May 1995 Frontispiece
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A DIPTEROLOGICAL PERSPECTIVE ON THE HOLOCENE HISTORY OF THE NORTH ATLANTIC AREA P. SKIDMORE Department of Archaeology and Prehistory. The University of Sheffield May 1995 Frontispiece "The Viking House-fly" Heleomyza serrata (Linnaeus), male. .A. DIPTEROLOGICAL PERSPECTIVE ON THE HOLOCENE HISTORY OF THE NORTH ATLANTIC AREA SUMMARY Whiht a copious literature testifies to the value of subfonil insect analyus in the interpretation of Holoc.ne deposits (Buckland Ind Coope, 1991; Elias, 1994), lost of thit mults frol Itudin of Coleopterous uterhl. Although Dipterous traglentl are often abundant in the 511. deposits, they have receivld little attention. This Thuil il concerned priurily .ith establishing the great valul of Dipterous subfossill and the potential for advancel in this field, Dipterous lorphology is considered and features of prillry valul in the identification of subfollil uttrhl HI highlighted. Problns ,ith the traditional taxonolic criteria, insofar .. idenU fication of such uterial is concerned, arl discussed, and nil approaches arl recollended. Thus, a brief survey of the lorphology of Tipuloid larval head-capsulls, and a revilional paper on thl puparia of British Sphaeroceridlt, are included. The study includes Iiny case-studies frol .xcavations across the region, spanning the last 5,000 years. Although ther, is an inevitable bias in favour of archaeological sites, and hence of the lore synanthropic eleMents of the Dipterous fauna, situations unassociated lith hu.an s.ttlel.nts are also discussed. A lajor objective in this .ork was to Ixalin. the role -of Diptera in thl insect colonisation of lands left in a stat. of tibuJi rUi by reclding glaciations, The geographical arn concerned here cO'prittS the entire North Atlantic continental seaboard and ishnds frOI Franc. and Labrador northwards. Thil area IIW the IIjor western expansion of thl Cilts, Ronns and the Vikings, frOI whose settlelents Iuch of the Dipteroul I.terial frol archaeological deposits WilS obtained for this Thuis. It aho SilW the listern ligration of Inuit cultures frOI the Canadian Arctic into Greenland. Dipterous subfossils tell us luch of the living conditions and econolies of these ploples. In zoogeographical teras the Nearctic and Palaearctic aeet in the wutern Athntic, and a discussion on the 6reenlandic "uscid flun. wholly endorses the trlditional Aaerican view that 6reenland's fauna is of Ne.rctic origin, The .re. considered here also includes a large proportion of the globll Arctic llndllss, I study of which lust lid in understanding early post-glaCial envi ronaentl, TABLE OF CONTENTS Frontispiece Heleomyza eerr et s (Lfnnaeua) the "Viking Housefly" Summary 1 Table of Contents 2 Index of Tables and Illustrations 5 Index of Tables 5 Index of Text-figures 7 Index of Plates 8 Acknowledgements 11 1 Historical Background 12 a The History of Dipterology 12 b The past use of subfossil Diptera in Site Evaluation 15 c The advantages and disadvantages of Dipterous evidence 15 d Implications for Dipterology of Research in subfossil Diptera 17 e Archaeological implications of Research in subfossil Diptera 19 2 Methodology 22 3 List of Sites 24- .. Dipterology 27 a General Classification and Morphology 27 b The taxa encountered during the current research programme treated in detail:- 2 TIPULOIDEA 41 (T1pu11dae, 41; Pedic11dae, 47; Limon11dae; 48) PSYCHODOIDEA 54 (Psychodidae, 56; Ptychopteridae, 56) CULICOIDEA 57 (Culicidae, 57; Ceratopogonidae, 60; Chironom1dae, 60; SimulUdae, 61> TRICHOCEROIDEA (Trichocer1dae) 62 ANISOPODOIDEA (An1sopodidae)' 69 BIBIONOIDEA (Bibionldae) 70 SCIAROIDEA 73 (Sciarldae, 73; Scatopsidae, 75; Cecidomylidae, 78) TABANOIDEA 79 (Stratiomyi1dae, 79; Tabanidae, 81) EMPIDOIDEA 81 (Hybotidae, 81; Empidldae, 82; Dolichopodidae, 85) LONCHOPTEROIDEA (Lonchopteridae) 87 PHOROIDEA (Phoridae) 88 SYRPHOIDEA (Syrphldae) 90 TEPHRITOIDEA (Otitidae) 92 MTCROPEZOIDEA (Micropezidae) 92 TANYPEZOIDEA (Ps111dae) 94 SCIOMYZOIDEA 94 (Sepsidae, 94; Sc1omyzidae, 97) HELEOMYZOI DEA 97 (Coelopidae, 97; Helcomyzidae, 99; Heleomyzldae, 99; Sphaeroceridae, 104) 3 OPOMYZOIDEA 106 (Lonchaeidae, 106; Piophilidae, 106; Opomyzidae, 109j Agromyzidae, 111) ASTEIOIDEA (Anthomyzidae) 111 DROSOPHILOIDEA 112 <Ephydridae, 112; Drosoph1lidae, 115; Mil1chiidae, 117; Carn1dae, 117; Chloropidae, 118) MUSCOIDEA 118 (Scathophagidae, 118; AnthomyUdae, 124; Fann11dae, 131; Muscidae, 133i Hippoboscidae, 140; Tach1nidae, 140i Calliphoridae, 141> 5 Case studies froa Site Excavations 146 a ENGLAND 1 Brigg Raftj 2 Catterick Campi 3 Leicester Causeway; 4,5 Lindow Man II and III; 6 London Ministry; 7 Newark Ditch; 8 OwsLebur y; 9 Oxford (St.Budoc' s Church) i 10 Plymouth ditch; 11 Rudston Well; 12 Shirley Pool; 13 Southampton; 14 Taunton; 15 Thorne Moor; 16 Worcester Barrel Latrine; 17 a and b York. 146- 175 b WALES 18 Hen Domen. 175 c SCOTLAND 19 Buiston Crannogi 20 Dun Vulan; 21 Loch Druldlbeg (South Uist)j 22 Loch Olabhat (North Uist); 23 Papa Stour (Shetland); 24 Pluscarden Priory; 25 Skara Brae (OrkneY)j 26 South Loch Boisdale (South Uist)j 27 Tuquoy (Orkney), 177- 206 d ICELAND 28 Bessasta~ir; 29 Eynhyrnlngur; 30 Finnb6gasta~ir; 31 31 Gjegurj 32 Holtj 33 6sabakkij 34 Papeyj 35 Reykholtj 36 St6raborg. 206- 237 e WEST GREENLAND 37 Qegertasussuk; 38 Egaliugialik; 39 NiAqussat; 4 40 Nipaatsoqj 41 Sandnes, 42, 43 Tumeralik, and lake site nearby. 237- 276 6 Zoogeography of the Muscidae in North Atlantic area 277 1 The Muscidae of the North Atlantic Area 279 2 Arctic Muscidae 283 3 The Muscidae of Greenland 293 * Zoogeography of the Muscidae of Greenland 302 5 Conclusions 311 7 Conclusion to Thesis 313 8 Appendices 315 1 Revision of Sphaeroceridae Puparia (Publ.) 315 2 Analyses of Sites 332 3 The Muscidae of the North Atlantic Region 404 9 Bibliography 432 INDEX OF TABLES AND ILLUSTRATIONS Tables Brigg Boat Diptera 148 2 The Occurrence of Cess-pit Diptera in Leicester Excavations 153 3 The Abundance of Eight species of Cess-pit Diptera in Leicester Causeway Excavations 154 4 The Occurrence of putrid conditions in the Leicester Causeway Site on Dipterous evidence 155 5 Distribution of Taxa in the Leicester Causeway Samples 155 6 List of Dipterous Taxa in the Lindow Man III Samples 158 7 Diptera from London Ministry Samples 160 8 Diptera from Newark Ditch 162 5 9 Diptera from Rudston Well 166 10 Diptera from Shirley Pool 167 11 D1ptera from Taunton Excavations 169 12 Diptera of the Thorne Trackway site 171 13 Diptera of Hen Domen 176 14a Summary of numbers of fragments per sample from Buiston Crannog 181 14b Diptera of Buiston Crannog 182 15 Diptera of Loch Druidibeg 186 16 Diptera of Loch Olabhat 190 17 Diptera of South Loch Boisdale 198 18 Analysis of Dipterous Taxa from Tuquoy in terms of habitat 200 19 The Coprophilous Sphaeroceridae of Tuquoy 201 20 Diptera of Tuquoy 204 21 Diptera of Bessasta~ir 213 22 Diptera of Finnb6gasta6ir 217 23 Diptera of Gjagur 219 24 Diptera of Reykholt 227 25 Diptera of St6raborg 233 26 Diptera of Qeqertasussuk 240 27a Diptera of Egalugial1k 242 27b Percentage Representation of Trophic Groups at Egalugialik 242 28a Diptera of NiaquBsat (C 9, samples 1- 10) 245 28b Diptera of Niaqussat (C 10, 11 and D9) 246 28c Complete List of Dipterous Taxa from Niaqussat 248 29 Percentage Representation of Trophic Groups at Niaqussat 249 30 Diptera from the Nipaatsoq Farm floors 254 31 Percentage Representation of Trophic Groups at Nipaatsoq 255 6 32 The Dipterous Trophic Groups in the Sandnes Middens 257 33 Percentage Representation of nine Trophic Groups in the Sandnes Dipterous Material 259 34a Sandnes Dipteraj Felt 1 (Au 1- IV) 269 34b Sandnes Dipteraj Felt 1 (Au V- VI) 270 34c Sandnes Dipteraj Felt 11- V 272 34d Sandnes Dipteraj Felt VI- VII 272 35 Complete List of Dipterous Taxa from Sandnes 273 36a Diptera of Tumeralik 275 3Gb Percentage Representation of Dipterous Trophic Groups at Tumeralik 275 37 Summary of the Muscidae of the North Atlantic Area 280 38 The Zoogeography of the Arctic Muscidae of Greenland 306 39 The Zoogeography of the non-Arctic Muscidae of Greenland 308 40 Diptera of the Leicester Causeway Excavations 330 41 The Muscidae of the Canadian Arctic Archipelago, Greenland and the coastal areas of north-western Quebec and Labrador 402 42 The Muscidae of the North Atlantic Seaboard and Islands of the Pelaearctic 409 Text-figures Frontispiece Adult Heleomyz~ serrata (Linnaeus) 1 The numbers of Dipterous families in various regions of the North Atlantic area 16 2 Occurrence of three species of necrophagous Diptera in two archaeological sites in West Greenland 21 3 Wing of male Lonchopter~ lutea Panzer 28 4 Wing of Winter Gnat, Trichocer~ maculipennis Meigen 30 7 5 Obtect pupae of Empid species 31 6 Anterior parts of pupa of crane-fly, Eriopter~ lute~ 32 7 Fragments of head of subfossil Sc~thoph~B~ 33 8 Wing of Limosinine Sphaerocerid species 35 9 Wing of C~lliphor~ species 35 10 Unhatched male Heleomyza serr~t~ dissected from puparium 36 11 Pediciidaei larval head-capsule of Pedici~ (Tricyphon~) iJIJJIJ~cul~t~ 47 12 Map 1; sites studied in the"United Kingdom 147 13 Map 2; sites studied in Iceland 207 14 Map 3; sites studied in Greenland 238 15. 16 Diptera of Nip6aatsoq floors 252 17 Map of Northern Hemisphere showing 5 and 10·e July Mean Isotherms, marking Low and High Arctic boundaries. 284 18 The proportional increase in Sptlogona sp'ecies with latitude 288 19 Map oJ North Allantie arn il'ltWIng Vlrl'I.lS ,sla.nits e.b:. ~ rrccl: to In ~eoS"'&f*"Ca.l Sccl:.lon 3' + Plates 1 Tipulidaej Tipula confusa and Prionocer~ turcic~ 43 2 Tf pu.l Ldaer Ct enopbore bl mecul et «, Ti pul e cbeethem: T. l et ere l t e 44 3 Pediciidae and Li.aniidae; Dtcrllnomy~ mitis, Eriopter~ lute~ taenionotll, Limnophilll sp., Pseudolimnophil~. 50 4 Limoniidaej DicranomYIl modesta, Limonill mllcrostiBm~ and Helius flavus 51 5 Limoniidaej Austrolimnophil~ ochr~cea, Epiphr~Bma ocell~ris, Hexiltomilbicolor, Limnophilll punctate, Ormosill heemorrhoidali~ Eriopterll pilipes. 53 6 Ptychopteridae; Ptychoptera conteminetll.