgehogs 2015. London and Ludlow: People’s Trust for Endangered Species and British Hedgehog Preservation Society.

AUTHOR  Nigel Reeve

Hedgehog 15

16  Atlas of the Mammals of Great Britain and Northern Ireland

European mole

Talpa europaea (LINNAEUS, 1758)

ANDREW FUSEK PETERS

DISTRIBUTION The mole is found throughout Great Britain, including on several islands where it was probably introduced. However, it is absent from Ireland. It is found in most habitats where the soil is deep enough to allow tunnelling, but is uncommon in coniferous forests, on moorlands and in sand dunes, probably because its prey is scarce here. There has been no change in distribution over time, even though it is regularly trapped. Most records are from molehills, with some live sightings and dead animals, including those killed by cats.

ECOLOGY The mole spends almost all of its life underground in a system of permanent and semi-permanent tunnels. Surface tunnels are usually short-lived and are employed in very shallow soils where prey is concentrated just below the surface. A system of permanent deep burrows, which form a complex network hundreds of metres long at varying soil depths, is more common. Permanent tunnels are used repeatedly for feeding over long periods

of time, sometimes by several generations of moles. Molehills are pushed up when redigging trampled tunnels or when establishing new territories. Moles are often seen in spring and autumn, associated with the onset of breeding and the dispersal of juveniles.

IDENTIFICATION The mole has short velvety fur. Its fur is usually black, silver, grey, cream or white, but shades of brown have also been recorded and can dominate in specific populations. It has spade-like forelimbs with large claws that face towards the rear of the animal. It has a pink fleshy snout and tiny eyes.

BIBLIOGRAPHY ATKINSON, R. 2013. Moles, 1st edn. London: Whittet Books. STONE, D. 1986. Moles. Oswestry: Anthony Nelson.

AUTHOR  Derek Crawley

European mole  17

18  Atlas of the Mammals of Great Britain and Northern Ireland

Common shrew

Sorex araneus (LINNAEUS, 1758) aggressive towards each other. The species is active almost continuously throughout the day and night.

IDENTIFICATION The common shrew has a long pointed nose, small eyes, small ears and red-tipped teeth. It has tricoloured fur: dark brown on the back, pale brown on the sides and whitish underneath. It can be distinguished from the water shrew Neomys fodiens by its brown (not black) upper surface, and smaller size. The common shrew differs from the smaller pygmy shrew Sorex minutus by having distinctively sharp contrast between the coat colours on the back and flank, as well as by having a relatively shorter, thinner tail.

BIBLIOGRAPHY CHURCHFIELD, S. & SEARLE, J. 2008. Common shrew Sorex araneus. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 257–265.

AUTHOR  Anna Champneys

MALCOLM WELCH

DISTRIBUTION The common shrew is widespread throughout much of Great Britain. However, it is absent from Ireland, as well as from Shetland, Orkney, the Outer Hebrides and the Isles of Scilly. Records come mainly from cat kills, animals live-trapped during small mammal surveys, and remains found in barn owl pellets.

ECOLOGY The common shrew is found in most terrestrial habitats where there is short vegetation cover such as thick grass, bushy scrub, hedgerows, bracken, deciduous woodland and roadside verges. It is an abundant species, with highest population densities being recorded in grassland habitats and bog. Its diet consists mainly of terrestrial invertebrates such as earthworms, slugs, snails, beetles, spiders and woodlice. Essentially solitary except during the breeding season, individuals are extremely

Common shrew  19

20  Atlas of the Mammals of Great Britain and Northern Ireland

Pygmy shrew

Sorex minutus (LINNAEUS, 1766)

BECKY HAYWOOD

DISTRIBUTION

IDENTIFICATION

The pygmy shrew is widespread throughout Great Britain and Ireland. However, it is absent from Shetland, Lewis, the Isles of Scilly and the Channel Islands. Records are scarce and come mainly from cat kills, animals live-trapped during small mammal surveys, and remains found in barn owl pellets.

The pygmy shrew is similar in appearance to the common shrew with a long pointed nose, small eyes, small ears and red-tipped teeth. It is, however, much smaller and possesses a relatively longer, thicker and more hairy tail. Its coat is also bicoloured, lacking the distinctive sharp contrast between coat colour on the back and flank seen in the common shrew.

ECOLOGY The pygmy shrew is found in most terrestrial habitats where there is plenty of ground cover. Population densities are typically higher in grassland than woodland with the exception of Ireland, where its competitor – the common shrew – is absent. The pygmy shrew is less abundant than the common shrew, and has a similar diet, although with a tendency for smaller-sized prey. Spiders, harvestmen, woodlice and beetles are its most common prey. The pygmy shrew, like the common shrew, is essentially solitary except during the breeding season and individuals are extremely aggressive towards each other. The species is active almost continuously throughout the day and night.

BIBLIOGRAPHY CHURCHFIELD, S. & SEARLE, J. 2008. Pygmy shrew Sorex minutus. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 267–271. MACDONALD, D.W., MACE, G.M. & RUSHTON, S. 1998. Proposals for Future Monitoring of British Mammals. London: Department of the Environment, Transport and the Regions with Joint Nature Conservation Committee, Peterborough.

AUTHOR  Anna Champneys

Pygmy shrew  21

22  Atlas of the Mammals of Great Britain and Northern Ireland

Water shrew

Neomys fodiens (PENNANT, 1771) The water shrew, like its terrestrial shrew counterparts, is solitary and territorial with an elusive nature. It exists in small localised populations and is active throughout the day and night.

IDENTIFICATION

KATIE NETHERCOAT

DISTRIBUTION Within Great Britain, the water shrew has a wide but patchy distribution. It is present on many islands including Skye, Mull, Anglesey and the Isle of Wight. However, it is more localised in Scotland and is absent from Ireland. It typically occurs at much lower population densities than its terrestrial counterparts, the common and pygmy shrews. The first national survey in 2004–05 identified water shrew presence in riparian habitats by the analysis of faecal samples collected in baited tubes. Of the 2,159 sites surveyed, 17% showed signs of water shrew occurrence. Although the presence of water shrews largely reflected the distribution of surveyors (the majority coming from England (82%), with 10% from Scotland and 8% from Wales), there was a notable concentration of positive records in central and eastern England. Records come mainly from cat kills, animals livetrapped during small mammal surveys, bait tube surveys, and remains found in barn owl pellets.

ECOLOGY The water shrew occupies a wide variety of riparian habitats including ponds, canals, reed beds and marshes as well as clear, fast-flowing streams and watercress beds – habitats with which it has been traditionally associated. However, it is also found regularly, if infrequently, away from water in agricultural habitats. Its diet consists mainly of terrestrial and aquatic invertebrates.

The water shrew is the largest of the six shrews inhabiting Britain; it can be distinguished from the terrestrial shrews by its short, dense, velvety black fur on the upper surface of the body and greyish-white underside. Most water shrews have tufts of white fur on the ears and around the eyes. There are distinctive stiff hairs on the edges of the feet and along the underside of the tail that form a keel. The water shrew has a long pointed nose, small eyes and small ears.

BIBLIOGRAPHY CARTER, P. & CHURCHFIELD, S. 2006. Distribution and Habitat Occurrence of Water Shrews in Great Britain. Environment Agency Science Report SC010073/SR. Bristol: Environment Agency. Mammal Society Research Report No. 7. London: The Mammal Society. CHURCHFIELD, S. 2008. Water shrew Neomys fodiens. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 271–275.

AUTHOR  Anna Champneys

Water shrew  23

24  Atlas of the Mammals of Great Britain and Northern Ireland

Lesser white-toothed shrew Crocidura suaveolens (PALLAS, 1811)

The species is active almost continuously throughout the day and night.

IDENTIFICATION The lesser white-toothed shrew has a long pointed nose, small eyes, short-haired prominent ears and wholly white teeth. Its coat is bicoloured with greyish or reddish brown fur on the back and paler fur on the underside. The tail of this species is covered with short hair interspersed with long, white hairs. It is the only shrew species on the Isles of Scilly. On Jersey it can be distinguished from the Millet’s shrew Sorex coronantus by the lack of red-tipped teeth and its long, scattered tail hairs.

BIBLIOGRAPHY

JULIE LOVE

DISTRIBUTION In the UK, the lesser white-toothed shrew is found only on the Isles of Scilly – where it is most abundant on St Mary’s and Tresco – and on the Channel Islands. Records come from animals live-trapped during small mammal surveys, faecal remains in bait tubes, cat kills, incidental sightings and individuals found under corrugated iron sheets.

ECOLOGY The lesser white-toothed shrew is found in most terrestrial habitats where there is adequate cover such as bracken, hedgerows and woodland. It can also be found in coastal habitats among boulders and vegetation on the shores. The most common prey of this species includes beetles, flies, insect larvae, centipedes, earthworms and gastropods. However, in coastal habitats, it feeds predominantly on crustaceans. The lesser white-toothed shrew is generally solitary but it has overlapping home ranges, so it is not as territorial as the other shrews.

CHURCHFIELD, S. & TEMPLE, R.K. 2008. Lesser whitetoothed shrew Crocidura suaveolens. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 276–280. CORNWALL MAMMAL GROUP. 2015. Lesser white-toothed shrew. Accessed at https://www.cornwallmammalgroup. org/lesser-white-toothed-shrew (30 July 2019).

AUTHOR  Anna Champneys

Lesser white-toothed shrew  25

26  Atlas of the Mammals of Great Britain and Northern Ireland

European rabbit

Oryctolagus cuniculus (LINNAEUS, 1758) has become recognised as a keystone ecosystem engineer providing environmental conditions required by a range of fauna and flora in UK habitats such as the East Anglian Breckland and southern chalk downland. It is also prey for a number of avian and mammalian predators.

IDENTIFICATION The species is easily distinguished from the brown hare Lepus europaeus by its shorter limbs and shorter ears, which lack the black tip of the latter. The brown pelage on the top of the body can be replaced by black coat in melanistic individuals in some regions. The adult body weight of a UK wild rabbit can reach 2 kg. PADRAIG KAVANAGH

DISTRIBUTION The rabbit is a naturalised species with a wide distribution across the UK. It is recorded in more hectads than any other species in both the current and historical atlas periods. However, there are currently concerns about its population status in parts of its range, particularly in Scotland, based on data collected by the British Trust for Ornithology’s Breeding Bird Survey. Mass mortalities have been caused by myxomatosis and rabbit viral haemorrhagic disease, but there remains uncertainty in the scale of the decline owing to the large natural variability in rabbit population sizes and the lack of systematic monitoring for this species.

ECOLOGY The rabbit has a herbivorous diet that can include a range of plant species. Where available, it prefers short grass swards where mammalian predators are visible. Tree bark may be consumed during winter months, and fallen leaves during summer droughts. The rabbit lives in warren-based territorial social groups of 1–3 males and 1–9 females in areas of burrowable substrate. The dispersal of juveniles is male-biased, with female offspring remaining in their natal social groups. The rabbit may be active above ground throughout the day, although activity peaks during crepuscular periods in areas of human disturbance. The species

BIBLIOGRAPHY HARRIS, S.J., MASSIMINO, D., GILLINGS, S., EATON, M.A., NOBLE, D.G., BALMER, D.E., PROCTER, D., PEARCEHIGGINS, J.W. & WOODCOCK, P. 2018. The Breeding Bird Survey 2017. The British Trust for Ornithology Research Report 706. Thetford: British Trust for Ornithology. LEES, A.C. & BELL, D.J. 2008. A conservation paradox for the 21st century: the European wild rabbit Oryctolagus cuniculus, an invasive alien and an endangered native species. Mammal Review 38 (4): 304–320. WEBB, N.J., IBRAHIM, K., BELL, D.J., HEWITT, G. 1995. Natal dispersal and genetic structure in a population of the European wild rabbit Oryctolagus cuniculus. Molecular Ecology 4 (2): 239–247.

AUTHOR  Diana Bell

European rabbit  27

28  Atlas of the Mammals of Great Britain and Northern Ireland

Brown hare

Lepus europaeus (PALLAS, 1778) Hares produce caecotrophs from the anus which they eat, thus ‘double-​digesting’ their food.

IDENTIFICATION The brown hare can be distinguished from the rabbit by its larger size and its longer ears with distinctive black tips. An adult brown hare weighs 3.5 kg. It is a tall animal with long legs that lopes or runs with a well-​ developed stride. At full gallop, a brown hare can reach speeds of 55–70 km/h (35–45 mph). When running, the white tail is usually held downwards showing a black upper surface. The coat is brown, ranging from russet to dark brown, with white on the belly, and the iris of the eye is orange. However, hair and eye colour mutations are found.

BIBLIOGRAPHY

IAN HULL

DISTRIBUTION There were no records of the brown hare in Britain before the Roman invasion in the first century ad, and it was introduced to Ireland for sport in the mid-late nineteenth century. Since its introduction, it has replaced the indigenous mountain hare Lepus timidus across most of Britain, and competes with it in Ireland. However, it is unable to eat coarse food and this has led to it staying below the heather line. Brown hare population sizes are thought to have dropped by 90% since the second half of the nineteenth century.

ECOLOGY The brown hare breeds throughout most of the year. The female hare, or jill, can have up to four litters a year, although most do not breed until their second year. Boxing, the jill rejecting the male, is most often noted in spring when the crops are low and most females come into season. The brown hare evolved on steppes and plains and spread as humans cut down forests. It prefers old meadow land with a mix of wild plants; leverets cannot eat farm crops when they get coarse.

RUSS, J. 2015. The Hare Book. Llangennech: Hare Preservation Trust and Graffeg Publishing. TAPPER, S. & YALDEN, D. 1987. The Brown Hare. Southampton: The Mammal Society.

AUTHOR  Jo Sharplin

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Mountain hare

Lepus timidus (LINNAEUS, 1758) ECOLOGY

BILLY STOCKWELL

DISTRIBUTION The mountain hare is the indigenous hare that has been in Britain since the last Ice Age. It is native to the Highlands of Scotland, but was introduced by the Victorians to the Peak District in Derbyshire. It is also found on some Scottish islands including Hoy (Orkney), Shetland, Mull and Skye. The subspecies L. timidus hibernicus is indigenous to Ireland, where it is known as the Irish hare. It is associated with heather moorlands and occupies higher ground than the brown hare: some 350 m up in the Lammermuir Hills in the Scottish Borders and 750 m to over 1,000 m in the Cairngorms. Population densities are known to vary at least tenfold, reaching a peak roughly every ten years; reasons for these variations are uncertain. There is concern that high culling rates in some areas may adversely affect the population status of the species, but the high variability in population densities, particularly in habitats where they are most abundant such as grouse-moors, makes understanding population trends challenging.

As well as heather, the mountain hare will browse gorse, and dwarf shrubs or trees such as birch, rowan and juniper. In summer, it prefers to eat grasses when available. In its upland and moorland habitat, it makes forms – depressions in snow, soil, or in the lea of a heather hummock – which provide shelter. It rests in these by day and at night, and in periods of snow cover it often gathers in large groups on leeward slopes to shelter or to feed. Its ‘runs’ typically pass up slopes as the mountain hare’s short front legs and powerful long back legs are ideal for running uphill at speeds of up to 60 km/h (37 mph). Mating begins at the end of January, and gestation lasts about 50 days. Most leverets are born between March and August inclusive, fully furred and with eyes open. There can be several litters a year, usually between one and three young. On average, the lifespan of the mountain hare is three to four years.

IDENTIFICATION The mountain hare is smaller and more compact than the brown hare; its total body length can range from 43 cm to 61 cm, and it weighs 2.5–3.5 kg. Its black-tipped ears, at 6–8 cm, are also shorter than those of the brown hare. In spring it loses its thick white coat, changing to a brownish one. The brown coat begins to shed in late October, and by December it again has thick white fur. The tail remains white, and the ear tips black, throughout the seasons. The moult is dependent upon temperature and appears to be initiated by shortening day lengths.

BIBLIOGRAPHY HESFORD, N., FLETCHER, K., HOWARTH, D., SMITH, A.A., AEBISCHER, N.J. & BAINES, D. 2019. Spatial and temporal variation in mountain hare (Lepus timidus) abundance in relation to red grouse (Lagopus lagopus scotica) management in Scotland. European Journal of Wildlife Research 65 (3): 33. RUSS, J. 2015. The Hare Book. Llangennech: Hare Preservation Trust and Graffeg Publishing. WATSON, A. & WILSON, J.D. 2018. Seven decades of mountain hare counts show severe declines where high-yield recreational game bird hunting is practised. Journal of Applied Ecology 55 (6): 2663–2672.

AUTHOR  Marion O’Neil and Jo Sharplin

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Red squirrel

Sciurus vulgaris (LINNAEUS, 1758) Recent survey and data review suggest that grey squirrel control may have stabilised, and in some localities begun to reverse, regional red squirrel declines. However, the pattern is complex, and most populations remain at risk from sympatric or adjacent grey squirrel populations.

ECOLOGY The red squirrel is an arboreal diurnal species typical of mature coniferous and broadleaved woodlands. Its diet is largely tree seeds, flowers and shoots, with a little animal matter and fungi. PADRAIG KAVANAGH

DISTRIBUTION The red squirrel is native to Britain and Ireland. Historically eliminated from some areas by culling, restocking in the nineteenth century included animals taken from populations elsewhere in Europe. It has recently been introduced to Tresco, Mersea and Caldey, and a series of conservation-​driven introductions have occurred in the north and north-west of Scotland, beyond its historical range. Over recent decades, widespread regional extinctions of the red squirrel have occurred as a result of resource competition with, and epizootic infection spread by, the grey squirrel. While absent from Scotland’s central belt, significant mainland populations remain in northern and border counties. The species also remains widespread in Cumbria and Northumberland, while small remnant populations persist in Durham, Yorkshire and Lancashire. Recent records in the southern part of England are likely to have originated from escapes. In Wales, isolated populations are found in northern Gwynedd, Clocaenog forest (Denbighshire) and conifer plantations surrounding Llyn Brianne (Tywi, Crychan forests) in mid-Wales. Notable island populations include Anglesey, Arran, Brownsea, Furzey and the Isle of Wight. In Ireland, the natural re-establishment of pine martens Martes martes is correlated with grey squirrel decline and red squirrel recovery. In the absence of this predator, Irish red squirrel populations continue to decline as the grey squirrel increases it range.

IDENTIFICATION The red squirrel has a long tail and typically prominent ear tufts. It has white chest hair, but the rest of the coat can vary from russet red to brown, black and occasionally grey. It can be confused with the larger and stockier grey squirrel, which seldom has ear tufts and has a characteristic outer band of silver on the tail hair. The red squirrel is usually smaller than the grey squirrel, with a body length of 18–23 cm.

BIBLIOGRAPHY GURNELL, J., LURZ, P. & BERTOLDI, W. 2014. The changing patterns in the distribution of red and grey squirrels in the North of England and Scotland between 1991 and 2010 based on volunteer surveys. Hystrix, the Italian Journal of Mammalogy 25 (2): 83–89. LAWTON, C., FLAHERTY, M., GOLDSTEIN, E.A., SHEEHY, E. & CAREY, M. 2012. Irish squirrel survey 2012. Irish Wildlife Manuals, No. 89. Dublin: National Parks and Wildlife Service, Department of Arts, Heritage and the Gaeltacht, Ireland. SHUTTLEWORTH, C.M., LURZ, P. & HAYWARD, M.W. 2015. Red Squirrels: Ecology, Conservation and Management in Europe. Kenilworth: European Squirrel Initiative.

AUTHOR  Craig Shuttleworth

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Grey squirrel

Sciurus carolinensis (GMELIN, 1788)

ROBIN MORRISON

DISTRIBUTION

ECOLOGY

The grey squirrel was introduced to the UK in 1876, and to Ireland in 1911, from North America. It is widely distributed and common throughout England, Wales, much of Ireland, southern Scotland and the central belt. Natural range expansion continues, with the exception of the Irish midlands, where population declines are correlated with the establishment of high-density pine marten populations. No populations exist on offshore islands since the species was eradicated from Anglesey in 2013. Individuals have, however, occasionally been found on the island of Skye (accidental transportation), the Isle of Wight (accidental or deliberate transportation) and Anglesey (limited natural dispersal from mainland). Systematic 4 km2 presence/absence surveys are carried out in Scotland and northern England as part of red squirrel conservation measures. These regional assessments are supplemented with records of culled animals and sighting reports. Elsewhere, because the grey squirrel lives at high density and is a highly visible species, distribution is not monitored through systematic study. Populations are known to be affected by both weather patterns and annual availability of tree seed crops; supplemental feeding (at bird tables) may thus buffer fluctuation in local population abundance. Deliberate or accidental transportation and release could accelerate range expansion. It is important that potential sightings of grey squirrels beyond the current established distribution are reported.

The grey squirrel is a diurnal species, less arboreal than the red squirrel and able to occupy more fragmented woodland habitats. Its diet is principally tree seeds, flowers and shoots, with some animal matter and fungi; it also strips bark and because of this is considered an economic pest.

IDENTIFICATION The grey squirrel is larger and stockier (body length 23–27 cm) than the native red squirrel and seldom has ear tufts. It also typically has a silver-coloured fringe to the tail hair and, although most individuals are grey in colour, many have reddish pelage on head and flanks, with some animals being very red. Melanistic (black) individuals can be found in south-east England.

BIBLIOGRAPHY GURNELL, J., LURZ, P. & BERTOLDI, W. 2014. The changing patterns in the distribution of red and grey squirrels in the North of England and Scotland between 1991 and 2010 based on volunteer surveys. Hystrix, the Italian Journal of Mammalogy 25 (2): 83–89. LAWTON, C., FLAHERTY, M., GOLDSTEIN, E.A., SHEEHY, E. & CAREY, M. 2012. Irish squirrel survey 2012. Irish Wildlife Manuals, No. 89. Dublin: National Parks and Wildlife Service, Department of Arts, Heritage and the Gaeltacht, Ireland. O’HARE, S. 2017. Results of the squirrel monitoring programme, Spring 2017. Carlisle: Red Squirrels Northern England.

AUTHOR  Craig Shuttleworth

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Eurasian beaver Castor fiber (LINNAEUS, 1758)

TONY HOUSE

DISTRIBUTION The Eurasian beaver is recovering from near global extinction and is now present throughout its former native range thanks to conservation efforts, including legal protection and translocations to over 25 European countries. Britain was one of the last EU states to consider restoring this species. Its numbers and distribution are currently increasing throughout Britain. Note that there is no evidence of this species ever being present outside of mainland Britain (on British islands, the Channel Islands, the Isle of Man or Ireland). The current British population is found mainly in Scotland and is widespread throughout the River Tay catchment, and Perthshire, including a small population officially released as part of the Scottish Beaver Trial (SBT), Knapdale forest and mid-Argyll. There are also smaller numbers of free-living individuals in England, mostly in the south-west, including the River Otter in Devon (now part of the Devon Beaver trial). Several locations in England have recently been granted licences to have beavers within extensive enclosures. There are reports of sightings of beaver signs in parts of Wales, but it is unclear whether there are any established populations. The individuals used for the recent reintroductions are confirmed as being from Norway (SBT) or Bavaria (Tayside and Devon).

ECOLOGY The Eurasian beaver is a semi-aquatic herbivore that can be found in fresh watercourses of varying size. It has been reported swimming in seawater but will not

reside there. It will dam narrower watercourses to create deeper pools for breeding and feeding. Tree felling is more pronounced during the autumn/winter and when beavers are first colonising an area. The beaver’s diet encompasses a wide range of riparian, emergent and aquatic vegetation. It is mainly nocturnal, though sightings during daylight hours are normal during spring/summer towards dusk and dawn.

IDENTIFICATION Adult beavers are large (approx. 25 kg) with rotund bodies, and are larger than otters Lutra lutra and badgers Meles meles. The key identification features include size, orange incisors and broad, flat tails, which are dark grey in colouration and appear scale-covered. The beaver’s fur is typically mid-brown in colour (variants are rare, but do exist across Europe, from black to light brown/ blonde), although there are reports of small numbers of black individuals having been observed within the Tayside catchment.

BIBLIOGRAPHY CAMPBELL-PALMER, R., GOW, D., NEEDHAM, R., JONES, S. & ROSELL, F. 2015. The Eurasian Beaver. Exeter: Pelagic Publishing and The Mammal Society. SCOTTISH NATURAL HERITAGE. 2015. Beavers in Scotland: A Report to Scottish Government. Inverness: Scottish Natural Heritage.

AUTHOR  Roisin Campbell-Palmer

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Hazel dormouse

Muscardinus avellanarius (LINNAEUS, 1758) nectar, fruits and invertebrates. The species hibernates in nests on or near the ground between November and April. It also exhibits torpor when in its active period as a means to conserve energy.

IDENTIFICATION

MATT BINSTEAD, BRITISH WILDLIFE CENTRE

DISTRIBUTION The hazel dormouse is native to England and Wales, but not to Scotland or Ireland. A hundred years ago it was thought to be present in almost all English counties. Having become extinct from 12 northern and Midland English counties over the past century, it is now recorded predominantly in more southerly parts of England and along the English–Welsh border. The most northerly county where there is still a native population is Cumbria. It remains well distributed throughout Wales, with higher concentrations towards the east and south of the country. Overall, in Britain, the species’ range is considered to be contracting. Data from the National Dormouse Monitoring Programme also indicates that the British population continues to decline. This is most likely the result of habitat fragmentation and inappropriate management.

ECOLOGY The hazel dormouse is a small arboreal rodent closely associated with the early successional stages of woody vegetation, such as that found in traditional coppiced and managed woodland and scrub. The species is considered relatively sedentary and a poor disperser, and it is unlikely to occupy suitable local habitat in the absence of good linkages. It also lacks a caecum (a pouch which is connected to the small and large intestine), which severely restricts its ability to digest cellulose; hence, it is a sequential specialist feeder on pollen,

The hazel dormouse is an agile arboreal rodent with large black eyes and prominent ears. The body length is 7–9 cm and its tail is well furred. Adults have sandy coloured coats on the back with paler bellies. Juveniles have more grey in the pelage. This species may be confused with the wood mouse Apodemus sylvaticus and bank vole, but its arboreal nature and furry tail are diagnostic. Hazelnuts are opened by dormice in a characteristic fashion, leaving a smooth opening, so feeding remains can be used to identify their presence.

BIBLIOGRAPHY GOODWIN, C., HODGSON, D.J., AL-FULAIJ, N., BAILEY, S., LANGTON, S. & McDONALD, R.A. 2017. Voluntary recording scheme reveals ongoing decline in the United Kingdom hazel dormouse Muscardinus avellanarius population. Mammal Review 47 (3): 183–197. WEMBRIDGE, D., AL-FULAIJ, N. & LANGTON, S. 2016. The State of Britain’s Dormice 2016. Report for the People’s Trust for Endangered Species. London: People’s Trust for Endangered Species.

AUTHOR  Ian White

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Edible dormouse Glis glis (BRISSON, 1762)

is often no reproduction at all, contrasting with single litter production in years with good tree flowering. The species hibernates typically for seven months, in rotten tree roots or burrows of other mammals, occasionally in drains, under floors and in outhouses. The edible dormouse is omnivorous, eating tree flowers, shoots and seeds, and predating hole-nesting bird species (especially in late spring/early summer). Infestations of edible dormice can cause bark stripping of trees and a severe nuisance (with health implications) in the built environment.

IDENTIFICATION

ROGER WASLEY

DISTRIBUTION The non-native edible dormouse escaped from Tring Park menagerie sometime after 1902. The species spread slowly but is now present more or less throughout the Chilterns area (bounded by the Chilterns chalk scarp and the M1, M25 and M40 motorways). Isolated records from many parts of England over the past 50 years are often lone unrepeated records, resulting from either misidentification or illegal translocations. Recently, dependable population records show populations venturing south and eastwards just outside the previous boundaries, including urban areas within the M25. There are also separate populations near Harlow in Essex and north of Reading.

ECOLOGY The edible dormouse is an arboreal nocturnal rodent that prefers old growth woodland and hibernates underground. It utilises fruiting-aged deciduous (especially beech), conifer or mixed woodlands, but travels along hedges and increasingly uses built-up environments as a refuge, both in rural and urban areas with nearby mature trees. It is considered a long-lived, generally sedentary species. In years with no or little tree flowering there

The edible dormouse has a typical blunt dormouse face, with large round ears and large eyes that are ringed with black fur. The head–body length is approximately 14–19 cm. The fur is grey over most of the body, but is white or buff on the underparts. The tail is bushy and flat on the underside. It has notable claws and ‘sticky’ feet adapted for climbing. The species also has cheek teeth, which are transversely ridged as is typical for dormice.

BIBLIOGRAPHY MORRIS, P.A. 2008. Edible dormouse Glis glis. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 82–85. THOMPSON, H.V. 1953. The Edible dormouse (Glis glis L.) in England, 1902–1951. Journal of Zoology 122 (4): 1017–1024. TROUT, R.C. & MOGG, A. 2017. Surveys of distribution and damage by the edible dormouse (Glis glis) in the built environment. Apodemus 14: 26–34.

AUTHOR  Roger Trout

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Bank vole

Myodes glareolus (SCHREBER, 1780)

GARY COX

DISTRIBUTION

IDENTIFICATION

The bank vole is present throughout mainland Britain and on the islands of Anglesey, Arran, Bute, Hayling, Handa, Ramsay, Scalpay, Ulva and the Isle of Wight, and with named subspecies on Mull, Raasay, Skomer and Jersey in the Channel Islands. It is not indigenous to Ireland, but was first recorded in County Kerry in 1964 (although probably accidentally introduced in the late 1920s) and is expanding its distribution, though it is not yet recorded in Northern Ireland. The bank vole is abundant throughout much of central and western mainland Europe.

The bank vole has a blunt nose, small ears, small eyes and a short tail. The back and top of the tail is chestnut brown, the underside is pale cream/grey. The head–body length is 8–12 cm. The tail is about half the length of the head and body. These features distinguish the bank vole from the field vole, which tends to be grey-brown in colour and has a tail about a third the length of the head and body. Characteristically, the bank vole feeds on hazelnuts, leaving a round hole with tooth marks across the inner edge but not on the surface of the nut. There are few other easy-to-spot field signs of its presence, although occasionally it may eat the bark of small trees.

ECOLOGY The bank vole is common in broadleaved woodland, scrub, field margins and hedgerows but is also found in conifer plantations, fenland and road verges. It is active night and day, with peaks at dawn and dusk. Although mainly herbivorous, it consumes a wide range of foods including green leaves, dead leaves (winter), flowers, fruits, seeds, roots, mosses, fungi and invertebrates. The bank vole exhibits annual cycles in densities, with peaks in the autumn. Multi-annual cycles are known to occur in northern Europe.

BIBLIOGRAPHY SHORE, R.F. & HARE, E.J. 2008. Bank vole Myodes glareolus. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 88–99.

AUTHOR  John Gurnell

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Field vole

Microtus agrestis (LINNAEUS, 1761) length. Relative tail length and colouration are the main ways to distinguish it from the bank vole. Its nests are made out of woven grass, are approximately 10 cm in diameter, and may be found in the base of grass tussocks or in burrows. Signs of presence also include runways, and tunnels through long grass, with clipped grass and droppings inside.

BIBLIOGRAPHY LAMBIN, X. 2008. Field vole Microtus agrestis. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 100–107.

AUTHOR  John Gurnell

DEREK CRAWLEY

DISTRIBUTION The field vole occurs throughout mainland Britain and many offshore islands, but is absent from Lewis, Barra and some Inner Hebrides islands, Orkney, Shetland, Lundy, the Isles of Scilly and the Channel Islands. It is not found in Ireland but is common throughout mainland Europe.

ECOLOGY The primary habitat of the field vole is rough, ungrazed grassland and young forest plantations. However, it may be found in woodland, hedgerows, bogs, dunes, moorland and road verges where grass is present. The field vole is mainly nocturnal, with activity peaks at dawn and dusk. It has a herbivorous diet, feeding on leaves and stems of grasses and occasionally mosses. At high densities, this species can cause damage to grassland and young plantations of broadleaved and fruit trees. It exhibits within-year and year-to-year changes in density, with some populations exhibiting three- to five-year multi-annual cycles.

IDENTIFICATION The field vole has a blunt, rounded nose, small furry ears, small eyes and a short tail. The head–body length is 8–13 cm and its tail is about a third of the head–body

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Common vole

Microtus arvalis (PALLAS, 1778) whether it colonised naturally before Guernsey became an island at the end of the last Ice Age.

ECOLOGY The common vole in Orkney (known as the Orkney vole) is active day and night, showing short-term cycles of activity and rest with a period of roughly three hours. It is herbivorous, feeding on the fleshy parts of a wide variety of grasses. There has been no evidence of regular cycles in abundance. It is present in most habitats in Orkney, but largely absent from improved grasslands and arable crops. Agricultural areas – ditches, fencelines and road verges – are important habitats. The Orkney vole is an important prey for hen harriers, shorteared owls and kestrels. In Guernsey, the common vole is more common in wet meadows than in drier areas.

IDENTIFICATION Unmistakable since these are the only voles present on Guernsey and the Orkney islands.

BIBLIOGRAPHY SHANE AUSTIN

DISTRIBUTION The common vole has a wide continental range from Spain to the Caucasus and central Siberia. In the British Isles, the species is restricted to Guernsey (M. arvalis sarnius) and to eight of the Orkney Islands (M. arvalis orcadensis): Mainland, Westray, Eday (introduced from Westray in 1987–88), Sanday, Burray, Hunda, South Ronaldsay and Rousay. The common vole was possibly present on Shapinsay until 1906. Neolithic remains from Holm of Westray might indicate an extinct population or transport of kills by raptors. Records of the species have been gathered from sightings, trapping, field signs and raptor pellets. Recent comparative genetic studies of Orcadian and mainland European populations are consistent with a human introduction in the Neolithic period, at least 5,100 years bp (confirmed by radiocarbon dating), from coastal regions of what is now Belgium. It is not clear whether the Guernsey vole was introduced, or

BOOTH, C. & BOOTH, J. 1994. The Mammals of Orkney: A Status with an Appendix on Amphibians and Reptiles. Kirkwall, Orkney: The Orcadian. MARTÍNKOVÁ, N., BARNETT, R., CUCCHI, T., STRUCHEN, R., PASCAL, M., PASCAL, M., FISCHER, M.C., HIGHAM, T., BRACE, S., HO, S.Y., QUÉRÉ, J.P., O’HIGGINS, P., EXCOFFIER, L., HECKEL, G., HOELZEL, A.R., DOBNEY, K.M. & SEARLE, J.B. 2013. Divergent evolutionary processes associated with colonization of offshore islands. Molecular Ecology 22 (20): 5205–5220.

AUTHOR  Martyn Gorman

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Water vole

Arvicola amphibius (LINNAEUS, 1758) Most records are from systematic field sign surveys (national, local or countywide, and proposed development sites) supplemented by reported sightings from the public.

ECOLOGY The water vole is a herbivorous rodent, found in a wide variety of wetland habitats. It has a general preference for still or slow-flowing water, with grassy or herbaceous bankside vegetation, as well as emergent or in-channel vegetation. Terrestrial/fossorial populations are known in some areas of Scotland and parts of north-east England. It eats a variety of plant species during summer, and roots, bark, berries and stored food during winter. It is diurnal, with peak periods of activity around dawn and dusk. MATT BINSTEAD, BRITISH WILDLIFE CENTRE

DISTRIBUTION In England, the water vole is widespread in the Midlands, East Anglia and southern England, the Isle of Wight and the Isle of Sheppey. It is patchily distributed in northern England, the Welsh Borders and the rest of Wales, with significant populations in Ceredigion, Monmouthshire and Anglesey. In Scotland, the Cairngorms, parts of Caithness and Sutherland, the north-west coast, the central belt (including within urban areas of Glasgow), Dumfries and Galloway, and the Scottish Borders have large water vole populations. The species has a very restricted distribution in Devon and is absent from Cornwall, the Isles of Scilly, the Isle of Man, the Channel Islands and the Scottish islands (with the exception of terrestrial/fossorial populations on small islands in the Sound of Jura). The species has faced significant decline over the twentieth century as identified by a national survey in 1989–90. A repeat survey in 1996–97 showed a continuing contraction of range as well as a decline in numbers, attributable to predation from the American mink Neovison vison as well as to the loss and fragmentation of habitat. Reintroductions of the water vole have occurred in some areas.

IDENTIFICATION The water vole is the largest species of vole in Britain, weighing 200–350 g. It has a rounded body, blunt muzzle and small ears, and the fur is chestnut brown. The head–body length is approximately 20 cm and the tail is approximately 60% of body length. A smaller melanistic (black) form occurs in parts of Scotland and has been recorded in parts of East Anglia. It can be confused with the larger brown rat Rattus norvegicus, but the latter has a more pointed muzzle and a pinkish scaly tail, whereas the tail of the water vole is brown and furry.

BIBLIOGRAPHY STRACHAN, R. & JEFFERIES, D.J. 1993. The Water Vole Arvicola terrestris in Britain 1989–90: Its Distribution and Changing Status. London: The Vincent Wildlife Trust. STRACHAN, C., STRACHAN, R. & JEFFERIES, D. 2000. Preliminary Report on the Changes in the Water Vole Population of Britain as Shown by the National Surveys of 1989–1990 and 1996–1998. London: The Vincent Wildlife Trust.

AUTHOR  Mike Dean

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Harvest mouse

Micromys minutus (PALLAS, 1771) Most records are of the characteristic woven nest, which is easiest to find in autumn and winter. Occasionally, populations are estimated through findings in owl pellets, and harvest mice can be trapped in Longworth live traps, particularly in winter when they abandon the stalk zone and forage at ground level.

ECOLOGY The harvest mouse is the smallest rodent in the UK. It has excellent climbing ability and occupies the stalk zone within a diverse range of grasses in summer and autumn. It is found in a wide range of habitats including rough grassland, reed beds, riparian margins, roadside verges, arable field margins and wild birdseed crops. The harvest mouse has a varied diet that includes seeds, berries and insects. Populations fluctuate greatly through the year with highest mortality in late winter.

IDENTIFICATION

MATT BINSTEAD, BRITISH WILDLIFE CENTRE

DISTRIBUTION Most records of harvest mouse populations lie to the south of a line drawn from south-east Wales to the North York Moors. The harvest mouse is also present in Pembrokeshire, north Wales, Cheshire, Anglesey and the Isle of Wight. Established populations are generally considered to be absent from Scotland, though there are a few records scattered across the country. The harvest mouse is unobtrusive and is easily overlooked by inexperienced surveyors, so its current status is not fully understood. For example, surveys in Essex, Suffolk and Devon have found the harvest mouse to have a widespread distribution within these counties and this may be the case across much of its range. However, there has been a national decline in the availability of high-quality habitat for the species.

The harvest mouse is a tiny mouse with golden-russet fur, pale belly and a semi-prehensile tail. Compared with other mice, the muzzle of a harvest mouse appears blunt and its eyes and ears are relatively smaller. The head–body length is 5–7 cm, and the tail is a similar length. The woven nest, constructed from split grass leaves that are still attached to their stalks, is a reliable field sign. These can be found as aerial nests in taller grasses, but in tussock-forming grasses they are usually hidden within the clump.

BIBLIOGRAPHY DOBSON, J. 2012. The Harvest Mouse Micromys minutus Pallas, 1771 in Essex. Essex Naturalist 29: 129–135. HARRIS, S. 1979. History, distribution, status and habitat requirements of the harvest mouse (Micromys minutus) in Britain. Mammal Review 9 (4): 159–171. MEEK, M. & BULLION, S. 2012. Can the harvest mouse survive in a modern arable landscape? A Suffolk case study. British Wildlife Magazine 23 (6): 419–423.

AUTHOR  Simone Bullion

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Wood mouse

Apodemus sylvaticus (LINNAEUS, 1758) ranges but size depends on habitat, whereas breeding female ranges are smaller and generally do not overlap with those of other breeding females. The wood mouse is primarily a nocturnal species, but pregnant females may also be diurnal. It is a good climber and an important prey species for many avian and mammalian predators.

IDENTIFICATION

GARY COX

DISTRIBUTION The wood mouse is ubiquitous throughout mainland Britain and Northern Ireland. It is also present on many islands such as Anglesey, the Isle of Man and the Isle of Wight, and on most small ones, including Fair Isle, Fetlar, Foula, Mainland, West Burra, Yell (Shetland); Copinsay, Eday, Hoy, Graemsay, Lings Holm, Mainland, North Ronaldsay, Sanday, Shapinsay, Stronsay (Orkney); Barra, Caenn Ear, Benbecula, Harris, Lewis, North Uist, St Kilda, South Uist (Outer Hebrides); Canna, Colonsay, Coll, Eigg, Gigha Iona, Jura, Lismore, Muck, Mull, Raasay, Rhum, Scarba, Shuna, Skye, Tiree, Islay (Inner Hebrides); Arran, Bute, Great Cumbrae (Clyde); Alderney, Guernsey, Herm, Jersey, Jethou, Sark (Channel Islands); St Marys, Tresco (Scilly Isles); Rathlin (Antrim); Tory Island (Donegal); Brownsea, Hayling Island, Hilbre, Skomer, Walney (misc.). Records are derived from live-trapping studies of small mammals, pest control, cat kills and bird pellets. There is a scarcity of records from open mountainous areas, possibly reflecting a lack of survey effort, but the species has been recorded at 1,250 m in the Cairngorms and 1,000 m in County Kerry.

ECOLOGY The wood mouse is present in most habitats, including woods, hedgerows, gardens, grassland and even on ploughed fields. Its diet consists of nuts, seeds, insects and green plant material. Breeding males have extensive

The upper fur of the wood mouse is dark brown, while the underparts are white. There may be a yellow neck/ chest spot or streak (occasionally spreading down and across the chest, almost joining the brown upper fur on neck, thus causing difficulties distinguishing from the yellow-necked mouse Apodemus flavicollis). However, melanistic and other colour forms of wood mouse can occur. The species has protruding eyes and ears. The head–body length is 8–11 cm, and the tail is just slightly shorter. Adult weights range from 14 g to 33 g (mean 20.4 g).

BIBLIOGRAPHY FLOWERDEW, J.R. 2014. The study of British small mammals – 60 years on. Mammal News 169: 10–12. FLOWERDEW, J.R. & TATTERSALL, F.H. 2008. Wood mouse Apodemus sylvaticus. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 125–137. GURNELL, J. & FLOWERDEW, J.R. 2006. Live Trapping Small Mammals: A Practical Guide, 4th edn. London: The Mammal Society.

AUTHOR  John Flowerdew

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Yellow-necked mouse Apodemus flavicollis (MELCHIOR, 1834) ECOLOGY The yellow-necked mouse is present in mature deciduous woodland, hedgerows, gardens and orchard habitats, and it is also common in outbuildings. It is associated with ancient woodland, and with mature diverse nut/fruit-bearing trees with complex structure and much fallen timber. Its diet consists of fruits, seeds, green plant material and insects. It is a good arboreal climber with nocturnal habits. Low summer temperatures possibly limit its distribution due to their impact on tree seed masting and woodland diversity.

IDENTIFICATION

MATT BINSTEAD, BRITISH WILDLIFE CENTRE

DISTRIBUTION The yellow-necked mouse is widespread in east and south-east England and around the Welsh Marches, with only sporadic isolated records in south-west England, south-west Wales and north Norfolk. It is absent from Ireland, Scotland and northern England, with no records after the year 2000 being reported north of mid-Cheshire. Over 20 deciduous woodlands that were surveyed in northern and south-west England, west Wales and northern/central England failed to provide records in 1998. This species often occurs at low density, except for local ‘hot spots’ such as Gwent, and it is less common than the wood mouse in all but 15% of surveyed deciduous woodlands. Historical and fossil evidence suggests a range extending north to Tyne and Wear/Northumberland. The yellow-necked mouse is absent from all islands, except for a recent record from Anglesey. Records of this species come from small mammal surveys using live-trapping, cat kills, bird pellets and pest control.

The yellow-necked mouse has dark brown upper fur with cream/white underparts. There is a distinctive yellow strip or ‘collar’ on the neck touching brown on either side, often extending centrally/posteriorly, which is equally clear in juveniles with grey fur. However, there is some potential confusion with the wood mouse, which also has yellow/buff markings. In the wood mouse these vary from being completely absent, to being a spot or star, to being an anterior–posterior stripe. The yellow-necked mouse has protruding eyes and ears similar to the wood mouse. The head–body length is 9–13 cm and the tail is slightly longer than this. Adults have a body mass 1.5 times that of an adult wood mouse, reaching 45 g or more (range 20–50 g). This species is more likely than the wood mouse to emit a high-pitched ‘scream’ when it is held.

BIBLIOGRAPHY MARSH, A.C.W. & MONTGOMERY, W.I. 2008. Yellow-necked mouse Apodemus flavicollis. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 137–141. MARSH, A.C.W., POULTON, S. & HARRIS, S. 2001. The yellow-necked mouse Apodemus flavicollis in Britain: status and analysis of factors affecting distribution. Mammal Review 31 (3–4): 203–227. MONTGOMERY, W.I. 1985. Interspecific competition and comparative ecology of two congeneric species of mice. In L.M. Cook (ed.) Case Studies in Population Biology. Manchester: Manchester University Press. pp. 126–187.

AUTHOR  John Flowerdew

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House mouse

Mus musculus (LINNAEUS, 1758) IDENTIFICATION The house mouse is a dull greyish brown dorsally, with a variable amount of white spotting ventrally. It is possibly confused with juvenile Apodemus spp., particularly for records derived from pest control operations.

BIBLIOGRAPHY BERRY, R.J. & TRICKER, B.J.K. 1969. Competition and extinction: the mice of Foula, with notes on those of Fair Isle and St Kilda. Journal of Zoology 158 (2): 247–265. MEEHAN, A.P. 1984. Rats and Mice: Their Biology and Control. East Grinstead: Rentokil.

AUTHOR  Sam Berry SHUTTERSTOCK, ERNI

DISTRIBUTION The house mouse is recorded throughout most of Great Britain and Northern Ireland. However, it is less ubiqui­tous than commonly thought. For example, it is rarely recorded in open field habitats, apparently excluded by other small rodents. Although often associated with human activity, it can live independently of people if other small mammals are absent (e.g. on islands: Skokholm, May, Lunga, Faray). However, it seems unable to tolerate interference from other small mammals (most commonly Apodemus spp., as seems to have occurred on St Kilda). The population status of the species is unclear, and it is highly likely to be under-recorded. Nevertheless, changes in housing construction and maintenance practices, together with alterations to agriculture, such as the use of combine harvesters and the introduction of more biosecure storage facilities for grain, are likely to have reduced the availability of habitat and food for the species since the 1960s.

ECOLOGY The house mouse is omnivorous and eclectic in its environment. It avoids wet places and is largely nocturnal. Records come from live-trapping studies of small mammals and pest control. The species tends to be under-represented in owl pellets or carnivore diet surveys.

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Brown rat

Rattus norvegicus (BERKENHOUT, 1769)

MALCOLM WELCH

DISTRIBUTION

IDENTIFICATION

The brown rat is believed to have originated in central Asia but now has a worldwide distribution. It spread across Europe in the eighteenth century, and has been present in Great Britain and Northern Ireland since the 1720s. The brown rat is found throughout Britain except in some mountain regions in Scotland and on some smaller offshore islands, but it is highly under-recorded.

Larger than the black rat, the brown rat has a blunter nose, smaller eyes, and smaller, hair-covered ears. The head–body length is 15–27 cm, and its almost hairless pinkish tail is shorter than its body. Body colours range from brown to black, but its underparts are paler. Its presence can be detected by droppings, smear marks, footprints, trails and holes, but these may not be easy to distinguish from those of the black rat. The brown rat can swim well and can be confused with the water vole, which has a shorter, blunter nose, smaller ears and a shorter, hairy tail that is half the length of its body. However, the body of the water vole is more buoyant in the water.

ECOLOGY The brown rat is mainly nocturnal, omnivorous, and lives in colonies consisting of several smaller social groups. It is a commensal rodent but can be found in some places away from humans, such as salt marshes in coastal areas and along watercourses. The brown rat occupies a wide range of urban and agricultural habitats: buildings, warehouses, farms, barns, waterways, sewers, crop fields and field margins – especially areas with good ground cover near water. When the brown rat arrived in Britain, it largely displaced the black rat. It is regarded as a pest because it eats and contaminates stored food, damages buildings and wiring, fouls surfaces, and acts as a vector of disease.

BIBLIOGRAPHY QUY, R.J. & MACDONALD, D.W. 2008. Common rat Rattus norvegicus. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 149–155.

AUTHOR  John Gurnell

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Black rat

Rattus rattus (LINNAEUS, 1758) groups each dominated by a male. Black rats are agile climbers and are often found in lofts or roof areas. Just like the brown rat, the black rat is regarded as a pest. On islands, it can have serious impacts on seabird colonies and endemic species.

IDENTIFICATION

MATT BINSTEAD, BRITISH WILDLIFE CENTRE

DISTRIBUTION The black rat is thought to have originated in South-East Asia and has been spread to many parts of the world by seafaring humans since the post-glacial period. After being introduced to Britain, probably with the Romans (first century bc to fifth century ad), it became widespread in Britain and across Europe. However, since the eighteenth century it has been almost entirely replaced by the brown rat. On the mainland, there are occasional records in and around dockyards or harbours, accidentally introduced in ships’ merchandise (e.g. Port of Tilbury and other London docks). Most former island colonies (e.g. Lundy, Shiant Isles) have now been eradicated. Overall there has been a dramatic decline (99%) in the distribution of this species between the historical and current atlas period: there were 82 positive hectads between 1960 and 1992, 16 between 2000 and 2009 (although some records may be caused by misidentifications) and one (the Shiants, which has subsequently been eliminated) between 2010 and 2016. Nevertheless the species is not yet classified as functionally extinct in the UK owing to the lack of systematic surveys.

ECOLOGY A commensal rodent, on the mainland it is almost always found in and around buildings (e.g. warehouses, restaurants, supermarkets, department stores), but can live among rocks and cliffs on islands. The black rat is mostly nocturnal, omnivorous, and lives in social

The black rat is similar in appearance to the brown rat, but it is slightly smaller with a more slender body, larger eyes and ears, and a longer tail. The head–body length is 15–24 cm. Coat colours vary from brown to black, although three different colour morphs have been described. Droppings, smear marks, footprints, trails and holes are signs of presence, but these may not be easy to distinguish from the brown rat.

BIBLIOGRAPHY TWIGG, G.I., BUCKLE, A.P. & BULLOCK, D.J. 2008. Ship rat Rattus rattus. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 155–158.

AUTHOR  John Gurnell

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Wildcat

Felis silvestris (MILLER, 1907) on the landscape, individuals may move seasonally between upland areas in the spring and summer to lower forest habitat over winter. Home range size can vary from 1 km2 to 27 km2 depending on prey availability and time of year: males range further in the late winter breeding season. The species is largely solitary except when breeding. It tends to be more active at night and around dusk and dawn, but can also be active during the day. The diet varies markedly across its distribution: there is a preference for rabbits where they are available, and elsewhere mice and voles are primary prey items.

IDENTIFICATION WENDY ELLISON

DISTRIBUTION Although once found across Britain, the current distribution of the wildcat is restricted to Scotland. Systematic surveys in 1983–87 and 2006–08 have reported populations in Caithness and Sutherland, Easter Ross, the Cairngorms, Moray and Aberdeenshire, Morvern, Ardnamurchan and Argyll. Since 2013, no reliable records have been reported in the northern Highlands and very few records in the west outside Morvern, Ardnamurchan and Kintyre. The wildcat is extremely rare with recent population estimates indicating very low densities (0.03–0.99 per km2). Recent declines may have been driven by a number of factors including rabbit population declines, a series of unusually hard winters in 2009–11, hybridisation with domestic cats and persecution. Currently, as part of the national action plan, in situ conservation efforts are focused on five priority areas, each in the range of 200–500 km2. As part of the same action plan, there is also a conservation breeding programme for future release to supplement populations in the wild. Records are obtained from questionnaires and public sightings, but mainly from camera-trap footage.

ECOLOGY The wildcat in Scotland is typically a species of woodland edge or scrub and adjacent rough grasslands, riparian habitats and moorland fringes. Depending

The wildcat displays a striped tabby-coloured coat pattern, with a thick, blunt, black-tipped tail with clear black bands and a dorsal stripe down the back that stops at the base of the tail. It also has distinctive stripes on the nape (four thick stripes) and shoulder (two thick stripes). On average, it is a little larger than the domestic cat, although there is considerable overlap in size. The wildcat generally has a broader face than the domestic cat. Hybridisation with the domestic cat is a major threat to the wildcat in Scotland, with most, if not all, remaining wildcats having at least some domestic cat ancestry. Hybrids can show a range of intermediate coat characteristics. Genetic tests and coat marking are both indicative of the level of hybridisation and both are correlated.

BIBLIOGRAPHY DAVIS, A.R. & GRAY, D. 2010. The Distribution of Scottish Wildcats (Felis silvestris) in Scotland (2006–2008). Scottish Natural Heritage Commissioned Report No. 360. Inverness: Scottish Natural Heritage. EASTERBEE, N., HEPBURN, L.V. & JEFFERIES, D.J. 1991. Survey of the Status and Distribution of the Wildcat in Scotland, 1983–1987. Edinburgh: Nature Conservancy Council for Scotland. KITCHENER, A.C., YAMAGUCHI, N., WARD, J.M. & MACDONALD, D.W. 2005. A diagnosis for the Scottish wildcat (Felis silvestris): a tool for conservation action for a critically-endangered felid. Animal Conservation 8 (3): 223–237.

AUTHORS  Ruairidh Campbell and Jenny Bryce

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Red fox

Vulpes vulpes (LINNAEUS, 1758) resident. The red fox is more common and abundant in habitats with suitable food resources and sites for dens.

ECOLOGY The red fox is an adaptable generalist found in a variety of different habitat types. Omnivorous and opportunistic in diet, this species eats a range of small mammals, birds, insects, fruit and scavenged items. It is mainly nocturnal but can also be crepuscular and active during the day.

IDENTIFICATION The red fox is a medium-sized canid, with slender legs and typical reddish-orange coat colouration. It has white fur on its upper lip and chin, extending to the chest and underside of the body. Its large pointed ears are black on the upper side. It has a long brush-like tail, typically with a white tip, and usually has black fur on its legs. Pelage can vary in colouration and is thicker in winter.

BIBLIOGRAPHY SCOTT, D.M., BERG, M., TOLHURST, B., CHAUVENET, A., SMITH, G., NEAVES, K., LOCHHEAD, J. & BAKER, P. 2014. Changes in the distribution of red foxes (Vulpes vulpes) in urban areas in Great Britain: findings and limitations of a media-driven nationwide survey. PLoS ONE 9 (6): e99059. DANIEL TRIM

DISTRIBUTION The red fox is common and widespread throughout almost all of mainland Britain and Northern Ireland. There is a lack of records for parts of Northern Ireland, the Scottish Highlands and more remote areas, which may reflect lower occurrence in these areas and/or under-recording. The species is also widespread and common on the Isle of Wight. It is not known to occur on Scottish islands, except Skye and Harris, although occasional reports can make the status unclear, and there have been previous hoax reports from islands. The red fox is currently considered absent from the Isles of Scilly and the Channel Islands. Recently, there has been an increase in the number of urban areas across the UK reporting foxes as being

AUTHOR  Dawn Scott

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Badger

Meles meles (LINNAEUS, 1758)

PADRAIG KAVANAGH

DISTRIBUTION

IDENTIFICATION

The badger is widely distributed in all counties of England, Scotland, Wales and Northern Ireland, having recovered from a decline prior to the early 1900s. It is also present on some offshore islands, including Anglesey, Arran, Canvey, the Isle of Wight, the Isle of Sheppey and Skye. Badger density is greatest in the south and west of England and Wales, and lowest in the north of Scotland. In Northern Ireland, it appears to be evenly distributed across the country. During the most recent national survey of badgers in Northern Ireland, 75% of the surveyed 1 km2 grids contained at least one sett. In England and Wales, over 50% contained an ‘active’ sett of any type, and in Scotland, only 15% contained a sett of any type.

The badger is a large, well-built animal with a medium-​ length muzzle, short legs and tail. The face is white with black stripes from ears to muzzle; the back and sides appear grey and the underside is usually dark. The badger is unlikely to be confused with any other mammal.

ECOLOGY The badger is a terrestrial carnivore found in a wide range of rural and urban habitats, with a preference in rural areas for mixed farmland (for foraging and cover) and well-drained soils (for setts). Omnivorous and opportunistic, the badger feeds on a wide range of invertebrates (earthworms, beetle larvae), vertebrate prey (small mammals, carrion), fruit (blackberries, elder), roots and cereals. It is nocturnal.

BIBLIOGRAPHY DELAHAY, R., WILSON, G., HARRIS, S. & MACDONALD, D.W., 2008. Badger Meles meles. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 425–436. JUDGE, J., WILSON, G.J., MACARTHUR, R., DELAHAY, R.J. & McDONALD, R.A. 2014. Density and abundance of badger social groups in England and Wales in 2011–2013. Scientific Reports 4: 3809. RAINEY, E., BUTLER, A., BIERMAN, S. & ROBERTS, A.M.I. 2009. Scottish Badger Distribution Survey 2006–2009: Estimating the Distribution and Density of Badger Main Setts in Scotland. Innerleithen: Scottish Badgers and Biomathematics and Statistics Scotland. REID, N., ETHERINGTON, T.R., WILSON, G., McDONALD, R.A. & MONTGOMERY, W.I. 2008. Badger Survey of Northern Ireland 2007/08. Report prepared by Quercus and Central Science Laboratory for the Department of Agriculture & Rural Development (DARD), Northern Ireland, UK. Belfast: Department of Agriculture & Rural Development. SMAL, C. 1995. The Badger and Habitat Survey of Ireland. Dublin: Government Stationery Office.

AUTHOR  Penny Lewns

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Otter

Lutra lutra (LINNAEUS, 1758) ECOLOGY The otter is a semi-aquatic carnivore found on inland watercourses, including still water, and also along the coast. It feeds mainly on fish, amphibians and crayfish. In freshwater habitats it is mainly nocturnal, but is more diurnal when living on the coast.

IDENTIFICATION The otter is a long-bodied animal, much larger than a cat, with a stout tapering tail. It is mid- to dark brown in colour. It may be confused with the American mink, but the latter is much smaller, very dark brown in colour, and has a fluffy tail. MARK BALDWIN

DISTRIBUTION The otter is present more or less throughout the UK, having recovered from a catastrophic British decline during the 1950s to the 1970s. Systematic national surveys have been carried out a number of times since 1977. The latest national surveys show it is still recovering in parts of England with few records from the south-east and parts of the Peak District, Cheshire and Lancashire. It was found at more than 75% of survey sites in all catchment areas in Wales other than Anglesey (68%) and Mid Glamorgan (71%). The otter population is stable in many parts of Scotland. Recovery of populations continues in Strathclyde, Ayrshire, Forth and the Borders, but more than 80% of sites were occupied in all counties in the latest survey. It is present throughout the whole of Ireland and is recorded on all Scottish islands of any size including the Orkneys and Shetland Isles. The otter is absent from the Isle of Man, the Isles of Scilly, Lundy and the Channel Islands. Most records are from systematic sign surveys supplemented by systematic collection of road casualties.

BIBLIOGRAPHY CRAWFORD, A. 2010. Fifth Otter Survey of England 2009–2010. Technical report. Bristol: Environment Agency. REID, N., HAYDEN, B., LUNDY, M.G., PIETRAVALLE, S., McDONALD, R.A. & MONTGOMERY, W.I. 2013. National Otter Survey of Ireland 2010/12. Irish Wildlife Manuals No. 76. Dublin: National Parks and Wildlife Service, Department of Arts, Heritage and the Gaeltacht. STRACHAN, R. 2007. National Survey of Otter Lutra lutra Distribution in Scotland 2003–04. Scottish Natural Heritage Commissioned Report No. 211 (ROAME No. F03AC309). Inverness: Scottish Natural Heritage. STRACHAN, R. 2015. Otter Survey of Wales 2009–10. Cardiff: Natural Resources Wales.

AUTHOR  Paul Chanin

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Pine marten

Martes martes (LINNAEUS, 1758) are also supplemented by road casualties, live sightings, and photos and videos from camera traps.

ECOLOGY The pine marten favours three-dimensional habitats, preferentially woodland, and dens in tree cavities, branches, wind-blown trees, squirrel dreys and bird nests. It has a varied omnivorous diet, principally comprising small mammals, fruits, birds and invertebrates. Pine martens are mostly nocturnal and crepuscular. They are solitary and mate during July–August with 1–5 young (kits) born the following March–April. PADRAIG KAVANAGH

DISTRIBUTION The pine marten has expanded its range in Scotland significantly over the past few decades, following an extensive population decline during the nineteenth century. It is now widespread in northern and central Scotland north of the industrialised central belt. It is also present on the islands of Skye and Mull. There are isolated populations in southern Scotland, in Dumfriesshire and the Scottish borders, originating from reintroductions. It has a scattered distribution in Northern Ireland. The pine marten remains absent from most of England where the population has not recovered from its historical decline. In 2015, the presence of a small number of pine martens was confirmed in Shropshire, and in 2016 footage of a pine marten was recorded in the New Forest. The origins of these animals is currently unclear though they may derive from covert releases. Elsewhere, there are occasional records from Cumbria, Northumberland, North Yorkshire and the Peak District. A pine marten population reinforcement was carried out in Wales in 2015 and 2016 by translocating pine martens from Scotland. A population is now established in mid-Wales in parts of Ceredigion and Powys, with occasional records from Carmarthenshire and Snowdonia. Most records from recent distribution surveys in Scotland are derived from scats, verified by DNA analysis, collected from systematic transect surveys. Records

IDENTIFICATION The pine marten is the size of a small domestic cat. It has a slim body with brown fur and a distinctive cream ‘bib’ on the throat and chest, which can be used to identify individuals. The pine marten has a long bushy tail, prominent rounded ears and relatively long legs.

BIBLIOGRAPHY BIRKS, J.D.S. & MESSENGER, J. 2010. Evidence of Pine Martens in England and Wales 1996–2007: Analysis of Reported Sightings and Foundations for the Future. Ledbury: The Vincent Wildlife Trust. CROOSE, E., BIRKS, J.D.S. & SCHOFIELD, H.W. 2013. Expansion Zone Survey of Pine Marten (Martes martes) Distribution in Scotland. Scottish Natural Heritage Commissioned Report No. 520. Ledbury: Scottish Natural Heritage. CROOSE, E., BIRKS, J.D.S., SCHOFIELD, H.W. & O’REILLY, C. 2014. Distribution of the Pine Marten (Martes martes) in southern Scotland in 2013. Scottish Natural Heritage Commissioned Report No. 740. Ledbury: Scottish Natural Heritage.

AUTHOR  Elizabeth Croose

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Stoat

Mustela erminea (LINNAEUS, 1758)

MIKE RAE

DISTRIBUTION The stoat is present throughout mainland Britain. Gaps in the mapped distribution of stoats here (Kintyre, southern and western Wales, western and northern Scotland) probably reflect under-recording rather than true absences. The stoat is found on more islands than the weasel, though under-recording again means that stoat populations are not well mapped on islands. It is present on Bute, Islay, Jura, the Isle of Man, Mull, Raasay, Sheppey, Skye, the Isle of Wight and Anglesey. It has been introduced to Shetland and, since 2010, to Orkney, where a project to remove it is beginning. It is also present in Northern Ireland, where it is recognised as a subspecies the Irish Stoat (M. ermina hibernica). The stoat has never been the subject of a large-scale survey and survey methodology is not well developed. Gamekeepers’ trapping records (game bags) have been used to capture major changes over time, principally the major decline in numbers during the 1950s myxomatosis outbreak that saw reduction in rabbit abundance and then subsequent recovery.

ECOLOGY The stoat is a small terrestrial carnivore that feeds mainly on lagomorphs (rabbits and hares; 65%) and small rodents (16%). The remainder of its diet is predominantly birds and bird eggs. It is a habitat generalist and is commonly found in close association with rabbit warrens, but is also frequently recorded

in diverse habitats from lowland arable farmland to marginal uplands and on moorland. Stoat population sizes are highly variable in time and space, due to high fertility when food is abundant. It exhibits intra-sexual territoriality though many individuals are transient. This species is active during the day and night.

IDENTIFICATION The stoat is small, long and thin. It is usually seen dashing or moving very rapidly, such as when crossing a road, moving or foraging with an undulating movement. It is clearly distinguished from all other British mammals apart from the weasel. The coat is sandy brown above with a creamy belly, and there is a regular margin between the back and belly. In northern Britain, its coat can turn white in winter, or become piebald brown and white. A stoat’s tail is conspicuous – approximately as long as the animal’s body – and has a marked black tip in all but the youngest juveniles. The tip of the tail is black throughout the year.

BIBLIOGRAPHY McDONALD, R.A. & KING, C.M. 2008. Stoat Mustela erminea. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 456–467.

AUTHOR  Robbie McDonald

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Weasel

Mustela nivalis (LINNAEUS, 1758) ECOLOGY The weasel is a small terrestrial carnivore and feeds mainly on small rodents (67%) and lagomorphs, predominantly small rabbits (25%). The remainder of the diet is made up of other mammals, birds and eggs. It thrives in rank grassland with large field vole populations. Weasel population sizes are highly volatile, owing to high mortality and high fertility when food is abundant. It exhibits intra-sexual territoriality though many individuals are transient. This species is active through the day and night.

IDENTIFICATION

MARK BALDWIN

DISTRIBUTION The weasel is present throughout mainland Britain. Gaps in the mapped distribution of this species on the mainland (Kintyre, Lly^ n, western and northern Scotland) probably reflect under-recording rather than true absence. Island distribution is genuinely one of presence/absence. It is recorded on Bute, the Isle of Sheppey, Skye, the Isle of Wight and Anglesey. The weasel is not found in Northern Ireland. Like the stoat, the weasel has never been the subject of a large-scale survey, and survey methodology is not well developed. Gamekeepers’ trapping records (game bags) have been used to derive trends, but are complicated by variation in trapping effort, which has changed markedly over time. Where trapping effort has been accounted for, south-west England appeared in the mid-1990s to have fewer weasel captures than other parts of Great Britain. Game bag records capture major changes over time, principally growth in numbers during the 1950s myxomatosis outbreak. At this time, there was a reduction in grazing pressure from rabbits and flourishing vole abundance. As well as directly increasing food supply, these changes may also have reduced the competition from stoats. Subsequently there was a decline up to the mid-1980s.

The weasel is very small, long and thin, roughly twice the size of a mouse. It is usually seen moving very rapidly, such as when crossing a road. It is clearly distinguished from all other British mammals apart from the stoat. It has chestnut-brown fur above with a white belly. There is an irregular margin between the back and the belly, with distinctive brown patches on the white throat. The species has a short, inconspicuous tail, lacking a black tip. By comparison the stoat’s tail is conspicuous, approximately as long as the animal’s body, and has a marked black tip in all but the youngest juveniles.

BIBLIOGRAPHY McDONALD, R.A. & KING, C.M. 2008. Weasel Mustela nivalis. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 467–476.

AUTHOR  Robbie McDonald

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Polecat

Mustela putorius (LINNAEUS, 1758) ECOLOGY The polecat occupies a wide range of habitats but is most numerous in lowland landscapes, where it dens in rabbit burrows, haystacks and log piles. Much of the polecat’s diet consists of rabbits, and it also takes small mammals, amphibians and birds. Polecats are mostly nocturnal and solitary. Mating takes place in March–May, and an average of 4–6 young (kits) are born between May and July.

IDENTIFICATION

DEREK CRAWLEY

DISTRIBUTION The polecat has undergone a considerable range expansion in England in recent decades, following a severe historical decline in the nineteenth century. It is widespread in Wales, the English Midlands and much of central and southern England, and has recently recolonised parts of south-west England and East Anglia. The species is also established in north-west England, following reintroductions in Cumbria during the 1960s to the 1980s, and is beginning to recolonise parts of western Northumberland and South Yorkshire. The polecat is absent from Northern Ireland and from most of Scotland. A population is present in Perthshire and Angus, originating from covert releases, and populations of uncertain status have been recorded in Argyll and Caithness and Sutherland. Recent records in Dumfriesshire suggest the polecat is recolonising parts of south-west Scotland from north-west England. Towards the periphery of the polecat’s core range, and in reintroduced populations, there is a higher occurrence of polecat–ferret hybrids. Feral ferrets are established on several offshore islands, including the Isle of Man, Mull, Bute, Islay, the Uists and Shetland. Most records from recent distribution surveys are derived from observations of road casualties, supplemented with live sightings and photographs and videos from camera traps.

The polecat is ferret-sized with a long, slim body, short legs and a long tail. It has dark brown-black fur with paler underfur, which gives the body a two-tone appearance. The polecat has a distinctive ‘bandit’-like mask on the face, with pale fur above the eyes and around the mouth. It may be confused with the American mink, but the latter can be distinguished by its uniformly black fur and the absence of the facial mask. Polecat–ferret hybrids can be identified from polecats by white paws, a white/cream throat patch, scattered white guard hairs over the body and a facial mask that does not reach the nose.

BIBLIOGRAPHY BIRKS, J.D.S. & KITCHENER, A.C. 1999. The Distribution and Status of the Polecat Mustela putorius in Britain in the 1990s. London: The Vincent Wildlife Trust. BIRKS, J.D.S. 2008. The Polecat Survey of Britain 2004–2006: A Report on the Polecat’s Distribution, Status and Conservation. Ledbury: The Vincent Wildlife Trust. CROOSE, E. 2016. The Distribution and Status of the Polecat (Mustela putorius) in Britain 2014–2015. Ledbury: The Vincent Wildlife Trust.

AUTHOR  Johnny Birks

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American mink

Neovison vison (SCHREBER, 1777)

DEREK CRAWLEY

DISTRIBUTION

ECOLOGY

The American mink is native to North America, but is now widespread in Britain and Ireland following escapes and releases from mink farms since the mid-twentieth century. The establishment, expansion and coalescence of feral populations was facilitated by the multiple locations of mink farms, and the initial scarcity of native competitors – the otter and the polecat (the latter is not native to Ireland). Now, the American mink is widespread and present in most counties, though it is scarce or absent in some areas such as northern Scotland. The species is present on some offshore islands such as Mull, Skye and Arran. Since the 1970s, field signs of American mink have been routinely recorded during national surveys of otters in Britain and Ireland (and, more recently, of water voles in Britain). These have provided evidence of changes in American mink distribution and, possibly, abundance. In Britain, post-2000 surveys have generally recorded a decline in the number of sites positive for American mink. For example, the fifth otter survey of England found that between 2000–02 and 2009–10 the number of survey sites positive for American mink had declined in ten out of twelve catchment areas, with six catchments recording declines of more than 50%. Naturalists also report declines in American mink abundance in some areas such as the Midlands and south-west England, and there are co-ordinated and effective control efforts over large areas of Scotland.

The American mink is a semi-aquatic generalist carnivore found on, or close to, a wide variety of freshwater bodies and some coastal habitats (e.g. sheltered, gently shelving rocky coastlines). It feeds on a wide range of prey including birds, mammals, fish, amphibians and crustaceans. Recent research suggests that the American mink in England has become more diurnal in its activity where it faces competition with otters and/or polecats.

IDENTIFICATION The American mink is a medium-sized slim-bodied mustelid, very dark brown in colour with a fluffy tail. It is much smaller than the otter and more buoyant when swimming. Paler coat colours occur occasionally and the species often has a pale patch under the chin.

BIBLIOGRAPHY CRAWFORD, A. 2010. Fifth Otter Survey of England 2009–2010. Technical report. Bristol: Environment Agency. McDONALD, R.A., O’HARA, K. & MORRISH, D.J. 2007. Decline of invasive alien mink (Mustela vison) is concurrent with recovery of native otters (Lutra lutra). Diversity and Distributions 13 (1): 92–98.

AUTHOR  Johnny Birks

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Wild boar

Sus scrofa (LINNAEUS, 1758) Groups of 2–5 reproductive females form sounders with their most recent litters and subadults from previous litters. Solitary males join the females during the main rutting period of November–January. Once a boar has reached maturity, it has a life expectancy of approximately ten years. However, mortality is high in youngsters. It has no significant predators in Britain.

IDENTIFICATION

JEFF LACK

DISTRIBUTION The wild boar was once native to Great Britain. Its history in Ireland is more contentious, with some arguing that the species is naturalised. The last freeliving wild boar in Britain were extirpated approximately 700 years ago, though various attempts at re-establishment within hunting preserves and parks over the centuries has led to confusion as to the exact date of their disappearance. It is, however, generally accepted that free-living boar were absent from the UK from the end of the sixteenth century. By 1998, there were viable populations of free-living wild boar back in the British countryside. The species is likely to have been absent from Ireland for a much longer period (up to 5,000 years bp). Populations have appeared in numerous locations; however, most of these have been temporary. The two best-known locations are on the Kent–Sussex border and in the Forest of Dean. There is also at least one population in Scotland, though the provenance and genetic signature are unknown. There are no confirmed records in Northern Ireland. Many re-emerged populations have disappeared, either through natural declines or active management.

ECOLOGY The wild boar is normally nocturnal, although in some locations it can be active during the day. It is omni­ vorous, rooting with its snout to feed on roots, rhizomes, tubers, invertebrates, small mammals and tree mast.

The wild boar is a large muscular animal with heavy-set shoulders, a large head with a long narrow snout, small erect ears, and tusks in males. Adult animals can measure over 2 m in length and are 70–90 cm high at the shoulders. They have relatively short legs and small hindquarters with a tail that is straight (15–40 cm in length) with long tassels at its end. Mature boars, both males and females, have a hair ‘mane’ down the centre line from the head to mid-back. The coat colour varies from dark grey and brown to sandy brown and black (never a mixture of colours). Young are striped, losing those stripes at 4–5 months. Breeding can occur throughout the year.

BIBLIOGRAPHY CARDEN, R.F. 2012. Review of the Natural History of Wild Boar (Sus scrofa) on the Island of Ireland. Report prepared for the Northern Ireland Environment Agency, Northern Ireland, UK, National Parks & Wildlife Service, Department of Arts, Heritage and the Gaeltacht, Dublin, Ireland and the National Museum of Ireland – Education & Outreach Department. DEFRA. 2008. Feral Wild Boar in England: An Action Plan. London: Department for Environment, Food and Rural Affairs. WILSON, C.J. 2014. The establishment and distribution of feral wild boar (Sus scrofa L.) in England. Wildlife Biology in Practice 10 (3): 1–6.

AUTHOR  John Dutton

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Red deer

Cervus elaphus (LINNAEUS, 1758) In harsher, upland habitats, or when overpopulated, females (hinds) may not breed until 3–4 years old and mature hinds may not ovulate when lactating.

IDENTIFICATION

LIZ BRACKEN

DISTRIBUTION The red deer is common in the Scottish Highlands and Islands, Dumfriesshire, the Lake District, East Anglia and the south-west of England. It is also present in parts of Northern Ireland. Further populations, largely descended from park escapes and deliberate releases, are found in the north of England, north and west Midlands, East Anglia, the New Forest, Sussex, Surrey and limited areas of Wales. Records are derived from the British Deer Society five-yearly Deer Distribution Survey (2016 data), together with other sightings, camera-trap images and road casualties.

ECOLOGY A predominantly grazing herbivore that prefers forest and woodland habitats, the red deer has adapted successfully to open moors and hills. Behaviour can vary between habitats: on open ground, large herds can form, while in woodland, deer tend to live in smaller groups or singly. In Scotland, animals tend to spend the days on the open hills and descend to lower ground at night. The sexes tend to live apart for most of the year and only come together for the rut, which takes place from the end of September to November. A single calf is born between mid-May and July; twins are very rare.

The largest British land mammal, a red deer stag can stand as high as 140 cm at the shoulder; hinds are typically at least a third smaller and lighter. Different climatic and environmental conditions lead to wide variations in species size. Southern, lowland and woodland deer tend to be notably larger than those living in the north in open environments. The summer coat is a reddish brown, turning darker and greyer in winter, with a pale caudal patch that extends above a relatively short tail. Most mature stags develop a shaggy mane as the rut approaches and carry wide, branched antlers. In wild animals, antlers may have up to 18 points (tines), including two brow tines, and sometimes more, especially in lowland or park specimens.

BIBLIOGRAPHY BRITISH DEER SOCIETY. 2017. 2016 deer distribution survey. Accessed at https://www.bds.org.uk/index.php/ news-events/297-2016-deer-distribution-survey-results​ -now-available (30 July 2019). PARLIAMENTARY OFFICE OF SCIENCE AND TECHNOLOGY. 2009. Wild deer. POSTnote No. 325. Accessed at http://researchbriefings.parliament.uk/ ResearchBriefing/Summary/POST-PN-325 (30 July 2019).

AUTHORS  Hugh Rose and Charles Smith-Jones

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Sika deer

Cervus nippon (TEMMINCK, 1838) IDENTIFICATION

SIMON JOHNSON

DISTRIBUTION The sika deer is native to Japan, where the species is considered abundant and increasing. Deliberate and accidental introduction into southern and northern England and into Scotland occurred from 1870 onwards. The species is now widespread and is expanding in Scotland, particularly in the west, while patchy populations are present across England, with locally strong presences in Dorset, Lancashire and Cumbria. The species is also present in much of Northern Ireland, particularly County Fermanagh and County Tyrone. Its presence in Wales is restricted and extremely localised. Records are derived from the British Deer Society fiveyearly Deer Distribution Survey (2016 data), together with other sightings, camera-trap images and road casualties.

ECOLOGY The sika deer is a browsing and grazing herbivore that takes a wide variety of food plants. It prefers coniferous woodlands and heathlands associated with acid soils, though other habitats are used as populations expand. The sexes tend to form separate herds for much of the year, coming together only for the rut, and are frequently encountered singly or in small groups at other times. Where they are locally numerous, large herds can form. Most activity is nocturnal or very close to the hours of darkness. The rut takes place between late September and November, and a single calf is born in the following May or June.

Similar in size to the fallow Dama dama, the sika deer stands about 100 cm high at the shoulder though males (stags) are significantly larger. The summer coat is comparable to that of the fallow deer, being chestnut brown with creamy spots and a black dorsal stripe, turning a darker grey/brown with less apparent spots in winter. The tail is shorter than that of the fallow deer and has a less distinct black stripe down the centre; the white rump patch is bordered by an indistinct black edge, which extends to the hocks. There is normally a distinct pale metatarsal gland marking on the hind legs. Males have branched antlers similar to those of red deer, but typically with no more than eight tines (points) and with a single brow tine leaving the main beam at an acute angle. The sika and red deer are capable of hybridisation. The first cross between the species has the appearance of both parents, but subsequent backcrossing makes many hybrids hard to detect in the field.

BIBLIOGRAPHY BRITISH DEER SOCIETY. 2017. 2016 deer distribution survey. Accessed at https://www.bds.org.uk/index.php/ news-events/297-2016-deer-distribution-survey-results​ -now-available (30 July 2019). HARRIS, R.B. 2015. Cervus nippon. The IUCN Red List of Threatened Species 2015: e.T41788A22155877. Accessed at http://dx.doi.org/10.2305/IUCN.UK.2015-2.RLTS. T41788A22155877.en (30 July 2019). PARLIAMENTARY OFFICE OF SCIENCE AND TECHNOLOGY. 2009. Wild deer. POSTnote No. 325. Accessed at http://researchbriefings.parliament.uk/ ResearchBriefing/Summary/POST-PN-325 (30 July 2019).

AUTHORS  Rory Putman, Josephine Pemberton and Charles Smith-Jones

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Fallow deer

Dama dama (LINNAEUS, 1758) IDENTIFICATION Similar in size to the sika deer, the fallow deer stands up to 100 cm high at the shoulder, though sexual dimorphism is apparent with the bucks being significantly larger. The fallow deer is the only deer in the UK with naturally occurring coat variations, ranging from black to white, some of which predominate according to location. The common variety is a chestnut colour with white spots in summer coat, changing to grey in winter, with a white rump bordered by a black inverted horseshoe marking. The black and white varieties are uniform in colour with no distinct rump markings. The tail is long and mobile. The buck grows large palmated antlers, which most animals shed during April. New sets of antlers are progressively larger with age. DES HAWLEY

DISTRIBUTION The fallow deer is naturalised in Great Britain and Northern Ireland. The current population is largely descended from deliberate introductions and park escapes. Now, the species is widely distributed and locally abundant across much of England and Wales, while it is patchier and more localised in Scotland and Northern Ireland. Records are derived from the British Deer Society five-yearly Deer Distribution Survey (2016 data), together with other sightings, camera-trap images and road casualties.

ECOLOGY The fallow deer is a grazing and browsing herbivore that prefers mature broadleaved woodland with understorey, open coniferous woodland and open agricultural land. It is largely diurnal but can become more nocturnal in disturbed areas. The fallow deer is a herding species, tending to spend most of the year in single-sex groups which can occasionally number as many as a hundred animals, though smaller groups are more usual. In larger populations, mature males (bucks) and females (does) usually come together only for the annual rut, which peaks in mid-October. A single fawn is born in June or July; twins are rare.

BIBLIOGRAPHY BRITISH DEER SOCIETY. 2017. 2016 deer distribution survey. Accessed at https://www.bds.org.uk/index.php/ news-events/297-2016-deer-distribution-survey-results​ -now-available (30 July 2019). PARLIAMENTARY OFFICE OF SCIENCE AND TECHNOLOGY. 2009. Wild deer. POSTnote No. 325. Accessed at http://researchbriefings.parliament.uk/ ResearchBriefing/Summary/POST-PN-325 (30 July 2019).

AUTHORS  Jochen Langbein and Charles Smith-Jones

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Roe deer

Capreolus capreolus (LINNAEUS, 1758) each other during the spring and summer. The rut takes place in late July and early August, and young are born in May and June. Twin kids are common, and triplets are occasionally recorded.

IDENTIFICATION A medium-sized deer, the roe deer is approximately 75 cm high at the shoulder. It is notably larger than the Reeves’ muntjac Muntiacus reevesi and Chinese water deer Hydropotes inermis. It is distinguished by a black nose with white spots, a white chin, no visible tail, and a white rump patch of erectile hair, which is flared when alarmed. It lacks the black rump markings usual in the fallow deer. The summer coat is a rich foxy red, whereas the coat is thicker and greyer in winter. Males typically have simple antlers with up to six tines (points); females have a short ‘tush’ of downward pointing white hair at the base of the rump patch. MIKE RAE

DISTRIBUTION The roe deer is common and widespread across most of England and Scotland, and is spreading into many parts of Wales. The species is either scarce or not present in eastern Kent and much of the Midlands. It is absent from Ireland, though there are rumours of illegal introductions. Records are derived from the British Deer Society five-yearly Deer Distribution Survey (2016 data), together with other sightings, camera-trap images and road casualties.

ECOLOGY The roe deer is a selective browsing herbivore that prefers forestry, or a mosaic of woodland and farmland, while also adapting to open hill, grassland, peri-urban and other habitats. It can be active throughout the day where undisturbed; otherwise, it is nocturnal with peak activity occurring at dawn and dusk. The species is usually seen alone or in small family groups, although larger groups can be encountered during the winter months. At other times, the species is solitary and adult males can be highly territorial and aggressive towards

BIBLIOGRAPHY BRITISH DEER SOCIETY. 2017. 2016 deer distribution survey. Accessed at https://www.bds.org.uk/index.php/ news-events/297-2016-deer-distribution-survey-results​ -now-available (30 July 2019). PARLIAMENTARY OFFICE OF SCIENCE AND TECHNOLOGY. 2009. Wild deer. POSTnote No. 325. Accessed at http://researchbriefings.parliament.uk/ ResearchBriefing/Summary/POST-PN-325 (30 July 2019).

AUTHORS  Alastair Ward and Charles Smith-Jones

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Chinese water deer Hydropotes inermis (SWINHOE, 1870)

and young are born in May and June. The Chinese water deer can give birth to up to six fawns, but generally produces two or three. Mortality can be very high during the first few weeks of life.

IDENTIFICATION

MARK BALDWIN

DISTRIBUTION The Chinese water deer is native to eastern China, the Republic of Korea and the Democratic People’s Republic of Korea, where populations are considered to be fragmented and decreasing (listed as Vulnerable on the International Union for the Conservation of Nature Red List). The UK population forms a high proportion of the global total. Deliberate and accidental introduction to locations in England occurred during the last century. The Chinese water deer is mainly found in Bedfordshire, Buckinghamshire and East Anglia, with some range expansion mainly to the west and south. There are small isolated populations elsewhere, probably attributable to escapes from collections and deliberate releases. There has been no confirmed record of wild Chinese water deer in Scotland, Wales or Northern Ireland. Records are derived from the British Deer Society five-yearly Deer Distribution Survey (2016 data), together with other sightings, camera-trap images and road casualties.

ECOLOGY A selective browsing and grazing herbivore, the Chinese water deer prefers wetland habitats but also uses parks and farmland in the UK. It is usually seen alone or in small groups, and is mostly active from dusk until dawn. The rut takes place in late November and December,

The Chinese water deer is a small deer, 50–55 cm high at the shoulder. It is slightly larger than the Reeves’ muntjac deer but is smaller than the roe deer. The coat is a uniform russet brown in summer and is paler in winter. It has large, hairy and rounded ears, and the face is said to resemble a teddy bear. The Chinese water deer has no distinctive rump markings and has a very short stubby tail. The rump is held higher than the shoulders. This species never grows antlers; instead, mature males have long, visible canine tusks, which are used for fighting with other males. It can be difficult to distinguish between the sexes in the field.

BIBLIOGRAPHY BRITISH DEER SOCIETY. 2017. 2016 deer distribution survey. Accessed at https://www.bds.org.uk/index.php/ news-events/297-2016-deer-distribution-survey-results​ -now-available (30 July 2019). HARRIS, R.B. & DUCKWORTH, J.W. 2015. Hydropotes inermis. The IUCN Red List of Threatened Species 2015: e.T10329A22163569. Accessed at http://dx.doi. org/10.2305/IUCN.UK.2015-2.RLTS.T10329A22163569. en (30 July 2019). COOKE, A.S. 2012. Chinese puzzle. Deer 16 (2): 10–14.

AUTHORS  Arnold Cooke and Charles Smith-Jones

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Reeves’ muntjac deer Muntiacus reevesi (OGILBY, 1839)

MIKE RAE

DISTRIBUTION

IDENTIFICATION

The Reeves’ muntjac deer is native to south-eastern China and Taiwan. Deliberate and accidental introduction into central England occurred during the 1890s and later. It is now present throughout most of central and southern England, extending into Wales and southwest England. Smaller, generally localised, populations exist in northern England and Northern Ireland, largely because of deliberate translocation. There are small numbers of records from Scotland, but it is thought that there is no established population there. The only island records in the UK are from the Isle of Wight. Records are derived from the British Deer Society five-yearly Deer Distribution Survey (2016 data), together with other sightings, camera-trap images and road casualties.

A small, porcine deer, the Reeves’ muntjac deer is about the size of a springer spaniel dog, with short legs and a hunched appearance. Short, simple antlers, and V-shaped markings extending up the long antler pedicles (the attachment points of the antlers), distinguish the males. Females have a diamond-shaped dark patch on the forehead. The tail is raised when alarmed to display a white underside. In comparison to Reeves’ muntjac, other small UK deer species are longer legged and appear more lightly built: the roe deer has very short pedicles and no visible tail; and the Chinese water deer has a face likened to a teddy bear and has a very short tail.

ECOLOGY The Reeves’ muntjac deer is a browsing herbivore that prefers areas of dense cover. It is usually seen alone or in small family groups. Often found in close proximity to human habitation, it is becoming increasingly common in urban areas. It is mostly nocturnal but can be diurnal in areas where it is undisturbed. Unlike other British deer species, the Reeves’ muntjac is an aseasonal breeder.

BIBLIOGRAPHY BRITISH DEER SOCIETY. 2017. 2016 deer distribution survey. Accessed at https://www.bds.org.uk/index.php/ news-events/297-2016-deer-distribution-survey-results​ -now-available (30 July 2019). TIMMINS, J & CHAN, B. 2016. Muntiacus reevesi. The IUCN Red List of Threatened Species 2016: e.T42191A22166608. Accessed at http://dx.doi.org/10.2305/IUCN.UK.2016-2. RLTS.T42191A22166608.en (30 July 2019).

AUTHOR  Charles Smith-Jones

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Greater horseshoe bat

Rhinolophus ferrumequinum (SCHREBER, 1774)

STEPHANIE COOLING-GREEN

DISTRIBUTION Following population declines in the early and mid-​ twentieth century, by the 1960s the greater horseshoe bat was restricted to south-west England and the southern coastal counties of Wales. It now shows signs of recovery with population numbers in the core of its range doubling in the past 20 years. In the same period, its distribution has expanded to include north and mid-Wales and the remaining southern counties of England. It has also spread up into southern areas of the west Midlands. Records in these areas are generally limited to individual animals, although a small number of summer colonies have also been discovered and there is evidence of some breeding. It is not recorded in Scotland or Northern Ireland.

ECOLOGY During the summer, the greater horseshoe bat forms maternity colonies, with roosts that are generally located in the attics of old buildings, although occasionally it will also breed in underground sites. During winter, this bat will hibernate in caves, mines and other underground sites. It is usually associated with landscapes of deciduous woodland with grazing pastures, in areas of the country where there are caves and mines. The species displays seasonal variation in its preferred foraging habitat. In the summer, cattle-grazed pasture is the

favoured habitat where it feeds along field margins on emerging dung beetles. During the spring and autumn, it is more associated with semi-natural ancient woodland. The greater horseshoe bat feeds preferentially on larger insect prey such as beetles or moths but will take smaller insect prey if the availability of the larger ones is limited. This species avoids flying in open or well-lit habitat, consequently commuting to its foraging grounds along linear landscape features such as well-grown hedgerows and avoiding areas with artificial lighting.

IDENTIFICATION One of the largest British bat species with a body the size of a pear, and easily identified by its horseshoe-shaped nose-leaf. It has buff-brown fur often tinged red. In torpor or hibernation, the wings are loosely wrapped around the body.

BIBLIOGRAPHY BRIGGS, P., HAWKINS, C., SHEPPARD, T. & WILSON, B. 2018. The State of the UK’s Bats 2017 (National Bat Monitoring Programme Population Trends). London: Bat Conservation Trust. DIETZ, C. & KIEFER, A. 2016. Bats of Britain and Europe. London: Bloomsbury.

AUTHOR  Henry Schofield

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Lesser horseshoe bat

Rhinolophus hipposideros (BECHSTEIN, 1800)

PATRICK WRIGHT

DISTRIBUTION The lesser horseshoe bat underwent a major decline and range contraction during the twentieth century. Populations in the south-east of England and as far north as Yorkshire were lost, and by the 1970s the species was confined to Wales, south-west England and the west Midlands. From the 1990s onwards it has started to recover, with populations more than doubling since then. Although most of this recovery in numbers has been in the areas it was restricted to in the 1970s, there are early signs of range expansion. Recently the species has been recorded in Cheshire for the first time since the early 1900s, and there is evidence of spread eastwards along the southern counties of England: small colonies have been found as far east as Buckinghamshire. It is not recorded in Scotland or Northern Ireland.

restricts it to karstic areas or to where human mining activity has created suitable sites. It feeds on small flies and moths and forages in deciduous woodland, riparian corridors and scrubby habitats. It avoids flying across open landscape by commuting along linear features such as hedgerows and treelines.

IDENTIFICATION One of the smallest British bats, the lesser horseshoe bat has a characteristic horseshoe-shaped nose-leaf and a body the size of a plum. Its size makes it easily distinguishable from the larger greater horseshoe bat. In torpor or hibernation, the wings are wrapped tightly around the body and, unlike the greater horseshoe bat, it does not form winter clusters in the UK, preferring instead to roost separately.

ECOLOGY

BIBLIOGRAPHY

The lesser horseshoe bat forms maternity roosts during the summer in the attics of old buildings and heated cellars. During winter, this species hibernates in caves, mines and other underground sites. However, in climatically mild areas in the south of its distribution, it may remain in buildings all year around. It is a lowland species, associated with pastoral landscapes with smaller field sizes and well-developed networks of hedgerows. Its requirement to hibernate underground usually

BRIGGS, P., HAWKINS, C., SHEPPARD, T. & WILSON, B. 2018. The State of the UK’s Bats 2017 (National Bat Monitoring Programme Population Trends). London: Bat Conservation Trust. DIETZ, C. & KIEFER, A. 2016. Bats of Britain and Europe. London: Bloomsbury. SCHOFIELD, H. 2008. The Lesser Horseshoe Bat Conservation Handbook. Ledbury: The Vincent Wildlife Trust.

AUTHOR  Henry Schofield

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Alcathoe bat

Myotis alcathoe (VON HELVERSEN & HELLER, 2001)

CHRISTOPHER DAMANT

DISTRIBUTION The Alcathoe bat was only confirmed in the UK in 2010 as it was previously not distinguished from whiskered Myotis mystacinus or Brandt’s Myotis brandtii bats. Therefore no historical datasets or distribution records exist. This highly cryptic species is difficult to identify and survey effort is very low, so it is likely to be under-recorded and its true distribution is likely to be larger than that currently known. Nevertheless, like populations elsewhere in Europe, its distribution appears to be patchy. Good populations are present throughout Sussex and Surrey where a number of maternity roosts have been identified. Breeding females have also been caught in Yorkshire, west Kent and Jersey. Individuals have most recently been discovered in Hampshire (2017) and at an autumn swarming site in Wiltshire (2018). It has not been recorded in Wales, Scotland or Northern Ireland. Most records are from trapping surveys, and commonly require DNA analysis to confirm species identification of animals judged likely to be Alcathoe bat on the basis of morphometric features.

woodland edges and wet woodlands, and, to a lesser extent, scrub and parkland habitats. The maternity roosts identified to date are almost exclusively located in the splits, cracks and loose bark of trees within woodlands, but one building roost is known in Surrey. It has not been recorded using bat boxes. Colonies appear to fragment into a number of small satellite roosts, and it is common for only low numbers to roost together at any one time (peak colony count of 96 individuals). Although it is captured at swarming sites, there is only one record of this species hibernating underground.

IDENTIFICATION The Alcathoe bat is the smallest of the Myotis bat species, with the whiskered and Brandt’s bats being very marginally larger. It has the typical pattern of fur colouration for Myotis bats, having pale underparts, darker fur on the upper surfaces and a pointed tragus. The Alcathoe bat can be distinguished from the whiskered or Brandt’s bat by its paler face, ears and tragus, its blunter muzzle and its shorter tragus to ear notch height.

ECOLOGY

BIBLIOGRAPHY

The Alcathoe bat appears to be a woodland specialist. It is mainly found foraging in broadleaved woodland, notably ancient and mature oak woodland, but also in

WHITBY, D. pers. obs.

AUTHOR  Daniel Whitby

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Whiskered bat

Myotis mystacinus (KUHL, 1817)

PAUL KENNEDY

DISTRIBUTION The whiskered bat is considered widespread throughout England and Wales although it is rare in East Anglia and in parts of the north-east coast. Its distribution is patchy in both Scotland and Northern Ireland, with most records coming from the southern areas. It has a wider range throughout the UK than the Brandt’s bat and is more common in southern England where the Brandt’s bat is rarer. The Brandt’s and whiskered bats cannot be distinguished using acoustic analysis and most confirmed species records are from trapping surveys, or from DNA analysis of droppings.

ECOLOGY The echolocation and morphology of the whiskered bat is very similar to the Brandt’s and Alcathoe bats. It is highly manoeuvreable in flight and has a broad dietary range, feeding as a hawker and gleaner of flies, moths and spiders. The species is caught in a wide variety of habitats and is commonly found in woodlands, orchards, riparian habitats, grassland with hedgerows or woodland edges, and scrub. However, it generally avoids arable and urban areas. Most identified maternity roosts are in buildings, but there are also roosts in bat boxes and trees. Under-recording of tree roosts is highly likely. The

whiskered bat uses swarming sites in late August and early September. It is commonly found hibernating underground and in old structures.

IDENTIFICATION As a cryptic small Myotis species, the whiskered bat is difficult to distinguish from the Alcathoe and Brandt’s bats and misidentifications are common. There are no morphological features that categorically separate these species, even in the hand, and DNA analysis is needed to definitively identify species. The whiskered bat is the most frequently encountered of the three small Myotis species, and its body size is intermediate between the Alcathoe and Brandt’s bats (though measurements overlap). It is also commonly darker in the face than the other small Myotis species, has a thin penis of uniform width, and can be distinguished on the basis of dentition.

BIBLIOGRAPHY BERGE, L. 2007. Resource partitioning between the cryptic species Brandt’s bat (Myotis brandtii) and the whiskered bat (M. mystacinus) in the UK. Unpublished PhD thesis, University of Bristol. VAUGHAN, N. 1997. The diets of British bats (Chiroptera). Mammal Review 27 (2): 77–94. WHITBY, D. pers. obs.

AUTHOR  Daniel Whitby

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Brandt’s bat

Myotis brandtii (EVERSMANN, 1845) areas. This species is frequently caught commuting along linear features and has been recorded foraging up to 3.2 km from the roost. Most identified maternity roosts are in buildings, but they also occur in bat boxes, bridges and trees. Under-recording of tree roosts is highly likely. It uses swarming sites in late August and early September, and is commonly found hibernating underground.

IDENTIFICATION

PAUL KENNEDY

DISTRIBUTION As a cryptic small Myotis species, the Brandt’s bat was only recognised as a species separate from the whiskered bat in 1970 in the UK. It is considered widespread throughout England, although is largely absent from East Anglia and patches along the north-east coast. It is more common in northern England and becomes rarer moving south, with no confirmed maternity colonies in many southern counties. The Brandt’s bat is present throughout Wales and has very recently been identified in southern Scotland. It appears to be absent elsewhere in Scotland and in Northern Ireland. The Brandt’s bat cannot be distinguished from the whiskered bat using acoustic analysis, and most confirmed species records are from trapping surveys, or from DNA analysis of droppings.

ECOLOGY The echolocation and morphological characteristics of the Brandt’s bat are very similar to those of the whiskered and Alcathoe bats. It is highly manoeuvreable in flight and has a broad dietary range, feeding as a hawker and gleaner of flies, moths and spiders. The species uses a wide range of habitats including coniferous, mixed and broadleaved woodland as well as grassland and scrub, but it generally avoids urban

The largest of the three small Myotis species, the Brandt’s bat has similar acoustic and morphological characteristics to the whiskered and Alcathoe bats. There are no morphological features that categorically separate the small Myotis species, even in the hand, and DNA analysis is regularly used to identify these species. However, the Brandt’s bat commonly has a large bulbous penis and a high protocone in the dentition (fourth premolar, upper jaw) which are the most reliable and commonly used identification features.

BIBLIOGRAPHY BERGE, L. 2007. Resource partitioning between the cryptic species Brandt’s bat (Myotis brandtii) and the whiskered bat (M. mystacinus) in the UK. Unpublished PhD thesis, University of Bristol. VAUGHAN, N. 1997. The diets of British bats (Chiroptera). Mammal Review 27 (2): 77–94. WHITBY, D. pers. obs.

AUTHOR  Daniel Whitby

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Bechstein’s bat

Myotis bechsteinii (KUHL, 1817)

CHRIS DAMANT

DISTRIBUTION The Bechstein’s bat is found in the southern part of the UK, from west Kent across southern England to east Devon. Its distribution extends northwards to Gloucestershire, Herefordshire and Worcestershire. The most northerly hectad is a lone record and is likely to be unreliable. It is found just inside south Wales (with the first maternity colony being identified in 2018) and is common throughout the Isle of Wight. It is absent from northern England, Scotland and Northern Ireland. As the Bechstein’s bat is difficult to study and catch and cannot be identified conclusively acoustically, it was historically considered exceptionally rare. Extensive studies over the last two decades, coupled with modern technology increasing survey success, have dramatically increased our know­ledge of the distribution of this species.

ECOLOGY The Bechstein’s bat is mainly associated with deciduous woodlands, particularly semi-natural ancient woodland with a high proportion of oak and a well-structured understorey. It favours foraging in wooded riparian corridors, with foraging distances being around 700 m on average, which is small compared with most UK bat species. There is evidence that the species sexually segregates, with females utilising the optimum foraging areas.

Although the Bechstein’s bat does catch insects on the wing, it is largely a gleaning species that feeds on diurnal non-volant species. Its diet mainly consists of moths and woodland-associated flies, as well as beetles and numerous woodland-associated insects. Maternity and solitary roosts are almost exclusively in woodland trees, most commonly in oak tree woodpecker and rot holes. Roosts are occasionally located in other tree species, and there is one confirmed building roost. Bat boxes can also be used for breeding animals, although this is not common and may be more likely where only less suitable habitat remains. The Bechstein’s bat is caught in low numbers swarming (peak in mid-August) at both underground and man-made structures. Few individuals are found hibernating, and both tree holes and underground sites are used.

IDENTIFICATION The Bechstein’s bat is a medium size Myotis species. It has large ears – much longer than the muzzle – which do not meet in the middle of the head. The tragus is spear-shaped, and the fur is pale on the underparts and darker on the upper surfaces. The calcar is straight and there is no fringe of hairs along the trailing edge of the tail membrane.

BIBLIOGRAPHY GREENAWAY, F. & HILL, D.A. 2004. Woodland Management Advice for Bechstein’s Bat and Barbastelle Bat. English Nature Research Reports, Report Number 658. Peterborough: English Nature. PALMER, E., PIMLEY, E., SUTTON, G. & BIRKS, J. 2013. A Study on the Population Size, Foraging Range and Roosting Ecology of Bechstein’s Bats at Grafton Wood SSSI Worcestershire. London: People’s Trust for Endangered Species and Worchester Wildlife Trust. SCHOFIELD, H. & MORRIS, C. 2000. Ranging Behaviour and Habitat Preferences of Female Zechstein’s Bat, Myotis bechsteinii (Kuhl, 1818), in Summer. With a Review of its Status, Distribution, Behaviour and Ecology in the UK. Ledbury: The Vincent Wildlife Trust.

AUTHOR  Daniel Whitby

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Daubenton’s bat

Myotis daubentonii (KUHL, 1817) and the sexes are usually segregated. The species is frequently recorded swarming at underground sites in the autumn (August), a behaviour likely to be associated with mating. The species is usually found hibernating in caves and other underground sites.

IDENTIFICATION The Daubenton’s bat is medium-sized relative to other UK bats (6–10 g). It has short ears for a Myotis species and a relatively blunt tragus and muzzle. Its dorsal fur is brown, and it has whitish grey fur on its ventral side. The calcar is straight and extends more than two-thirds along the length of the tail membrane. It also has large feet, which are covered with stiff bristles. As with other Myotis bats, it can be difficult to identify the species conclusively on the basis of acoustic records alone.

BIBLIOGRAPHY

HENRY SCHOFIELD

DISTRIBUTION The Daubenton’s bat is widespread across the UK. Its distribution appears patchier in Scotland and Northern Ireland, although it is not clear whether this represents true absence or is due to lower recording effort. Records are derived from a combination of visual surveys of bats in flight (identified by their characteristic hunting behaviour over water surfaces), roost visits, acoustic surveys, grounded animals and the capture of individuals in the field. There is no evidence of a change in distribution over time.

ECOLOGY The Daubenton’s bat usually forages low over smooth water, taking insects (mainly aquatic flies) from the air, but also ‘gaffing’ prey from the water’s surface using its large feet or tail membrane. Identified summer roosts are typically close to water, in bankside trees, buildings and bridges, but the species is highly under-recorded. Roost switching is common during the breeding season,

ANGELL, R.L., BUTLIN, R.K. & ALTRINGHAM, J.D. 2013. Sexual segregation and flexible mating patterns in temperate bats. PLoS ONE 8 (1): e54194. SENIOR, P., BUTLIN, R.K. & ALTRINGHAM, J.D. 2005. Sex and segregation in temperate bats. Proceedings of the Royal Society B 272 (1580): 2467–2473.

AUTHOR  Anita Glover

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Greater mouse-eared bat Myotis myotis (BORKHAUSEN, 1797)

DISTRIBUTION The distribution of the greater mouse-eared bat is currently restricted to a single hectad in England, where several different hibernation sites are used by the same individual. The same area was formerly used by a hibernating population of up to 30 bats, but this reduced to one male by 1985 and this individual was recorded annually until 1990. A small hibernating population discovered in Dorset in 1956 was no longer present by 1980. There are also isolated records of two other individuals: one male recorded in Kent in the winter of 1985, which was considered a vagrant; and one old female found grounded in Bognor Regis, West Sussex, in January 2001, which is excluded because the roosting location was not identified. There are no records for Scotland, Wales or Northern Ireland.

ECOLOGY In continental Europe, maternity roosts are located in large roof spaces and caves, and colonies can comprise many hundreds of animals. It hibernates in tunnels and caves. Nothing is known of its foraging ecology in England; elsewhere in Europe its diet is comprised of large beetles, caterpillars, cockchafers and crickets. Because most prey are caught on the ground, it prefers to forage in deciduous woodlands with little ground vegetation, or in recently mown or grazed meadows.

IDENTIFICATION The species is the largest of all British bats. It has extremely long ears that are not joined at the base, and is unlikely to be confused with any other species.

BIBLIOGRAPHY BLACKMORE, M. 1956. An occurrence of the mouse-eared bat, Myotis myotis (Borkhausen), in England. Journal of Zoology 127 (2): 201–203. PHILLIPS, W.W.A. & BLACKMORE, M. 1970. Mouse-eared bats Myotis myotis in Sussex. Journal of Zoology 162 (4): 520–521.

AUTHOR  Fiona Mathews

HENRY SCHOFIELD

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Natterer’s bat

Myotis nattereri (KUHL, 1817) ECOLOGY The Natterer’s bat is commonly associated with trees, particularly broadleaved woodlands, tree-lined river corridors, trees in parkland, and hedgerows adjacent to pasture. It has also been observed foraging over grass and thistles on roadsides, in the open over pasture and meadows, and using mature Corsican pine plantations in Scotland. Maternity roosts are located in trees, bat boxes and buildings – largely barns, churches and old dwelling houses. Roosts tend to be close to woodlands, with most being within 500 m; however, the size of the woodland does not appear important. Winter roosts may be found in underground sites such as canal and railway tunnels, caves, mines and ice houses.

IDENTIFICATION The Natterer’s bat is a medium-sized Myotis bat with brown-grey dorsal fur and grey-white under and ventral fur. The main distinguishing characteristics are the S-shaped calcar, the conspicuous fringe of short stiff hairs along the edge of the tail membrane, and the long, sharply pointed tragus. The ears are relatively long, reaching a little beyond the tip of the muzzle if folded forwards. As with other Myotis bats, it can be difficult to identify the species conclusively on the basis of acoustic records alone.

BIBLIOGRAPHY CHRIS DAMANT

DISTRIBUTION The Natterer’s bat is widespread in the UK. Its distribution is sparser in Scotland and Northern Ireland than in England and Wales, but some under-recording is likely. It is also found on the island of Arran, the Isle of Man and the Isle of Wight. There is a single recent record from Islay. Records are derived from roost visits, grounded bats and trapping surveys.

SMITH, P.G. & RIVERS, N.M. 2008. Natterer’s bat Myotis nattereri. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 323–328. STEBBINGS, R.E. 1993. The Greywell Tunnel: An Internationally Important Haven for Bats. Newbury: English Nature.

AUTHOR  Peter Smith

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Serotine bat

Eptesicus serotinus (SCHREBER, 1774)

DANIEL HARGREAVES

DISTRIBUTION The serotine bat occurs across southern England, with scattered records in Wales. It is absent from Scotland and Northern Ireland. A high proportion of the total records are derived from acoustic surveys, and the increasing use of broadband acoustic detectors over time is likely to explain the apparent increase in occupancy of this species. Because of overlap in call parameters, acoustic records from noctule Nyctalus noctula and Leisler’s Nyctalus leisleri bats may be confused for those of the serotine bat. Other records come from roost visits, trapping surveys and grounded animals. Expert opinion suggests a decline in roost occupation in south-east England with a corresponding increase in the south-west.

ECOLOGY The serotine bat is associated with pastures and parklands. It is an early emerging species and feeds on moths, beetles and flies. Summer roosts are usually in the roofs and walls of large houses, and maternity

colonies tend to be small. The animal is quiet and so roosts in buildings are likely to be under-recorded. Few winter roosts are known, though they are occasionally found at underground sites where they are also sometimes captured during swarming.

IDENTIFICATION One of the larger British bats, the serotine bat has a rounded tragus and a dark muzzle, ears and wing membrane. A post-calcarial lobe is present and there is a free tail tip. The fur is dark and long compared with that of the noctule bat – the only likely confusion in the hand. Droppings are large and may be confused with those of greater horseshoe bats, but are oval in profile.

BIBLIOGRAPHY HUTSON, A.M. 2008. Serotine Eptesicus serotinus. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 356–360.

AUTHOR  Fiona Mathews

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Leisler’s bat

Nyctalus leisleri (KUHL, 1817) and serotine bats, and partly because it is mainly a tree-roosting species and so is less likely to be recorded than species that are regularly found in buildings. The exception is in Northern Ireland, where the species often occurs in large numbers in buildings. Records are derived from acoustic surveys, grounded animals, and bat box and other roost surveys.

ECOLOGY

PAUL KENNEDY

DISTRIBUTION The Leisler’s bat occurs widely in England, Wales and Northern Ireland, although its distribution is patchy. In Scotland its main population area is the south-west, north to the Highland boundary and including the islands of Arran and Bute. It is apparently absent in south-east Scotland where the noctule bat population is larger. Precise details of its distribution over much of Great Britain are uncertain because of confusion with the larger noctule bat, at least where records are dependent on acoustic identification (bat detectors). In Northern Ireland the situation is simpler since the noctule bat is absent. While its occurrence largely overlaps with the noctule bat, there are areas, such as Greater London, where the Leisler’s bat is the more common species. The Leisler’s bat is under-recorded, partly as a result of confusion with acoustic records of noctule

The Leisler’s bat is a forest and woodland species, but it is adaptable and can also be found in mixed rural and suburban areas. It prefers to roost and form maternity/ nursery roosts in trees. Even where it is not uncommon in buildings, such as in Northern Ireland, studies have shown that most roosts are in tree holes and crevices. Tree species used for roosting include oak, ash, beech, Scots pine and even a Norway spruce. Locations used for nursery roosts are typically rot holes resulting from damage or deformation. Hibernation is likely to be mainly in trees, but they have been found in bat boxes in the winter, as well as in tree-like sites in buildings such as behind wooden fascias. This bat is an aerial feeder, emerging early and returning late to roost, i.e. sometimes shortly before sunset and soon after sunrise. Its diet consists of a variety of insects, which varies according to habitat.

IDENTIFICATION The Leisler’s bat is a medium-sized species, which can be distinguished from the similar noctule bat by size (forearm less than 47 mm) and fur colour (darker base of fur, whereas noctule fur is one colour). Its echolocation calls, heard on a bat detector, can be separated, with experience, from the similar noctule and serotine bat calls, though there is some overlap in call parameters.

BIBLIOGRAPHY SHIEL, C.B., DUVERGÉ, P.L., SMIDDY, P. & FAIRLEY, J.S. 1998. Analysis of the diet of Leisler’s bat (Nyctalus leisleri) in Ireland with some comparative analyses from England and Germany. Journal of Zoology 246 (4): 417–425.

AUTHOR  John Haddow

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Noctule bat

Nyctalus noctula (SCHREBER, 1774) ranging from crickets and dung beetles to non-biting midges, invariably caught on the wing. Generally it is the first bat to emerge in the evening, flying high and straight towards foraging areas, which are preferentially open areas, woodlands, wetlands or grasslands.

IDENTIFICATION Britain’s largest bat (except for the greater mouse-eared bat, which has only a single known individual), with long narrow wings and a broad muzzle. The fur is a bright ginger colour, though darker just after the moult. Ears are rounded, and the tragus mushroom-shaped. This species has a distinctive musky smell, particularly the breeding males.

BIBLIOGRAPHY

STEPHANIE COOLING-GREEN

DISTRIBUTION The noctule bat is widely distributed throughout England and Wales, though it is absent from the uplands and in some poorly wooded areas. It is present in southwest Scotland (Dumfries and Galloway), with scattered records mainly south of the central belt and a few vagrants north of this. It is not present in Northern Ireland. This species has a strong association with tree roosts, which means that its distribution and density of records closely mirror the distribution of broadleaved woodlands. An increasing proportion of records come from acoustic surveys as the species has loud and characteristic echolocation calls that can be easily identified, though there is overlap in call parameters with serotine and Leisler’s bats. Other records come from grounded animals, bat box surveys and roost visits.

ECOLOGY The noctule bat is a large, fast-flying bat that is strongly associated with tree-hole roosts, though it is occasionally found in buildings. It takes a wide variety of insect prey,

DIETZ, C. & KIEFER, A. 2016. Bats of Britain and Europe. London: Bloomsbury. MACKIE, I.J. & RACEY, P.A. 2008. Noctule Nyctalus noctula. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 338–342.

AUTHOR  Tony Mitchell-Jones

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Common pipistrelle bat Pipistrellus pipistrellus (SCHREBER, 1774)

PAUL KENNEDY

DISTRIBUTION The common pipistrelle bat is present across the UK. Gaps in the distribution will usually reflect a lack of survey effort, since this bat is found across all habitats even at high altitudes (e.g. upland wind energy sites) and in highly urban environments. Records are derived from acoustic surveys, roost visits, grounded animals and capture. Time trends in distribution are difficult to assess because the species was only recently distinguished from the soprano pipistrelle bat Pipistrellus pygmaeus. Although all records used here are for P. pipistrellus sensu stricto, and those submitted with a generic term (e.g. pipistrelle bat) are excluded, older records will be a combination of common and soprano pipistrelle bats. Even now, misidentification is likely because of similarities in morphology, habitat and geographical ranges. The increasing use of acoustic detectors, coupled with extensive survey effort for built developments over the last 15 years, is likely to explain the apparent range expansion between the historical and current atlas periods.

ECOLOGY The species is found in all habitat types. It emerges from the roost early and feeds mainly on flies. Summer roosts are mainly in buildings, including barns, churches and domestic houses. Maternity colonies are large, sometimes including several hundred animals, and are noisy. Roost switching within a season is common. Few

winter roosts are known, though they are very rarely found at underground sites and in buildings.

IDENTIFICATION The common pipistrelle bat is one of the smallest British bats, weighing just 4–6 g. Like other pipistrelle species, it has a rounded tragus and short, blunt ears. The fur is mid-brown, with little differentiation between the dorsal and ventral side. A suite of characteristics including smell, nostril shape, wing veination and call profile can aid separation from the soprano pipistrelle bat. Droppings are small, and may potentially be confused with those of small Myotis bats. It can be distinguished acoustically from other pipistrelle bats reasonably reliably on the basis of the peak frequency of its call.

BIBLIOGRAPHY BARRATT, E.M., DEAVILLE, R., BURLAND, T.M., BRUFORD, M.W., JONES, G., RACEY, P.A. & WAYNE, R.K. 1997. DNA answers the call of pipistrelle bat species. Nature 387 (6629): 138–139. LINTOTT, P.R., BARLOW, K., BUNNEFELD, N., BRIGGS, P., GAJAS ROIG, C. & PARK, K.J. 2016. Differential responses of cryptic bat species to the urban landscape. Ecology and Evolution 6 (7): 2044–2052. MATHEWS, F., RICHARDSON, S.M. & HOSKEN, D.J. 2016. Understand the Risks to Bat Populations Posed by Wind Turbines – Phase 2 – WC0753. London: DEFRA.

AUTHOR  Fiona Mathews

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Soprano pipistrelle bat Pipistrellus pygmaeus (LEACH, 1825)

IDENTIFICATION The soprano pipistrelle is one of the smallest British bats (4–6 g). Like other pipistrelle species, it has a rounded tragus and short, blunt ears. The fur is mid-brown, with little differentiation between the dorsal and ventral side. A suite of characteristics including smell, nostril shape, wing veination and call profile can aid separation from the common pipistrelle bat. Droppings are small, and may potentially be confused with those of small Myotis bats. It can be distinguished acoustically from other pipistrelle bats reasonably reliably on the basis of the peak frequency of its call.

BIBLIOGRAPHY

HEATHER CAMPBELL

DISTRIBUTION The soprano pipistrelle bat is present across the UK. Gaps in the distribution will usually reflect a lack of survey effort, since this species is found even at high altitudes (e.g. upland wind energy sites) and highly urbanised habitats. Records are derived from a combination of acoustic surveys of bats in flight, roost visits, grounded animals and the capture of individuals in the field. Changes in distribution over time are difficult to assess because identification of the species occurred after the historical atlas period.

ECOLOGY The species is found in all habitat types. It emerges from the roost early and feeds mainly on flies. Summer roosts are mainly in buildings, including barns, churches and domestic houses. Maternity colonies are large, sometimes including several hundred animals, and are noisy. Roost switching within a season is common. Few winter roosts are known, though they are very rarely found at underground sites and in buildings.

BARRATT, E.M., DEAVILLE, R., BURLAND, T.M., BRUFORD, M.W., JONES, G., RACEY, P.A. & WAYNE, R.K. 1997. DNA answers the call of pipistrelle bat species. Nature 387 (6629): 138–139. LINTOTT, P.R., BUNNEFELD, N., FUENTES-MONTEMAYOR, E., MINDERMAN, J., MAYHEW, R.J., OLLEY, L. & PARK, K.J. 2014. City life makes females fussy: sex differences in habitat use of temperate bats in urban areas. Royal Society Open Science 1 (3): 140200. STONE, E., ZEALE, M.R.K., NEWSON, S.E., BROWNE, W.J., HARRIS, S. & JONES, G. 2015. Managing conflict between bats and humans: the response of soprano pipistrelles (Pipistrellus pygmaeus) to exclusion from roosts in houses. PLoS ONE 10 (8): e0131825.

AUTHOR  Fiona Mathews

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Nathusius’ pipistrelle bat Pipistrellus nathusii (KEYSERLING & BLASIUS, 1839) ECOLOGY

PAUL KENNEDY

This species is associated with riparian habitats, favouring large water bodies with associated woodlands, estuaries, and large managed gardens with ponds and lakes. It is an aerial hawker catching and eating prey in flight, with flies and moths comprising the major prey species. Maternity colonies have been found in houses, with records of colonies mixed with soprano and common pipistrelle bats. Solitary roosts have been found in bat boxes, trees and buildings, and mating roosts have been identified in bat boxes and buildings. Hibernation sites are unknown, but a few individuals have been found hibernating in log piles and behind shutters on boarded windows.

DISTRIBUTION

IDENTIFICATION

The Nathusius’ pipistrelle bat has a widespread distribution across the UK, with records occurring from Cornwall to the Scottish islands and Northern Ireland. Nevertheless, most records – which are largely derived from acoustic surveys, live-trapping studies and grounded animals – are for solitary individuals. Fewer than ten maternity colonies have been discovered in Britain. All of these are from the east coast (Kent, Norfolk and Northumberland). Northern Ireland is a stronghold for the species and stable maternity colonies have been recorded in old stone buildings and have existed for over 20 years. The Nathusius’ pipistrelle is a long-distance migrant with breeding areas in the north-east of Europe and wintering sites in the south-west. To date, six long-​ distance migration records, through ringing and recaptures, have been recorded between southern England and the European countries of Latvia, Lithuania and the Netherlands. This species migrates along coastlines, and activity peaks in these areas in the UK during September and October. There has been a significant increase in survey effort between the historical and current atlas period which may partly explain the apparent range expansion, though some true expansion is also likely.

The Nathusius’ pipistrelle is a medium-sized pipistrelle bat (5.5–11 g) that has rapid and direct flight patterns. The fur is dark brown to chestnut on the back but lighter on the belly, and extends down the tail membrane. The wings are broad, sometimes bordered with a thin pale lateral edge. The wing cell that runs from the first joint of the fifth finger to the forearm is divided with a distinct bar. Males have a bulbous penis fringed with white hair. Its echolocation calls can be distinguished fairly reliably from those of other pipistrelle bats.

BIBLIOGRAPHY BAT CONSERVATION TRUST. 2019. National Nathusius’ pipistrelle project. Accessed at https://www.bats.org.uk/ our-work/national-bat-monitoring-programme/surveys/ national-nathusius-pipistrelle-survey (30 July 2019). DIETZ, C. & KIEFER, A. 2016. Bats of Britain and Europe. London: Bloomsbury. KRÜGER, F., CLARE, E.L., SYMONDSON, W.O., KEIŠ S, O. & PĒ TERSONS, G. 2014. Diet of the insectivorous Bat Pipistrellus nathusii during autumn migration and summer residence. Molecular Ecology 23 (15): 3672–3683.

AUTHOR  Daniel Hargreaves

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Barbastelle bat

Barbastella barbastellus (SCHREBER, 1774)

CHRISTOPHER DAMANT

DISTRIBUTION The barbastelle bat is found throughout the southern half of the UK and is absent from northern England, Scotland and Northern Ireland. Records reach up to north Wales and across to Lincolnshire, with occasional older records stretching up to North Yorkshire. There are good populations throughout much of East Anglia, Sussex, Hampshire and the Isle of Wight, the south Wales border counties and much of south-west England. However, it has a low population density and it is infrequently encountered in trapping surveys. It is likely to be under-recorded throughout its range as it rarely roosts in buildings and so detection relies on specialist acoustic and trapping surveys.

ECOLOGY Maternity roosts are almost exclusively found in trees (cracks, splits, hazard beams, wounds and lose bark), particularly oaks in ancient woodland but also in individual parkland trees. The species only rarely has maternity roosts in buildings in the UK, though solitary individuals are sometimes found. The barbastelle bat usually hibernates in trees, particularly in areas with a dense understorey, so these roosts are rarely found. However, occasional individuals are recorded in

underground sites including caves, tunnels and grottos. It is caught in low numbers at swarming sites, although it is rarely found hibernating at these same locations. The barbastelle bat forages over wide areas and can use a range of habitats. During the breeding season it forages in woodlands and close to large hedgerows early in the night, but later in the night it can use floodplains, meadows and unimproved grasslands, and readily crosses open areas. Radio tracking in southern England has shown that the mean core range of females from maternity colonies is 8 km, but they can fly long distances rapidly, frequently crossing very open habitat including downland and moorland, to reach foraging areas up to 20 km away. This species is an aerial hawker that feeds almost exclusively on hearing moths, and it is likely that its extensive foraging range is linked with food availability. During cooler periods, it is commonly found foraging in riparian and old broadleaved woodland, notably oak with a good understorey.

IDENTIFICATION The barbastelle bat is a medium-sized bat impossible to confuse with other European species, so is easily identified in the hand. It has darkly coloured skin and a short ‘pug’-shaped nose, with the nostrils opening upwards. The fur is dark, often with light frosted tops on its back and a paler ventral side. The eyes appear to be set inside the base of the ears. The ears are triangular and meet in the middle of the head, a feature shared only with the long-eared bats Plecotus spp.

BIBLIOGRAPHY GREENAWAY, F. 2001. The Barbastelle in Britain. British Wildlife 12 (5): 327–334. ZEALE, M.R.K. 2011. Conservation biology of the barbastelle (Barbastella barbastellus): applications of spatial modelling, ecology and molecular analysis of diet. Unpublished PhD thesis, University of Bristol. ZEALE, M.R.K., DAVIDSON-WATTS, I. & JONES, G. 2012. Home range use and habitat selection by barbastelle bats (Barbastella barbastellus): implications for conservation. Journal of Mammalogy 93 (4): 1110–1118.

AUTHOR  Daniel Whitby

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Brown long-eared bat Plecotus auritus (LINNAEUS, 1758)

DANIEL HARGREAVES

DISTRIBUTION The brown long-eared bat is widespread across the UK. It is unclear whether the patchier distribution observed in Scotland and Northern Ireland is the result of lower recording effort or reflects true absences. Records are largely derived from roosts, where the species is identified by the presence of individuals, from droppings, or from grounded animals. Because of its quiet call, the species is recorded infrequently on bat detectors relative to its true abundance. A small number of records are derived from trapping surveys at swarming sites and other locations, and from bat box schemes. There is no evidence of a change in distribution over time.

ECOLOGY Most summer roost records in Great Britain are from buildings such as barns, churches and large roof spaces in dwelling houses. Here, this species often roosts along the ridge beam where it is readily visible, as are its droppings that accumulate beneath the roosting sites. However, the species also makes extensive use of tree roosts (as identified in radio-tracking projects) but these are severely under-recorded because of the practical difficulties of locating them. The brown long-eared bat is also one of the species most commonly encountered in bat boxes. Little is known about the hibernation sites of the species, but it is occasionally found in underground sites and in fallen or felled trees. It is caught regularly

at swarming sites, with two peak swarming activity periods, August to October and February to April. Maternity colonies are generally small. The species emerges late and forages primarily in deciduous and mixed woodlands. It preys largely on moths, beetles and other large insects.

IDENTIFICATION The brown long-eared bat is distinguished by its extremely long ears, which are not joined at the base. It is unlikely to be confused with any species except the very rare grey long-eared bat Plecotus austriacus, from which it can only be reliably separated using DNA analysis. Its droppings have a twisted appearance. The species is difficult to detect acoustically because its calls are quiet. Recordings made with broadband detectors can be identified (though not distinguished from those of the grey long-eared bat with certainty), but in some circumstances there is potential for confusion with Myotis species.

BIBLIOGRAPHY SPEAKMAN, J.R., RACEY, P.A., CATTO, C.M.C., WEBB, P.I., SWIFT, S.M. & BURNETT, A.M. 1991. Minimum summer populations and densities of bats in N.E. Scotland, near the northern borders of their distributions. Journal of Zoology 225 (2): 327–345.

AUTHOR  Fiona Mathews

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Grey long-eared bat Plecotus austriacus (FISCHER, 1829)

PATRICK WRIGHT

DISTRIBUTION There are very few confirmed records of the grey longeared bat and these are all derived from areas close to the coast in lowland areas of southern England and the Isle of Wight. The species is absent from Scotland and Northern Ireland and a single genetic record from Pembrokeshire, Wales, could not be fully confirmed. The grey long-eared bat is cryptic and cannot be separated reliably from the brown long-eared bat on the basis of acoustic records. It is possible that there is some under-recording of roosts owing to misidentification, because the brown long-eared bat is common and morphologically similar. Identification is therefore heavily reliant on DNA analysis. There are currently ten known maternity roosts. A small number of other locations used by the species have been identified through the genotyping of droppings, but it is unclear whether these are maternity sites. Follow-up surveys of roosts known historically suggest that a high proportion have been lost over recent decades.

ECOLOGY All maternity roosts are in large loft spaces of residential buildings, and no hibernation sites have been identified.

Maternity colonies are generally small. The species emerges late and feeds mainly on moths, foraging over meadows and along woodland edges. Loss of traditional meadows and wet pasture is thought to be a primary cause of the species’ decline.

IDENTIFICATION The grey long-eared bat is a cryptic species, very similar in morphology and flight pattern to the brown longeared bat, but it has a darker face and muzzle, wider tragus and shorter thumbs. Droppings can only be identified with certainty using genotyping, and acoustic records cannot be reliably distinguished from those of the brown long-eared bat.

BIBLIOGRAPHY RAZGOUR, O.N. 2012. From genes to landscapes: conservation biology of the grey long-eared bat, Plecotus austriacus, across spatio-temporal scales. Unpublished PhD thesis, University of Bristol.

AUTHOR  Fiona Mathews

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Grey seal

Halichoerus grypus (FABRICIUS, 1791) although it can feed up to several hundred kilometres offshore. Foraging trips can therefore last anywhere between one and 30 days. The grey seal spends longer periods hauled out during the annual moult (December–April) and during the breeding season (August–December). It breeds in the autumn in the UK, and females give birth to a single white-coated pup, which they suckle for 17–23 days. It has a polygynous breeding system, with dominant males monopolising access to females and mating with them as they come into oestrus after the end of lactation.

IDENTIFICATION TOM McDONNELL

DISTRIBUTION Approximately 38% of the world’s grey seals breed in the UK, and 88% of these breed at colonies in Scotland, with the main concentrations being in the Outer Hebrides and in Orkney. There are also breeding colonies in Shetland, on the north and east coasts of mainland Britain, in south-west England and Wales, Northern Ireland (particularly Strangford Lough) and the Isle of Man. Although the number of pups throughout the UK has grown steadily since the 1960s when records began, there is clear evidence that the population growth is levelling off in all areas except the central and southern North Sea where growth rates remain high.

ECOLOGY The grey seal is a large, sexually dimorphic phocid (true seal); adult males can weigh over 300 kg and live for over 20 years, while females typically weigh 150–200 kg and can live for over 30 years. It is a generalist feeder, foraging mainly on the seabed at depths of up to 100 m. It feeds on a wide variety of prey fish species including sandeels, gadoids (cod, whiting, haddock, ling) and flatfish (plaice, sole, flounder, dab). Of these, the sandeel is typically the main prey species, but the diet varies seasonally and regionally. While the grey seal forages in the open sea, it returns regularly to haul out on land to rest, moult and breed. Most foraging occurs within 100 km of a haul-out,

The pelage colour and pattern are individually variable in the grey seal, but most are shades of grey with numerous irregular, large, darker blotches and spots across the body. The head is large with an elongated, wide nose, parallel nostrils and widely separated eyes.

BIBLIOGRAPHY HAMMOND, P.S., HALL, A.J. & PRIME, J.H. 1994. The diet of grey seals around Orkney and other island and mainland sites in north-eastern Scotland. Journal of Applied Ecology 31 (2): 340–350. McCONNELL, B.J. FEDAK, M.A. LOVELL, P. & HAMMOND, P.S. 1999. Movements and foraging areas of grey seals in the North Sea. Journal of Applied Ecology 36 (4): 373–590. SCOS, 2016. Scientific Advice on Matters Related to the Management of Seal Populations: 2016. SCOS Main Advice 2016. Swindon: Scientific Committee on Seals.

AUTHOR  David Thompson

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Harbour seal

Phoca vitulina (LINNAEUS, 1758)

DEREK CRAWLEY

DISTRIBUTION Approximately 30% of the harbour seal population of Europe is found in the UK, but this proportion has declined from approximately 40% in 2002. The harbour seal is widespread around the west coast of Scotland and throughout the Hebrides and Northern Isles. It is regularly found in Northern Ireland and the largest breeding population is in Strangford Lough. On the east coast of the UK, its distribution is more restricted, with concentrations in the major estuaries of the Thames, the Wash, the Firth of Tay and the Moray Firth. Approximately 79% of the UK harbour seal population is in Scotland, with 16% in England and 5% in Northern Ireland. Major declines have now been documented in several populations around Scotland since 2000. The causes of the recorded declines are unknown, but are not thought to be linked to the 2002 phocine distemper virus epidemic. However, the Wash and eastern England were not severely affected by this disease until 2009.

ECOLOGY The harbour seal shows little sexual dimorphism with adults typically weighing 80–100 kg. It is a long-lived animal, and reaches 20–30 years of age. The species comes ashore in sheltered waters, typically on sandbanks and in estuaries, but also in rocky

areas. Most haul-out sites are used daily, although foraging trips can last for several days. The harbour seal is primarily a coastal species and normally feeds within 40–50 km of its haul-out sites. It is a generalist feeder that takes a wide variety of fish, cephalopods and crustaceans that are obtained from surface, mid-water and benthic habitats. Its diet varies seasonally and geographically but includes sandeels, gadoids, herring and sprat, flatfish, octopus and squid. It gives birth to pups in June and July. The pups are born having shed their white coat in utero and can swim almost immediately. Females come into oestrus about a month after giving birth, and mating takes place in the water with males defending underwater calling territories. The moult in August follows the pupping and mating season, and the timing of the moult varies by age and sex class.

IDENTIFICATION The harbour seal has a small, catlike head with V-shaped nostrils. The eyes are relatively large and set close together. Prominent, light-coloured vibrissae are characteristic of the species, and it has a fine, spot-patterned pelage that can vary in colour from grey to brown, with some ring-like markings and larger blotches.

BIBLIOGRAPHY BONESS, D.J., BOWEN, W.D., BUHLEIER, B.M. & MARSHALL, G.J. 2006. Mating tactics and mating system of an aquatic-​mating pinniped: the harbor seal, Phoca vitulina. Behavioural Ecology and Sociobiology 61 (1): 119–130. SCOS, 2016. Scientific Advice on Matters Related to the Management of Seal Populations: 2016. SCOS Main Advice 2016. Swindon: Scientific Committee on Seals. TOLLIT, D.J., BLACK, A.D., THOMPSON, P.M., MACKAY, A., CORPE, H.M., WILSON, B., VAN PARIJS, S.M., GRELLIER, K. & PARLANE, S. 1998. Variations in harbour seal Phoca vitulina diet and dive-depths in relation to foraging habitat. Journal of Zoology 244 (2): 209–222.

AUTHOR  David Thompson

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North Atlantic right whale Eubalaena glacialis (MÜLLER, 1776)

FERNANDO UGARTE, SEA WATCH FOUNDATION

DISTRIBUTION The North Atlantic right whale, along with its recently separated close relative, the North Pacific right whale Eubalaena japonica, are the rarest of all the great whales. With a population of c. 500 individuals it occurs almost entirely along the eastern US coast from Texas, Florida and the Gulf of Mexico north to the Gulf of St Lawrence and coasts of Nova Scotia, Newfoundland, Labrador and south Greenland. In the eastern North Atlantic, it once ranged from north-west Africa, the Azores and the Mediterranean, north to the Bay of Biscay, western Ireland, the Hebrides, Shetland, the Faroe Islands, Iceland and Svalbard. Since the 1920s, however, sightings have been sporadic, with isolated records from the Canaries, Madeira, Spain, Portugal, the UK, Ireland and Iceland. In UK and Irish waters, seven records have occurred over the last half-century: two off south-west Ireland (June 1964 and August 1970), one near Rockall Bank, 600 km west of Scotland, sighted twice (September 1974), one (of two individuals) in the northern Irish Sea (May 1979), one west of the Outer Hebrides (August 1980), one north-west of Rockall (May 2000), and one north of Shetland (July 2000). There is also a photographic record thought to be of this species in the southern North Sea off Schouwen, Zealand in the Netherlands (July 2005) and a recent sighting off Penmarc’h, French Biscay (June 2019).

between 20° and 70°N, in sea temperatures of 0–21.8°C (averaging 12.3°C).

IDENTIFICATION The species has a very stout body, 13.5–17 m in length, a massive head that forms 25% of the total body length in adults, and up to 35% in juveniles, and a narrow rostrum with highly arched jaw-line. It has very long, narrow, dark-brown, grey or black baleen plates in each side of the upper jaw, and both the head and jaws have several large white, grey or yellowish callosities, which are infested with parasites. Numerous hairs also occur on the upper jaw and the chin. Unlike all other great whales except the bowhead, it lacks a dorsal fin altogether. Instead, the dorsal surface is smooth and ridgeless. On diving, the broad, black, deeply notched tail flukes are raised above the water. The body colour is generally black, occasionally with a white belly and chin patches. The blow is diagnostic, being V-shaped as a result of the widely separated blowholes, and may reach 7 m in height.

BIBLIOGRAPHY

ECOLOGY

KRAUS, S.D. & EVANS, P.G.H. 2008. Northern right whale Eubalaena glacialis. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 658–662. KRAUS, S.D. & ROLLAND, R.M. (eds). 2007. The Urban Whale: North Atlantic Right Whales at the Crossroads. Cambridge, MA: Harvard University Press.

The North Atlantic right whale normally occurs in coastal temperate regions with high copepod abundance

AUTHORS  Peter Evans

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Bowhead whale

Balaena mysticetus (LINNAEUS, 1758)

CAROLINE WEIR

DISTRIBUTION

IDENTIFICATION

The bowhead whale is an Arctic species with a circumpolar distribution. It favours the ice edge around the Arctic Ocean and the waters surrounding Greenland and Svalbard across to Novaya Zemlya, migrating northwards in summer into the high Arctic as the ice retreats. In recent years, however, the species has been recorded much further south – in Cape Cod Bay, New England (eastern USA), as well as off the coast of Cornwall (Isles of Scilly in February 2015, Marazion in May 2016), Northern Ireland (County Down in May 2016), France (Brittany in May 2016), Belgium (Ostende in March–April 2017), the Netherlands (Vlissingen in April 2017) and Ireland (County Cork in April 2017). The European sightings could all represent the same lone individual. These extralimital records may reflect the break-up and southward drift of ice that the Arctic is experiencing.

The bowhead whale only superficially resembles its relative, the North Atlantic right whale. It has a very rotund body, 18–20 m in length, a large (up to c. 40% of its body) though relatively narrow head and a strongly arched lower jaw. Like other right whales, it is black in colour and has no dorsal fin. A distinctive feature is the prominent muscular bulge in the area of the blowhole with an obvious depression behind. It has a very thick skin and blubber layer, and the longest baleen plates of any whale. The plates are dark grey to brown or black, generally with lighter fringes, which from the front may show as a white patch in front of the lower jaw. The flippers are large and fan-shaped with blunt tips. The tail flukes are wide and tapered at the tips with no central notch, and there is often a light grey or white band across the tailstock.

ECOLOGY

GEORGE, J.C., RUGH, D. & SUYDAM, R. 2017. Bowhead whale Balaena mysticetus. In B. Würsig, J.G.M. Thewissen & K.M. Kovacs (eds) Encyclopedia of Marine Mammals, 3rd edn. London: Academic Press. pp. 133–135.

The bowhead whale occupies Arctic and subarctic regions of the northern hemisphere, generally between 55° and 85°N.

BIBLIOGRAPHY

AUTHORS  Peter Evans

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Humpback whale

Megaptera novaeangliae (BOROWSKI, 1781) ECOLOGY The humpback whale is strongly migratory, feeding in summer in high latitudes, and mating and calving in winter in tropical waters, although a few overwinter on the feeding-grounds. It frequently may be found in coastal waters.

IDENTIFICATION

PETER EVANS, SEA WATCH FOUNDATION

DISTRIBUTION The humpback whale has a worldwide distribution in all seas, occurring even occasionally to the ice edge. The main feeding areas in the North Atlantic include the Gulf of Maine, Gulf of St Lawrence, Newfoundland/ Labrador, Greenland, Iceland and Norway. The eastern North Atlantic population migrates primarily to the West Indies, although some animals overwinter in the Cape Verde Islands. Despite fidelity to specific feeding-​ grounds, however, whales from all North Atlantic areas appear to mix spatially and genetically in the West Indies in winter. In the eastern North Atlantic it occurs around Iceland, Norway, Great Britain and Ireland. Sightings from around Britain and Ireland have increased markedly since the early 1980s, occurring in three main areas: the Northern Isles south to eastern England; the northern Irish Sea north to west Scotland; and the Celtic Sea between southern Ireland, south-west Wales and south-west England, with some sightings and strandings also being recorded in the southern North Sea. In the shelf waters of north-west Europe, it has been recorded in every month of the year, but with most sightings occurring between June and January.

One of the most distinctive of all rorquals, the humpback whale is 13–15 m in length, with a more robust body. It has a relatively short head, the top of which is flattened and covered by a number of fleshy knobs or tubercles. These extend over the lower jaw, which also has a rounded protuberance near the tip. The dorsal fin is very variable in both size and shape, ranging from a small triangular knob to a larger distinctly recurved fin, placed nearly two-thirds along the back. The species has a broad, bushy 2.5–3 m high blow. The flippers are very long, nearly one-third the total body length, scalloped with knobs or bumps along the leading edge, and white particularly on the underside. The tail flukes are very broad and distinctly notched, but are commonly scalloped with knobs along the trailing edge giving an irregular appearance. The under-surface is partially or completely white, often forming a unique pattern that enables different individuals to be recognised. It frequently breaches right out of the water, particularly in its tropical breeding grounds.

BIBLIOGRAPHY CLAPHAM, P. & EVANS, P.G.H. 2008. Humpback whale Megaptera novaeangliae. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 663–665.

AUTHORS  Peter Evans

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Blue whale

Balaenoptera musculus (LINNAEUS, 1758)

PETER EVANS, SEA WATCH FOUNDATION

DISTRIBUTION

IDENTIFICATION

The blue whale is found worldwide in all seas. Although rare after a century of exploitation, the species occurs regularly in deep waters of the North Atlantic, from the Caribbean to the Davis Strait/southern Greenland in the west, and from the Canaries, Cape Verde islands and West Africa to Jan Mayen, Svalbard and the Barents Sea in the east. There have been no sightings in the shelf seas of Britain and Ireland. All records are from deep waters offshore between May and October. Sightings and acoustic monitoring both reveal small numbers of animals in the deep waters of the Faroe–Shetland Channel and Rockall Trough, south to the Bay of Biscay.

At 23–27 m in length, the blue whale is the largest of all mammals. The head is broad, flat and U-shaped, with a single ridge extending from a raised area forward of the blowhole towards the tip of the snout. The head and most of the body is characteristically pale bluish-grey, mottled with grey or greyish white. It has a very small dorsal fin that varies in shape from nearly triangular to moderately recurved, and is situated distinctly more than two-thirds along the back so that it is seen only just prior to a dive, and sometimes after the blow. The blow itself is tall and slender, rising vertically to a height of 9–12 m. On diving, the species lifts its tail at a slight angle, whereas in the fin Balaenoptera physalus, sei Balaenoptera borealis and minke Balaenoptera acutorostrata whales, the tail is rarely visible above the surface. The tail is broad and triangular in shape, with slender, pointed tips to the flukes and only a slight central notch.

ECOLOGY The blue whale is usually found in deep waters of 400–1,000 m in depth. It is thought to spend the winter in tropical and subtropical seas where it breeds, and then migrates to feed during summer months in cold temperate and polar waters. However, acoustic recordings in the mid-Atlantic suggest that some individuals, at least, remain at high latitudes throughout winter.

BIBLIOGRAPHY EVANS, P.G.H. 2008. Blue whale Balaenoptera musculus. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 675–678.

AUTHORS  Peter Evans

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Fin whale

Balaenoptera physalus (LINNAEUS, 1758)

PETER EVANS, SEA WATCH FOUNDATION

DISTRIBUTION

IDENTIFICATION

The fin whale has a worldwide distribution, mainly in temperate and polar seas of both hemispheres. It is uncommon but widely distributed in deep waters of the North Atlantic from Baffin Bay, Iceland and Norway, south to the Gulf of Mexico and Greater Antilles in the west and the Iberian Peninsula and Mediterranean in the east, where a separate resident population exists. Large numbers occur in the Bay of Biscay. In UK and Irish waters, the species occurs mainly along the edge of the continental shelf, in the Faroe–Shetland Channel south to the South West Approaches to the Channel, as well as along the south coast of Ireland east into the Celtic Deep within the St George’s Channel. There are only a few records in the North Sea. Although it may show seasonal latitudinal migration, remaining in polar seas only during summer, those which are further south at the latitude of the UK and Ireland have been recorded in all months of the year, though mainly between July and February.

The fin whale is the second largest of all mammals, with adults reaching a length of 17.5–24 m. Like all rorquals, it has a slender streamlined body. The head resembles that of the blue whale, but is narrower and more V-shaped. Like the blue whale, there is a single prominent ridge along the middle of the top of the head but it is not quite so flat. There is a characteristic extension of white (sometimes yellowish-tinged) on the right side to include the front baleen plates, mouth cavity and lower lip. Sometimes a light grey streak or chevron extends from here onto the top of the neck. Otherwise, the head, back and flanks are uniformly dark grey or brown. On surfacing, its blow resembles an elongated inverted cone, rising to a height of 4–6 m, followed by a long shallow roll. The dorsal fin is small, though taller than in the blue whale, much more obvious and backwards-pointing, and placed two-thirds along the back. There is a prominent ridge along the tailstock between the dorsal fin and the flukes. The flippers are long and tapered.

ECOLOGY The species is most commonly recorded in deep waters (400–2,000 m in depth) off the edge of the continental shelf. However, it is much more likely to occur over the continental slope than blue or sei whale, and in some localities (e.g. southern Ireland and St George’s Channel) it occurs in areas of less than 200 m in depth. It appears to favour localities with high topographic variation – underwater sills or ledges, upwellings and frontal zones between mixed and stratified waters.

BIBLIOGRAPHY NOTARBARTOLO DI SCIARA, G. & EVANS, P.G.H. 2008. Fin whale Balaenoptera physalus. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 669–672.

AUTHORS  Peter Evans

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Sei whale

Balaenoptera borealis (LESSON, 1828)

CAROLINE WEIR

DISTRIBUTION

ECOLOGY

The sei whale has a worldwide distribution, occurring mainly offshore in deep waters from the tropics to polar seas of both hemispheres, with seasonal latitudinal migrations. In the central North Atlantic, summering populations are concentrated in deep waters north to Iceland, with a concentration of animals observed in June just north and south-west of the Charlie Gibbs Fracture Zone (53°N) over the Mid-Atlantic Ridge. It is also seen regularly in small numbers in the Azores and Madeira. In the western North Atlantic, the species is reported in summer mainly over the Nova Scotia shelf and in Labrador, and in winter from Florida, the Gulf of Mexico and the Caribbean. In the eastern North Atlantic, it is thought to winter off north-west Africa, Spain and Portugal and in the Bay of Biscay, migrating north to summering grounds off Shetland, the Faroes, Norway and Svalbard. The sei whale is uncommon in UK and Irish waters, occurring mainly beyond the shelf edge between the Faroes and Northern Isles of Scotland, in the Rockall Trough south towards the Porcupine Bight. The species occurs occasionally in the shelf seas in the Hebrides, off Shetland and Orkney, north-east Britain, the south-west of England and western Ireland. All sighting records have been between June and October.

Appears to occur mainly in depths of 500–3,000 m.

IDENTIFICATION The sei whale is a large slender rorqual typically 12–17 m in length with a relatively slender head. It has a slightly arched forehead, similar to that of the fin whale, but rounder than that of the blue whale. A single prominent ridge occurs along the middle of the top of the head, and the tip of the upper jaw tends to be down-turned. The head, back and flanks are usually a dark steelyblack or dark grey colour. The blow of the sei whale resembles that of the fin whale but is lower and less dense, typically rising to a height of 3 m. The dorsal fin and blow tend to show almost simultaneously, and both remain in view for relatively long periods before a normally shallow dive. The dorsal fin is nearly erect and strongly recurved. It is taller than those of other large rorquals, and is placed slightly less than two-thirds along the back and thus further forward than that of the blue or fin whale.

BIBLIOGRAPHY EVANS, P.G.H. 2008. Sei whale Balaenoptera borealis. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 672–675.

AUTHORS  Peter Evans

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Minke whale

Balaenoptera acutorostrata (LACEPEDE, 1804)

PETER EVANS, SEA WATCH FOUNDATION

DISTRIBUTION

IDENTIFICATION

The minke whale has a cosmopolitan distribution from the tropics to the ice edge in both hemispheres, though more uncommon in equatorial waters. It is widespread along the Atlantic seaboard of Europe from Norway south to the southern tip of Portugal, as well as in the North Sea, although abundance is greatest in the north. Around Great Britain and Ireland, highest numbers occur off the north and west coasts of Scotland and the Hebrides, the west and south coasts of Ireland, the central part of the Irish Sea including the Celtic Deep, and in the northern and central North Sea including around the Dogger Bank. It is rare in the southernmost North Sea and eastern half of the English Channel. In the western English Channel south to south-west Portugal it is present, but uncommon, out to the edge of the continental shelf, but it is largely absent from the deeper parts of the Bay of Biscay. The species is a casual visitor to the inner Danish waters, Baltic and Mediterranean (mainly the western part). Most sightings in UK and Irish waters are between May and October.

The minke whale, at 6.5–8.5 m in length, is the smallest and commonest of the baleen whales in north-west Europe. The head is more pointed and triangular than other rorquals. The body is slimmer, and it has a relatively tall, recurved dorsal fin, nearly two-thirds of the way along the back. Only under particular atmospheric conditions is the blow seen clearly, occurring almost simultaneously with the appearance of the dorsal fin. There is a single prominent ridge along the middle of the top of the head forward of the blowhole. Occasionally there is a light chevron on the back behind the head, as in the fin whale. Otherwise the head, back and upper flanks are dark grey to black, but with areas of light grey often on the flanks, one just above and behind the flippers and the other in front of and below the dorsal fin. The slim, pointed flippers each have a distinctive transverse white band, which sometimes extends over almost the entire upper surface.

ECOLOGY In the North Atlantic, the minke whale inhabits cold temperate shelf seas. Those from high latitudes are thought to migrate southwards to winter in lower latitudes; at mid-latitudes, however, such as around Great Britain and Ireland, at least some have been recorded in every month of the year.

BIBLIOGRAPHY ANDERWALD, P., EVANS, P.G.H., HOELZEL, A.R. & PAPASTAVROU, V. 2008. Minke whale Balaenoptera acutorostrata. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 665–669.

AUTHORS  Peter Evans

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Northern bottlenose whale Hyperoodon ampullatus (FORSTER, 1770)

IDENTIFICATION

SAANA ISOJUNNO

DISTRIBUTION The northern bottlenose whale is confined to the North Atlantic from temperate to Arctic seas. Its main range extends from Baffin Island and west Greenland south to New England in the west, and from Svalbard to the southern tip of the Iberian Peninsula in the east, including around the oceanic archipelago of the Azores. It occurs casually further south (to c. 15°N), having been recorded in the Caribbean in the west and the Canaries in the east. Main regions of concentration, identified from former whaling activities, appear to be west of Norway, west of Svalbard, north of Iceland, in the Davis Strait off Labrador, off the Faroes, and in The Gully off eastern Canada. In north-west Europe, the species is seen primarily in waters exceeding 1,000 m, such as the Faroe–Shetland Channel, Rockall Trough and southern Bay of Biscay. In summer, it may move onto north-west European shelf waters, such as around the Hebrides, where most records occur between July and September.

ECOLOGY The northern bottlenose whale typically inhabits deep ocean abysses of depths greater than 500 m, with greatest numbers in cold temperate to Arctic seas.

Being a deep diver (regularly to depths greater than 800 m), and able to remain under the surface for more than 70 minutes, the northern bottlenose whale is infrequently observed. Its most distinctive feature is the bulbous forehead and short dolphin-like beak that can be seen if the animal spy-hops or breaches as it occasionally does. The species frequently approaches vessels and has also been observed to lob-tail. The body is long (7–8.5 m in females; 8–9.5 m in males), robust and cylindrical in shape. Colouration varies from chocolate-​brown to greenish-​brown above, often lighter on the flanks, lightening to buff or cream all over with age. The blow is bushy, rising to a height of about 2 m, and is slightly forward pointing. The species has a strongly recurved dorsal fin two-thirds along the back, which may lead to confusion with the minke or sei whales unless other more distinctive features such as head shape are seen. Its fin is generally more erect and hooked than that of the minke whale, whereas an adult sei whale is almost twice the length of a bottlenose whale.

BIBLIOGRAPHY HOOKER, S.K., GOWANS, S. & EVANS, P.G.H. 2008. Northern bottlenose whale Hyperoodon ampullatus. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 685–690.

AUTHORS  Peter Evans

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Cuvier’s beaked whale Ziphius cavirostris (CUVIER, 1823)

ECOLOGY The Cuvier’s beaked whale is a deep-water species (500– 3,000 m in depth) occurring mainly over continental or island slopes in warm temperate to tropical seas.

IDENTIFICATION

FRAZER COOMBER, CIMA RESEARCH FOUNDATION

DISTRIBUTION The Cuvier’s beaked whale has the most widespread distribution of the family Ziphiidae, probably occurring worldwide in tropical to warm temperate seas. Favouring deep, warm waters, Cuvier’s beaked whale is the most common beaked whale in southern Europe – off the Iberian Peninsula, in the Bay of Biscay and in the Mediterranean. It is relatively rare in the UK and Ireland (although the number of records has increased in recent years) and further north and east, with isolated records from Iceland, Sweden and the Netherlands. It is a regular inhabitant of the waters around the Azores, Madeira and the Canaries, where it is seen all year round. In the western North Atlantic, it occurs in the Caribbean, Gulf of Mexico and south-east USA. Around Britain and Ireland, there have been only a handful of sighting records in contrast to the number of sightings further south in the Bay of Biscay. On the other hand, there have been around 100 strandings on the coasts of Britain and Ireland over the last 100 years. Two-thirds of these strandings have occurred since 1963, almost all coming from the Atlantic coasts, mainly in late winter or spring. The few sightings in UK and Irish waters have been mainly in July, with none between September and December.

Although reaching lengths of 5.1–6.9 m, it is, however, difficult to observe except in calm seas. Usually it provides only a brief view of its back, and the small, triangular or hooked fin two-thirds of the way along. In some older adults the head and back of the neck are light or almost white, with darker crescent-shaped areas around the eyes. Its head is small, with a sloping or slightly bulbous forehead, short indistinct beak, and up-curved mouth line. A single pair of conical teeth is situated at the tip of the lower jaw, erupting only in adult males and then protruding forward from the mouth. It has small, narrow flippers with a pointed tip, located low down on the flanks and fitting into ‘flipper pockets’. Back colouration can be dark, rust brown, slate-grey or fawn. The head and belly are generally paler, particularly in older males. The back and sides are usually covered with linear scars and with white or cream-coloured oval blotches. Sometimes yellow diatoms may occur in patches over the body.

BIBLIOGRAPHY EVANS, P.G.H., SMEENK, C. & VAN WAEREBEEK, K. 2008. Cuvier’s beaked whale Ziphius cavirostris. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 690–692.

AUTHORS  Peter Evans

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Sowerby’s beaked whale Mesoplodon bidens (SOWERBY, 1804)

ECOLOGY The Sowerby’s beaked whale is a deep-water species inhabiting ocean basins and trenches of 500–3,000 m in depth in warm to cold temperate seas, though mostly north of 30°N.

IDENTIFICATION

JUSTIN HART

DISTRIBUTION Known only from the temperate North Atlantic, the Sowerby’s beaked whale seems to occur more commonly in European seas than in eastern North America. Its distribution is centred upon deep waters of the midand eastern North Atlantic, somewhat north of other Mesoplodon species. In northern European waters, there have been confirmed sightings south of Iceland, in the Norwegian Sea, west of Norway, around the Faroe Islands, north and west of Great Britain and Ireland, in the Channel Approaches and in the Bay of Biscay. Further south and west, it has been observed regularly around the Azores and Madeira. Most reported strandings in Europe are from Great Britain and Ireland. These have occurred mainly in the Northern Isles of Scotland and along the coast of eastern Britain, although there are a number of records also from the English Channel, western coasts of Britain and Ireland, and those European countries bordering the North Sea. Strandings within the relatively shallow North Sea, however, may reflect passive drift from further north. Live sightings in shelf seas of Britain and Ireland are very few, with most being off the edge of the continental shelf north and west of Scotland and west of Ireland. Sightings have occurred in most months of the year but with the great majority in July.

The Sowerby’s beaked whale is the most frequently recorded species of the genus Mesoplodon in European seas. However, like others of this genus, it is rarely seen alive, occurring typically in very deep waters, and usually coming to the surface only briefly. It has a long (up to 5.5 m in males; 5.1 m in females), slender tapering body that is usually dark grey in colour. It has relatively small narrow flippers often tucked into ‘flipper pockets’, and a small and slender recurved dorsal fin situated almost two-thirds along the back. Light spots and scratches scattered over the back may also be seen. The head is small with a slightly concave forehead, and a well-defined and relatively long beak. It has a protruding, moderately arched, lower jaw and, in adult males, a single pair of conspicuous teeth extrude from the middle of the beak, which distinguishes it from other related species. A prominent bulge often shows in front of the crescent-shaped blowhole.

BIBLIOGRAPHY EVANS, P.G.H., HERMAN, J.S. & KITCHENER, A.C. 2008. Sowerby’s beaked whale Mesoplodon bidens. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. p. 694.

AUTHORS  Peter Evans

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Pygmy sperm whale Kogia breviceps (BLAINVILLE, 1838)

IDENTIFICATION

PATRICK GRIFFIN, OCEANSOUNDS (MARINE MAMMAL RESEARCH & CONSERVATION)

DISTRIBUTION The pygmy sperm whale has an apparently worldwide distribution, occurring in warm waters of both hemispheres. It is rarely sighted, and information is instead largely derived from strandings, most of which have occurred on the coasts of North America. Records in the North Atlantic range from equatorial waters north to Nova Scotia in the west, and Great Britain and Ireland in the east. The species generally appears to favour more temperate waters than its cousin, the dwarf sperm whale Kogia sima. In north-west Europe, most strandings have been from Atlantic European coasts, with 16 from the UK and ten from the Republic of Ireland. There has been an increase in strandings in northern Europe since 2000, possibly reflecting increased sea temperatures in the region. Although not easy to see and identify at sea, there has also been a handful of sightings in the Bay of Biscay, and in waters around Britain and Ireland including the North Sea, the furthest north being off north-west Ireland and east Scotland. All UK sightings have been between August and November.

ECOLOGY The pygmy sperm whale typically occurs beyond the shelf edge in deep tropical, subtropical and warm temperate seas.

The pygmy sperm whale is difficult to detect except in very calm conditions, and is very poorly known. The size and position of the dorsal fin are among the more diagnostic characteristics for distinguishing it from its close relative, the dwarf sperm whale. The species has a small stout body reaching lengths of only 2.7–3.8 m. It surfaces relatively slowly, and in calm seas may lie still for a period, showing a blunt squarish head, dark, slightly humped, back, and a small recurved dorsal fin. The dorsal fin is proportionately smaller than that of the dwarf sperm whale, is low, hooked and placed a little way behind the centre of the back. The conical-​shaped head, one-sixth of its body length, becomes squarer with age. There is no beak but the species has an underslung jaw. The blowhole is situated on the top of the head as in most other toothed whales, but generally more than 10% along from the snout tip. Like the sperm whale Physeter macrocephalus and dwarf sperm whale, it has an oil-filled spermaceti organ above the skull behind a large melon of fatty tissue. There are short ill-defined grooves in the throat region, which give the superficial appearance of gills. The body colour is dark blue-grey on the back, outer margins of the flippers and upper surface of the tail flukes, lightening to pale grey on the flanks and dull white belly (sometimes with a pinkish tinge). The flippers are relatively long, wide at the base tapering to a rounded point, and are set far forward on the sides near the head. The tail has a concave trailing edge with a distinct median notch.

BIBLIOGRAPHY LEAPER, R. & EVANS, P.G.H. 2008. Pygmy sperm whale Kogia breviceps. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 683–685.

AUTHORS  Peter Evans

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Dwarf sperm whale Kogia sima (OWEN, 1866)

ROBIN BAIRD

DISTRIBUTION The dwarf sperm whale has a worldwide, largely tropical, distribution, although its range is known mainly from strandings. In the North Atlantic, it has been recorded from equatorial waters to Virginia (USA) in the west, and to Great Britain in the east. In north-west Europe, the species has been confirmed off the coast of Portugal, where there are also several sightings of unidentified Kogia species. Elsewhere, there have been just seven records – five from France, one from Spain and one from the UK. There are also two extralimital records from the Mediterranean, both from Italy. The UK record was of a live animal that came ashore in Cornwall in October 2011, but was successfully refloated.

ECOLOGY This deep-water species appears to be more restricted to tropical and subtropical seas than the pygmy sperm whale.

IDENTIFICATION The dwarf sperm whale was not recognised as a distinct species until the mid-1960s and our know­ledge of it is rudimentary. It is very similar in appearance to the

pygmy sperm whale, having a triangular or squarish head, a narrow underslung lower jaw, and a small robust body that tapers rapidly behind the dorsal fin. As with the pygmy sperm whale, the head shape and light-coloured false gill slit give it a rather shark-like appearance. It is smaller than the pygmy sperm whale, with adults generally reaching lengths of just 2.5 m. The dorsal fin is proportionately larger and more erect – typically 9–16% of the snout to fin length – and located nearer the middle of the back, giving an overall appearance more like a dolphin. The blowhole is also positioned further forward than in its close relative, generally less than 10% of the distance from the tip of the snout. There is usually a pair of short throat grooves similar to those in beaked whales. The flippers are small with bluntish tips, and are situated close to the head. It is brownish-grey in colour over the back and flanks, with a white or pinkish belly.

BIBLIOGRAPHY McALPINE, D.F. 2017. Pygmy and dwarf sperm whales Kogia breviceps and K. sima. In B. Würsig, J.G.M. Thewissen & K.M. Kovacs (eds) Encyclopedia of Marine Mammals, 3rd edn. London: Academic Press. pp. 786–788.

AUTHORS  Peter Evans

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Sperm whale

Physeter macrocephalus (LINNAEUS, 1758) they move increasingly to higher latitudes. Only large males are found at the highest latitudes, sometimes occurring even close to ice edge, but generally in the most productive deep waters.

IDENTIFICATION

PETER EVANS, SEA WATCH FOUNDATION

DISTRIBUTION The sperm whale has a worldwide distribution, occurring in deep waters of all seas. In the North Atlantic, the species is widely distributed mainly off the continental shelf, along the Mid-Atlantic Ridge and around oceanic archipelagos (Azores, Madeira, Canaries and Cape Verde islands). It can be found from Davis Strait and Newfoundland in the west, and Norway and Iceland in the east, south to breeding grounds in the Caribbean in the west, and the Iberian Peninsula, Macaronesia and the Mediterranean Sea in the east. In north-west Europe, known preferred feeding areas for males include deeper waters along the continental slope west of Portugal and north of Spain within the Bay of Biscay, west and north of Great Britain and Ireland, including the edge of the Porcupine Bank adjoining the Porcupine Seabight, the Rockall Trough and the Faroe–Shetland Channel. Most sperm whales that venture far into the North Sea tend to strand. Most sightings in waters around Britain and Ireland occur between May and September, in contrast to strandings which occur mainly between November and March.

ECOLOGY The species favours waters exceeding 200 m in depth, and usually 500–2,000 m. Females and juvenile males have a more limited range than adolescent and mature males, being confined more or less to warmer waters (generally above 15°C) between c. 45°N and c. 45°S. Young males accompany females in tropical and subtropical waters, but from the age of 14–21 years

The sperm whale is the largest of all the toothed whales, with adult males reaching lengths of 11–18 m and adult females 8–12 m. It has a very distinctive shape, its large square-shaped head occupying at least one-third of the body, and being particularly pronounced in adult males. This readily distinguishes it from the more streamlined body of rorquals. The sperm whale is also distinctive in lacking a dorsal fin, having instead a triangular dorsal hump two-thirds of the way along the body, followed by a spinal ridge. Corrugations to the skin give it a shrivelled appearance. Another diagnostic feature is the single bushy blow directed forwards at a sharp angle to the left, rising to a height of c. 1.5 m. The lower jaw is narrow and underslung, and there are 2–10 short deep throat grooves. The flippers are short and spatulate in shape. Colouration is brownish-grey to black, with white areas around the mouth and frequently on the belly. There are often scratches and scars over the head and back. When it makes a steep dive, it characteristically throws its broad, triangular and deeply notched tail flukes into the air.

BIBLIOGRAPHY GORDON, J.C.D. & EVANS, P.G.H. 2008. Sperm whale Physeter macrocephalus. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 678–683.

AUTHORS  Peter Evans

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Beluga

Delphinapterus leucas (PALLAS, 1776)

CHRISTOPHER SWANN

DISTRIBUTION The beluga has a circumpolar Arctic distribution. In the North Atlantic, the main populations occur off northeast Canada south to the Gulf of St Lawrence, around the coasts of Greenland (mainly in Baffin Bay), and in the Barents Sea including around Svalbard. In northwest Europe, the species may wander southwards west of Norway, with records from southern Scandinavia and the Baltic and North seas (Denmark, Sweden, Finland, Lithuania, Poland, Germany, the Netherlands and Belgium). There are fewer than a dozen records, mainly sightings, from Britain and Ireland, for the most part in the central and northern North Sea along the east coast of Britain between July and September.

ECOLOGY The beluga has an Arctic distribution, normally occurring at or near the ice edge. Its seasonal movements are largely dictated by the annual sea-ice cycle.

IDENTIFICATION The beluga is the only medium-sized cetacean (up to 5.5 m in length) lacking a dorsal fin and with a uniform white body colour, although the latter is only achieved in adulthood. It is unique among cetaceans in having

all, or almost all, of the neck vertebrae unfused, giving the neck great flexibility. It has a stout body with a small head and pronounced melon. The head has a slight beak, and looks unusually small compared with the body owing to the great thickness of blubber covering the thorax and abdomen. The melon is bulbous and malleable. The skin is soft, often with small transverse ridges, and is frequently scarred. Adults are pure white or (in early summer) yellowish, whereas calves are grey or grey-brown, and often blotched. Juveniles become progressively lighter with age, although females often are still light grey at sexual maturity. The flippers are short and rounded, and are progressively turned up at their tips in adulthood. The tail fluke is deeply notched and also changes shape with age, developing a lobe on each side of the trailing edge. A distinct ridge takes the place of a dorsal fin.

BIBLIOGRAPHY MARTIN, A.R. & EVANS, P.G.H. 2008. Beluga Delphinapterus leucas. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 699–702.

AUTHORS  Peter Evans

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Killer whale or Orca Orcinus orca (LINNAEUS, 1758)

ECOLOGY The species has a global distribution that almost certainly exceeds all other cetacean species. In most regions, it is uncommon but numbers are greatest in subpolar and polar waters, usually in waters exceeding 50 m in depth.

IDENTIFICATION

COLIN BIRD, SEA WATCH FOUNDATION

DISTRIBUTION The killer whale has a cosmopolitan distribution. Although most abundant at high latitudes, its North Atlantic distribution is wide and extends south to the Caribbean, the Azores, Madeira, the Canaries and the western Mediterranean. It occurs in coastal northern European waters, particularly around Iceland, the Faroe Islands and western Norway. It is rare south of Britain and Ireland although a small population inhabits the Strait of Gibraltar. The species only occasionally enters inner Danish waters and the Baltic Sea. In the waters around Britain and Ireland, it is most frequently seen in northern and western Scotland, and western Ireland; it is rare in the Irish Sea, central and southern North Sea, and English Channel. Although recorded all year round, most sightings in near-shore waters off Scotland and Ireland occur between May and September. Individuals feeding particularly on herring between Norway, Iceland and the Northern Isles do so largely from October to February, then come closer around the Northern Isles mainly between May and August where they will predate seals. These seem to be separate from the small community that inhabits Atlantic waters largely west of Britain (particularly the Hebrides) and western Ireland.

The killer whale is a robust medium-sized whale – the largest member of the family Delphinidae. Varying in size from 5.7 to 9.5 m, adult male killer whales are about 25% larger than adult females, which range from 4.5 to 6.6 m. The males develop a very tall (up to almost 2 m), triangular and erect dorsal fin, which is sometimes tilted forwards, and may even lie flat on the back of the animal. Immature animals and adult females have a smaller, more recurved dorsal fin and can only be distinguished by overall size and height of the dorsal fin. The colouration of killer whales is very striking – black on the back and sides, with a white belly extending as a backwards-pointing lobe up the flanks and less markedly around the throat, chin and undersides of the flippers. There is a white oval patch above and behind the eye, and a less distinct grey saddle on the back behind the dorsal fin, which shows up clearly when the animal surfaces. The species has a conical-shaped black head with indistinct beak, large paddle-shaped flippers and broad tail flukes with a straight or slightly convex trailing edge and tips that may curl down.

BIBLIOGRAPHY BORAN, J.R., HOELZEL, A.R. & EVANS, P.G.H. 2008. Killer whale Orcinus orca. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 743–747.

AUTHORS  Peter Evans

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False killer whale

Pseudorca crassidens (OWEN, 1846)

CHRISTOPHER SWANN

DISTRIBUTION The false killer whale has a worldwide distribution, occurring mainly offshore in deep, warm waters. In the North Atlantic, it occurs from the Equator north to Maryland (USA) in the west, and Norway, Britain and Ireland in the east. In Europe, the species occurs only occasionally north of the Bay of Biscay, although records are largely confined to a few mass strandings from the UK (in 1927, 1934 and 1935), and a handful of sightings from the South West Approaches to the English Channel, west of Ireland, the Hebrides and east of Orkney, all between June and November.

Feresa attenuata and melon-headed Peponocephala electra whales. The flippers are long, narrow and tapered, with a distinctive broad hump on the front margin near the middle of the flipper. The dorsal fin, situated slightly behind the middle of the back, is tall and recurved, although its shape can vary from rounded to sharply pointed at the tip. The false killer whale is a fast active swimmer often forming large herds and frequently approaches vessels to bow-ride although, living in deep waters mainly far from the coast, it is only occasionally encountered.

BIBLIOGRAPHY

The species inhabits mainly tropical to warm temperate seas, greater than 200 m in depth.

BORAN, J.R. & EVANS, P.G.H. 2008. False killer whale Pseudorca crassidens. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 738–740.

IDENTIFICATION

AUTHORS  Peter Evans

ECOLOGY

The false killer whale has a long slender body, 5–6 m in length. It is almost all-black in colour except for a blaze of grey on the chest between the flippers, and a grey area that is sometimes present on the sides of the head. The head is small and narrow, and tapers to overhang the lower jaw unlike the similar pygmy killer

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Long-finned pilot whale Globicephala melas (TRAILL, 1809)

MASSIMILIANO ROSSO, CIMA RESEARCH FOUNDATION

DISTRIBUTION The long-finned pilot whale is widespread in temperate regions of the world, occurring in the seas of the North Atlantic and Mediterranean, particularly in deep waters off the continental shelf, where it can form groups numbering hundreds of animals. In the north-east Atlantic, the species is common and widely distributed from the Faroe Islands and Iceland south to the Bay of Biscay and Iberian Peninsula; it is also common in the Mediterranean. In north-west Europe, the main areas that appear to be favoured include the Faroe–Shetland Channel, Rockall Trough, Porcupine Bight, the South West Approaches to the English Channel, and the northern part of the Bay of Biscay. The species is rare in the North Sea except in the northernmost sector, but regularly enters the English Channel. The species occurs in the waters around Britain and Ireland in all months of the year, but with a strong peak of records in July.

ECOLOGY The long-finned pilot whale usually occurs in deep temperate and subpolar waters of 200–3,000 m in depth (particularly around the 1,000 m isobath), seaward and along edges of the continental shelf where bottom relief is greatest. It may venture occasionally into coastal waters, entering fjords and bays. This behaviour was exploited by humans operating drive fisheries in the

Northern Isles and Hebrides up to the early twentieth century, continuing in the Faroe Islands to the present time.

IDENTIFICATION The species has a robust body with long, pointed, sickle-​ shaped flippers. Adult males can reach 5–5.9 m and females 3.8–4.8 m. The head is distinctive in shape, being rather square and bulbous, particularly in old males, and with a slightly protruding upper lip. Both the head and back are black to dark grey in colour but an anchor-shaped patch of greyish white can be seen on the chin, which is lighter in younger individuals. The dorsal fin is fairly low, long-based and situated slightly forwards of the midpoint of the back, varying from recurved in immature animals and adult females to flag-shaped in adult males. There is a thick keel on the tailstock, which is more pronounced on adult males. The tail flukes have a concave trailing edge, and are deeply notched in the centre.

BIBLIOGRAPHY BORAN, J.R., EVANS, P.G.H. & MARTIN, A.R. 2008. Longfinned pilot whale Globicephala melas. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 735–738.

AUTHORS  Peter Evans

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Risso’s dolphin

Grampus griseus (CUVIER, 1812)

MASSIMO D’INCÀ, CIMA RESEARCH FOUNDATION

DISTRIBUTION The Risso’s dolphin is uncommon but widely distributed, occurring mainly in warm temperate and subtropical seas of all oceans. In the North Atlantic, the main range of the Risso’s dolphin is from the tropics north to the Georges Bank (north-east USA) in the west and the Shetland Isles in the east. Major populations exist around the oceanic archipelagos of the Azores, Madeira and the Canaries. The species has been recorded occasionally north to Sable Island, Nova Scotia and Newfoundland in the west, and the Faroe Islands and west Norway in the east. Although not common anywhere, the major populations in northern European waters occur in the Hebrides, and the species is regular in small numbers in Shetland, Orkney and north-east Scotland, in the Irish Sea (particularly off the coasts of County Wexford, west Pembrokeshire, Anglesey and the Lly^n Peninsula, and the Isle of Man), and around western Ireland, particularly the south-west. It is rare in the North Sea but in recent years it has been seen regularly off east Scotland. In the English Channel, it occurs mainly in the western part, although it is also a regular visitor to the Channel Islands. Further south, the species is present in west France, the southern Bay of Biscay, all around the Iberian Peninsula, and in the Mediterranean Sea.

Around Britain and Ireland, it is seen in all months, but mainly between April and October.

ECOLOGY The species favours continental slope waters of 10–28°C, in depths of 200–1,200 m, although in Britain and Ireland it is frequently found over slopes of 50–100 m in depth.

IDENTIFICATION It is a large, stout dolphin, about 3.5 m in length. Although the young are dark grey in colouration, as they grow older the head, back and flanks become light grey, and even white in the more mature animals. Many scratches and scars typically show around the head and back, particularly in front of the dorsal fin. The fin is relatively tall and recurved, lightening with age, particularly along the leading edge. The blunt, rounded head with slight melon and no beak, and the long, pointed flippers, show similarities to the pilot whales.

BIBLIOGRAPHY EVANS, P.G.H. 2008. Risso’s dolphin Grampus griseus. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 740–743.

AUTHORS  Peter Evans

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Atlantic white-sided dolphin

Lagenorhynchus acutus (GRAY, 1828) ECOLOGY

The species occurs in cold waters generally at temperatures of 3–12°C, favouring the continental slope (mainly around 100–300 m in depth) and deeper waters, particularly areas of high bottom relief and around deep submarine canyons.

IDENTIFICATION

ELLIOTT HAZEN

DISTRIBUTION The Atlantic white-sided dolphin is confined to temperate and subpolar seas of the North Atlantic, mainly found offshore from south-west Greenland, Iceland and the western Barents Sea southwards to the Georges Bank (north-east USA) and south-west Ireland. However, the species occurs casually south to Virginia (USA) in the west and the Bay of Biscay (47°N) in the east. It is rare in the Irish Sea, English Channel and southern North Sea, and occurs only occasionally further east in Danish waters and the Baltic. The following appear to represent areas of relatively high density: south-west Gulf of Maine (40–42°N) and continental slope at c. 39°N; the Irminger Basin between south-east Greenland and Iceland; the Iceland Basin south and east of Iceland; the Faroe Bank Channel and Faroe–Shetland Channel; Halten Bank, west of Norway; and the Rockall Trough west of Scotland and Ireland. Over the north-west European continental shelf, it is usually recorded in groups numbering 5–50 individuals during the months of June to September. In UK and Irish waters, there is some indication that the range of the species has shifted northwards since the 1990s, possibly linked to climate change.

The species superficially resembles the white-beaked dolphin Lagenorhynchus albirostris in size, shape and markings. However, it is slightly slimmer and smaller at 2.1–2.6 m, and can best be identified by a distinctive white blaze on the flank, and behind it a yellow or ochre band. Neither extends over the dorsal surface, in contrast to the pale area on the flanks of the whitebeaked dolphin. It has a sloping forehead with a short beak, and in the centre of its black back is a slender recurved fin. The tailstock has a thickened keel particularly in males, which then narrows sharply close to the flukes.

BIBLIOGRAPHY EVANS, P.G.H. & SMEENK, C. 2008. Atlantic white-sided dolphin Leucopleura acutus. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 727–731.

AUTHORS  Peter Evans

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White-beaked dolphin

Lagenorhynchus albirostris (GRAY, 1846)

200 m isobath. In west Greenland, it occurs in much deeper waters of 300–1,000 m, and in the Barents Sea commonly at 150–200 m and 400 m depths.

IDENTIFICATION

CHIARA GIULIA BERTULLI, SEA WATCH FOUNDATION

DISTRIBUTION The white-beaked dolphin is confined to the North Atlantic, occurring from south-west and central east Greenland, Svalbard and the Barents Sea, south to around Cape Cod (USA) in the west and the Bay of Biscay in the east. It is occasionally sighted around the Iberian Peninsula. It is the most common dolphin inhabiting the cold temperate and low Arctic shelf waters of the North Atlantic and North Sea. Four principal centres of high density can be identified: the Labrador Shelf including south-west Greenland; Icelandic waters; the waters around Scotland and northeast England, including the central and northern North Sea and north-west coast of Scotland; and the narrow shelf along the Norwegian coast, extending north into the Barents and White seas. In the UK, the species is most abundant in the northern Hebrides and central and north-western North Sea, occurring also in the southern North Sea, while a small population inhabits the waters of south Devon and Cornwall. Although recorded in all months of the year, most sightings are between June and September.

ECOLOGY The species occurs in temperate and subpolar seas (generally in temperatures of 2–13°C) of the North Atlantic, including a large part of the north-west European continental shelf, mainly in waters of 50–100 m in depth, and almost entirely within the

It is a much larger, stouter species than the striped Stenella coeruleoalba or common Delphinus delphis dolphins, being 2.4–2.8 m in length, with a centrally placed, taller and more recurved dorsal fin. It has a rounded snout and short thick beak that is generally tipped light grey or white. There can often be areas of white also on the head, and generally a black thoracic patch, surrounded by lighter areas. Most characteristic is the pale grey or white area extending along the flanks and over the otherwise dark grey or black dorsal surface behind the fin. The white on the flanks can lead to confusion with the Atlantic white-sided dolphin, although the latter never has white over the back behind the dorsal fin. Juveniles have more uniform colouration, the white areas being more indistinct. The thick tailstock gradually tapers towards the slightly notched tail flukes.

BIBLIOGRAPHY EVANS, P.G.H. & SMEENK, C. 2008. White-beaked dolphin Lagenorhynchus albirostris. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 724–727. GALATIUS, A. & KINZE, C.C. 2016. Lagenorhynchus albirostris (Cetacea: Delphinidae). Mammalian Species 48 (933): 35–47.

AUTHORS  Peter Evans

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Common dolphin

Delphinus delphis (LINNAEUS, 1758)

KATRIN LOHRENGEL, SEA WATCH FOUNDATION

DISTRIBUTION

IDENTIFICATION

The common dolphin has a worldwide distribution in oceanic and shelf-edge waters of tropical, subtropical and temperate seas, occurring in both hemispheres. It is abundant and widely distributed in the eastern North Atlantic, mainly occurring in deeper waters from the Iberian Peninsula north to approximately 65°N latitude (though rare north of 62°N), west of Norway and the Faroe Islands. It occurs westwards at least to the Mid-Atlantic Ridge, and eastwards it enters the western Mediterranean, with a distinct isolated population in the Black Sea. Sightings are rare in the eastern English Channel and the southern North Sea, but are abundant in the Bay of Biscay. On the UK continental shelf, the species is common in the western English Channel and the southern Irish Sea, and further north in the Sea of Hebrides and southern part of the Minch. It is also common south and west of Ireland. In some years, the species occurs further north and east in shelf seas – in the northern Hebrides, around Shetland and Orkney, and in the northern North Sea. Since the 1990s, the species has become regular in the North Sea and even entered the Baltic. Most sightings in coastal waters are between June and October.

The common dolphin is among the smallest of the true dolphins, being around 1.7–2 m in length. It is an active, fast-moving species, frequently bow-riding boats and jumping clear of the water. The common dolphin has a slender body with a falcate to erect, centrally placed, dorsal fin. In temperate waters, it is the only dolphin species likely to be seen with a long, narrow beak – dark in colour, though sometimes tipped white. On its flanks, there is an hourglass pattern of tan or yellowish tan becoming pale grey behind the dorsal fin. This pale patch may reach the dorsal surface. Underwater, the hourglass pattern shows up very clearly, as does the creamy white belly, contrasting with the brownish black back and upper flanks.

ECOLOGY

AUTHORS  Peter Evans

In the offshore North Atlantic it seems to favour waters over 15°C and shelf-edge features at depths of 400–1,000 m between 49° and 55°N, especially between 20° and 30°W.

BIBLIOGRAPHY MURPHY, S., EVANS, P.G.H. & COLLET, A. 2008. Common dolphin Delphinus delphis. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 719–724. MURPHY, S., PINN, E.H. & JEPSON, P.D. 2013. The shortbeaked common dolphin (Delphinus delphis) in the northeastern Atlantic: distribution, ecology, management and conservation status. Oceanography and Marine Biology: An Annual Review 51: 193–280.

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Striped dolphin

Stenella coeruleoalba (MEYEN, 1833) ECOLOGY An oceanic species of warm and temperate waters (mainly 12.5–19.0°C sea surface temperatures), usually occurring well beyond the continental shelf in depths of more than 1,000 m, although it will occasionally come onto the shelf, where it can be recorded in waters of 60 m or less in depth.

IDENTIFICATION

FRAZER COOMBER, CIMA RESEARCH FOUNDATION

DISTRIBUTION The striped dolphin has a worldwide distribution, occurring mainly in tropical and warm temperate seas. The species is the most common and widely distributed delphinid in the Mediterranean. In the eastern North Atlantic, it generally occurs further offshore than the common dolphin, with highest densities in the deep waters of the western Bay of Biscay beyond the continental shelf of Spain, Portugal and France. Further north, around Great Britain and Ireland, it is an occasional visitor, recorded mainly from the south-west. However, there has been a sharp increase in records in this region since the 1980s, with the species occurring also in the northern North Sea and even straying into the western Baltic, possibly reflecting warming sea temperatures in the region. The striped dolphin occurs in the mid-Atlantic to at least 62°N, suggesting that its distribution offshore may be extended northward by the Gulf Stream. The species has also been recorded recently from Icelandic, Danish, Swedish and Norwegian waters (with sightings up to 66.5°N). Sightings in UK and Irish waters have all been between May and November, with most in July and August.

Superficially the striped dolphin resembles the common dolphin, but with a more robust body and a shorter beak (to c. 10–12 cm). It lacks yellow patches and has two distinct black stripes on the flanks, one from the eye to the flipper, and the other to the anus. A distinct groove separates the beak from the forehead. Also evident is a white or light grey V-shaped blaze, one branch of this narrowing to a point below the dorsal fin, the other extending backwards towards the tail. Adults are between 1.9 and 2.4 m in length. The colouration of the body is variable, dark grey or bluish grey on the back, with lighter grey flanks, the posterior part of which is light grey and sometimes extends upwards over the dorsal surface of the tailstock. The belly is white. The dorsal fin is slender, recurved and centrally placed. The tailstock is narrow with no obvious keel, and the dark tail flukes have a median notch in them.

BIBLIOGRAPHY EVANS, P.G.H. & COLLET, A. 2008. Striped dolphin Stenella coeruleoalba. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 715–719.

AUTHORS  Peter Evans

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Bottlenose dolphin Tursiops truncatus (MONTAGU, 1821)

ECOLOGY In coastal waters, the bottlenose dolphin often favours river estuaries, headlands or sandbanks where there is uneven bottom relief and/or strong tidal currents. Offshore, the species tends to range along the shelf edge.

IDENTIFICATION

DAVE POWELL, SEA WATCH FOUNDATION

DISTRIBUTION The bottlenose dolphin has a worldwide distribution in tropical and temperate seas in both hemispheres. In the North Atlantic, it occurs mainly from Nova Scotia in the west and the Faroe Islands in the east, southwards to the Equator and beyond. Along the Atlantic seaboard of Europe, it is locally fairly common near-shore off the coasts of Spain, Portugal, north-west France, western Ireland (particularly the Shannon Estuary and Connemara), north-east Scotland (particularly the Moray Firth south to the Firth of Forth), south-west Scotland, in the Irish Sea (particularly north and west Wales, including all of Cardigan Bay), and in the English Channel (particularly around Cornwall and the Channel Islands/Normandy coast). Smaller groups have also taken up residence at other localities – for example, around the Outer Hebridean island of Barra, and in the Inner Hebrides (Islay, Mull, Coll, Tiree and southern Skye) in west Scotland. Although present all year round, most sightings are between May and September. The species also occurs offshore in the eastern North Atlantic, especially along the shelf edge, as far north as the Faroe Islands and even Svalbard. In the Bay of Biscay, there are particularly high numbers over the outer shelf and shelf break. During the summer, some pelagic groups may enter near-shore waters around the Faroe Islands, northern and western Scotland, western Ireland, in the Bay of Biscay, and around the Iberian Peninsula.

A relatively large, stout dolphin, the bottlenose dolphin reaches adult lengths of 3–3.8 m. Unlike many other dolphin species, it lacks distinctive markings, being dark grey on the back, lighter grey on the flanks, and grading to white on the belly. When breaching backwards, it displays its white throat and belly and bottle-shaped nose. The rounded head has a distinct short beak, often tipped with white on the lower jaw. The flippers are fairly long and pointed and the centrally placed dorsal fin is tall, slender and recurved, although its size and shape can be very variable. Nicks along the edge of the dorsal fin, and scratches on the fin and back, are commonly used to identify individuals.

BIBLIOGRAPHY WILSON, B. 2008. Bottlenose dolphin Tursiops truncatus. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 709–715.

AUTHORS  Peter Evans

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Harbour porpoise

Phocoena phocoena (LINNAEUS, 1758) ECOLOGY The species is restricted to temperate and subarctic (mainly 11–14°C) seas, mainly over the continental shelf at depths of 20–200 m. Although the harbour porpoise can be found in deep waters off the edge of the European continental shelf (for example within the Faroe Bank Channel), it is comparatively rare in waters exceeding 200 m in depth. Radio-tagged porpoises from west Greenland have been tracked seasonally to the mid-Atlantic at depths of 1,000–3,000 m. In coastal waters, the species frequently uses tidal conditions for foraging.

IDENTIFICATION

PETER EVANS, SEA WATCH FOUNDATION

DISTRIBUTION The harbour porpoise is confined to the North Atlantic and North Pacific. In the North Atlantic, the species occurs mainly from central west Greenland and Novaya Zemlya in the north to North Carolina and Senegal in the south. There is a geographically isolated population in the Black Sea. It is the commonest and most widely distributed species of cetacean in northern European seas, occurring over the continental shelf from the Barents Sea and Iceland, south to the coasts of France and Spain. In the 1970s it became scarce in the southernmost North Sea, English Channel and Bay of Biscay, but the species has returned to the southernmost North Sea, English Channel and French Biscay coast since the 1990s. There is some evidence for a southward shift from the north-western to the south-western North Sea. The harbour porpoise occurs all year round in UK and Irish waters, although most sightings are made between June and September.

It is the smallest cetacean in European seas, with adults averaging a length of around 1.5 m. It only occasionally leaps clear of the water, and usual views are of a small, centrally placed triangular dorsal fin and a glimpse of the back. It has a small rotund body, and a small head with no forehead or beak. The flippers are short and rounded. Its back is dark grey with a paler patch on the flanks that extends up the sides in front of the dorsal fin. From the flippers, a dark grey line extends to the jaw-line.

BIBLIOGRAPHY EVANS, P.G.H., LOCKYER, C.H., SMEENK, C., ADDINK, M. & READ, A.J. 2008. Harbour porpoise Phocoena. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 704–709.

AUTHORS  Peter Evans

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CETACEANS KNOWN IN BRITAIN AND IRELAND ONLY FROM STRANDINGS Blainville’s beaked whale

Mesoplodon densirostris (BLAINVILLE, 1817)

ROBIN BAIRD

In the eastern North Atlantic, the Blainville’s beaked whale has been recorded from Spain, Portugal, Madeira and the Canaries, with strandings recorded in the Netherlands, Iceland, and France. There are no live sightings from UK or Irish waters. However, there has been a single extralimital stranding record from Wales in July 1993. The Blainville’s beaked whale is a deep-water species inhabiting ocean basins and trenches of 500–5,000 m in depth. It is a medium-sized beaked whale (4–5 m) with a body shape typical of the genus, which is bluish grey or black in colour and often marked with pale ovals and linear scars. This species has a strongly arched jaw, and in males a massive pair of flattened triangular teeth erupt at the apex (often encrusted with barnacles).

Gervais’ beaked whale

Mesoplodon europaeus (GERVAIS, 1855)

GONZALO MUCIENTES SANDOVAL

The Gervais’ beaked whale is only known from the Atlantic and recently several strandings have been recorded in the eastern North Atlantic. These include western Ireland (January 1989), France, Portugal, southern Spain, the Azores, Mauritania and Guinea-Bissau, although most records have come from the Canaries. Except for the type specimen found floating in the English Channel in 1848, there are no sightings or strandings from UK waters. This species is a medium sized (4.5–5 m) beaked whale, with a dark grey or indigo back that becomes light grey on the lower flanks and belly. It is distinguished from other beaked whales by the short, but clearly defined, slender beak, and relatively straight mouth line. In adult males one pair of laterally compressed, triangular teeth erupt in the lower jaw about one-third of the way along the gape.

Cetaceans known in Britain and Ireland only from strandings  183

True’s beaked whale Mesoplodon mirus (TRUE, 1913)

BRIAN CLASPER, ORCA

Since 1899, there have been 11 records of stranded True’s beaked whale, ten of which were from the west of Ireland. There are no definite records from UK waters but sightings based on appearance in the eastern North Atlantic (not all confirmed by genetics) have been made in the Canaries, Azores and Bay of Biscay. The most diagnostic feature of this species is a short but clearly defined beak, sloping into a slightly bulbous forehead. There is a single pair of slightly laterally compressed teeth, oval in cross section, directed forward and upward at the extreme tip of the lower jaw, exposed above the mouth line only in adult males. The body is typical of the genus (4–5.5 m), slate grey in colour and often accompanied by pale spots and linear scars. A dark eye patch may also be present.

Narwhal

Monodon monoceros (LINNAEUS, 1758)

SHUTTERSTOCK, WILDESTANIMAL

The narwhal rarely occurs outside of the artic, however, since the sixteenth century there have been seven records from Britain and Ireland. The only and last sighting of a narwhal in UK waters was of two individuals off Orkney in June 1949. Elsewhere in western Europe, the only records are from Germany (1736), the Netherlands (1921), Sweden (1992) and Belgium (2016). The narwhal is a distinctive 5 m-long cetacean (excluding tusk). Its body is stout with a small rounded head with bulbous forehead and very slight beak. It is mottled grey-green, cream and black in colour but older males appear lighter, partly because of the accumulation of white scar tissue. The left tooth of the male narwhal is greatly extended (up to 2.7 m long) in all but the youngest animals and erupts through the upper lip as a spiral tusk.

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Melon-headed whale Peponocephala electra (GRAY, 1846)

The only records of the melon-headed whale in north-west Europe are from a stranding in Charlestown, Cornwall, in September 1949, and from three individuals that live-stranded near La Rochelle in France (two in 2003, and another in 2008). This species is generally found in waters deeper than 1,000 m. The species reaches 2.3–2.7 m in length, with a torpedo-​ shaped body that is uniformly dark in colour but often has white lips and a subtle grey cape pattern over the back. The dorsal fin is relatively tall, recurved and centrally placed on an arched back.

ROBIN BAIRD

Fraser’s dolphin

Lagenodelphis hosei (FRASER, 1956)

ROBIN BAIRD

In the eastern North Atlantic Fraser’s dolphin occurs mainly off the coast of north-west Africa and in the Canaries, the Azores and Madeira. However, there was a mass stranding of 11 animals in north Brittany on the Atlantic French coast in 1984, and a single stranding record from the Outer Hebrides, north-west Scotland, in 1996. With characteristics of both the common and white-beaked dolphin, it has a robust body (2.4–2.7 m in length) with a short but well-defined beak and small curved dorsal fin and flippers. The body is bluish grey on the back and light pink or white on the belly. A creamy white band and parallel black band extend from the eye along the flanks towards the anus.

BIBLIOGRAPHY EVANS, P.G.H. 2008. Fraser’s dolphin Lagenodelphis hosei. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 731–733. EVANS, P.G.H. 2008. Melon-headed whale Peponocephala electra. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 733–735. EVANS, P.G.H., AGUILAR DE SOTO, N., HERMAN, J.S. & KITCHENER, A.C. 2008. Blainville’s beaked whale Mesoplodon densirostris. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 697–699. EVANS, P.G.H., HERMAN, J.S. & KITCHENER, A.C. 2008. Gervais’ beaked whale Mesoplodon europaeus. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 696–697. EVANS, P.G.H., HERMAN, J.S. & KITCHENER, A.C. 2008. True’s beaked whale Mesoplodon mirus. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 694–696. MARTIN, A.R. & EVANS, P.G.H. 2008. Narwhal Monodon monoceros. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 702–704.

VAGRANT SPECIES AND THOSE WITHOUT ESTABLISHED POPULATIONS IN THE UK

Raccoon

Procyon lotor (LINNAEUS, 1758)

MARK BALDWIN

The raccoon is a cat-sized carnivore with grey fur, a distinctive black band across its eyes, and a series of black bands on its tail. It has a plump body with a hunched back and pointed nose and ears. This species has been spotted in a few locations and these records probably relate to escaped individuals. No breeding populations are known and there are no presence records of this species in the databases used to create this Atlas. However, the high potential for this species to become invasive means that ongoing surveillance is essential.

Red-necked wallaby

Macropus rufogriseus (DESMAREST, 1817)

Between 2000 and 2016 there were 132 verified sightings of the red-necked wallaby from 23 different hectads. This includes 105 records from the Peak District, 11 records from the Isle of Man, 11 records in Buckinghamshire and Bedfordshire, and two in Yorkshire. There have also been individual records from Norfolk, Devon and Loch Lomond. The red-necked wallaby is the only species of large marsupial known to be present in the UK. Feral colonies of this species began from escaped zoo stock in the 1940s with colonies in the Peak District, the Weald, Sussex, and Inchconnachan Island in Loch Lomond.

MARK BALDWIN

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Reindeer

Rangifer tarandus (LINNAEUS, 1758)

EUGENE BUTTERWORTH

There are 208 records from seven hectads since the year 2000 of reindeer in the UK. Most of these are from within a few hectads in the Cairngorms National Park. There is also a single record east of Edinburgh. The reindeer is a domesticated species that was introduced to the UK in 1952 from Sweden (wild reindeer being extinct in the UK since the Mesolithic period) and, despite being free-ranging, the Scottish population is privately owned and managed. The reindeer is a robustly built medium-sized deer, standing around 85–150 cm tall at the shoulders. Both sexes have multi-branched antlers. It has a thick dense coat (thicker on the shoulders) that is greyish brown, and often lighter upper parts.

Pond bat

Myotis dasycneme (BOIE, 1825)

HENRY SCHOFIELD

The pond bat has a distribution stretching from northwestern France and southern Scandinavia south to Serbia and Montenegro, Ukraine, and north Kazakhstan, east to central Russia. However, within north-west Europe, records are patchy. There is currently only one record of this species in the UK. That was of an adult male found in Kent in 2004. The pond bat is a medium sized bat (13–18 g) with dense pale grey-brown dorsal fur and well-demarcated white/light grey ventral fur. The skin is reddish to light brown on the face, whilst the skin elsewhere is grey-brown. The feet are noticeably large and hairy, and the tragus – which is unusually short for a Myotis species – has a blunt tip and is bent slightly inwards.

Geoffroy’s bat

Myotis emarginatus (GEOFFROY, 1806)

HENRY SCHOFIELD

The Geoffroy’s bat is distributed throughout the Mediterranean region. In the UK there are a very small number of records from swarming sites. These include two individuals captured in 2012 and 2013 in West Sussex; and an animal with the morphological characteristics of the species but lacking genetic confirmation, captured in 2013 in Wiltshire. This medium sized bat (6–9 g) is very similar in appearance the Natterer’s bat, so some under-recording is possible. The key characteristics distinguishing it from the Natterer’s bat are the long rust-brown coloured woolly fur and a distinctive right angled notch on the outer ear margin. The tragus is long and thin but does not reach the ear notch.

Vagrant species and those without established populations in the UK  187

Northern bat

Eptesicus nilssonii (KEYSERLING AND BLASIUS, 1839)

JENS RYDELL

The northern bat is distributed across central and eastern Europe, through to France and Northern Italy in the west, and into the Arctic Circle in the north. It is generally considered to be a sedentary species, though there are occasional records of very long distance movements (>450km). In the UK there are three isolated records for the species that appear to derive from natural movements. These include a hibernation record in Surrey (1986), one on a North Sea oil platform (1996) and one in Berkshire (2014). The northern bat is similar in appearance to the serotine, with dark brown skin, broadly rounded ears, and a tragus that is broadest in the middle. It is smaller (9–13 g) than the serotine bat. The ventral fur is characteristic, being dark with golden-tips, and is well-demarcated from the yellow-brown ventral fur, particularly around the throat. The ears are broadly rounded, do not meet in the middle and have a tragus that is broadest in the middle.

Parti-coloured bat

Vespertilio murinus (LINNAEUS, 1758)

SALLY-ANN HURRY

The parti-coloured bat is distributed throughout most of northern Europe with a scattering of records from the UK. Since 2000 there have been 15 records from 12 different hectads, six of which have come from the Shetland Isles. The other records are derived from across Britain, including two on the Isle of Wight, four near Eastbourne, one in Oxfordshire and one on the Isle of Arran. The parti-coloured bat is a medium-sized bat (10–15 g) with short, broad, rounded ears. It has dark brown or black fur that is longer on the back and tipped with silver, giving a frosted appearance. The face and ears are blackish brown in colour.

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Kuhl’s pipistrelle bat Pipistrellus kuhlii (KUHL, 1817)

DANIEL WHITBY

Kuhl’s pipistrelle is a Mediterranean bat species and is distributed throughout southern Europe and northern Africa. Since 2000, there have been eight records of Kuhl’s pipistrelle in the UK. These are mainly derived from south-east England, with four records from the Isle of Wight, one in the Thames Valley, two near Colchester and one in Bourne. The species is also regularly recorded along the coast of parts of south-east England in acoustic surveys. Records of Kuhl’s pipistrelle in the UK have been attributed by some as translocations and by others as colonisation from a species with an increasing northward range expansion. This species is very similar in appearance to other European pipistrelle bats (5–10 g), but is distinguished by its dentition: the incisors have a single cusp whereas all other pipistrelle bats have two cusps. It may therefore be under-recorded.

Savi’s pipistrelle bat Hypsugo savii (BONAPARTE, 1837)

The Savi’s pipistrelle has a wide distribution stretching from the Mediterranean and North Africa through to the Middle East and Caucasus to northern India. Over the last 30 years, it has expanded northwards in central Europe, with some individuals being found to migrate to Germany. In the UK, individuals that appear to have arrived naturally have been recorded in Eastbourne (1993) and Merseyside (1996), but there are no records since 2000. This species has the typical general characteristics of a pipistrelle bat (5–9 g). However the dorsal fur is longer, and the golden-yellowish tip to the brown fur are noticeable. The ears are relatively short with a tragus that broadens at the apex, being much broader than that of a common or soprano pipistrelle. The face and ears are very black, and somewhat shiny. The penis has a right-angled bend near the base, and the tail projects 4–5mm from the tail membrane. HENRY SCHOFIELD

Vagrant species and those without established populations in the UK  189 There are five species of phocid or true seals that are normally distributed in the higher latitudes of the Atlantic Ocean but which have occasionally been recorded around the UK.

Bearded seal

Erignathus barbatus (ERXLEBEN, 1777)

L. JACKSON

Since 2000 there have been eight records of the bearded seal from six different hectads. Records are mainly on the Shetland Isles, but two sightings were also made in the Moray Firth and one on the Northumberland coast. Adults of both sexes are around 2.2–2.7 m in length from nose to tail and weigh around 235–340 kg. The bearded seal has a distinctively small head with characteristic vibrissae, giving the bearded appearance. The colour is generally grey with touches of brown, especially around the head.

Harp seal

Pagophilus groenlandicus (ERXLEBEN, 1777)

SHUTTERSTOCK, TODD BOLAND

Since 2000 there have been ten records of harp seal sightings from six different hectads. Records include six in Cornwall, two in Liverpool Bay, one near Newcastle and one from the Pembrokeshire coast. Adults of both sexes are up to 1.7 m in length from nose to tail and weigh around 130 kg. It is easily distinguished by its dark face-mask and by the large ‘harp’-shaped dark band along each flank starting at the shoulders.

Hooded seal

Cystophora cristata (ERXLEBEN, 1777)

SHUTTERSTOCK, ENRIQUE AGUIRRE

Since 2000, there have been 11 records of hooded seals from ten different hectads. The records include two in Cornwall, two in Devon, two from Norfolk and the Wash, four between Newcastle upon Tyne and Whitby, and one from the Shetland Isles. Males are larger than females, reaching 2.2–2.5 m in length and weighing over 400 kg compared with 2.2 m and 300 kg for females. Males also have a very obvious inflatable hood ornament. Both sexes are generally grey in colour and have numerous dark patches of irregular sizes across the body.

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Ringed seal

Pusa hispida (SCHREBER, 1775)

SHUTTERSTOCK, CHONLASUB WORAVICHAN

Since 2000 there have been two sightings of ringed seals, one in Shetland and one on the Norfolk coast. Adults from both sexes are around 1.5 m in length from nose to tail and weigh around 45–95 kg. Females are generally smaller than males. This species is very similar in appearance to the common seal. It is grey with black spots that often have pale edges, giving the species its common name.

Walrus

Odobenus rosmarus (LINNAEUS, 1758)

Since 2000 there have been two records of walruses in British waters. They both occurred in Orkney in 2013 and in 2018. The walrus is a large seal species growing up to 3.65 m and weighing in at 1,270 kg for an adult male, females being much smaller at 3 m and 850 kg. This species is very distinctive with unique dentition of large tusks – which are present in both sexes – and its flushed red colour when hauled out. L. JACKSON

BIBLIOGRAPHY BAKER, S.J. & HILLS, D. 2008. Species that have survived for more than 1 year, but not bred. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 783–793. DANSIE, E., PUTMAN, R.J. & YALDEN, D.W. 2008. Reindeer Rangifer tarandus. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 604–605. DIETZ, C. & KIEFER, A. 2016. Bats of Britain and Europe. London: Bloomsbury. HALL, A.J. 2008. Vagrant seals. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 547–550. HUTSON, A.M. 2008. Northern bat Eptesicus nilssonii In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. pp. 360–361. HUTSON, A.M. 2008. Pond bat Myotis dasycneme In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. p. 323. HUTSON, A.M. 2008. Savi’s pipistrelle bat Hypsugo savii. In S. Harris & D.W. Yalden (eds) Mammals of the British Isles, 4th edn. Southampton: The Mammal Society. p. 356.

AUTHORS  Fiona Mathews and Frazer Coomber

FERAL COLONIES AND POPULATIONS

There are several feral animals recorded in the UK. However, because of their domestic background there are many caveats that relate to these records. A true feral population is a wild self-sustaining population of escaped or released individuals descended from domesticated stock. A good example of a feral population is the Soay sheep Ovis aries that live on the islands of St Kilda. Moreover, the feral species’ records that were collated during the Atlas are likely to include a large number of domestic or managed individuals that are impossible to distinguish without additional information.

Feral ferret

Mustela putorius fero

MARK BALDWIN

Since 2000 there have been 610 records from 306 different hectads of the feral ferret, and 460 records from 26 hectads of ferret–polecat hybrids. Almost all counties of Great Britain and Northern Ireland have records of polecat-ferrets and feral ferrets, but the records are most numerous for southern England and the east Midlands. There are also records from many of the offshore islands including the Isle of Wight, Anglesey, the Isle of Man, Arran, Jura, Mull, Eigg, Skye, the Uists and Benbecula, Shetland and Rathlin.

Feral goat

Capra aegagrus hircus

Since 2000 there have been 515 records of the feral goat across the UK from 89 different hectads, with notable record clumps in Gwynedd, Wigtown, Stirling and Falkirk, Inverness, and Ross and Cromarty. The feral goat is also present on a number of islands including the Isle of Wight, Lundy, Islay, Jura, Kerrera, Seil, Mull, Rum and Skye. JAMES MILLER

192  Atlas of the Mammals of Great Britain and Northern Ireland

Feral sheep Ovis aries

Since 2000 there have been 42 records of the feral sheep from 27 different hectads. These are mainly from Britain, with noticeable groupings of records in the Cairngorms, Cumbria and South Yorkshire. There are also records from Lundy and the Isle of Wight.

VIVI BOLIN