Discerning beetles, an entomo-archaeological study of coleopteranfaunas in relation to place and time Katherine Jane Grove A thesis submitted to the Faculty of Science for the degree of Doctor of Philosophy. Departmentof Archaeology and Prehistory, University of Sheffield, Northgate House, West Street, Sheffield S 14ET March 2001 Discerning beetles,an entomo-archaeologicalstudy of coleopteran faunasin relation to place and time CONTENTS VOLUME 1- Text ABSTRACT i ACKNOWLEDGEMENTS ii LIST OF CONTENTS iii VOLUME 2- Illustrative material and bibliography LIST OF FIGURES i LIST OF TABLES iv LIST OF PLATES v APPENDIX 156 BIBLIOGRAPHY 157 ABSTRACT This work initially examinesthe origins and methods of entomo-archaeologicalstudies and reviews the current stateof this discipline. Original work is presentedon coleopteranfaunas, mainly from medieval pits, recovered and analysed from sites in Winchester, Southampton and Chichester. The faunas resemble those recovered from deposits of similar provenancefrom other sites. They also contain the earliest records of some species in Britain and the earliest medieval occurrencesof others which, were common in Roman Britain, but missing from the Saxonurban record. A modem analogue of a medieval cesspit was set up in order to investigate the coleopteranfaunas, which develop in that environment. Further experimental work was carried out using a choice chamber, to determine the preferred pabulum of certain speciesofAphodius dung beetles. The findings are placed in a wider context as a representative sample of all work, carried out on Roman and post-Roman coleopteran thanatocoenoses,were included in the following investigations. Methods of standardising data from different sourcesare discussed. The distribution of synanthropic species through time was studied, with special emphasis on Tipnus unicolor and Cryptolestesferrugineus. Investigations into the characteristicsof pit faunas and into the relationship between the assemblageand the physical properties of the feature, in which it formed, were undertaken.Correspondence analysis, from the CANOCO computer program, was used as an aid to interpretation, in both exercises.Definite faunal types were discovered in pit assemblages,which could be related to the known archaeologicaldetails and certain propertiesof the feature were shown to influence the development of the fauna. These exercises proved that comparing work by different authors can be productive and that CANOCO is a powerful tool in analysis. ACKNOWLEDGEMENTS Firstly, I would like to thank John Magilton, former Director of Chichester District Council Archaeological Unit and my boss at the time, who suggested the idea and provided practical help and encouragement. More thanks are due to the Booth Museum of Natural History in Brighton who allowed me free access to their beetle collections for identification purposes. David Hemingway, an amateur entomologist, who gave me my copy of Joy's Practical Handbook of British Beetles and my husband, Chris, who has been very supportive and uncomplaining throughout. Many changes have occurred while I have been working on this project. The Archaeology Department in Sheffield moved from Glossop Road to West Street, the supertram schemehas been set up and the Museum of Popular Music, was conceived, built, openedand closed. I acknowledge all the help my supervisor Paul Buckland has given me over this long period and especially for developing the database"BUGS" which has been an invaluable aid. Many thanks also to my second supervisor Mark Dinnin for all his helpful comments and for all the miles of red ink he deployed on my behalf. The staff and studentsin the Department of Archaeology, who put aside their own work to help me, on my visits to Sheffield, are much appreciated.I would especially like to mention Pat Wagner, who has provided help and encouragementall the way through from my first sessionof identifying beetlesto the writing up stage. During most of my visits to Sheffield I have stayed with Pat, as well as offering helpful advice, her generous hospitality and friendship have been greatly appreciated. And finally, more thanks to Paul and his wife Joan, for welcoming me into their home and for many enjoyable meals and discussionsover the years. ii CONTENTS CHAPTER 1: INTRODUCTION, AIMS AND OBJECTIVES 1 .................. 1.1 INTRODUCTION 1 ................................................................................ 1.2 TYPES OF EVIDENCE OBTAINED 1 .................................................. 1.2.1 CHANGES IN CLIMATE 2 ....».......... » ................................. »..... ».................. ». 1.2.2 INSTRUMENTS FOR ARCHAEOLOGICAL INTERPRETATION 2 .»».. ». 1.23 UNDERSTANDING THE DEVELOPMENT OF THE BRITISH FAUNA. 3 1.3 AIMS AND OBJECTIVES OF THIS STUDY 4 .................................... CHAPTER 2: THE SIGNIFICANCE OF FOSSIL BEETLES AND THE PROCESSOF RECOVERY 6 ...................................................................... 2.1 INTRODUCTION 6 ................................................................................ 2.2 THE USE OF BEETLES AS A RESEARCH TOOL 6 ........................... 2.2.1 PRESERVATION OF REMAINS 6 ...............»................ »... ».... »................. »».. 2.2.2 EVOLUTIONARY STABILITY OF COLEOPTERA 7 ............». »........ ».. ».... 2.3 COLLECTION OF DATA 8 ............................. ...................................... 2.3.1 FORMATION OF THE THANATOCOENOSIS..... 8 ».».... »............ »». »»»»». 23.2 CHANGES AFTER BURIAL 10 ..»..... »..... »......... »... ».......... » .............. »..... »..... 23.2.1 Recording the 11 state of preservation »». »............ »»»... »»» .............. » . 2.3.3 SAMPLING PROCEDURE...... 13 »». ». » ....................... »»»...... ».. ».. ».... »...... ».. 2.33.1 Sample 14 size .N.. N ............. H....N.. ». »........ »... NNN...N........ NN.... ». NN..NN. N....... 2.3.3.2 Measurement 15 of sample sizeN.. N..»............ NHN.N. N..... N..... N»NN. H..... »».... 2.3.3.3 Comparison 15 of assemblages by percentage composition ... N..». N..».. »... 2.3.3.4 Standardisation of sample collection ............... NH.......N». .»..... »NN»»16 2.3.3.5 Sampling frequency.... 16 N»...NN. N..... »N..... HN........ ». N.......N.. N.. N...... N.......... N. N... N17 2.3.4 EXTRACTION METHODS NNN.... N ............... H.. N....... ......... N.. N....... H. 2.3.4.1 Efficacy of method ............. N. N.......... N.......... »............ ....... ........ ...... NNNN. 18 2.3.4.2 The 18 process of sorting... NN.N. N.... NNN..... ».. N.». N............. NHN..NN»..... ..... N19 2.3.5 THE IDENTIFICATION PROCESS. N.... N....... NN....... N ................. NN..... 2.3.6 THE RECORDING PROCESS 20 ....e .................. N..... N..... N.................. ........... 2.4 CONCLUSIONN... 21 NNN. .NN. N. N» .N. N. N... NNNNNN. H........... ......N ............. CHAPTER 3: THE INTERPRETATION OF THE DEATH ASSEMBLAGE... 22 N22 N... N. NNN.. N. N. N. 3.1 EARLY WORK NN.. N. N. N. N.. N.. N... NN.. N....... N.. N... NNN....... .NNN. 3.2 MOSAIC METHOD OF INTERPRETATION 23 ........................NN................... iii 3.3 THE WHOLE ASSEMBLAGE APPROACH 24 .................................... 3.3.1PRESENTATION OF DATA 24 .NN. NN.. NN. ». » .............. N...... »........... N. NN.. NN.... 3.3.1.1. Rank 27 order curves . N.»...... N..N....... N. N............ NN. NN.. N.... »».. N..NN......... 3.3.1.2 MacArthur's broken stick method N.N». »... ». N...»NNN ............... -27 3.3.2 HABITAT GROUPS. N. N..... NN. N............. N...... NN........ N......... .... ................... 28 3.3.3 ASSEMBLAGE STATISTICS 29 ....N... »NN. NN. N NNNN................. ». NNN........ 3.3.3.1 Measurement of diversity ... N.. N...... N..... »N...... N................. »...... N. N. N..... N30 3.3.4 CRITICISM OF THE WHOLE ASSEMBLAGE METHOD 31 ................... 3.3.4.1 Sample 31 size NNNNN»».N.. NN.. N. NN... »N... »NNN. »NN.. ».. ». N...»N». N. N.. 3.3.4.2 The figures». 32 calculation of percentage NN»N........NN... ».... NN»N.»»NN. »... 3.3.4.3 Use the indoor/outdoor 33 of category. ». »N»N............N.. ».... N..N..... ».... 3.3.4.4 The the index diversity....... 34 use of of N.» ............. N.»........ NN.N. N... ». »NN.». 3.3.4.5 Importance feature type. 34 of »NN.N. »N»N. »NN... »N.. »....... NN..........N.. N»...... 3.3.4.6 Difficulties in interpretations 35 verifying . ».. ».. »». »N»..N...... »»»N»».. »...... 3.3.4.7 Conclusions- 36 »NNN.....»».. ».... N...N. NNNN»». » ............. N.....».... »NN.».... 3.4 METHODS OF COMPARING FAUNAL DIVERSITY 36 .................... 3.4.1 JACCARD'S AND SORENSON'S MEASUREMENTS' OF DIVERSITY .N». ».. »»... »... »N. NNN.. ».. »»N»».... »N»N». N»»». N36 3.4.2 A STUDY BASED ON THE THEORY OF ISLAND BIOGEOGRAPHY. 37 3.4.3 MULTIVARIATE ANALYSIS. 38 ....»»..... »NN»».. N. N.......... ». 3.4.3.1 Cluster 39 analysis, a classification technique ....N...... .......... N............»....... 3.4.3.2 Ordination techniques......N. N.. ». N... N. N.. N... NN.................... N......... N». NN.. 40 CHAPTER 4: ECOLOGICAL STUDIES FOR ARCHAEOLOGICAL E4TERPRETATIONS 42 .............................................................................. 4.1 MODERN ANALOGUE STUDIES 42 ................................................... 42 4.1.1 CHARACTERISTICS OF THE DEATH ASSEMBLAGE...... ...»... »...... OF SPECIES 42 4.1.2 DETERMINATION ASSOCIATIONS »..........»». » ............... 4.1.2.1 Beetles in cess..... »».. »».. »..... ».. ».......... »........ ». ». »». ».... »»....