Fossil fishes of Great Britain

D.L. Dineley Department of Geology University of Bristol Bristol, UK

and

S J. Metcalf Humber Estuary Discovery Centre, North East Lincolnshire Council Cleethorpes, UK

GCR Editor: D. Palmer

Published by the Joint Nature Conservation Committee, Monkstone House, City Road, Peterborough, PE1 1J' UK

First edition 1999

© 1999 Joint Nature Conservation Committee

Typeset in 10/12pt Garamond ITC by JNCC

Printed in Great Britain by Hobbs the Printers Ltd. on 100gsm Silverblade Matt.

ISBN 1 86107 470 0.

Apart from any fair dealing for the purposes of research or private study, or criticism or review, as permitted under the UK Copyright Designs and Patents Act, 1988, this publication may not be reproduced, stored, or transmitted, in any form or by any means, without the prior permission in writing of the publishers, or in the case of reprographic reproduction only in accordance with the terms of the licences issued by the Copyright Licensing Agency in the UK, or in accordance with the terms and licences issued by the appropriate Reproduction Rights Organization outside the UK. Enquiries concerning reproduction outside the terms stated here should be sent to the GCR Team, JNCC.

The publisher makes no representation, express or implied, with regard to the accuracy of the information contained in this book and cannot accept any legal responsibility or liability for any errors or omissions that may be made.

British Geological Survey Copyright protected materials

1. The copyright of materials derived from the British Geological Survey's work is vested in the Natural Environment Research Council. No part of these materials (geological maps, charts, plans, diagrams, graphs, cross-sections, figures, sketch maps, tables, photographs) may be reproduced or transmitted in any form or by any means, or stored in a retrieval system of any nature, without the written permission of the copyright holder, in advance.

2. To ensure that copyright infringements do not arise, permission has to be obtained from the copyright owner. In the case of BGS maps this includes both BGS and the Ordnance Survey. Most BGS geological maps make use of Ordnance Survey topography (Crown Copyright), and this is acknowledged on BGS maps. Reproduction of Ordnance Survey materials may be independently permitted by the licences issued by Ordnance Survey to many users. Users who do not have an Ordnance Survey licence to reproduce the topography must make their own arrangments with the Ordnance Survey, Copyright Branch, Romsey Road, Southampton SO9 4DH (Tel. 01703 792913).

3. Permission to reproduce BGS materials must be sought in writing from the Copyright Manager, British Geological Survey, Kingsley Dunham Centre, Keyworth, Nottingham NG12 5GG (Tel. 0115 936 3100).

Reproduction of the following figures is by kind permission of The Royal Society of Edinburgh, from Transactions of The Royal Society of Edinburgh: Earth Sciences,

(Figure 6.18) A and B, from Pearson and Westoll, (1979), 70, 337–399, fig. 16.

(Figure 6.18) D, E and F, from Miles and Westoll, (1968), 68, 372–476, figs. 24a and b, 25a.

(Figure 9.11) A and B, from Andrews, (1985), 76, 67–95, fig. 1.

(Figure 9.15) A and B, from Dick, (1981), 72, 99–113, figs. 7 and 12.

(Figure 9.15) C, from Dick, (1978), 70, 1–108, fig. 26.

(Figure 15.7) from Coates, (1994), 84, 317–327, figs. 1,7 and 10.

(Figure 15.9) from Milner and Sequeira, (1994), 84, 331–361, fig. 5 and 18.

(Figure 15.10) from Milner, (1994), 84, 363–368, fig. 1.

(Figure 15.11) from Clack, (1994), 84, 369–376, fig. 2.

(Figure 15.12) from Smithson, (1994), 84, 377–382, fig. 3.

(Figure 15.13) A, B and C, from Smithson et al., (1994), 84, 387–412, fig. 6.

(Figure 15.13) C, from Smithson et al., (1994), 84, 387–412, fig. 21.

Reproduction of the following figures from Cappetta (1987) is by kind permission of Verlag Dr. Friedrich Pfeil:

(Figure 13.15)

(Figure 13.24)

(Figure 14.05)

(Figure 14.07)

(Figure 14.09)

(Figure 14.12)

Reproduction of the following figures from Moy-Thomas and Miles (1971) is with kind permission of Kluwer Academic Publishers:

(Figure 06.15)

(Figure 06.22)

(Figure 09.2)

(Figure 09.15)

(Figure 09.21)

(Figure 10.6)

(Figure 10.7)

(Figure 10.9)

A catalogue record for this book is available from the British Library.

Contents

Acknowledgements

Access to the countryside

Conserving our fossil heritage — JNCC Policy Statement

Preface

Museum abbreviations

1 British fossil fish and amphibian sites

Introduction

Amphibians

The geological background

Palaeontology

Classification

Fishes of the Palaeozoic Era

Fishes of the Mesozoic and Cenozoic

Taphonomy

Palaeoecology

Biostratigraphy

History of research

Palaeontological conservation

The choice and distribution of the GCR sites

Sites of British fossil amphibians

2 Silurian fossil fishes sites of Scotland

Introduction: Silurian palaeogeography and stratigraphy

Environments

Fish faunas

Fish sites

Comparison with the faunas of other regions

Birk Knowes

Dunside

Shiel Burn

Dippal Burn

Slot Burn

Birkenhead Burn The Toutties

Ardmore–Gallanach

3 Late Silurian fossil fishes sites of the Welsh Borders

Introduction: Palaeogeography and stratigraphy

Environments

Fish faunas

Fish sites

Cwar Glas

Church Hill Quarry

Ludford Lane and Ludford Corner

Ledbury Cutting

Temeside, Ludlow

Tite's Point (Purton Passage)

Lydney

Downton Castle area: Downton Castle Bridge, Tin Mill Race,

Forge Rough Weir and Castle Bridge Mill

Bradnor Hill Quarry

4 Early Devonian fossil fishes sites of the Welsh Borders

Introduction: Palaeogeography and stratigraphy

Environments

Fish faunas

Fish sites

Devil's Hole

Oak Dingle, Tugford

Cwm Mill

Wayne Herbert Quarry

Besom Farm Quarry

Hoel Senni Quarry

5 Early Devonian fossil fishes sites of Scotland

Introduction: Palaeogeography and stratigraphy

Environments

Fish faunas

Fish sites

Tillywhandland Quarry

Aberlemno Quarry

Wolf's Hole Quarry

Whitehouse Den

6 Mid-Devonian fossil fishes sites of Scotland

Introduction: Palaeogeography and stratigraphy

Environments

Fish faunas

Fish sites

Westerdale Quarry

Achanarras Quarry

Cruaday Quarry

Black Park, Edderton Den of Findon, Gamrie

Tynet Burn, Elgin Melby

Papa Stour

Dipple Brae

Spinal Quarry

Banniskirk Quarry Holborn Head Quarry

Weydale Quarry Pennyland

John o'Groats

The Cletts, Exnaboe Sumburgh Head

7 Mid- and Late Devonian fossil fishes sites of England and Wales

Introduction: Palaeogeography and stratigraphy

Environments

Fish faunas

Fish sites

Bedruthan Steps

Mill Rock

Portishead

Prescott Corner

Afon y Waen

Comparison with other regions

8 Late Devonian fossil fishes sites of Scotland

Introduction: Palaeogeography and stratigraphy

Environments

Fish faunas

Fish sites

Oxendean Burn

Hawk's Heugh

Boghole, Muckle Burn

Scaat Craig

Comparison with other regions

9 British Carboniferous fossil fishes sites

Introduction: Palaeogeography and stratigraphy

Environments

Fish faunas

Fish sites

Foulden

Wardie

Glencartholm

Cheese Bay

Inchkeith

Ardross Castle

Abden

Steeplehouse Quarry

Bearsden

10 British Permian fossil fishes sites

Introduction: Palaeogeography and stratigraphy

Environments

Fish faunas

Fish sites

Middridge

Amphibian faunas and sites

11 British Triassic fossil fishes sites

Introduction: Palaeogeography and stratigraphy

Environments

Fish and amphibian faunas

Mid-Triassic of central and southern England

Fish and amphibian sites

Sidmouth

Late Triassic of central and South-West England

Fish sites

Aust Cliff

Vertebrate-bearing fissure deposits of South-West England

Fish sites

12 British Jurassic fossil fishes sites

Introduction: Palaeogeography and stratigraphy

Environments

Fish faunas

Early Jurassic or Lias

Fish sites

Lyme Regis coast (Pinhay Bay–Charmouth)

Blockley Station Quarry

Whitby coast (East Pier–Whitestone Point)

Mid-Jurassic or Dogger

Fish and tetrapod sites

Stonesfield

Kirtlington Old Cement Works Quarry

Watton Cliff

Late Jurassic or Maim

Fish and amphibian sites

Kimmeridge Bay (Gaulter Gap–Broad Bench)

Durlston Bay

13 British Cretaceous fossil fishes sites

Introduction: Palaeogeography and stratigraphy

Environments

Fish faunas

Early Cretaceous: Wealden Group (Berriasian–Barremian)

Fish sites

Hastings

Brook–Atherfield Point

Early ('Mid-') Cretaceous (Aptian–Albian)

Fish sites

East Wear Bay Late Cretaceous: the Chalk

Fish sites

Blue Bell Hills Pits

Totternhoe (Chalk Quarry)

Southerham (Machine Bottom Pit)

Southerham Grey Pit

Southerham (Lime Kiln Quarries)

Boxford Chalk Pit

14 British Cenozoic fossil fishes sites

Introduction: Palaeogeography and stratigraphy

Environments

Fish faunas

Pre-London Clay Tertiaries of the London Basin

Fish sites

Pegwell Bay

Herne Bay

Upnor

Abbey Wood

London Clay Formation

Fish sites

Bognor Regis

Maylandsea

Sheppey

Burnham-on-Crouch

Late Palaeogene of the Hampshire Basin

Fish sites

Bracklesham Bay

Lee-on-Solent

Barton Cliff

Hordle Cliff

King's Quay

15 Sites for British stem Tetrapoda and Amphibia

Introduction

Classification and evolution

Stem tetrapods and amphibians in the British palaeontological record

East Kirkton, Bathgate

Headon Hill

References

Glossary

Index

Acknowledgements

The initial task to identify, inspect and assess the condition of sites from which notable fossil fishes had been collected in the past began under the auspices of the Nature Conservancy Council (NCC) in the 1980s. Miss S. Turner and, later, Miss M.A. Rowlands set the work in motion for the NCC, visiting localities and corresponding with many individuals with local knowledge or expertise in the field of palaeoichthyology. An appreciable fund of data was established for Palaeozoic sites, and the task of GCR site selection from the list of potential localities had been largely completed by the late 1980s. Work towards publication of the results of this part of the GCR was initiated and Miss Rowlands prepared the draft of a text on the pre-Mesozoic sites. The work was then delayed for some time. In 1995 the present senior author (D.L.D.) was engaged by the Joint Nature Conservation Committee ONCC) to revise and extend the account of the Palaeozoic sites and, with the assistance of SJ.M., to complete the task of examining localities within the Mesozoic and Cenozoic formations and to prepare this report. Subsequently it was agreed to add an account (Chapter 15) of fossil amphibian tetrapod sites.

Almost all of the sites described in this volume have been visited and assessed relatively recently, but it is possible that the condition of some of them may have changed in the ensuing two years. Many sites may be visited only with the express permission of third parties. Whenever sought by the present authors, this was immediately forthcoming, and it is pleasant to record our gratitude for this.

A large number of people have contributed to and helped in the work at all stages, and to them sincere thanks are extended. Professor M.J. Benton has been staunch in his support throughout. He and the following friends and colleagues have read parts of the draft text: Dr M.I. Coates, Mr R. Davidson, Dr C. Duffin, Dr Forey, Prof. B.G. Gardiner, Dr E.J. Loeffler, Dr A.R. Milner, the late Dr C. Patterson, Dr N.H. Trewin, Dr CJ. Underwood and Dr DJ. Ward. Their corrections and comments have much improved the outcome. At the University of Bristol much kind help has also been provided by Mrs P. Baldaro, Dr E. Cook, Dr P. Orr and Mr S. Powell.

Staff of the country conservation agencies and JNCC, Dr. N. Clark of the Hunterian Museum, Glasgow and Drs R. Paton and M. Taylor at the Royal Scottish Museum, Edinburgh, have responded most kindly to requests for information and assistance of one sort or another. Dr C.F. Pamplin and Dr S. White of Xipress IT Solutions have applied their enthusiasm, skill and care to producing the diagrams. Special thanks are due also to Dr D. Palmer, to Mr N. Ellis, Miss AJ. Carter and Mr N. Cousins of the GCR Unit, for their unstinting help throughout.

As well as those organizations who have given their kind permission for the reproduction of illustrations and photographs, thanks also go to the many authors named in the figure captions.

Access to the countryside

This volume is not intended for use as a field guide. The description or mention of any site should not be taken as an indication that access to a site is open or that a right of way exists. Most sites described are in private ownership and their inclusion herein is solely for the purpose of justifying their conservation. Their description or appearance on a map in this work should in no way be construed as an invitation to visit. Prior consent for visits should always be obtained from the landowner and/or occupier.

Information on conservation matters, including site ownership, relating to Sites of Special Scientific Interest (SSSIs) or National Nature Reserves (NNRs) in particular counties or districts may be obtained from the relevant country conservation agency headquarters listed below:

Countryside Council for Wales, Plas Penrhos, Ffordd Penrhos, Bangor, Gwynedd LL57 2LQ.

English Nature, Northminster House, Peterborough PE1 1 UA.

Scottish Natural Heritage, 12 Hope Terrace, Edinburgh EH9 2AS.

Conserving our fossil heritage JNCC policy statement

Fossils are a key part of our natural heritage and form a major scientific, educational and cultural resource. They are fundamental to understanding the evolution of life and the character of ancient environments. Fossils also provide a basis for comparing the ages of rocks the world over.

The discovery, collection and study of the fossilized remains of ancient life can be enjoyable and stimulating activities that give people a fascinating insight into the geological and biological history of the Earth. However, the available fossil resource is finite. It is only through maintaining a prudent approach to the management of important fossil sites that future generations will be able to experience, study and enjoy this resource.

Responsible fossil collecting

In most circumstances, responsible fossil collecting is not harmful to the conservation of fossil sites. It can actually benefit our understanding of geology. This is particularly true where the fossils are relatively common or the sites in which they are found are subject to high levels of natural or artificial degradation, such as coastal cliffs that are being eroded or quarries that are being actively worked. In such situations collecting fossil specimens that might otherwise be destroyed can be beneficial to science, provided that they are properly documented and made available for study. Responsible fossil collecting can therefore be a valuable activity in the sustainable management and safeguard of our fossil heritage.

Irresponsible fossil collecting

Irresponsible collecting provides no scientific or educational gain and is therefore an unacceptable activity resulting in irreparable damage to our fossil heritage. It will pose a clear threat where fossils are rare or the fossil source is limited in extent, for example in a cave or a river channel deposit. Collecting without proper recording and curation, inexpert collecting, over-collecting and inappropriate use of power tools and heavy machinery are likely to reduce or even destroy the scientific value of such sites. Unless the activity is undertaken in an appropriate manner, the statutory nature conservation agencies, the Countryside Council for Wales, English Nature, Environment and Heritage Service and Scottish Natural Heritage, will oppose fossil collecting on the small number of Sites of Special Scientific Interest / Areas of Special Scientific Interest where this activity would cause significant damage to the features of special interest.

Code of good practice

Adopting a responsible approach to collecting is essential for conserving our fossil heritage. The basic principles set out below should be followed by all those intending to collect fossils.

Access and ownership — permission to enter private land and collect fossils must always be gained and local bylaws should be obeyed. A clear agreement should be made over the future ownership of any fossils collected.

Collecting — in general, collect only a few representative specimens and obtain these from fallen or loose material. Detailed scientific study will require collection of fossils in situ.

Site management — avoid disturbance to wildlife. Many invertebrates and lower plants live on or under loose rocks that should be replaced in their original positions whenever possible. Do not leave the site in an untidy or dangerous condition for those who follow.

Recording and curation — always record precisely the locality at which fossils are found and, if collected in situ, record relevant details of the position of the rock layer from where the fossil was collected. Ensure that these records can be directly related to the relevant specimens. Where necessary, seek specialist advice on specimen identification and care. Fossils of prime scientific importance should be placed in a suitable repository, normally a museum with adequate curatorial and storage facilities.

Achieving positive management

In order to achieve the successful management of the fossil heritage of the United Kingdom, the statutory nature conservation agencies will:

• Promote the responsible approach outlined in the Code of Good Practice, above.
• Encourage the placement of scientifically important fossils into a suitable repository (such as a museum) in order to ensure their proper curation, longterm security and accessibility.
• Recognize the contribution that responsible fossil collectors can make to geological and palaeontological study.
• Encourage collaboration within the geological community to ensure that maximum educational and scientific gain is made from our fossil resource.
• Support and encourage initiatives that increase awareness and understanding of the value of our fossil resource and the need to conserve it.
• Increase awareness and understanding of the differing management needs of fossil sites. In particular, encourage landowners and occupiers to become advocates for conservation of the fossil resource.
• Review the need for export and import controls on the international trade in fossil specimens.

JNCC, 1997.

Preface

This book summarizes the results of part of the Geological Conservation Review (GCR), an extensive research programme that aimed to assess the scientific significance of Britain's geological and geomorphological localities so that a representative set of the most important ones could be protected by law. Ultimately, the GCR sites were selected with a view to their designation as Sites of Special Scientific Interest (SSSIs) under the Wildlife and Countryside Act (1981).

In this volume the scientific importance of the set of fossil fish GCR sites is described. The surveys of fossil fish sites were carried out initially for three GCR 'Blocks' — Silurian-Devonian Chordata, Carboniferous-Permian Fish/Amphibia and Mesozoic-Tertiary Fish/Amphibia. In each block, a list of candidate GCR sites was established on the basis of previous research and published material; the list was refined to contain only the most scientifically important localities after consultation with as many people as possible and visits to as many sites as possible. The refined list of GCR sites comprised those nationally and internationally important sites that were needed to reflect the diversity of the fossil fishes of Britain and the history of research and investigation already undertaken.

Because there is potentially a great wealth of sites from which to choose, inevitably reliance has to be placed on those that have already been discovered, documented and researched. Also, while some of the sites described have been the subject of research or study very recently, others have been known for over 100 years, and there may be other classic sites emerging as a result of research under way at the present time. This emphasizes that the sites included in this volume represent what might be thought of as a snapshot at a particular point in time reflecting the way in which the need for a range of sites of different types is reconciled with the background of the information that has become available. It is also important to remember that some potential sites may overlap with sites described in other volumes of the Geological Conservation Review Series which were selected for the GCR for other special interests, such as stratigraphy, fossil mammals (Benton et al., in prep.) or fossil reptiles (Benton and Spencer, 1995).

Most of the SSSI proposals made as a result of the Geological Conservation Review have already been translated into site designations by the appropriate country conservation agencies (the Countryside Council for Wales, English Nature and Scottish Natural Heritage).

This volume is not intended to cover the practical problems involved in future site conservation, but rather to record the scientific justification for conserving particular sites and to demonstrate the character and significance that the sites have against the background of a wider palaeontological context. Although some of the sections necessarily use some technical terms, the accounts (particularly the 'conclusions' sections) have been constructed to be accessible to the non-specialist as far as possible; also the glossary at the end of the volume is compiled with this in mind.

We hope that readers will appreciate this volume as a foundation to describe the sites included but also bear in mind that these are only some of the sites that could have been designated. The purpose of the volume is not only to ensure that the selected GCR sites are available and documented for future generations but to acknowledge that, as further research is undertaken, additional knowledge can be added to that contained in this volume.

Museum abbreviations

BATM, Bath Museum.

BGS(GSM), British Geological Survey, Keyworth (old Geological Survey Museum collection, London).

BMB, Bedford Museum.

BRSMG, Bristol City Museum Geology Collections.

BRSUG, University of Bristol Geology Museum.

CAMMZ, Cambridge University Museum of Zoology.

CAMSM, Sedgwick Museum, Department of Earth Sciences, Cambridge University.

DORCM, Dorset County Museum, Dorchester.

EXEMS, Royal Albert Memorial Museum, Exeter.

GCM, Gloucester City Museum.

GLAHM, Glasgow Hunterian Museum.

LEICSM, Leicester City Museum.

MAIDM, Maidstone Museum.

NHM, Natural History Museum, London.

OUM, University Museum, Oxford.

RSM, Royal Scottish Museum.

SM, Stroud District Museum.

SMLU, Ludlow Museum.

UCL, University College London.

UMXC, Museum of Department of Zoology, University of Cambridge.

WHIMS, Whitby Museum.

YORMS, York Museum.

References