THE LIBRARY 
 
 OF 
 
 THE UNIVERSITY 
 OF CALIFORNIA 
 
 LOS ANGELES 
 
 The RALPH D. REED LIBRARY 
 
 O 
 
 DEPARTMENT OF GEOLOGY 
 UNIVERSITY OF CALIFORNIA 
 
 LOS ANGELES, CALIF.
 
 THE 
 
 PAL^ONTOGRAPHICAL SOCIETY. 
 
 INSTITUTED MDCCCXLVII, 
 
 VOLUME FOE 1920. 
 
 LONDON: 
 
 MDCCCCXXII.
 
 MONOGKAPH OF THE FOSSIL INSECTS OF THE 
 BRITISH COAL MEASURES. 
 
 ORDER OF BINDING AND DATES OF PUBLICATION. 
 
 PARTS AND PAGES 
 
 PLATES 
 
 ISSUED IN VOL. 
 FOB TEAK 
 
 PUBLISHED 
 
 General Title and Index 
 
 
 1920 
 
 July, 1922 
 
 180 
 
 I IV 
 
 1919 
 
 July, 1921 
 
 81156 
 
 V X 
 
 1920 
 
 July, 1922
 
 A MONOGRAPH 
 
 FOSSIL INSECTS 
 
 BRITISH COAL MEASURES 
 
 HERBERT BOLTON, M.Sc., F.R.S.E., F.G.S., 
 
 DIRECTOR OF THE BRISTOL MUSEUM. 
 
 LONDON: 
 
 FEINTED FOil THE PAL^ONTOGKAPHIC AL SOCIETY. 
 
 19211922.
 
 PRINTED BY ADLARD AND SON AND WKST NEWMAN, LTD., LONDON AND DORKING.
 
 Geology 
 Library 
 
 SYSTEMATIC INDEX. 
 
 PAGE 
 
 INTRODUCTION 1 
 
 HABITS AND MODE OF OCCUR- 
 RENCE OF FOSSIL INSECTS ... 5 
 CONDITIONS OF LARVAL INSECT 
 
 LIFE ... 7 
 
 FOOD OF COAL MEASURE INSECTS 10 
 
 CLASSIFICATION ... ... 11 
 
 FAUNAL ASSOCIATION 18 
 
 SYSTEMATIC DESCRIPTIONS 24 
 
 Order PAL.EODICTYOPTERA ... ... "... 24 
 
 Family DICTYONEURID^E ... ... 24 
 
 Dictyoneura ... ... ... 25 
 
 higginsii ... ... 25 
 
 Family ORTHOCOSTID^E ... ... 27 
 
 Orthocosta 27 
 
 spleiidens ... ... 27 
 
 Family PTERONIDIID^E ... ... 29 
 
 Pteronidia 30 
 
 plicatula 30 
 
 Family HYPERMEGETHIDJE ... ... 31 
 
 Hyperinegethes .. ... ... 32 
 
 northumbriae ... 32 
 
 Family CRYPTOVENIID^E 34 
 
 Cryptovenia ... ... ... 35 
 
 moyseyi ... ... 35 
 
 Family MECYNOPTERID.E ... ... 36 
 
 Mecynoptera ... 36 
 
 tuberculata ... 37 
 
 INCERT^S SEDIS .. ... ... ... 40 
 
 Palaeomantis ... ... ... 40 
 
 macroptera ... 40 
 
 Family LITHOMANTID^: ... ... 42 
 
 Litbomantis ... ... ... 42 
 
 carbonarius ... ... 43 
 
 Lithosialis ... . . ... 45 
 
 brongniarti ... ... 46 
 
 Pruvostia ... ... ... ... 48 
 
 spectabilis ... ... 48 
 
 Family BREYERIID.S... r .. ... 50 
 
 Breyeria ... ... ... ... 50 
 
 woodwardiana . ... 50 
 
 Order PAL^ODICTYOPTERA (cont.) 
 
 Family SPILAPTERID^ ... ... ... 53 
 
 Spilaptera ... ... ... ... 53 
 
 sutcliffei ... ... 54 
 
 Family LAMPROPTILID^ ... ... 56 
 
 Boltoniella ... ... 56 
 
 tenuitegminata ... 56 
 
 Family BRODIID^: ... ... ... 57 
 
 Brodia ' 58 
 
 priscotincta ... ... 59 
 
 j uvenis ... 63 
 
 furcata ... ... ... 66 
 
 " Pteronepionites " ... ... 67 
 
 johnsoni ... 68 
 
 ambigua ... 68 
 
 lepus ... 69 
 
 Family ^ENIGMATODID-S: 70 
 
 jEnigmatodes ... ... ... 71 
 
 (?) regularis ... 71 
 
 Pseudofouquea ... 72 
 
 cambrensis ... 72 
 
 INCERT^E SEDIS... .. ... ... 74 
 
 Archeoptilus ... ... ... 74 
 
 ingens ... ... 74 
 
 Order MIXOTERMITOIDEA ... ... ... 76 
 
 Greroneura ... ... ... ... 76 
 
 (?) ovata 77 
 
 Order PROTORTHOPTERA ... ... ... 78 
 
 jEdceopliasma ... ... ... 78 
 
 anglica ... ... 78 
 
 Family COSELIID^: ... ... ... 81 
 
 Coselia 81 
 
 palmiformis ... ... 81 
 
 Family <EDISCHIID.SC ... ... ... 84 
 
 Grenentomum ... ... ... 84 
 
 (?) subacutum ... 84 
 
 Xeroptera ... ... .^ ... 85 
 
 obtusata ... ... 85 
 
 Scalseoptera ... ... ... 88 
 
 oo 
 
 recta ... 
 
 Order PROTOBLATTOIDEA ... ... 90 
 
 Ptenodera ... 90
 
 VI 
 
 SYSTEMATIC INDEX. 
 
 Order PROTOBLATTOIDEA (cont.) 
 
 Ptenodera dubius 
 INCERT^E SEDIS ... 
 
 Plesioidischia sp. 
 Order BLATTOIDEA ... 
 
 Family ARCHIMTLACRID^E 
 Aphthoroblattina ... 
 
 johnsoni 
 eggintoni 
 Archimylacris 
 
 hastata 
 woodwardi 
 incisa 
 (Schizoblatta) 
 
 vata 
 sp. ... 
 sp. .. 
 
 Phyloblatta 
 
 sulcata 
 
 transversalis . . . 
 (Archimylacridse) kirkbyi 
 BLATTOIDS INCERTJE SEDIS 
 Family MYLACRID^E 
 Hemimylacris 
 
 obtusa 
 convexa 
 
 obo- 
 
 PAGE 
 
 
 PAGE 
 
 
 Order BLATTOIDEA (cont.) 
 
 
 91 
 
 Family MYLACRID.E (cont.) 
 
 
 92 
 
 Phylomylacris 
 
 ... 124 
 
 92 
 
 maiitidioid.es 
 
 ... 125 
 
 93 
 
 Soomylacris 
 
 ... 128 
 
 96 
 
 deanensis 
 
 ... 128 
 
 96 
 
 stocki 
 
 ... 130 
 
 96 
 
 burri ... 
 
 ... 131 
 
 100 
 
 Ortliomylacris 
 
 ... 133 
 
 103 
 
 lanceolata . . . 
 
 ... 133 
 
 103 
 
 LARVAI, BLATTOIDS 
 
 ... 134 
 
 106 
 
 (Blattoidea) peaclii 
 
 ... 134 
 
 107 
 
 Leptoblattina exilis 
 
 ... 136 
 
 
 Order PROTODONATA 
 
 ... 139 
 
 109 
 
 Family MEGANEURID^E 
 
 ... 139 
 
 112 
 
 Meganeura 
 
 140 
 
 113 
 
 Boltonites ... 
 
 ... 140 
 
 113 
 
 radstockensis , . . 
 
 ... 140 
 
 113 
 
 INCERT^E SEDIS . . . 
 
 ... 144 
 
 115 
 
 Tillyardia 
 
 ... 144 
 
 118 
 
 multiplicata 
 
 ... 145 
 
 119 
 
 ADDENDUM 
 
 ... 146 
 
 120 
 
 Archimylacris 
 
 ... 146 
 
 121 
 
 pringlei . . . 
 
 ... 146 
 
 122 
 
 BIBLIOGEAPHY 
 
 ... 148 
 
 123 
 
 INDEX 
 
 ... 154
 
 LIST OF TEXT- FIGURES. 
 
 FIG. 
 
 PAGE 
 
 PIG. 
 
 1. Orthocosta splendens 
 
 28 
 
 24. 
 
 2. Pteronidia plicatula 
 
 ... 30 
 
 25. 
 
 3. Hypermegethes northumbrias 
 
 ... 32 
 
 26. 
 
 4. Cryptovenia moyseyi 
 
 ... 36 
 
 27. 
 
 5. Mecynoptera tubercidata 
 
 ... 37 
 
 28. 
 
 6. Ditto 
 
 ... 37 
 
 29. 
 
 7. Palfeomantis macroptera 
 
 ... 41 
 
 30. 
 
 8. Ditto 
 
 41 
 
 31. 
 
 9. Lithomantis carbonarius 
 
 43 
 
 32. 
 
 10. LitJwsialis brongniarti ... 
 
 ... 47 
 
 33. 
 
 11. Pruvostia spectabilis 
 
 ... 48 
 
 34. 
 
 12. Breyeria woodwardiana . . . 
 
 ... 51 
 
 35. 
 
 13. Ditto ... 
 
 ... 51 
 
 36. 
 
 14. Spilaptera sutcliffei 
 
 ... 54 
 
 37. 
 
 15. Brodia priscotincta 
 
 ... 59 
 
 38. 
 
 16 Ditto 
 
 62 
 
 39. 
 
 17. Ditto, forma juvenis 
 
 ... 64 
 
 40. 
 
 18. Ditto 
 
 ... 65 
 
 41. 
 
 19. Brodia furcata ... 
 
 66 
 
 42. 
 
 20. " Pteronepionites " johnsoni 
 
 ... 68 
 
 43. 
 
 21. jEnigmatodes (/) regularis 
 
 ... 71 
 
 44. 
 
 22. Pseudofouquea cambrensis 
 
 ... 73 
 
 45. 
 
 23. Archxoptilus ingens 
 
 ... 75 
 
 
 PAGE 
 
 JEdaopliasma anglica ... ... ... 79 
 
 Coselia palmiformis ... ... ... 82 
 
 Xeroptera obtusata ... ... ... 85 
 
 Scalieoptera recta ... ... ... 88 
 
 Ptenodera dubius ... ... ... 91 
 
 Plesioidischia sp. ... ... ... 92 
 
 Aphthoroblattina johnsoni ... ... 97 
 
 Ditto 97 
 
 Aphthoroblattina eggintoni ... ... 101 
 
 Archimylacris hastata ... ... ... 104 
 
 Archimylacris woodwardi ... ... 106 
 
 Archimylacris (Schizdblatta) obovata ... 110 
 
 Phyloblatta sulcata 114 
 
 Phyloblatta transversalis ... ... 116 
 
 Hemimylacris obtusa ... ... ... 122 
 
 Phylomylacris mantidioides ... ... 125 
 
 Soomylacris deanensis ... ... ... 129 
 
 Soomylacris stocki ... ... ... 131 
 
 Soomylacris burri ... ... ... 132 
 
 Orthomylacris lanceolata ... ... 134 
 
 Boltonites radstockensis ... ... ... 141 
 
 Tillyardia multiplicata ... ... ... 145
 
 PREFACE. 
 
 This monograph owes its existence to the friendly encouragement of Dr. 
 A. Smith Woodward, F.R.S., Keeper, and of Dr. F. A. Bather, F.R.S., Deputy- 
 Keeper of the Geological Department of the British Museum (Natural History), 
 and to these gentlemen I am indebted for the loan of the undescribed Coal Measure 
 fossil insects in their charge, and for much helpful assistance in other ways. 
 
 Others to whom my thanks are due are the Director of the National Museum, 
 Washington, U.S.A. : Dr. F. L. Kitchin, of the Museum of the Geological Survey ; 
 Mr. G. W. Lee, of the Scottish Geological Survey ; Dr. J. A. Clubb, of the Derby 
 and Mayer Museums, Liverpool; Dr. W. E. Hoyle, of the National Museum of 
 Wales, Cardiff ; Prof. W. S. Boulton, of the Birmingham University ; Dr. W. M. 
 Tattersall, of the Manchester Museum, Victoria University ; Mr. L. Gill, M.Sc., of 
 the Hancock Museum, Newcastle ; Mr. H. Y. Simpson, of the Kilmarnock 
 Museum ; and Messrs. Eltringham, S. Priest, W. Egginton, and D. Davies. 
 
 To Mr. W. J. Tutcher I am deeply indebted for his ready assistance in 
 photographing all the various specimens submitted for examination. His remark- 
 able skill has been ahvays at my service, and by means of it I have been able to 
 determine many details of structure which otherwise would have remained 
 obscure. 
 
 I am indebted to the Royal Society for several grants towards the cost of the 
 work. 
 
 H. BOLTOX.
 
 THE FOSSIL INSECTS OF THE BRITISH COAL 
 
 MEASURES. 
 
 INTRODUCTION. 
 
 THE first recorded Palaeozoic insect of any country appears to have been a 
 British specimen, Lithosialis Irongniarti (Mantel!), which was discovered in the 
 Coal Measures of Coalbrookdale in the early part of last century. It was sent by 
 Mantell to Brongniart as a leaf impression. Brongniart in turn submitted the 
 fossil to Mons. Audouin, who (1833, Audouin, 'Ann. Soc. But. France/ ii, Bull., 
 p. 7) described it as " d'un insecte inconnu," and allied to the Hemerobiidae, Semblis, 
 and especially to Corydalis and Mantis. The specimen was afterwards figured and 
 named by Mantell (1854, * Medals of Creation,' vol. ii, p. 575, fig. 2). 
 
 According to Parkinson, however, Lhuyd first recognised fossil insects in the 
 British Coal Measures. Parkinson (' Organic Remains,' vol. iii, p. 258, 18041811) 
 states that Lhuyd in a postscript to a letter to Dr. Richardson wrote as follows : 
 " Scripsi olim suspicari me Araneorum quorundam icones, una cum lithophytis, 
 in schisto carbonario observasse ; hoc jam ulteriore experientia edoctus aperte 
 assero. Alias icones habeo, quaa ad Scarabaeorum genus quam proxime accedunt. 
 In posterum ergo non tantum Lithophyta, sed et quaBdam insecta in hoc lapide 
 investigare conabimur." (' Lithophylacii,' p. 113.) 
 
 ["I have formerly written that I believed I had observed certain impressions of 
 spiders identical with Lithophytes in carbonaceous shales ; this I now, taught by 
 later experience, openly assert. I have other impressions which approach nearest 
 to the family of beetles. For the future, therefore, we will endeavour to investi- 
 gate not only Lithophyta, but also certain insects in these shales."] 
 
 Parkinson reprints four figures given by Lhuyd in his ' Iconograph,' tab. 4. 
 Two of these figures show eight legs and must therefore represent the remains of 
 Arachnids. None of the figures show wing-structure. 
 
 Interest in the occurrence of fossil insects was stimulated in 1837 by the 
 publication of Dean Buckland's 'Bridge water Treatise' on Geology, in which he 
 described and figured two fossils found at Coalbrookdale by Mr. Anstice (1837, 
 Buckland, ' Geology and Mineralogy,' 2nd ed., vol. ii, p. 76). He determined both 
 specimens to be the remains of coleopterous insects a determination since 
 corrected by H. Woodward (1871, ' Geol. Mag.,' vol. viii, p. 386, pi. xi), by 
 Scudder, and finally by Pocock, who referred them to the Araclmida (" Terrestrial 
 Carboniferous Araclmida," ' Mon. Pal. Soc.,' 1911, pp. 39, 77). 
 1
 
 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 Attention was afterwards diverted from the Coal Measures by the remarkable 
 discoveries of insect-remains made by Brodie in the Purbeck and Liassic rocks, 
 and by similar discoveries on the continent. 
 
 Mr. E. W. Binney (1867, ' Proc. Lit. and Phil. Soc. Manchester/ vol. vi, p. 59) 
 exhibited a specimen which "bore some resemblance to the pupa state of a 
 coleopterous insect," and had been found in the Cinderford Dyke Pit at Bradley,' 
 near Huddersfield. 
 
 A second specimen exhibited by Binney at the same time was referred to 
 Xt/lolrius sigillaride, Dawson. Binney added : " "We must expect great additions to 
 be made to the Carboniferous fauna, as doubtless the rich and luxurious 
 vegetation of that remote period would afford food and shelter for numerous 
 insects." 
 
 Binney's notes on the Huddersfield specimens caused the Rev. P. B. Brodie to 
 record (1867, * Geol. Mag.,' vol. iv, pp. 285 286) that he had in his collection " a 
 wing of a gigantic Neuropterous insect in ironstone from the Derbyshire Coal 
 Measures." 
 
 The same year Kirkby (1867, 'Geol. Mag.,' vol. iv, pp. 388390) reported the 
 finding of clearly defined insect-remains in the Durham Coal Measures. One 
 example consisted of " portions of the fore wing or tegmina of an orthopterous 
 insect nearly allied to Blatta or Cockroach," and the other " of an orthopterous 
 insect, apparently the abortive wing of a specimen related to the Pliasmidae." 
 Kirkby 's first specimen is the small but very fine wing described here under the 
 name of Phyloinylacris mantidiuides (Sternberg). The second specimen is not 
 determinable as an insect-fragment, and may prove to be a fossil fruit, referable 
 to one of the higher plants of the Coal Measures. 
 
 From 1867 onwards the finding of fossil insects in the British Coal Measures 
 occurred at long intervals until in 1908, the date of publication of Handlirsch's 
 ' Fossilen Insekten,' the following had been recorded : 
 
 Phylomylacris mantidioides (Groldenberg). 
 
 olim, " allied to Blatta," Kirkby. 
 
 ohm, " Blatiina mantidioides" 
 
 Groldenberg. 
 Lithosialis brongniarti (Mantell). 
 
 olim, " Gryllacris 'brongniarti," Mautell. 
 Lithomantis carbonarius, Woodward. 
 Archxoptilns ing ens, Scudder. 
 Brodia priscotincta, Scudder. 
 JEdceophasma anglica, Scudder. 
 Aphthoroblattina johnsoni (Woodward). 
 
 olim, " Etoblattina johnsoni," Woodward. 
 
 (Blattoidea) peachii (Woodward). 
 
 olim, " Etoblattina peachii" Woodward. 
 Leptoblattina exilis, Woodward. 
 Lithomylacris kirkbyi, Woodward. 
 Soomylacris deanensis (Woodward). 
 
 olim, " Etoblattina deanensis," Woodward. 
 Pseudofouquea cambrensis (Allen). 
 
 olim, " Fouquea cambrensis," Allen. 
 Breyeria woodivardiana (Handlirsch). 
 
 olim, " aflBnity Avith Lithomantis carlonarius," 
 Stobbs. 
 
 olim, " Stobbsia ivoodivardiana," Handlirsch. 
 
 The numerous discoveries of insect-remains in the Coal Measures of Commentry 
 (Allier), France, and the remarkable series made known by Handlirsch from the
 
 INTRODUCTION. 3 
 
 continent generally, and from the United States, overshadowed the limited British 
 series, which seemed almost trivial by comparison. The present monograph shows, 
 however, that the fossil insect-remains of the British Coal Measures are far more 
 abundant than was supposed, and that they are by no means unimportant. About 
 seventy specimens are known, of about sixty species, and they represent five of 
 the great groups of fossil insects. 
 
 Palaeodictyoptera are the dominant forms, and are closely followed by Blattoids, 
 several of which are referable to genera occurring in the French and Belgian 
 coalfields. The British examples of Soomylacris are represented near Lens and 
 Lievin by Soomylacris lievinensis, Pr., while Pliylomylacris mantidioides has its 
 counterparts in Phijlomijhicris godoni, Pr., and Phylomylacris Infittci, Pr., from Lens. 
 The great Protodonata of Commentry are represented in the Bristol coalfield by 
 Boltonites radstockensis. 
 
 The generic identity of French and British Coal Measure insects implies that 
 they formed part of a general and wide-spread fauna, a view which is strengthened 
 by the fact that while Soomylacris deanensis and 8. stocld occur in the Forest of 
 Dean coalfield, to the west, 8. burri occurs in the Kent coalfield to the east, and 
 only separated by the Straits of Dover from the Coal Measures of Northern 
 France, in which Pruvost finds other species of the same genus. 
 
 Pruvost has also shown that in the Coal Measures of Lens and Lievin there is 
 present a well-defined horizon of Authmcomya philUpsii, in which that species 
 passes through the same developmental changes as in the Kent coalfield. 
 
 It is extremely likely that the Kent coalfield will later yield numerous insect- 
 remains closely allied to those of France, and that systematic search will amplify 
 the list of forms already known from all the British coalfields. 
 
 The British Palreodictyoptera, on the whole, are more varied than the French, 
 few forms showing the primitive condition of Stenodictya, while certain examples, 
 such as Mecynoptera tuberculata, Palseomantis macroptera, and the three genera of 
 Lithomantids, are highly specialised. A similar degree of specialisation is seen in 
 the British examples of the Protorthoptera, while the Blattoids, by their numerous 
 genera and species, indicate that the group had a long history and a wide 
 geographical range in the British coal period. 
 
 The fossil insects already found in the British Coal Measures form probably 
 but a small fraction of those which remain to be discovered when attention is more 
 fully directed to them. The insect-fauna, however, is not usually associated with 
 the general fauna in the Coal Measures, .but occurs in beds of lighter coloured rock 
 than the ordinary carbonaceous shales, and with abundant ironstone nodules, or in 
 the case of the Blattoids, in association with masses of drifted vegetation in the 
 black shales, where the neuration of the wings so closely simulates the pinnules of 
 Neuropterts as to be mistaken for the latter and cast aside. 
 
 While insect-remains are usually regarded as wholly restricted to the West-
 
 4 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 phalian and Steplmnian stages of the Coal Measures, the fauna with which they 
 are most often associated in this country may indicate a greater age, as it is known 
 to occur as low down as the Calciferous Series of Scotland. Any statement, 
 therefore, of the range in time of British Palaeozoic insects based on the present 
 known forms may have to be set aside by later discoveries. 
 
 The fauna with which fossil insects are usually associated in Great Britain is 
 one in which arachnids and certain of the more primitive arthropods are the 
 dominant forms. Arachnids are kno\\n to occur in the Calciferous Sandstone 
 Series of Scotland at Redhall, near Slateford, Edinburgh, and in the Cement-stone 
 Group of the Lower Carboniferous at Langholme, Dumfriesshire (1911, Pocock, 
 ' Mon. Pal. Soc.,' p. 18), and elsewhere, the genus Archaeoctonus being represented by 
 A. glaber and A. tiiberculatus, and the genus Cycloplitlialmus by C. euyli/ptus at 
 
 Redhall, Blair Point, near Dysart, and Cramon near Edinburgh. 
 
 No insect-remains are known from any of these horizons, but if the faunal 
 
 association seen in the Coal Measures is a trustworthy guide, they may be looked 
 
 for with some prospect of success. 
 
 The faunal association existing in the " Soapstone Bed " of the Lower Coal 
 
 Measures at Carre Keys, Colne, Lancashire (1905, Bolton, ' Geol. Mag. ' [5], vol. ii), 
 
 is so similar in character to the typical insect-fauna elsewhere, that it is likely 
 
 that insects lived in the Lower Coal Measure period in Lancashire. 
 
 This faunal association at Carre Heys is as follows, and may be compared 
 
 with the faunal association in which insects have been found to occur in other 
 
 coalfields : 
 
 PISCES. 
 
 Hybodopsis wardi, Barkas. 
 Acanthodes wardi, Egerton. 
 Elonichthys aitkeni, Iraq. 
 
 AMPHIBIA. 
 
 Eugyrinus ivildi (A. S. Woodw.). 
 Microsaurian remains. 
 
 ARTHKOPODA. 
 
 Pygocephalus cooperi, Huxley. 
 Anthrapalxmon serratus, Woodw. 
 
 ,, woodwarcU, Etheridge. 
 
 traquairi, Peach. 
 
 Prestwichia rotundata, Woodw. 
 Architarbus subovalis, Woodw. 
 Euphoberia browni, Woodw. 
 Xylobius monilifer, Woodw. 
 
 The oldest known fossil insect in the British Carboniferous appears to be a 
 fragmentary wing (Genentomum subacutum), described by the author from shales 
 at a depth of 637 feet below the Bedminster Great Vein in the Bristol Coalfield, 
 and therefore at a considerable depth below the Pennant Grit. 
 
 Pseudofouquea cambrensis (Allen) was obtained from the top of the Four-foot 
 Seam in the Lower Coal Measures at the Llanbradach Colliery near Cardiff ; while 
 the shales over the No. 2 Rhondda Seam have yielded a wing-fragment of 
 Bolloniella temiitegminata (Bolton) ; and the shales over the Graigola Seam have 
 yielded the wings of two Blattoids, Hemimylacris convexa and Orthomylacris 
 lanceolata.
 
 HABITS AND MODE OF OCCURRENCE. 5 
 
 The No. 2 Rhondda Seam and the Graigola or Six-foot Vein of Swansea both 
 occur in the Pennant Grits, the former Hear the base of the series and the latter 
 at 200 yards below the Swansea Four-foot Seam, which forms the base of the Middle 
 Coal Measures in South Wales. 
 
 In Monmouthshire, insect-remains occur in shales over the Mynddislwyn 
 Vein, a seam at the base of the Upper Coal Measures. 
 
 The Durham and South Lancashire Coalfields have yielded insect-remains in 
 measures near the top of the Middle Series, while those recorded from the Derby- 
 shire Coalfield are on a still lower horizon in the Middle Series. Few fossil insects 
 are known from the Upper Coal Measures. 
 
 HABITS AND MODE OF OCCURRENCE OF FOSSIL INSECTS. 
 
 The bodies, and more particularly the wings, of insects, have been entombed 
 in various deposits under conditions difficult to determine. Whatever the condi- 
 tions, they must have been closely related to the habits of life. The older writers 
 claimed that wind-dispersal and water-carriage were the chief agents. Buck! and, 
 for example (' Anniv. Address to the Geol. Soc.,' 1842), supposed " that multitudes 
 of insects have been occasionally drifted by tempests to the sea." Mantell 
 ('Wonders of Geology,' 7th ed., 1857) pointed out that Westwood had drawn 
 attention to the fact that " the streams brought down innumerable insects at 
 certain periods, perhaps those of heavy rain." 
 
 Alfred Russel Wallace (' The Geographical Distribution of Animals,' 1876) and 
 Heilprin ('The Distribution of Animals,' 1887) alike drew attention to the wide- 
 spread occurrence of living insects far out at sea, in some instances still flying 
 strongly. Members of the British Association on their voyage to Australia in 1914 
 verified these statements by the capture of locusts as their vessel proceeded down 
 the Red Sea and into the Indian Ocean. More than a score of locusts were 
 captured on the vessel by which the writer travelled, and many more must have 
 been driven down into the water by the fringe of a simoon into which the vessel 
 entered beyond Aden. 
 
 Many insects are destroyed yearly by falling into streams and rivers after the 
 deposition of their eggs in the water, and by becoming entangled in the 
 surface film. 
 
 The occurrence of whole, or almost whole, insects is more likely to furnish 
 surer proof of the conditions under which life was passed than is the occurrence of 
 wings only, because the bodies, being more compact and much heavier than the 
 wings, are' less likely to have drifted to great distances. Sometimes, as we shall 
 see later when we consider special cases, such as the Coal Measures of Commentry, 
 France, or the remarkable faunal associations of certain of the British fossil insects,
 
 6 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 valuable information is supplied by the deposits, or by the nature of the associated 
 forms of life. 
 
 The great group of the Palseodictyoptera and certain of the Protorthoptera 
 and Protodonata had large wings, and Avere powerful fliers. We should therefore 
 expect to find their remains widely dispersed in deposits of varied nature. This 
 seems to be the case. Compact heavy-bodied insects like the Blattoids would have 
 a more limited range, and their bodies after death could not be carried to great 
 distances. Larval forms would in most cases be included in the deposits in the 
 immediate neighbourhood of the area in which they lived. 
 
 M. Henri Fayol, in his description of the Coal Measures of Coinmentry, 
 France, shows that these deposits were laid down in narrow land-locked lakes of a 
 trough-like form lying in depressions of older schistose and crystalline rocks. 
 The tranquil waters received only the finest mud in suspension, and the resultant 
 inudstones have yielded a large insect-fauna, in which Blattoids are most numerous. 
 The bodies of the insects are preserved in many cases. Certain of the insects were 
 strong fliers, and their occurrence with the bodies intact indicates that they, in 
 all probability, haunted the vicinity of the lakes and flew over them. When 
 strongly-flying insects like Boltonites radstockensis or Litliosialis brongniarti died 
 upon the land, the wings, because of their membranous and chitinous nature, 
 would persist after the destruction of the softer body, and be swept off into streams 
 after heavy rains or flooding of the land-surface, their great superficial area 
 combined with their lightness making flotation easy. 
 
 The transference of insect-wings from the land into water would be accom- 
 panied by the drifting of plant-material, and the two would be buried together in 
 the deposit then forming. The wing of Boltonites from Radstock was found with 
 plant-remains in deposits of this sort, and may be taken as a proof, supported as 
 it is by other examples of Protodonate wings, that these insects lived over the 
 land and died upon it. 
 
 The Palseodictyoptera, with their wings capable only of an up-and-down move- 
 ment in one plane at right-angles to the body, and, when in a position of rest, 
 disposed straight outwards, are not likely to have frequented the ground, except 
 in the open. These insects, like most of the Palaeozoic forms, were all of large size, 
 as contrasted with living types. Pruvost assumes that the characters of the wing 
 unfitted these insects for a forest life, and that they must have been restricted to 
 flight in the open neighbourhood of swamp pools. I do not wholly agree with 
 this assumption, for the branches and leaves of the Coal Measure plants do not 
 seem to have had so great a density and interlacing of foliage as seriously to 
 impede the flight of powerful winged insects. There seems no reason why these 
 insects should not have lived among the brakes of Lepidodendroid and Calamitean 
 trees, and after death fallen or been blown into adjacent waters. The fact that 
 isolated wings are often found in perfect condition and without any signs of wear
 
 CONDITIONS OF LARVAL INSECT LIFE. 7 
 
 and tear such as the wings of aged insects show to-day, raises the question whether 
 in some cases the wings were not shed, as in certain species of recent ants, the 
 shorn insect continuing its life as a ground-feeder. 
 
 The Protorthoptera Avere, judging from the structure of their mouth-parts, 
 somewhat general feeders or carnivorous, and the presence of strong walking legs 
 suggests that they spent much of their life on the ground, possibly along the 
 margins of swamps, where food would be especially abundant. They had, never- 
 theless, powerful wings, and some members, such as the (Edischiidae, had legs 
 adapted for leaping. Ortlioptera, represented chiefly by Blattoid forms, were all 
 fitted for flight by means of their large membranous hind-wings, and equally well 
 fitted by powerful walking legs for life on the ground. In repose the hind- 
 wings were hidden under the modified fore-wings. 
 
 I have elsewhere given my reason for a belief that the Blattoids were not 
 wholly phytophagous, but in all probability carnivorous also (' Quart. Journ. Geol. 
 Soc.,' vol. Ixvii, p. 153, 1911). 
 
 Blattoids may also have entered the water in search of food, for the hind 
 flying wings would be securely protected by the tegmina, whose broad muscular 
 bases of attachment were sufficiently powerful to compress them down on the back 
 and prevent water entering beneath, just as in the case of the living water-beetle, 
 Hydrophilus piceus. The chitinous surface of the body and of the tegmina would 
 not hinder progress in water, for their surfaces are no rougher than those of the 
 modern Dytiscus, nor would the insect on emerging bring with it so heavy a film 
 of water as to clog its movements. An objection may be found in the presence of 
 stout bristle-like hairs on the legs seen on such forms as Necymylacris lerickei 
 (Bolton) (1917, 'Mem. and Proc. Lit. and Phil. Soc. Manchester,' vol. Ixi, p. 15), 
 which might conceivably cause air-bubbles to cling in such profusion as to prevent 
 the insect being able to submerge. The presence of fine hairs on the swimming 
 legs of Hydrophiliis and Dijtiscus does not hinder the immersion of these insects in 
 water, so that this is not a valid objection. If no hindrance to immersion was 
 caused by the bristle-like hairs, they may have been useful in assisting the act of 
 swimming. 
 
 I think the probabilities are in favour of the Blattoids being at least semi- 
 aquatic as well as land insects. 
 
 CONDITIONS OF LARVAL INSECT LIFE. 
 
 The conditions under which larval life was passed are even more conjectural. 
 The Protodonata may be regarded as insects whose larvae must have been aquatic, 
 like the aquatic larvae of the Odonata now living, but Tillyard (1917, ' The Biology 
 of Dragonflies,' Camb. Univ. Press, p. 306) conjectures that since adult Prot- 
 odonata are found at Commentry without the occurrence of larval forms, the latter
 
 8 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 may have dwelt in damp eartli rather than in water, and that " the formation of 
 the larval tracheal system undoubtedly proves that this at one time was the case. 
 It may well have been so in Carboniferous times." 
 
 Tillyard's views are well worth quoting in full, especially as they support in 
 some measure those of Pruvost : " We may picture to ourselves the giant insects 
 of Commentry as inhabiting the shores of a large, shallow, nearly stagnant 
 lake. In the muddy ooze around its borders grew forests of the Giant Mare's 
 Tail, while further back on the sandy slopes the graceful Cycads and other extra- 
 ordinary plants formed a more diversified medley. There, amidst rotting vegetation, 
 these insects lived and bred. In such almost amphibious conditions it may well be 
 that the larvae of Protephemeroidea and Protodonata first began that series of 
 adaptive changes which finally led them to adopt a purely aquatic mode of life." 
 
 The larvae of Brodta and of other forms whose wings I describe under the 
 name of "Pteronepionites " must have lived under conditions fitted for their gradual 
 metamorphosis. The body was long, well segmented, and bore rudimentary 
 wings, which were carried well up over the thorax in an erect or semi-erect position. 
 Though rudimentary, the wings possess features which may have determined to a 
 large extent the mode of life. They are attached by broad, strong bases to the 
 thorax, and are very muscular, as shown by the stout ridges proceeding from the 
 point of attachment into the wings, and the anterior margins are also thickened. 
 The bodies with their lateral expansions of the terga are very suggestive of those 
 of the Diplopoda, and like them would offer no serious obstacle to progression 
 through rank and rotting vegetation. That these larval insects would also 
 penetrate soft muds, if necessary, in search of food is possible, since the 
 soft-bodied caterpillars of the Hawk-moths of to-day are able to enter hard soil 
 before pupation takes place. 
 
 The stout wing-bases and the strengthened margins of the wings would 
 prevent damage to these structures as the larvae crawled about, or sought to bury 
 themselves in the soil or muds. They were essentially adapted for a ground 
 habit. Whether they were capable of an aquatic or semi-aquatic habit can only 
 be settled by a knowledge of the mode of respiration. 
 
 Lubbock, Gegenbaur and others have adduced strong reasons in favour of an 
 aquatic origin of the insects, and in the Carboniferous types we should naturally 
 expect that the original habits had not had time to undergo any great modification. 
 Larval wings of the " Pteronepionites " type must have been living structures in 
 which metabolism was active, and very unlike the dried membranous sac-like 
 expansions of the adult insect. The growth of the larval wings was continued 
 throughout metamorphosis, and during this period their delicate nature, broad 
 expanse, and the thinness of the integument may have enabled them to assist in 
 the respiratory function. 
 
 The researches of Comstock and Needham show that larval wings of recent
 
 CONDITIONS OF LARVAL INSECT LIFE. 9 
 
 insects receive a plentiful supply of traclieal branches at an early stage, and it is 
 evident that these are much in excess of any aeration the wings are likely to 
 require. The traclieal development seems to be a persistence from an earlier 
 more active condition, when the larval wings may have played a part in assisting 
 the respiration. These considerations, and the presence of spiracle-like structures 
 in the interstitial neuration of the adult wings of many Palasodictyoptera, lend 
 support to the inference that the wings functioned as organs of respiration. These 
 spiracle-like structures are usually oval or rounded in outline, and thickened. In 
 some instances the} 7 show a series of raised lines radiating from the thickened 
 edge into the surrounding areas, as if they had been muscular strands and capable 
 of expansion and contraction. 
 
 Are these structures the atrophied remains of spiracles once functional, and 
 fitting the larva for a more or less aquatic existence ? During a recent 
 visit to this country Dr. Tillyard has suggested to me that they are 
 rudiments of sensory organs which may have been scent-glands. Scent- 
 glands are known to occur on the wings of many insects, as, for example, the 
 Green-veined White (Pieris napae), the Small "White (Pieris rapas) and others, 
 and their appearance is certainly a strong argument in favour of the view. Scent- 
 glands are, however, in all probability but specialised developments of previously 
 existing structures, and it is possible that the glandular-like organs to which I 
 give the name of "pseudo-spiracles," and Handlirsch the name of "pterostigmata," 
 are an earlier development connected with the " tracheoles " of Comstock, or that 
 primary tracheolation to which Tillyard has given the name of " archyodictyon." 
 Tillyard does not accept the view that they had ever any connection with 
 respiration. 
 
 The almost total absence of structures which can be accepted as functional 
 gills in these fossil insect-larvae may be accounted for by the perishable nature of 
 such organs. Before dismissing the question of the respiratory function in its 
 relation to the conditions under which larval life was passed, it is desirable to draw 
 attention to the larval Blattoid, Leptollattina ex His t Woodw. In this insect the 
 abdominal segments have the dorsal hinder margin extended into broad lamellar 
 expansions so filmy in texture that they may have served as organs of aeration. 
 The lamellar expansions were longer in life than they now are, the hinder borders 
 showing an irregular torn edge. Their extreme thinness would permit of a ready 
 osmo tic-like action, especially in damp vegetation, or in an aquatic or semi-aquatic 
 habitat. Scudder, Handlirsch, Lameere and others are all agreed in the belief 
 that the Blattoids frequented decaying vegetation in or near water, and under these 
 conditions the presence of organs of aeration similar in character to the abdominal 
 expansions of Leptoblattina exilis would be of the greatest value and offer no 
 difficulties to the habit of life. No similar structures are known in any other 
 larval Blattoid, so that the view cannot be pressed. 
 2
 
 10 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 "We shall not be far wrong in assuming that the larvae of some of the Coal 
 Measure insects were wholly aquatic, others semi-aquatic ; that the adult Blattoids 
 were indifferently aquatic or terrestrial; the adults of the non-Blattoid types spent 
 most of their life in the vicinity of the swamp-pools in which their larval stages 
 were passed, and to which they might need to return to lay their eggs. 
 
 Such a view seems to accord with the known facts, and will explain the special 
 character of the fauna of such deposits as those of Coseley, in Staffordshire, and 
 the brick-clays of Sparth Bottoms, Rochdale, Lancashire. These are evidently 
 true lagoon or swamp-pool deposits as contrasted with the ordinary shales and 
 binds of the Coal Measures. 
 
 FOOD OF COAL MEASURE INSECTS. 
 
 The nature of the food of Coal Measure insects has been much discussed, as it 
 is so closely associated with habits. Handlirsch considers that the Gymnosperms 
 and Pteridosperms of the Coal Measure forests were not likely to have been 
 frequented by insects in search of food, as these plants do not prove attractive to 
 living insects. Pruvost, on the other hand (1919, ? 1920, " La Faune Continental 
 du Terrain Houiller du Nord de la France," 'Mem. Carte Geol. France,' pp. 266 
 267), considers that many members of the Coal Measure flora possessed in their 
 spores, or in the case of the higher plants, in their cones, a plentiful food supply for 
 insects, and he finds in the association of a Phylloblaita at Lens with the Potonica 
 of Linopteris some support for his conclusions. 
 
 The contemporaneous rapid development of plants and insects is also quoted 
 by Pruvost in support of his views. 
 
 Several writers have argued that the powerful wings and consequent powers 
 of rapid flight of many of the insects are more in accordance with a predatory and 
 carnivorous habit than with a purely frugivorous or herbivorous one, and this 
 belief has led Lameere to write as follows (1917, ' Bull. Soc. Zool. France,' vol. xlii, 
 pp. 36 37) : "Over the lake of Commentry flew magnificent Epherneroptera and 
 splendid Odonatoptera, the carnivorous larvae of which were aquatic ; doubtless 
 the Odonatoptera, when fully grown, devoured the Ephemeroptera, of which the 
 most fully developed types, the Megasecopterida3, which have left no descendants, 
 must have made great slaughter among the smaller insects. 
 
 " On the ground, in the forests, swarmed innumerable Blattoids, which 
 frequented the detritus, and which had as enemies the ferocious and agile 
 Orthoptera, the varied counterparts of our Mantidae. These latter must have 
 attacked equally the large vegetarian Orthoptera, the counterparts of the Phasmae, 
 which probably climbed on trees, and the bulky Protohemiptera, which sucked the 
 sap. Some of the Orthoptera jumped, and there were some which by their 
 appearance recall our Acridians, but all these beings were mute.
 
 FOOD OF COAL MEASURE INSECTS. 11 
 
 " Small amphibians, and numerous arachnidians, came to limit the swarm of 
 this articulate world in a country without birds or mammals." 
 
 Pruvost does not accept the view of a primitive aquatic origin of insects, but 
 affirms his belief in a terrestrial origin, and thinks that even if an aquatic habit 
 be proved for certain of the larvae of the Coal Measure insects, the habit must 
 have been secondary, and derived from an earlier land ancestry (op. cit., p. 268). 
 
 Scudder^ has observed in the case of the Blattidae that the venation of the 
 tegmina very closely resembles the surface-features of the Neuropteris pinnule 
 so strongly in fact as to suggest mimetism. 
 
 Pruvost rightly urges that a mimetism is of little value unless the mimetic 
 insect frequents the plant mimicked. At the same time, it can be urged that the 
 stout compact bodies of the fossil Blattoids and their powerful walking legs were 
 equally admirably fitted for progression among rank and decaying vegetation, and 
 that in these conditions the Blattoids were quite as likely to have been omnivorous, 
 while finding some degree of protection among the Neuropteris pinnules lying on 
 the ground. 
 
 The writer has previously commented on the association of the wings of Blattoids 
 with the leaves of Gordaites (1911, ' Quart. Journ. Geol. Soc.,' vol. Ixvii, pp. 164 
 165), and has made the following comment : " While Carboniferous Blattoids 
 may have been wholly phytophagous, it is interesting to note that the leaves of 
 Cordaites (in the present case) are impressed with shallow pits, which show faint 
 traces of a spiral. I have in many previous instances found that such pits owed 
 their origin to attached shells of Spirorbis pusillus. Whether these leaves were 
 partially submerged in water during life is an open question ; but in all cases the 
 plant-tissues of the pittings are depressed, and are accurate impressions of 
 Spirorbis. If the Carboniferous Blattoids were not wholly vegetable feeders, the 
 occurrence of Spirorbis pusillus upon the Gordaites may supply a reason for their 
 frequent association." 
 
 CLASSIFICATION. 
 
 The classification of fossil insects has presented great difficulties, both to the 
 palaeontologist, and to the systematist of living forms. Palaeozoic insects show to 
 the systematist a series of forms not strictly referable to any modern grouping, 
 but presenting certain generalised details of structure which link two or more 
 now widely separated groups, besides other features not met with in living 
 forms. 
 
 The palaeontologist finds that he has not to deal with early and primitive 
 types, followed by a regular series showing a developmental progression, but with 
 an apparent sudden incursion of large series of highly modified and well-developed 
 insects, co-existent with others of more primitive type.
 
 12 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 Further discoveries will doubtless do much to eliminate these difficulties, but 
 present knowledge is such that recent entomology helps very little, and the 
 classification of Palaeozoic insects must be largely based on the known fossils, 
 realising that many of the intermediate forms are yet unknown. 
 
 The classifications of various authors vary widely, and even the broad general 
 facts of relationship are still uncertain. The earliest attempt at a classification 
 of fossil insects appears to have been that of Goldenberg (1873 1877, 'Fauna 
 Sarsepontana Fossilis : Die Fossilen Thiere aus der Steinkohlenformation von 
 Saarbriicken.') He recognised a new Order, Palaeodictyoptera, for the inclusion 
 of fossil insects differing in structure and the shape of the wings from 
 living representatives of the Orders Neuroptera and Orthoptera, while also 
 possessing characters which seem to link the two orders together. He did 
 not define the order, but arranged it with the other orders as follows : 
 
 Order. PAL.EODICTYOPTEKA, Goldenb. 
 
 Genera. Dictyoneura, Goldenb.; Eugereon, Dohrn; Miamia, Dana; Hemeristia, Dana; 
 
 HnplopTilebium, Scudder. 
 Order. ORTHOPTERA. 
 
 Sub-order. Orthoptera Pseudo-Neuroptera. 
 
 Genera. Termes, Goldenb. ; Termitidiiim, Goldenb. 
 Sub-order. Orthoptera vera. 
 
 Genus. Blattina, Germar. 
 Order. EHYNCHOTA. 
 
 Sub-order. Homoptera. 
 
 Genus. Fulgorina, Goldenb. 
 
 Scudder (1887, article " Insecta," ' Traite de Palasontologie,' by Karl von 
 Zittel, translated by Dr. Charles Barrois, vol. ii, ' Palasozoologie,' pp. 746833), in 
 his latest classification, considerably extended that of Goldenberg, while retaining 
 the primitive group of the Pala3odictyoptera. His complete arrangement is as 
 follows : 
 
 A. PAL^ODICTYOPTEEA, Goldenberg. 
 ORTHOPTEROIDEA, Scudder. 
 
 Family. Palaeoblattarise, Scudder. 
 Sub-family. My lacridae, Scudder. 
 
 Genera. Mylacris, Scd. ; Premylacris, Scd. ; Paramylacris, Scd. ; Lithomylacris, 
 
 Scd. ; jNecymylacris, Scd. 
 Sul -family. Blattinarise, Scudder. 
 
 Genera. EtoUattina, Scd. ; Spiloblattina, Scd. ; Archimylacris, Scd. ; Anthraco- 
 blattina, Scd. ; Gerablattina, Scd. ; Hermatoblattina, Scd. ; Progonoblat- 
 1ina,Sc<\.; Oryctoblattina,^d.; Petrablattina, Scd. ; Poroblattina, Scd. 
 Family. Protoph asm idae, Brong. 
 
 Genera. Titanopliasina, Brong. ; Litoneura, Scd. ; Dictyoneura, Goldeiib. ; 
 Polioptenus, Scd. ; Archxoptilus, Scd. ; Protophasma, Brong. ; Breyeria, 
 de Borre ; Meganeura, Brong. ; ^EdceopJiasma, Scd. ; Goldenbergia, 
 Scd. ; Haplophlebium, Scd. ; Paolia, Smith ; ArchegogryUus, Scd.
 
 CLASSIFICATION. 13 
 
 NEUROPTEROIDEA, Scudder. 
 
 Family. Palephemeridae, Scd. 
 
 Genus. Platephemem, Scd. 
 Family. Houiothetidae, Scd. 
 
 Genera. Acridites, Aud. ; Eucxnus, Scd. ; Genopteryx, Scd. ; Genentotnum, Scd.; 
 Didymojjlileps, Scd. ; Homothetus, Scd. ; Mixotermes, Sterzel ; Omalia, 
 van Ben. 
 Family. Palseopterina, Scd. 
 
 Genera. Miamia, Scd. ; Dieconeura, Scd. ; Strephecladus, Scd. 
 Family. Xenoueuridse, Scd. 
 
 Genus. Xenonenra, Scd. 
 Family. Hemeristina, Scd. 
 
 Genera. Lithomantis, Woodw. ; Lithosialis, Scd. ; Pucliytylepsis, de Borre ; 
 
 Lithentomum, Scd. ; Hemeristia, Dana. 
 Family. Gerarina, Scd. 
 
 Genera. Polyernus, Scd. ; Gerams, Scd. ; Adiphlebia, Scd. ; Megathentomum, Scd. 
 HEMIPTEROIDEA, Scudder. 
 
 Genera. Eugereon, Dohrn ; Fulgorina, Goldenb. 
 
 COLEOPTEROIDEA, Scudder. 
 
 Palseodictyoptera having a coleopterous aspect indicated by Geiuitz and Brongniart. 
 
 B. HETEEOMETABOLA, Packard. 
 ORTHOPTERA, Olivier. 
 
 Family. Forficulariee, Latreille. 
 Family. Blattariae, Latreille. 
 
 Genera. Neortliroblattina, Scd. ; Scutinoblattina, Scd. ; Blattidium, Westw. ; 
 
 Mesoblattina, Gein. 
 Family. Mantidae, Latreille. 
 
 Genus. Mantis, Limit'. 
 Family. Phasmidae, Leach. 
 
 Genera. Agathemera, Scd. ; Pseudoperla, Pictet. 
 Families. Acridii, Latreille ; Locustidae, Latreille ; Gryllidaa, Latreille. 
 
 NEUROPTERA, Linue. 
 
 Sub-order. Pseudoneuroptera, Erichson. 
 Sub-order. Neuroptera vera. 
 
 HEMIPTE^A, Linne. 
 
 Homoptera, Latreille. 
 Heteroptera, Latreille. 
 
 COLEOPTERA. 
 
 C. METABOLA, Packard. 
 
 DlPTERA. 
 
 LEPIDOPTERA. 
 HYMENOPTERA. 
 
 A modification of this classification was introduced by Handlirsch, in his 
 "Sub-phylum Insecta" in Eastman's translation of Zittel's 'Text-book of 
 Palaeontology,' 1913, as follows :
 
 14 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 ClassI.-PTERYGOGENEA, Brauer. 
 
 Order. PAL^ODICTYOPTERA, Goldeuberg. 
 
 Families. Dictjoueuridse, Megaptilidae, Hypermegethidae, Lithomantidse, Heolidae, 
 
 Fouqueidse, Spilapteridse, Lamproptilidse, Polycreagridse, Paolidse. 
 Order. MIXOTEKMITOIDEA, Haudlirsch. 
 Order. EECULOIDEA, Haudlirsch. 
 Order. PEOTORTHOPTEKA, Haudlirsch . 
 
 Families. Spanioderidee, Ischnoneuridae, Caloneuridse, Sthenaropodidse, (Ediscbiidse, 
 
 Geraridse, Cacurgidse. 
 Order. ORTHOPTERA, Olivier. 
 
 Sub-order. Locustoidea, Leach. 
 Sub-order. Acridioidea, Haudl. 
 Order. PHASMOIDEA, Leach. 
 Order. DERMAPTERA, De Geer. 
 Order. DIPLOGLOSSATA, De Saussure. 
 Order. THYSANOPTERA, Haliday. 
 Order. PROTOBLATTOIDEA, Haudlirsch. 
 
 Families. Stenoneuridse, Protophasmidse, Eoblattidse, Oryctoblattiuidae, J^tophlebidse, 
 
 Cheliphlebidae, Eucseuidse. 
 Order. BLATTOIDEA, Haudlirsch. 
 
 Families. Spiloblattiuidse, Mylacridae, Poroblattinidae, Neorthroblattinidae, Mesoblat- 
 tinidse, Pseudomylaeridse, Dictyomylacridae, Neomylacridae, Pteridomy- 
 lacridae, Idiomylacridae, Diechoblattinidte, Proteremidse. 
 Order. MANTOIDEA, Handlirsch. 
 
 Genus. Palasomantis, Bolton. 
 Order. SYPHAROPTEROIDEA, Haudlirsch. 
 Order. HAPALOPTEROIDEA, Haudlirsch. 
 Order. PROTOEPHEMEROIDEA, Haudlirsch. 
 Order. PROTODONATA, Brongniart. 
 Order. MEGASECOPTERA, Brongniart. 
 Order. PROTOHEMIPTERA, Handlirsch. 
 
 The publication of Handlirsch's great work, 'Die Fossilen Insekten,' 1906 
 1908, marked an important phase in the history of the study of fossil insects. 
 Handlirsch surveyed the whole field of fossil entomology, and brought the great 
 bulk of the known forms under a broad classification. The Order Palasodictyoptera 
 was much extended, denned, and made to include a large series of families, 
 several of which, however, are clearly widely divergent in type. This 
 was soon recognised by other workers, as doubtless by Handlirsch himself, 
 who may have considered it wiser to extend Goldenberg's order, even to 
 the inclusion of forms not definitely related, rather than to formulate a new 
 classification the components of which could not be rigidly defined. Knowing 
 that the field of research was rapidly widening, Handlirsch exercised a wise 
 restraint in not adding a new classification, which could only be of a temporary 
 character. Subsequent events have proved the wisdom of his action. Since 
 1908, the study of fossil insects has attracted more students, new localities and
 
 CLASSIFICATION. 15 
 
 in sect- horizons have been found, and many new types recorded. Some of these 
 linking forms already known, and others indicate relationships not fully under- 
 stood. The retention of the Order Palaeodictyoptera has therefore resulted in the 
 formation of a somewhat heterogeneous assemblage, all members of which have 
 one point of agreement. They are primitive co-types, more nearly related to each 
 other in various ways than they are -to recent insects, although that relationship is 
 not always as demonstrable as one could wish. 
 
 The most ambitious classification yet attempted is that of Prof. Lameere 
 (1917, 'Bull. Mus. Hist. Nat., Paris,' no. 1), who has published only a summary of 
 his conclusions. We are not able to determine how valid are his arguments, or 
 if he is justified by evidence in setting forth his new scheme. He sweeps the 
 Order Pala3odictyoptera wholly away, pointing out that it consists of a hetero- 
 geneous assemblage, and substitutes a more detailed classification as follows : 
 
 SUBULICORNIA. 
 
 EPHEMEROPTERA. 
 
 Family. Spilapteridae. 
 
 Genera. Lamproptilia, Brong. ; Epitethe, Handl. ; Becquerelia, Brong. ; 
 Palteoptihis, Brong. ; Compsoneura, Brong. ; Spiloptilus, Handl. ; 
 Homaloneura, Brong. ; Graphiptih(s,~Brong.; Spilaptera,~Brong. 
 Family. Megasecopteridse. 
 
 Genera. Aspidothorax, Brong. ; Corydaloides, Brong. ; Diaphanoptera, Brong. ; 
 Cycloscelis, Broiig. ; Sphecoptera, Brong. ; Psilothorax, Brong. ; Mis- 
 choptera, Brong. ; Ischnoptilus, Brong. 
 Family. Protephemeridse. 
 
 Genera. Apopappns, Handl. ; Triplosoba, Handl. 
 ODONATOPTERA. 
 
 Family. Fouqueidae. 
 
 Genei-a. Fouquea, Brong. ; Rhabdoptilus, Brong. 
 Family. Dictyoneuridae. 
 
 Genera. Microdict ya, Brong.; Stenodidya, Brong. 
 Family. Dictyoptilidae (Protodouata). 
 
 Genera. Arcli&megapiilus, Brong. ; Dictyoptilus, Brong. ; Poromaptera, Brong. ; 
 
 Protayrion, Broug. ; Gilsonia, Brong. ; Meganeura, Brong. 
 EHYNCHOTA. 
 PROTOHEMIPTERA. 
 
 Family. Homoiopteridse. 
 
 Genera. Lycocercus, Handl. ; Homoioptera, Brong. ; Lithoptilus, Brong. 
 Family. Megaptilidse. 
 
 Genus. Megapttius, Brong. 
 Family. Breyeridse. 
 
 Genus. Megaptiloides, Handl. ; Borrea, Brong. 
 Family. Mecynostomidoa. 
 
 Genus. Mecynostoma, Brong. 
 HEMIPTERA. 
 
 Family. Dictyocicadidae. 
 
 Genus. Dictyocicada, Brong.
 
 16 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 ORTHOPTERA. 
 
 Nomoiieura (Blattseformia, Handl.). 
 (a) Blattoidea. 
 
 Families. Hyaloptilidse, Protoperlidte, Fayoliellidae, Oryctoblattinida?. 
 (6) Mantoidea. 
 
 Families. Stenoneuritidae, Stenoneuridae, Isclmoneuridse. 
 
 Heteroneui'a. (Equivalent in part to Orthoptera Cursovia and Orthopteroidea, Handl.) 
 (a) Phasrnoidea. 
 
 Family. Sthenaropodidse. 
 
 (&) Locustidae. (Equivalent in part to Orthoptera Saltatoria.) 
 Families. (Ediscliiidae, Caloneuridse. 
 
 Lameere restricted his research to the French fossil insects, his studies being 
 based on the types described and figured by Brongniart (1893, ' Bull. Soc. 
 Industrie, Saint Etienne,' 3 ser., vii), the collections made by Fayol, and the 
 large series of fossil insects from the Upper Coal Measures (Stephanian) of 
 Commentry, now preserved in the National Museum of Natural History, Paris. 
 The following remarks may be made on his classification : 
 
 EPHEMKKOPTERA. The three families forming this .division are regarded as 
 closely related, the genus Becqnerelia of the Spilapteridae bearing certain characters 
 of the Megasecopteridas, while the family is also linked through the genus 
 Apopappus (which is taken to supply a natural transition between the Spilapteridse) 
 to the genus Triplosoba. 
 
 ODONATOPTUKA. The family Fouqueidae is held to differ from the Spilapteridae 
 in that transverse veins are numerous, close together, and form a network over 
 the inner margin, and in the anal area a feature which brings it nearer to the 
 Protodonata. The family DictyoneuridaB possesses a network of veins extending 
 over the whole wing, as in Microdictya. The remaining family, Dictyoptilidae, 
 contains Archdemegaptilus, which Lameere considers differs only from the Dictyo- 
 neuridae in the fusion of the median and radial veins at the base of the wing. 
 P rota gr ion is considered nearest to the true Odonata, while Meganeura and Gilsonia 
 are specialised types. 
 
 RHYNCHOTA. The presence of a rostrum in Lycocercus goldenbergi, and the 
 resemblance of the head and leg of Homoioptera gigantea to the same structures in 
 Eugereon, are considered sufficient proof of the Protohemipteroid characters of 
 Lycercus, Homoioptera, and the allied genera Lithoptilus, Megaptllus, Mecynostoma, 
 Arcltaeoptilus and Paramegaptilus. 
 
 ORTHOPTERA. Lameere regards Handlirsch's group of Protorthoptera as an 
 assemblage of two related but distinct types, which he classifies under Nomoneura 
 and Heteroneura. The genus Stenoneurites is regarded as the connecting link 
 between the Mantoidea and the ancestors of the Blattoidea, the genus Stenoneura 
 being also in some, measure transitional between Stenoneurites and the Ischno- 
 neuridae.
 
 CLASSIFICATION. 17 
 
 Nomoneura. This sub-division includes the Blattaeformia of Handlirsch, and 
 is distinguished by the wings having no precostal area, as contrasted with a second 
 division, Heteroneura, in which a precostal area is present, and in which the legs 
 are adapted for running and jumping. The Nomoneura include forms classified 
 by Handlirsch under the Protorthoptera and Protoblattoidea. Lameere separates 
 his Nomoneura into (a) Blattoidea, and (b) Mantoidea, the former characterised by 
 a sub-costal which joins the outer or costal margin, a more or less lengthened 
 radius, and a small cubitus. The Mantoidea have the sub-costa joining the radius, 
 and a large preponderating cubitus. 
 
 Heteronrura. This sub-division includes the Phasmoidea and the Locustoidea, 
 the former containing the family (a) Sthenaropodida3, in which the legs are long 
 and stout, the head prognathous, the pro thorax long and narrow in front and very 
 wide behind, and presenting two dorsal expansions. 
 
 The wing-venation of the Sthenaropodidas is such that they may have 
 been the ancestors of the Phasmida3. Lameere, however, does not regard the 
 Sthenaropodidas as ancestors of the Phasmidas, but as arising with them from a 
 common ancestor. 
 
 The members of the families (Edischiida3 and Caloneuridae possess legs fitted 
 for jumping, but differ considerably in their wing-venation. The (Edischiida3 are 
 possibly linked with the Locustidae, and the Caloneurida3 with the Acridida3. 
 
 Circumstantial and detailed as Prof. Lameere's classification is, the arguments 
 and deductions are not easily followed, the paper being only in abstract. A study 
 of the fossil insect-material from Commentry alone is not in itself likely to yield 
 all the facts and premisses upon which a classification can be built applicable to the 
 Palaeozoic insect-fauna of all coalfields and countries. Much more evidence is 
 wanted, and until the full paper is published, it is necessary to hold the classification 
 in suspense. 
 
 The most recent publication on the Palaeozoic insects is an extensive and 
 valuable memoir by Dr. P. Pruvost (1920, "La Faune Continentale du Terrain 
 Houiller du Nord de la France," 'Memoires pour servir a 1'Explication de la Carte 
 Geologique detaillee de la France/ Paris, 1 920) on the fossil insects recently found 
 by him, and others, in the neighbourhood of Lens and Lievin in the north of 
 France. Dr. Pruvost adopts the classification of Handlirsch with few emendations 
 as follows, and by his new material he has added considerably to our knowledge of 
 the Protoblattoidea and Blattoidea : 
 
 Order. PAL^ODICTYOPTERA. 
 
 Family. Stenodict.yopteridse, Broug. (Dictyoneuridae, Handlirsch). 
 
 Family. Spilapteridse, Broug., emend. Handlirsch. 
 Order. PROTORTHOPTERA, Handlirsch. 
 
 Families. (Edischiidse, Caloneuridae. 
 Order. HAPALOPTEROIDEA, Handlirsch. 
 
 Family. Hapalopteridse. 
 
 3
 
 18 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 Order. PROTOBLATTOIDEA, Hancllirsch, emend. Pruvost. 
 Sub-order. Archiblattidse, Pruvost. 
 Family. Cacurgidae, Handl. 
 Sub order. Archimanticlse, Pruvost, 
 
 Family. Cymenophlebidae, Pruvost. 
 Order. BLATTOIDEA, Haudlirscb. 
 
 Family. Arcbimylacridse, Handl. 
 
 Genera. Archimylacris, Asemoblatta, ManoUatia, Achnoblatta, Phyloblatta, Archxo- 
 
 iiplie, Barroisiblatta, Grypoblatta, Mesitoblatta. 
 Familv. Mylacridse, Scd. 
 
 Genera. Hemimylacris, Phylomylacris, Trilophomylacris, Soomylacris, Orthomylacris, 
 
 Stenomylacris, and LitJiomylacris. 
 Family. Poroblattinidae, Handl. 
 Genus. Premnoblatta, Pruv. 
 
 I have compared notes with M. Pruvost, and we have arrived independently 
 at the conclusion that for the present the classification of Handlirsch is, with few 
 emendations, the best to adopt, and most in keeping with the known facts. 
 
 FAUNAL ASSOCIATION. 
 
 Various collectors in British Coalfields have discovered not only insect-remains, 
 but a definite faunal association, of which the significance seems to have been 
 overlooked, and it has therefore not received the attention it deserves. 
 
 Most of the insect-remains are found in ironstone nodules, and the beds in 
 which these nodules occur are usually light-coloured rocks more similar to hardened 
 clay than to normal shales. The nodules are in vast numbers, ranging in size from 
 half-an-inch to ten and twelve inches in diameter. The beds seem more com- 
 parable to the fireclays or seat-earths than to the ordinary fissile shales, and both 
 in lithological character and fossil contents stand in some measure apart from the 
 ordinary Coal Measure rocks. They are not restricted to one coalfield, but have a 
 wide distribution. Where a systematic search of beds of this character has been 
 made, the insect-remains have been found accompanied by a fauna in which 
 arthropods of a more primitive type than insects are conspicuous. 
 
 The character of this fauna will be best understood by reference to the 
 following lists of fossils which have been recorded from certain localities : 
 
 DURHAM COALFIELD. " Zone of Anthracomya jrfiillipsii (Will.) " in upper part 
 of the Middle Coal Measures ; Claxheugh escarpment, two miles west of Sunder- 
 land, Durham. 
 
 PELECYPODA. j MEROSTOMATA. 
 
 " Ancylus vinti, Kirkby " (ef. "Spat" of Belinurus trechmanni (Woodw.): 
 
 Anthracomya pJiillipsii, Will.). DIPLOPODA. 
 
 Anthracomya minima (Ludvig). A. Isevia, Euphoberia, sp. 
 
 var. scotica.
 
 FAUNAL ASSOCIATION. 
 
 19 
 
 OsTRACODA. 
 
 Bei/richia, McCoy. 
 Cy there or Cypris. 
 
 INSECTA. 
 
 Litlwmylacris kirkbyi, Woodward. 
 
 Phylomylacris mantidioides (Groldenberg). 
 PISCES. 
 
 Rhizodopsis sauroides (Will.). 
 
 NOTTINGHAM AND DERBYSHIRE COALFIELD. Below the Top Hard Coal, Middle Coal 
 Measures (1911, Moysey, ' Geol. Mag.' [5], vol. viii, p. 50(5); Shipley Manor clay-pit, 
 one and a half miles north of Ilkeston, Derbyshire. 
 
 ANNELIDA. 
 
 Spirorbie, sp. 
 PELECYPODA. 
 
 Anthracotnya aiodiolaris (Sow.). 
 Carbonicola aquiUna (Sow.). 
 Naiaditeg modiolaris (Sow.). 
 OSTRACODA. 
 
 Bei/richia arcuata ! (Bean). 
 Estheria sp. 
 
 Leaia trigonioides, Moysey. 
 CRUSTACEA. 
 
 Preanaspides precursor, Woodw. 
 Arthroplaura armata, Jordan. 
 
 sp. uov., Moysey. 
 
 ARACHNIDA. 
 Cyclus, sp. 
 
 ., johnsoni, Woodw. 
 Belinnrue bellulus, Konig. 
 
 ,, kcenigianus, Woodw. 
 ,, longicaudatus, Woodw. 
 
 sp. 
 
 Presttricltia anthrax (Prestw.). 
 birtwelli, Woodw. 
 
 rotundata (Prestw.). 
 
 sp. 
 Eurypterus derbiensis, Woodw. 
 
 ARACHNIDA (continued). 
 
 Eurypterus moyseyi, Woodw. 
 
 Scorpion, post-abdominal segment, Moysey. 
 
 Geralinura britannica, Pocock. 
 
 Eobutlius holti, Pocock. 
 
 A nthracosiro fritschi, Pocock . 
 
 ,, woodwardi, Pocock. 
 
 sp. 
 
 Protolycosa, sp. 
 INSECTA. 
 
 Cryptovenia moyseyi, Bolton. 
 Orthocosta splendens, Boltou. 
 Pteronidia plicatula, Bolton. 
 PISCES. 
 
 Elonichthys, sp. 
 EGG CAPSULES OF FISHES (V). 
 Fayolia crenulata, Moysey. 
 cf. dentata, Zeiller. 
 Palxoxyris carbonarius (Schimper). 
 
 helicteroides (Morris). 
 ,, prendeli, Lesq. 
 Vetacapsula, johnsoni (Kidston). 
 
 cooperi, Mackie and Crocker. 
 PLANT EEMAINS. 
 
 Annularia; Calamocladus ; Sphenophyllum 
 Lepidophyllum ; Calamites ; ferns. 
 
 LANCASHIRE CoALFiELD.-Greyish-blue shales, 135-180 feet above the Royley 
 or Arley mine, the latter at the base of the Middle Coal Measures; Sparth Bottoms, 
 Rochdale, Lancashire. I am indebted for the following list to Mr. Walter 
 Baldwin, F.G.S., who, with Messrs. Sutcliffe, Parker, Platt and others, has < 
 years to the examination of these beds. 
 
 VERMES. 
 
 Spirorbis 
 BRACHIOPODA. 
 
 Lingula, sp 
 
 PELECTPODA. 
 
 Carbonicola acuta (Sow.). 
 robusta (Sow.). 
 
 turgida (Brown).
 
 20 
 
 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 PELECYPODA (continued). 
 
 Naiadites modiolaris (Sow.). 
 ,, iriangnlaris (Sow.). 
 ,, carinata (Sow.) 
 ,, elongata (Hind). 
 Anthracomya irilliamsoni (Brown). 
 
 EtJCRUSTACEA. 
 
 Dithyrocaris, sp. 
 Pygocephahis cooper i, Huxley. 
 
 (.' Anthrapaliemon) parkeri 
 
 (Woodw.). 
 
 Anthrapal&mon grossarti, Salter. 
 Eurypterus, sp. 
 Cyclus johnsoni, Woodw. 
 Rochdalia parkeri, Woodw. 
 Selinurus lunatus (Martin). 
 konigiamis, Woodw. 
 baldwini, Woodw. 
 longicaudatus, Woodw. 
 testudineus (Woodw.). 
 ARACHNIDA. 
 
 Prestwichia birtwelli, Woodw. 
 
 ., rotundata (Prestwich). 
 
 var. major, Woodw. 
 
 ,, minor, Woodw. 
 
 (Euproops] dance, Meek Worthen. 
 anthrax (Prestwich). 
 
 Eoscorpius (Mazonia) wardingleyi, Woodw. 
 Eobuthus holti, Pocock. 
 
 AEACHNIDA (continued). 
 
 Antliracoscorpio buthifortnis, Pocock. 
 sparthensis, Pocock. 
 
 Geralimira sutcliffei, Woodw. 
 Anthracomartus trilobitus, Scudder. 
 sp. 1 in Platt Coll. 
 sp. 2 in Platt Coll. 
 Anthracosiro woodwardi, Pocock. 
 Phalangiotarbus svbovalis (Woodw.). 
 
 ,, (Architarbnp) rotundata, 
 
 Woodw. 
 
 DlPLOPODA. 
 
 Xylobius platti, Woodw. 
 Euphoberia ferox, Salter. 
 
 ,, armigera (Baldwin) 
 ,, robiista (Baldwin). 
 ,, woodwardi (Baldwin). 
 Archiitlus, sp. 
 Acantherpestes major, Meek & Wort-hen. 
 
 ,, giganteus, Baldwin. 
 
 INSECTA. 
 
 Spilaptera sutcliffei, Bolton. 
 Mecynoptera tuberculata, Boltou. 
 PISCES. 
 
 Plalysomus tenuistriatus, Traquair. 
 Khizodopsis sauroides (Will.). 
 INCERTJS SEDIS. 
 
 Palxoxyris prendeli, Lesq. 
 
 SOUTH STAFFORDSHIRE COALFIELD. Binds between the " Brooch " and " Thick 
 coals ; Tip ton, Dudley, and Coseley. 
 
 EUCRUSTACEA. 
 
 Euphoberia ferox, Salter. 
 ARACHNIDA. 
 
 Anthracoscorpio biithiformis, Pocock. 
 Geralimira britannica, Pocock. 
 Grteophontu anglicvs, Pocock. 
 Eocteniza silvicola, Pocock. 
 Archgeometa nephilina, Pocock. 
 Curculioides ansticii, Bucklaud. 
 Poliochera alticcps, Pocock. 
 Plesiro madeleyi, Pocock. 
 Geraphrynus angulatus, Pocock. 
 
 hindi, Pocock. 
 
 tuberculatus, Pocock. 
 
 eggintoni, Pocock. 
 
 torpedo, Pocock. 
 
 ARACHNIDA (continued). 
 
 GerapJirynus angustus, Pocock. 
 Anthracomartus hindi, Pocock. 
 priesti, Pocock. 
 
 Anthracosiro ivoodwardi, Pocock. 
 
 ,, fritschi, Pocock. 
 
 Trigoiwtarbus johnsoni, Pocock. 
 
 INSECTA. 
 
 Palseod ict joptera. 
 
 Pruvostia spectabilis, Bolton. 
 Brodia prixcotincta, Scudder. 
 
 furcata (Handl.). 
 " Pteronepiunites" lepus, Bolton. 
 
 ,, ambiguus, Bolton, 
 
 Geroneura (/) ovata, Bolton.
 
 FAUNAL ASSOCIATION. 21 
 
 INSECTA (continued). INSECTA (continued). 
 
 Protorthoptera. Blattoidea (cont inued) . 
 
 Xeroptera obtusata, Bolton. Aphthoroblattina eggintoni (Bolton). 
 
 Scateoptera recta, Bolton. Archimylacris incisa (Bolton). 
 
 Coselia palmiformis, Bolton. Phylloblatta tramversalis (Bolton). 
 
 Blattoidea. LARVAL BLATTOID. 
 
 Aphthoroblattina johnsoni (Woodw.). Leptoblattina exilis, Woodw. 
 
 COALBROOKDALE COALFIELD, SHROPSHIRE. Ironstone Nodules of the Pennystone 
 Series. 
 
 EUCRUSTACEA. ARACHNIDA (continued). 
 
 Eitphoberia ferox, Salter. Curculioides ansticii, Buckland. 
 
 Anthrapaltemon (Ap^ls) dubius (Prestw.). EopJirynm prestvici (Buckland). 
 
 ARACHNIDA. j DIPLOPODA. 
 
 Prestivichia anthrax (Prestw.). Acantherpestes brodiei, Scd. 
 
 ,, rotundata (Prestw.) INSECTA. 
 ,, trilobitoides, Woodw. Lithosialia brongniarti (Mantell). 
 
 SOUTH WALES COALFIELD. Shales in the neighbourhood of the Mynddislwyn 
 Vein, base of the Upper Coal Measures. 
 
 ARACHNIDA. INSECTA. 
 
 Apliantomarttis areolatus, Pocock. Aphthoroblattina sulcata (Bolton). 
 
 Grxophonns anglicus, Pocock. r thorn ylacr is lanceolata (Bolton). 
 
 Mniocercus celticus, Pocock. Archimylacris hastata (Bolton). 
 
 Kreischeria vemtcosa, Pocock. ,, obovata (Bolton). 
 
 The arthropod association in the lists given is significant, for no other animals 
 enumerated are so readily water-borne as are insects. It may be assumed that 
 neither the more primitive arthropods, nor the insects, have been transported to 
 any great distance from their former habitat. Their preservation under similar 
 conditions supports the belief that their habits and habitats were the same, or 
 closely approximated to the same conditions of entombment. 
 
 The freedom of the deposits from comminuted carbonaceous matter, such as is 
 usually a chief constituent of the Coal Shales, may be accounted for by the beds 
 having been laid down in quiet lagoons or swamp lakes, into which only the finer 
 mud particles and floating pinnules and debris of coal-plants could pass and 
 accumulate. Such waters were probably fresh or brackish, shallow, and limited 
 in area. As we have indicated elsewhere (p. 10), it has been considered that the 
 larvae of many of the Coal Measure insects were semi- or wholly aquatic, and if 
 such was the case, they would be more likely to be found in the deposits accumu- 
 lating in quiet waters than in others exposed to movement. The presence of 
 Mollusca and the lower orders of Arthropoda, with such forms as Palxoxyris, and 
 even fish-remains, can be accounted for by the existence of occasional or 
 permanent passages leading to open waters, such as river channels and the 
 open sea.
 
 22 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 A succession of such lagunal lakes or swamp pools might be seasonal features 
 along a depression which, in a wet season, formed a water-course. 
 
 The paucity of Mollusca is noteworthy, only one species (" Unio ") having been 
 recorded by Moysey, notwithstanding his careful search of material from the 
 Shipley clay-pit at Ilkeston, Derbyshire; while Kirkby, and more recently 
 Trechmann and Woolacott have recorded Antkracomya pkillipsii, in addition 
 to " Ancylus vinti " (now known to be the larval form of A. phillipsii), from the 
 insect-bearing beds at Claxheugh, near Sunderland. 
 
 The deposits at Sparth Bottoms, Rochdale, Lancashire, are remarkable in that 
 they have yielded three species of Carlonicola, four species of Naiadites, and one 
 species of Antkracomya. 
 
 In regard to the presence of Crustacea, Thomson (1894, 'Trans. Linn. Soc. 
 London,' Zoology [2], vol. vi, p. 3), has shown that the recent Anasptdes tasmanise 
 lives in freshwater pools and lakes which are wholly cut off from the sea, and 
 Dr. H. Woodward (1908, < Geol. Mag.' [5] vol. v, p. 385) has described an allied 
 form, Preanaspides prascursor, from the Coal Measures of Shipley Clay-pit. 
 Further research may prove that not only Preanaspides, but such forms as 
 Belinurus, Prestwichia, Euryptems and Anthrapalaemon were also fresh-water in 
 habit, co-existing with insect larva? in the lagoons and swamp lakes. 
 
 Observations by the writer during the visit of the British Association to 
 Australia in 1914 bear on this point. While collecting in the " Bush country " at 
 St. Margaret's Bay, Western Australia, at Warburton, S. Australia, and else- 
 where, examination of almost every loose sheet of bark hanging to the gum trees 
 revealed a colony of scorpions, millipedes, spiders and cockroaches. In the 
 Australian localities mentioned true " bush " conditions prevailed, and seemed 
 much unlike those of low-lying swamps, such as are predicated for the Coal 
 Measure period. Subsequent experiences along the coast of North Queensland 
 modified these first impressions. It was found that notwithstanding the hot 
 tropical day, or perhaps by reason .of it, the nights brought an extremely heavy 
 dew, so that it was impossible to move four yards in the jungle before the clothing 
 was running with water discharged from the leaves of the jungle plants. In a 
 short time after sunrise the jungle was dry again, but much of the moisture must 
 have been caught up under projections capable of resisting the penetration of the 
 sun's heat. In the " bush " country, slabs of bark may give conditions which the 
 arthropod fauna find the most tolerable during the hot season. The Australian 
 winter with its heavy rains would more nearly accord with swamp conditions. 
 Long stretches of the coastline of Queensland, north of Brisbane, the country 
 along the Hinchinbrooke Channel, and in the neighbourhood of Townsville, are 
 low-lying, and covered by dense mangrove swamps growing out into the sea and 
 completely hiding the outlets of rivers. 
 
 The conditions seem similar to those of the low-level country, swamp and
 
 FAUNAL ASSOCIATION. 23 
 
 mud flat upon which the coal forests are believed to have flourished. During the 
 voyage along the coast, a landing was made at Lucinda Point, and an opportunity 
 was afforded of entering the swamp. Part of the swamp was awash on landing, 
 but as the tide receded, it was possible to reach the shore. The latter was found 
 to be a flat shelving beach, consisting of fine marine sand on the seaward side, 
 passing shorewards into a fine tenacious clayey mud in which the mangroves grew. 
 The sand bore numerous remains of echini, mollusca and marine debris, while the 
 mud below high-water mark and above it was penetrated in all directions by 
 the roots of mangroves, by the burrows of crabs, and by a burrowing gasteropod, 
 Telescopium fuscum. 
 
 Masses of leaves, branches and other vegetable material were mixed with the 
 mud, the latter in some places forming irregular lumps and boulder-like masses 
 around the vegetable material. A broad shallow depression, with a few inches of 
 water over a thick bed of mud littered with leaves, led into the swamp. It was 
 evidently the bed of a stream during the wet season, remaining as a series of 
 disconnected pools at other times, or drying up. 
 
 The resemblance of the physical conditions to those accepted as dominant in 
 the Coal Measure period was so evident that special attention was given to it with 
 a view to discovering discrepancies ; yet, had the mangroves been replaced by 
 dense groves of gum trees with their arthropod fauna, the circumstances would 
 have been well-nigh identical. Had the faunal association of scorpions, millipedes, 
 spiders and cockroaches of West Australia been transported to the swamp, their 
 remains would have become entombed in material not unlike that at Sparth 
 Bottoms or Coseley. 
 
 There is ample evidence to show that the coal plants grew on ground of this 
 character, and we know that the remains of fossil insects are associated with the 
 plant-remains. 
 
 The swamp conditions of the North Queensland coast reproduce with great 
 exactitude the presumed Coal Measure conditions, but lack the arthropod fauna 
 owing to the unsuitability of the vegetation. 
 
 Scudder had some such habitat and faunal association in mind when writing 
 his work " Archipolypoda : a Sub-ordinal Type of Spined Myriapods from the 
 Carboniferous Formation" ('Mem. Boston Soc. Nat. Hist.,' vol. iii, no. 5, pp. 
 143 182). Plate X of that work is described as "an attempted restoration of a 
 specimen of Acantherpestes major, Scd." The specimen is represented as leaving 
 the water, in which the hinder part of the body is still swimming by means of its 
 legs, while the fore part of the body is creeping up the trunk of a Lepidodendron 
 (L. vestitum). On the trunk crawls a cockroach (Etoblattina mazona, Scd.) ; and a 
 broken stem of Galamites cistii, Brong., lies partly fallen in a clump of Neuropteris 
 hescldi, Lesq. 
 
 We are, therefore, not without some justification in assuming that the faunal
 
 24 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 association which exists at Coseley, Sparth Bottoms, and elsewhere is not due 
 to the accident of deposition or transportation, but the natural result of the 
 conditions under which insect life was passed. 
 
 SYSTEMATIC DESCRIPTIONS. 
 Order PAL^ODICTYOPTERA (Goldenberg), Handlirsch. 
 
 1877. Goldenberg, Die Fossilen Thiere aus der Steinkoblenformation von Saarbruckeu. Fauna 
 
 Sarsepontana Fossilis, pt. ii, p. 8. 
 1906. Handlirsch, Die Fossilen Insekten, p. 61. 
 
 Slender insects with moderate-sized head and biting jaws, simple antenna?, 
 and two pairs of equal and similarly shaped wings. The wing-venation is not 
 unlike the hypothetical tracheation of the primitive nymph worked out by 
 Comstock and Needham ('American Naturalist,' vol. xxxii, no. 374, p. 85, fig. 4, 
 189899). 
 
 The wings could not be folded, being outstretched laterally in the position 
 of rest, and only moving in a vertical plane at right-angles to the body. The 
 thoracic segments are three in number, with wing-like pleurites in some cases 
 on the first segment. The abdomen has eleven segments, the eleventh segment 
 bearing cerci. Legs all similar, and fitted for running. 
 
 Goldenberg did not define the characters of the Order, but included in it a 
 group of Palaeozoic insects which, while somewhat related to the existing forms 
 of Neuroptera, are sufficiently unlike to prevent their inclusion in the latter 
 Order. 
 
 Handlirsch defined the Order as a primitive generalised group, probably the 
 ancestors of all later insects, and wholly confined to the Palaeozoic. He considered 
 that the larvas were predatory and aquatic, developing their wings gradually 
 without resting stages, and being in other respects similar to the imago. His 
 diagram 0f the primitive Pala3odictyopteroid wing shows a slight advance upon 
 the primitive nymph of Comstock and Needham, the primary tracheation being 
 increased by the development of cross-nervures, uniting to form a mesh work. 
 
 Family DICTYONEURID.E, Handlirsch. 
 
 1906. Handlirsch, Die Fossilen Insekten, p. 63. 
 
 1919. Handlirsch, Eevision der Palao/oischen Insekten, p. 3. 
 
 Palaeozoic insects in which the wings possess a close reticulated neuration 
 between the principal veins, the latter strong and parallel over the first third of 
 the wing. Branches of the radial sector, median and cubitus few in number, and 
 strongly curved back to the inner margin. 
 
 Handlirsch regards this family as closely related to Microdictya.
 
 DICTYONETJRA HIGGINSII. 25 
 
 Genus DICTYONEURA, Goldenberg. 
 
 1854. Dictyoneura, G-oldenberg, Palseontographica, vol. iv, p. 33. 
 
 Wing three times as long as wide. Base narrow, outer margin curving 
 regularly backwards beyond the middle of the wing into the wing-apex. Inner 
 margin rounded, and joining the apex in a blunt curve. Subcosta straight, 
 reaching the costal border in the outer third of the wing. Radius simple, parallel 
 to subcosta, and giving off a radial sector with few branches. Hinder branch 
 of median and first branch of cubitus uniting to form a " cubito-median Y-vein." 
 Anal veins with few branches. Interstitial neuration irregularly reticulate. 
 
 Dictyoneura higginsii (Handlirsch). Plate I, fig. 1. 
 
 1871. "Neuropterous Insect- Wing," Higgins, Proc, Liverpool Nat. Field Club, vol. ii, p. 18, fig. 15. 
 
 1906. (Pal&odictyopteron) higginsii, Handlirsch, Foss. Insekt., p. 125, pi. xiii, fig. 6. 
 
 1917. (Dictyoneuron) higginsii, Bolton, Quart. Journ. G-eol. Soc., vol. Ixxii, p. 46, pi. iii, fig. 2. 
 
 1919. Sherborniella higginsii, Handlirsch, Eevis. Paliiozoischen Insekten, p. 25. 
 
 Type. Basal portion of a left wing in counterpart ; Liverpool Museum. 
 
 Horizon aiul Locality. Middle Coal Measures; Ravenhead railway cutting, 
 near St. Helens, Lancashire. 
 
 Description. The wing-fragment lies on the surface of a greyish-purple 
 ironstone nodule, and has a length of 32 mm., with a greatest width of 22 mm. 
 The total length and width cannot be exactly estimated. 
 
 Handlirsch, who named the species, did not describe it, and it is doubtful 
 if he ever saw it. It was first described by the late Rev. H. H. Higgins thus : 
 "A second and smaller specimen of insect wing was obtained by myself j and 
 referred to the genus Gorudalu. Mr. J. P. G-. Smith compared it with Fulgora. 
 A slight sketch of it was seen by Mr. F. Smith, of the British Museum, whom 
 it reminded of Oryllotalpa. Mr. Benj. Cooke, of Manchester, after a careful 
 examination, says, ' I believe the fossil represents the basal portion, about 
 one-third only, of the forewing of a Chrysopa, Goldeneye, or Lace-wing fly, or 
 rather Nothochrysa, separated from Chrysopa by Mr. McLachlan on account of 
 the manner in which the third cubital cell is divided.' This cell is remarkably 
 well shown in the fossil, and though I could only judge from memory, I believe 
 it is sufficient to settle its relationship." 
 
 The "cubital cell" mentioned by Mr. Benj. Cooke is an elongated area 
 enclosed by a union of two of the main veins in the base of the wing. The 
 identity of the veins enclosing the " cell " will be considered later. 
 
 The costal border is preserved for a length of 29 mm., is strongly curved 
 proximally and less so distally. It would seem to have been directed somewhat 
 4
 
 26 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 backwards in its course to the wing-apex. The intercostal area is very wide 
 at the base, and diminishes in diameter towards the wing-apex, but it is doubtful 
 if it reached the latter. 
 
 The subcosta is straight, but not well defined. The radius is fairly parallel 
 to the subcosta and gives off the radial sector low down in the base of the wing ; 
 the radial sector comes off from the radius at an acute angle, and keeps parallel 
 with it as far as preserved. Its direction is such that it must have reached the 
 wing-apex. The median and cubital veins merit special consideration. In the 
 specimen three veins occupy the position of the normal median and cubitus. 
 Of these, two either arise from a common root, or so close together as to be 
 indistinguishable, the third having a separate origin. The innermost of the three 
 sends off from near its base a forwardly directed twig, which joins the second 
 of the two outer veins. 
 
 It is necessary to resolve these veins into median and cubitus. To do this 
 we must consider recent research on living insects. Comstock and Needham 
 concluded that the primitive median vein was a four-branched structure; more 
 recently Tillyard, from further studies of the wing-venation of recent nymphs, 
 concludes that the primitive median had an initial dichotomy, of which the outer 
 branch divided up into four (" M " of Tillyard, ' Proc. Linn. Soc. N.S. Wales,' 
 vol. xliv, 1919, p. 552), and the second branch remained simple. Comstock and 
 Needham concluded that the primitive cubitus was two-branched; Tillyard considers 
 it (loc. tit., p. 553) as three-branched, the outer or first branch having a distal 
 forking into two feeble twigs, and an inner feeble branch which remains simple. 
 The primary cubital fork is situated near the base of the wing. 
 
 Tillyard's views are not very different from those of Comstock and Needham, 
 as he admits a basal fork of the cubitus, but adds a secondary forking of the end of 
 the outer branch. 
 
 Tillyard has studied the relation of the two veins in the various Orders, and 
 finds that the " posterior arculus " of Comstock, which is supposed to be a cross- 
 vein from the median to the cubitus, is, as shown in the fossil Order Paramecop- 
 tera, not a cross-vein, but a true branch of the median. The various stages by 
 which the connection becomes established and afterwards developed into a 
 combined vein from the point of union, are fully stated (loc. cit., p. 357), and the 
 compound vein is named the " cubito-median Y-vein." An examination of the 
 conditions observable in Dictyoneum higginsii shows that the united veins and 
 their single-stemmed prolongation are in position and character identical with 
 Tillyard's " cubito-median Y-vein." The three veins of the specimen, therefore, 
 are the outer median vein, the inner median uniting with the first cubital vein, to 
 form a " cubito-median Y-vein," and lastly, the second (inner) branch of the cubitus. 
 
 Tillyard recognises the presence of the " cubito-median Y-vein " in the Permian 
 Order Paramecoptera, but its discovery in the Coal Measure genus Dictyoneum
 
 ORTHOCOSTA SPLENDENS. 27 
 
 shows that its origin is of much older date. The cubito-median Y-vein lies close 
 to the outer median for a short distance, and then bends obliquely inwards to the 
 margin of the wing. The inner cubital vein is fairly straight, and widely spaced 
 along almost its whole length from the cubito-median Y-vein. 
 
 Six anal veins are present, all going obliquely to the inner margin, the second 
 and third alone forking. The interstitial neuration, which I have been able to 
 study better by immersing the specimen in water, consists of numerous irregular 
 cross-nervures, occasionally uniting in crossing, or forming a loose meshwork, 
 especially between the median and cubital veins. 
 
 Affinities. With so small a wing- fragment, and with few wings of similar 
 character as guides, it is not possible to form an accurate idea of the whole wing. 
 The radius and radial sector entered the wing-apex, and the median-cubital 
 elements occupied the greater part of the inner margin of the wing. 
 
 The broad intercostal space and the course of the subcosta are very similar to 
 those in Polycreagra elegants, Handlirsch, but the general trend of the main veins, 
 and most of all the character of the interstitial neuration, are so suggestive of 
 Dicti/oneura libeUuloide*, Goldenberg, that I had referred the specimen to that 
 genus before Handlirsch's paper was published. 
 
 Handlirsch has now (1919) referred this species to a new genus, Sherboniiella, 
 giving the new name " for the sake of uniformity," but without diagnosis. 
 
 Family ORTHOCOSTID^E, Bolton. 
 
 1912. Boltou, Quart. Journ. Greol. Soc., vol. Ixviii, p. 313. 
 
 Wings with almost straight outer margin; inner margin well rounded; costa, 
 subcosta and radius closely approximated ; median, cubitus and anal veins 
 occupying nearly the whole of the inner half of the wing. 
 
 Genus ORTHOCOSTA, Bolton. 
 1912. Orthocosta, Bolton, toe. cit., p. 313. 
 
 Generic Characters. Radial sector diverging from radius, and with few 
 divisions. Median with two branches united by a commissure. Cubitus stout, 
 forked near base, the outer branch forking twice, and the inner once. Anal area 
 elliptical. Interstitial neuration forming an open polygonal meshwork. 
 
 Orthocosta splendens, Bolton. Plate I, fig. 2 ; Text-figure 1. 
 
 1912. Orthocosta splendens, Bolton, Quart. Journ. Geol. Soc., vol. Ixviii, p. 310, pi. xxxi, figs. 13. 
 
 Type. Incomplete wing, and impression; Moysey Collection, Museum of 
 Practical Geology, Jermyn Street (nos. 30,222 and 30,223).
 
 28 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 Horizon and Locality. Middle Coal Measures (below the Top Hard Coal) ; 
 Shipley Manor Claypit, Ilkeston, Derbyshire. 
 
 Specific Characters. Radial sector reaching the inner half of the wing-tip. 
 Outer branch of median four or five times divided, inner branch simpler and 
 forked. Cubitus dividing low down, the outer branch the stronger, and each 
 doubly forked. Anal veins one or two in number, alternately branched. 
 
 Description. The species is founded on a wing-fragment, the apex, a portion 
 of the inner margin, and the base being incomplete. The total length is 84 mm., 
 the width 33 mm. The complete wing must have had a length of at least 100 mm., 
 and a width of 3540 mm. The whole insect had in all probability a span of 
 wing measuring 25'5 cm. (10 in. or more). 
 
 The outer third of the wing is differentiated from the rest by its uniform and 
 gentle convexity, and by the character of the costa, subcosta, radius, and median 
 veins, which pass outwards towards the wing-apex in straight lines, and show no 
 trace of divisions until well beyond the middle of the wing, contrasting strongly 
 
 FIG. 1. Orthocosta splendens, Bolton; restoration of wing, showing portion preserved and the 
 character of the interstitial neuration, natural size Middle Coal Measures (below the Top 
 Hard Coal) ; Shipley Manor Claypit, Ilkeston, Derbyshire. Moysey Collection, Mus. Pract. Geol. 
 (nos. 30,222 and 30,223). 
 
 with the areas occupied by the marginal divisions of the median, cubital and anal 
 veins. The inner two-thirds of the wing is marked by deep furrows, in which lie 
 the marginal branches of the median vein, and the whole of the cubital and anal 
 veins. The areas between any two veins in this region are markedly convex. 
 The differences in character of the outer and inner portions of the wing are empha- 
 sised by a line of fracture which fairly accurately separates the two. Its occur- 
 rence suggests that it has followed a natural line of weakness, the more delicate 
 inner part of the wing breaking away from the outer stronger portion. 
 
 The costa, subcosta, radius and median are all well-developed veins, stout in 
 structure and standing out in relief. The first three retain this evidence of 
 strength over two-thirds of their length, the median vein showing signs of 
 attenuation beyond the proximal third. 
 
 The general structure of the Aving indicates considerable powers of flight. 
 The outer margin, of which only a portion is preserved, appears as a stout, slightly 
 elevated and well-rounded ridge. 
 
 The subcosta agrees in general character with the costal remnant, and is
 
 PTERONIDIID^E. 29 
 
 straight to the wing-tip, parallel with the outer margin, and not far removed from 
 it. Basally it appears to be united to the radius. 
 
 The radius gives off the radial sector at about 10 mm. from its base, and then 
 passes out to the wing-tip, at no point being more than 10 mm. from the subcosta. 
 The radial sector diverges widely from the radius, the two enclosing along, narrow 
 triangular area. At 50 mm. beyond its origin it divides into. two equal branches, 
 which diverge to a distance of 3 mm. and then become parallel. The direction of 
 the branches of the radial sector is such that they must have ended on the inner 
 half of the wing-apex, the outer branch probably forking again before the wing- 
 apex was reached. The median vein consists of two branches, the common origin 
 of which is not shown, owing to the absence of the base of the wing. To the 
 middle of the wing the two branches are parallel, and both pass beyond this 
 point before branching. The outer branch attains a length of 66 mm. before 
 branching, and then gives off four branches on its outer side, the last arising close 
 to the margin. 
 
 The second of the four branches bifurcates. The inner branch of the median 
 gives off one forward branch only, which forks into a feeble twig dying out in the 
 polygonal meshwork, and into a stronger division which reaches the margin. The 
 two branches of the median are united at the base of the wing by an oblique 
 commissural vein which comes off at an acute angle from the outer branch, and 
 passes obliquely to the inner branch. 
 
 The basal portion of the cubitus has been lost, so that its branching is not 
 readily determinable. The main vein seems to have divided near the base into 
 two equal branches, which curve down to the inner wing-margin, bifurcating 
 twice in each case before the margin is reached. The eight marginal twigs of the 
 cubitus thus produced enclose the middle third of the inner margin. The anal 
 area is wholly destroyed on the wing, and only a trace of one vein can be distin- 
 guished on the counterpart. This is a narrow deeply-sunk vein which gives off 
 feeble off-shoots on both sides. It diminishes in strength, so that the last portion 
 of its course can only be determined with difficulty. The anal area is comparatively 
 narrow and small, and exhibits few veins. 
 
 Family PTERONIDIID.E, Bolton (emend. Cockerell). 
 1912. Bolton, Quart, Journ. G-eol. Soc., vol. Ixviii, p. 314. 
 
 Wings long and narrow. Outer margin arcuate. Median and cubitus with 
 divergent branches. 
 
 The attempt to classify these wings is difficult. That they possess affinities 
 with the genus Polycreagra is certain, but they are more simple and more 
 Dictyoneurid in type. As Dr. Handlirsch observes, it is not possible to refer them 
 to the family Polycreagridge, and they must be taken as the type of a new family.
 
 30 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 Prof. T. D. A. Cockerell has pointed out to me that the family name was 
 originally wrongly written " Pteronidiae," and that the generic name was ante- 
 dated by Pteronidea, Rohwer, 1911. He thinks that the difference is sufficient to 
 justify retention, although it is only a single letter ('Proc. U.S. Nat. Mus.,' vol. xlix, 
 p. 469, 1915). Handlirsch ('Revision Palaozoischen Insekten,' 1919, p. 15) has 
 made the same correction, but seems unaware that Cockerell forestalled him. 
 
 Genus PTERONIDIA, Bolton. 
 1912. Pteronidia, Bolton, Quart. Journ. Geol. Soc., vol. Ixviii, p. 313. 
 
 Generic Characters. Wings three times as long as wide. Radial sector with 
 five simple, backwardly directed twigs. Median dividing lo\v down into two 
 principal branches, and ending on the inner margin in five twigs. Cubitus stout, 
 divided into two principal branches, the foremost with two twigs, and the hinder 
 ending in four. Interstitial neuration of thin oblique nervures. 
 
 Pteronidia plicatula, Bolton. Plate I, fig. 8 ; Text-figure 2. 
 
 1912. Pteronidia plicatula, Bolton, Quart. Journ. G-eol. Soc., vol. Ixviii, p. 313, pi. xxxii, figs. 1 3. 
 
 Type. Incomplete wing and impression ; Moysey Collection, Museum of 
 Practical Geology, Jermyn Street (nos. 30,224 and 30,225). 
 
 FIG. 2. Pteronidia plicatula, Bolton ; restoration of left wing, natural size. Middle Coal Measures 
 (below Top Hard Coal) ; Shipley Manor Claypit, Ilkeston, Derbyshire. Moysey Collection, Mus. 
 Pract. Geol. (nos. 30,224 and 30,225). 
 
 Horizon and Locality. Middle Coal Measures (below the Top Hard Coal) ; 
 Shipley Manor Claypit, Ilkeston, Derbyshire. 
 
 Specific Characters. Wings triangular in outline, with subacute tip, strongly 
 plicated. Radial sector dividing into five simple twigs. Median with two prin- 
 cipal branches and five marginal twigs. Cubitus wide-spaced; two principal 
 branches, the inner doubly forked. 
 
 Description. This is a long delicate wing, of which the outer marginal portion 
 and the base are missing. It was probably about 7075 mm. long, with a basal 
 width of 20 mm., reduced to 8 mm. near the tip. 
 
 The outer margin must have been flatly convex, while the inner margin is
 
 HYPERMEGETHID^E. 31 
 
 nearly straight. The tip of the wing is acute. The costal and subcostal veins are 
 not seen. The radius is represented by a single forked vein, not shown in the 
 figures. The radial sector is unbranched until well beyond the middle of the wing, 
 beyond which it gives off four (possibly five) backward branches, which occupy 
 the greater part of the tip of the wing and a portion of the distal inner margin. 
 
 The median is represented by two branches, the outer dividing into two twigs 
 and the inner into three. 
 
 The marginal area occupied by the median is a little longer than that of the 
 radial sector, and the two extend over the distal half of the inner wing-margin. 
 
 The cubitus is a relatively short and stout vein, dividing into two main 
 branches, the foremost having two twigs, and the hinder dividing into four by a 
 double bifurcation. Owing to the wide divergence of its branches the cubital 
 area is larger than that of the median. The anal area was large, but the whole 
 surface has been pitted by attempts to clear the matrix, and as a result the 
 presence and character of the veins cannot be distinguished with certainty. It is 
 possible that certain surface-indications are evidence of two simple widely 
 separated veins. The wing-surface exhibits a strong plication. 
 
 Viewed obliquely, the wing shows a series of ridges formed by the vein with 
 V-shaped intervening sulci, which only flatten out close to the wing-margin. 
 Springing from the principal veins is a close series of fine cross-nervures, obliquely 
 disposed in the direction of the wing-margin. The portion of the wing preserved 
 is sufficient to indicate that as a whole it was triangular in outline. 
 
 Family HTPERMEGETHID^E, Handlirsch. 
 
 1906. Handlirsch, Proc. U.S. National Museum, vol. xxix, p. 672. 
 
 Costa marginal, costal area broad ; radius simple, radial sector present ; the 
 median probably dividing at the base into two or more main branches, the first 
 of which may become united to the radius. Cubitus forked near the base, with its 
 branches widely spaced. Anal veins few, and anal area not exceeding one-third 
 of the inner margin. 
 
 Handlirsch established this family for the inclusion of a gigantic wing of which 
 only the basal half is known. The total length of the whole wing was estimated 
 to be 120 mm., the basal half having a length of 60 mm. The discovery in the 
 Coal Measures of Durham of a nearly similar wing adds to our knowledge of the 
 group, and enables the family to be defined with more accuracy than was at first 
 
 possible. 
 
 The general assemblage of characters found in the Hypermegethidae is, I 
 believe, highly suggestive of the Protodonata, but no definite conclusions can be 
 formulated until the whole wing is known.
 
 32 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 Genus HYPERMEGETHES, Handlirsch. 
 1906. Hypermegethes, Handlirsch, Proc. U.S. National Museum, vol. xxix, p. 672. 
 
 Generic Characters. Costal border feebly convex, subcosta and radius close 
 together along the greater part of their length. Ctibitus composed of two, 
 possibly more, parallel and simply forked branches, divided near the point of 
 origin. Anal veins few ; interstitial nenration of fine, irregularly anastomosing 
 nervures. 
 
 Hypermegethes northumbriae, Bolton. Plate I, fig. 4; Text-figure 3. 
 
 1917. Hypermegethes northumbriie, Boltou, Quart, Journ. G-eol. Soc., vol. Ixxii, p. 55, pi. iv, figs. 2 
 and 3, and woodcut in text. 
 
 Type. Portion of the basal half of a left wing, in counterpart; British 
 Museum (no. In. 18524). 
 
 FIG. 3. Hypermegethes northumbrise, Bolton ; suggested outline of left wing, restored, showing portion 
 preserved, slightly less than natural size. Coal Measures (shale above the Crow Coal) ; Phoenix 
 Brickworks, Crawcrook, Durham. Brit. Mus. (no. In. 18524). 
 
 Horizon and Locality. Coal Measures (shale above the Crow Coal); Phoenix 
 Brickworks, Crawcrook, Durham. 
 
 Specific Characters. -Costal area wide, and crossed by an irregular meshwork 
 of small veins. Median united with the radial sector basally, and giving off an 
 inner branch which unites with the cubitus. Cubitus with a short stem forking 
 into two equal and widely separated branches. Anal veins simple, and widely 
 spaced. 
 
 Description. The fragment is little more than a third of the whole wing, and 
 its characters can only be determined with difficulty. The two halves of the 
 nodule do not coincide, and the outline drawing of the wing is built up from 
 details supplied by both halves. The inner margin, anal area and base of the 
 wing have been lost, so that a little less than two-thirds of the outer portion of 
 the basal half of the wing is present. 
 
 The portion of wing remaining being 63 mm. long, with a depth of 31 mm. at 
 its widest part, the whole wing must have had a length of about 126 mm., or
 
 HYPERMEGETHES NORTHUMBRLE. 33 
 
 5 in., and the whole insect a span of wing of nearly 11 in. The wing-fragment 
 shows the basal portions of the costa, subcosta, radius, radial sector, median, and 
 cubitus, and possibly a trace of an anal vein. 
 
 Much of the finer detail of the wing is not seen until the fossil is immersed in 
 water a mode of treatment suggested to me by Dr. F. A. Bather, F.R.S., who had 
 previously photographed the wing in this manner, and brought out details of 
 which, by ordinary methods, I could not find a trace. 
 
 The costa is moderately convex from its base to a distance of about 30 mm., 
 beyond which it becomes straight. Separated from the outer or costal border by 
 a wide area basally is the subcosta, an extremely feeble and hardly distinguishable 
 vein. It passes straight to the outer margin some distance beyond the middle of 
 the wing. The radius arises close to the subcosta, and is parallel with it. It gives 
 off two branches posteriorly, the proximal branch passing obliquely towards the 
 inner side of the wing-apex, while the second or distal branch arises from the 
 radius a little further out, and keeps parallel with it. 
 
 I had formerly considered the proximal branch of the radius to be the radial 
 sector, and the distal a branch of the radius ; but as the interstitial neuration now 
 shows the specimen to be closely related to Hypermegefhes schucherti, the proximal 
 branching vein must be regarded as the main stem of the median which has entered 
 into union with the radius, and the distal branch as the radial sector. 
 
 Regarding the proximal offshoot of the radius as the median vein, it diverges 
 widely from the radius, giving off a forward twig parallel with the radial sector, 
 and then continues inwards and unites with the next vein, separating again a little 
 further out on the inner side. A comparison of this assumed median vein with 
 that of H. sclmcherti, Handl., is instructive. In the latter species the branching 
 of the main stem of the median arises nearer the base of the wing than does the 
 branching off of the radial sector. It is therefore somewhat in the position of 
 the starting-point of the median vein as a free structure in the present wing. The 
 median vein of 11. schucherti, Handl., has, however, no union with the radius, the 
 main stem running out towards the wing-apex parallel with the inner branch of 
 the radius, giving off a backward branch of the cubitus, which passes obliquely 
 inwards and unites with the anterior branch. In the middle of its length it gives 
 origin to a twig running parallel with the main stem, and midway between 
 that vein and the cubitus. The course of the median vein in the specimen 
 here described is exactly similar to that of the branch of the median vein in 
 H. schucherti, Handl., except that the inner branch not only unites with the 
 cubitus, but crosses it. 
 
 The condition may be summarised by saying that, in H. schucherti, Handl., the 
 median vein is entirely free and gives off an inner branch dividing into two twigs, 
 of which the inner unites with the cubitus. In the specimen here described the 
 median is united with the radius for some little distance, becoming free before the
 
 34 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 radial sector is reached, and then forking, the inner twig uniting with and crossing 
 the cubitus. 
 
 The cubitns has a short, stont basal stem, forking into two equal and somewhat 
 widely separated branches, the outer uniting with the inner branch of the median. 
 The inner branch is forked just before the broken edge of the wing is reached. 
 Nearer the inner margin of the wing are traces of two other veins. The first has 
 its basal portion missing, and follows a course parallel with the inner branch of 
 the cubitus. It is strongly forked. The remaining vein is represented by three 
 detached fragments. If Handlirsch's interpretation of the wing of H. sehucherti 
 is followed, we should regard both these veins as anal. I am, however, of opinion 
 that, while the innermost fragmentary vein may be anal, the forked vein by its 
 position, its forked character and stoutness, must be regarded as a portion of the 
 cubitus. I am likewise of opinion that the first, and possibly the second, of the 
 veins marked as anal in Handlirsch's figure of H. schncherti, ought also to be 
 classed as cubital. Both in H. schucherti, and in this specimen, the anal area 
 would have an enormous development and occupy most of the wing-margin, if the 
 veins alluded to were wholly anal in character. I feel sure that, if the vein nearest 
 the cubital had been better preserved, it would be found branching off from the 
 cubitus. 
 
 The interstitial neuration is typically that of Hypermegethes. The intercostal 
 area is filled with an irregular meshwork of fine nervures, with a tendency on the 
 outer and inner sides to a transverse arrangement. Between the median and the 
 cubital veins the interstitial neuration consists of short, straight and transverse 
 nervures, and feeble traces of similar nervures can be seen in the median area. 
 The cubital area is filled with a meshed neuration, larger and more regular than 
 that of the intercostal area, and this seems to continue into the anal area. 
 
 Family CRYPTOVENIID^;, Bolton. 
 1912. Bolton, Quart. Journ. Geol. Soc., vol. Ixviii, p. 315. 
 
 Wings short and broad. Apex rounded, costal vein marginal ; subcosta feeble, 
 and extending to near apex. Radius simple ; radial sector and median with few 
 divisions ; cubitus with two main branches. 
 
 Among the insect-remains discovered by the late Dr. L. Moysey at the Shipley 
 Manor Claypit is a small wing, 16 mm. long, unlike any previously known. It 
 is typically Palaeodictyopterous, and agrees remarkably well with Dr. Handlirsch's 
 type-figure ('Mitth. Geol. Gesell. Wien,' vol. iii, 1910, p. 505, fig. 1). It differs from 
 that form in the greater division of the cubitus, which ends in five twigs instead of 
 three. The greatest depth of the wing was also, in all probability, nearer the base 
 than in his figure. With the genus Athymodictya, Handlirsch (< Amer. Journ. Sci.'
 
 CRYPTOVENIA MOYSEYI. 35 
 
 [4], vol. xxxi, 1911, p. 298), the relationship is even closer, as in that genus the 
 costa and subcosta are close together, the radial sector arises low down and is 
 divergent from the radius, the median is a simple vein ending in three branches, 
 while the cubitus is almost identical in its divisions, the difference being that the 
 first forking arises at a higher point than that in Athymodicti/a parva, Handl., and 
 that the inner simple ramus of the outer branch comes off a little below the middle 
 of the wing. The anal veins number four in A. parva as against five in the wing 
 under consideration. The wings are almost equal in size. It is not possible, 
 however, to refer this wing to Athymodicti/a owing to the character of the interstitial 
 neuration. In Athymodictya this is reticulated, while in the British wing the 
 interstitial neuration is apparently made up of transverse nervures. If this point 
 can be clearly determined, the wing may have affinities with the Homoiopteridse 
 or with the Lithomantidas, but it cannot be referred to either of these families. 
 Xeither Dr. Handlirsch nor I could satisfactorily refer the wing to any known 
 family, and I therefore founded the family Cryptoveniidas to receive it. 
 
 Genus CRYPTOVENIA, Bolton. 
 1912. Gryptovenia, Bolton, Quart. Journ. Geol. Soc., vol. Ixviii, p. 315. 
 
 Generic Characters. Wings twice as long as wide. Median, cubitus and anal 
 veins curving sharply inwards. Interstitial neuration of straight cross-nervures. 
 
 Cryptovenia moyseyi, Bolton. Plate II, fig. 1 ; Text-figure 4. 
 
 1912. Cryptovenia moyseyi, Bolton, loc. cit., p. 315, pi. xxxii, figs. 4 6. 
 
 Type. Incomplete wing, in counterpart; Moysey Collection, Museum of 
 Practical Geology, Jermyn Street (nos. 30,226 and 30,227). 
 
 Horizon and Locality. Middle Coal Measures (below the top Hard Coal) ; 
 Shipley Manor Claypit, Ilkeston, Derbyshire. 
 
 Specific Characters. Costa feebly curved distally, subcosta reaching outer 
 margin near wing-apex. Radius close to and parallel with the subcosta; radial 
 sector arising from radius in basal half of the wing, the five branches occupying 
 the inner half of the wing-tip. Median with the first branch undivided, and the 
 second forking twice. Cubital vein strongly arcuate, two-branched, and ending m 
 five twigs. Anal veins five or more in number, and curving sharply inwards. 
 Wing plicated, and the interstitial neuration obscured by a mass of wrinkles. 
 
 Description. A small delicate wing, incomplete at the base, which must have 
 been very narrow and with the main veins crowded together. The total length of 
 wing preserved is 16 mm., and its maximum diameter, in the cubito-anal region, 
 is 8 mm.
 
 36 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 The costal vein is stout and raised above the general level of the wing in the 
 outer third of its length, where it curves backwards into the wing-tip. The 
 subcostal is a weaker vein, parallel with and very close to the costa, dying out or 
 joining the outer margin at the apex. The radius is a powerful undivided vein, 
 parallel with the subcosta and reaching the middle of the wing-apex. The radial 
 sector is given off about the middle of the length of the wing and forks into two 
 equal branches, which again fork before reaching the margin ; the outermost branch 
 of the second forks again and divides just before the wing-apex is reached. The 
 
 FIG. 4. Crypiovenia moyseyi, Bolton ; restoration of left wing reconstructed from the wing-fragment 
 and counter-impression, enlarged two-and-a-half times. Middle Coal Measures (below the Top 
 Hard Coal) ; Shipley Manor Claypit, Ilkeston, Derbyshire. Moysey Collection, Mus Pract. Geol 
 (nos. 30,226 and 30,227.) 
 
 inner half of the wing-apex is occupied by the five branches of the radial sector. 
 The median vein is a comparatively simple structure, forking low down below the 
 middle of the wing into two nearly equal branches. The outer branch remains 
 undivided, and gently curves to the inner margin. The inner branch divides 
 twice, first at a point near the middle of the wing, and again before the margin 
 is reached. The median vein therefore ends on the margin in four twigs, three 
 members of which arise from the inner of the two main branches. 
 
 The cubital vein is strongly arcuate, dividing near the base into two branches, 
 the outer forking twice and the inner once. The cubital vein therefore ends in five 
 twigs. 
 
 Family MECYNOPTERIDJ;, Handlirsch. 
 1906. Handlirsch, Die Fossilen Insekten, p. 82. 
 
 Handlirsch established this family to receive a large wing from the Middle 
 Upper Carboniferous of Belgium, and placed it between the families Hyperme- 
 gethidae and Lithomantidae. The characters of the family are based on those of 
 the type-species, Mecynoptera splendida, Handl. 
 
 Genus MECYNOPTERA, Handlirsch. 
 
 1904. Mecynoptera, Handlirsch, Mein. Mus. Boy. Hist. Nat. Belg., vol. iii, p. 7. 
 1906. Mecynoptera, Handlirsch, Die Fossilen Insekten, p. 82.
 
 MECYNOPTERA TUBERCULATA. 37 
 
 Generic Characters. Wing three to four times as long as broad, veins of costal 
 region specially compact, and thickened basally. Costa, subcosta and radius 
 closely approximated and tuberculated. Radial sector and median well developed, 
 and occupying a considerable portion of the wing-surface. Cubitus small. Inter- 
 stitial neuration of transverse nervures in junction areas of principal veins, and 
 with a meshwork further out. 
 
 Mecynoptera tuberculata, sp. nov. Plate II, fig. 2 ; Text-figures 5 and 6. 
 1911. Stenodictya lobata, Baldwin (errore), Greol. Mag. [5], vol. viii, p. 75. 
 
 Type. Portions of two fore-wings, and the cubito-anal portion of a hind-wing 
 contained in a nodule of ironstone, 3 in. long, and 1^ in. wide; British Museum 
 (no. In. 18,576). 
 
 Horizon and Locality. Middle Coal Measures (grey-blue shales at 135 180 
 feet above the Royley or Arley Mine) ; Sparth Bottoms, Rochdale, Lancashire. 
 
 Specific Characters. Principal veins thickened at base and finely tuberculated. 
 Costal margin almost straight, much thickened at base, and covered with a fine 
 
 FIG. 5. 
 
 FIG. 6. 
 
 FIG. 5. Mecynoptera tuberculata, sp. nov. ; diagram of remains of the two fore-wings, and cubito-anal 
 portion of a hind-wing (the sub-costa appearing as a fine line close to the radius, and sending off 
 three branches to the costal margin), enlarged one-and-a-half times. Middle Coal Measures 
 (above the Royley or Arley Mine) ; Sparth Bottoms, Rochdale, Lancashire. Brit. Mus. (no 
 In. 18,576). 
 
 FIG. 6. Mecynoptera tuberculata, sp. nov. ; diagram of fragmentary impression of the two fore-wings, 
 enlarged one-and-a-half times. Middle Coal Measures (above the Royley or Arley Mine) ; Sparth 
 Bottoms, Rochdale, Lancashire. Brit. Miis. (no. In. 18,576). 
 
 tuberculation. Subcosta sunken, close to costa, and extending to the wing-apex. 
 Radius close to costa, tuberculated, and elevated. Radial sector arising low down, 
 much branched. Median with two main branches, the first simple, the second 
 with three twigs. Cubital vein small. 
 
 Description. The type-specimen comprises remnants of the two fore-wings, 
 60 mm. long and 18 mm. wide, and a portion of the cubital-anal area of a single 
 hind-wing in one ironstone nodule. A similar ironstone nodule from the same 
 horizon and locality shows the radius and radial sector areas of the two wings of a 
 second insect.
 
 38 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 When complete, the wings must have had a length of 05 70 mm. They were 
 originally identified with those of Stenodictya lubata, Brongniart, and are quoted as 
 such in the faunal lists of the Sparth Bottoms deposits, but with no detailed 
 description or figures. 
 
 When the specimens were first examined they presented an apparently anoma- 
 lous union or suppression of certain of the principal veins. Careful development 
 has since shown that what was formerly considered to be the outer or costal 
 margin of the wings was really the thickened and tuberculated radius vein, and 
 that the outer margin, with the costa and subcosta, had been hidden under the 
 matrix. A little of the outer marginal costa and the subcosta are now uncovered, 
 and the wing-structure is therefore proved to be of a normal type. 
 
 The three wings in the larger, more nearly complete specimen are superposed 
 and fragmentary. The uppermost fore-wing is represented by two portions, one 
 having a little of the outer or costal margin, the subcosta, and the greater part of 
 the radius and radial sector. The second part consists of the median,, and a 
 portion of the cubitus. The second fragment is displaced backwards, allowing 
 the radius and radial sector of the second fore- wing to be seen. The median- 
 cubital area of the hind-wing is very thin and closely pressed on the rest, but the 
 course of the veins is clearly discernible, and by their sharp inward turn indicates 
 that the hind-wings were much broader than the fore-wings. 
 
 Although the wings are thus broken up, displaced, and superposed, the parts 
 missing in the one wing are present in the other, and it is possible to reconstruct 
 their general character. The outer margin is almost straight, gently rounded into 
 the base, and into the wing-apex. The subcosta is weak, and lies in a deep sulcus, 
 extending into the apex of the wing. The radius is a little elevated, thickened, 
 tuberculated in the basal third, and in close proximity to the subcostal vein. The 
 radius is a strong vein, raised above the level of the rest of the outer margin of 
 the wing, so that when the latter was hidden, it naturally appeared to be the outer 
 marginal costal vein. It is much thickened in its basal third, tuberculated, and 
 remains parallel with the subcosta in the apex of the wing. 
 
 The wing-space taken up by these three veins is very small, and their close 
 approximation and the coriaceous thickening of the costa and radius serve to give a 
 considerable degree of strength and rigidity to the outer margin of the wing. The 
 radial sector passes straight outwards to the apex of the wing, giving off three inner 
 branches in one wing and two in the other, the first branch only forking. The first 
 branch soon forks, the outer fork dividing into two twigs, and the inner into three 
 upon the distal inner half of the wing-apex. The median is a large vein, occupying 
 the greater part of the inner half of the wing. It divides at its point of origin into 
 an outer undivided branch which traverses the middle of the wing, and an inner 
 branch which is widely separated from its fellow, and gives off three twigs to the 
 wing-margin, the first forking in the middle of its length. Cubital veins are repre-
 
 MECYNOPTERA TUBERCULATA. 39 
 
 sen ted by two undivided elements which are directed obliquely to the margin. 
 The inner margin of the wing is more curved than the outer, and merges into the 
 apex. The interstitial neuration is remarkable. Between the costa, subcosta and 
 radius it consists of short stout nervures crossing the areas somewhat obliquely, 
 and in some cases arranged in V-shape. The area between the radius and radial 
 sector is crossed by a numerous S-shaped series of 'nervures, which are joined up 
 into a meshwork in the outer or radial half of the area. The area itself is very 
 wide, and only equalled by that separating the two branches of the median. 
 
 The radial sector and the median occupy the greater part of the wing- 
 surface. 
 
 The remaining areas are crossed by transverse nervures near the junction of 
 branches with the principal veins, and further out by nervures which are joined up 
 by zig-zag longitudinal branches which occasionally enclose polygonal cells. The 
 interstitial neuration is very well developed, and must have added materially to 
 the strength of the wings. 
 
 Affinities. The close approximation of the costa, subcosta and radius, and the 
 coriaceous thickening and tuberculations of the principal veins, are characters 
 which may rank as of generic importance. Less distinctive, but also characteristic, 
 is the interstitial neuration of the transverse nervures and meshwork, with its 
 minute monilation. 
 
 The characters of the genus Mecynoptera are in closest agreement with those 
 of this insect. There is the same approximation of the costa, subcosta and radius, 
 the radial sector is widely spaced from the latter, and the interstitial neuration 
 consists of transverse nervures near the junctions of the principal veins and their 
 branches, and of a meshwork further out. The general outline of the wing is also 
 the same. 
 
 M. s-plendida, Handlirsch, throws light on one point which had proved a 
 difficulty in the determination of the wings. It has the radial sector of large 
 size and much branched, the divisions occupying all the inner half of the apex of 
 the wing, and extending out on the inner margin into the area usually occupied by 
 the outer branches of the median. Prior to noting this feature in M. splencluli, 
 I had formed the opinion that the first division of the radial sector was the median, 
 which had united with the radial sector and the radius. 
 
 The broken condition of the Rochdale wings prevents absolute determination 
 of this point, but by analogy I conclude that the whole vein which branches off 
 from the radius basally is the radial sector, and the next vein the median. This 
 conclusion also removes the insect from the neighbourhood of ^Enigmatodes danielsi, 
 Handlirsch, a wing discovered in the Coal Measures of Mazon Creek, near Morris, 
 Illinois, U.S.A., which possesses the same character of interstitial neuration, but 
 with the divisions of the median vein stretching out to the wing-apex, and the 
 costa, subcosta and radius more widely separated.
 
 40 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 Unless it can be shown by the discovery of a whole wing that the reconstruction 
 now attempted is faulty, the balance of evidence is in favour of the provisional 
 reference of the Sparth Bottoms wing to the genus Mecynoptera. 
 
 Family INCERTJ: SEDIS. 
 Genus PAUEOMANTIS, Bolton, 
 1917. Palxomantis, Bolton, Quart, Jouru. Geol. Soc , vol. Ixxii, p. 52. 
 
 Generic Characters. Wings short, twice as long as broad. Apex well rounded. 
 Radius, radial sector and median all powerful veins with few divisions. Median 
 with two main branches. Anal veins directed almost straight inwards. Interstitial 
 neuration forming an irregular meshwork. 
 
 Palseomantis macroptera, Bolton. Plate II, fig. 3 ; Text-figures 7 and 8. 
 
 1871. "Wing of large insect," Higgins, Pres. Add. Liverpool Naturalists' Field Club, vol. ii, p. 18. 
 1917. Palxomantis macroptera, Bolton, Quart. Jouru. Geol. Soc., vol. Ixxii, p. 48, pi. hi, figs. 34, 
 text-figs. 23. 
 
 Type. Remains of two wings in nodule ; Liverpool Museum. 
 
 Horizon and Locality. Middle Coal Measures ; Ravenhead railway cutting, 
 near St. Helens, Lancashire. 
 
 Specific Characters. Divisions of radial sector occupying almost the whole tip 
 of the wing. Divisions of median and cubital veins occupying distal two-thirds of 
 inner margin. 
 
 Description. The larger half of the nodule has lost a portion which contained 
 the tip of the right wing. The proximal third of the left wing remains, with its 
 dorsal surface uppermost, and its ventral surface closely applied to that of the 
 right wing. It is evident that the whole of the two wings was contained in the 
 nodule, but, when the latter was split open, a thin film of ironstone carried away 
 the middle and distal portions of the left wing. 
 
 One unusual feature in the position of the wings is that they lie with their 
 ventral surfaces apposed. To bring them into this position one must have become 
 bent under the body, instead of falling sideways across the thorax of the insect. 
 The body of the insect would thus, if the wings still remained attached, lie between 
 the two. No trace of the body can be seen, but the left wing has a deep inward 
 flexure, such as it would naturally acquire if the body had been carried round with 
 the right wing and pushed into the anal area of the left wing. Had the body been 
 carried round in this way the right wing would not coincide in position with the 
 left, but be thrust further out, which is actually the case, the outward displacement
 
 PALuEOMANTIS MACROPTERA. 
 
 41 
 
 of the right wing as compared with that of the left being at least 20 mm. The 
 wings are remarkably wide a feature which we usually associate with hind-wings. 
 
 The outer third of each wing is supported by strong and fairly rigid veins, 
 becoming more slender as they pass towards the wing-tip. The inner margins of 
 the wings are more membranous, and were evidently lacking in a rigidity equal to 
 that of the outer margins. 
 
 A little more than the proximal third of the left wing is present and in good 
 preservation. It is 42 mm. long and about the same in width. The impression 
 on the other half of the nodule shows apparently all but the wing-apex. The main 
 stems of the costa, subcosta, radius and median are all stout, and in relief on the 
 wing-surface. The cubital and anal veins are but. half the thickness of the former and 
 lie in shallow grooves. The costal vein forms the outer margin of the wing, which 
 is slightly convex forwards, and slopes gradually into the apex. The subcosta is 
 
 FIG. 7. 
 
 Fia. 8. 
 
 FIG. 7. Palseomantis macroptera, Bolton ; restoration of left wing, with restored outline, natural size. 
 Middle Coal Measures; Eavenhead Eailway Cutting, near St. Helens, Lancashire. Liverpool 
 
 FIG. 8. Palxomantis macroptera, Bolton; restoration of right wing, natural size. Middle Coal 
 Measiires ; Eavenhead Eailway Cutting, near St. Helens, Lancashire. Liverpool Museum. 
 
 separated from the outer margin by a costal area 4 mm. broad. It passes almost 
 in a straight line to the outer edge of the wing-apex, and is joined to the costa by 
 a series of oblique transverse branches nearly parallel with each other. The radius 
 arises close to the base of the subcosta, and diverges a little from it in its course 
 to the apex, forking once just before the broken edge of the wing is reached. The 
 main stem of the radial sector probably arises from the radius close to the base of 
 the latter, but the exact point cannot be determined, owing to the base of the wing 
 being broken away. The radial sector forks three times, and each of the resultant 
 twigs again divides on the wing-apex. The median vein, which is stout basally, 
 divides very low down into two branches, the outer of which has a feeble forking 
 near the inner margin, while the inner branch forks three times. The cubitus is 
 a much weaker vein than its fellows, and divides near its point of origin into two 
 branches. The first branch forks once and the second twice. Seven anal veins 
 are present, all undivided except the third, which forks twice. The interstitial 
 6
 
 42 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 neuration is in the form of a meslnvork, except in the intercostal area, where are 
 the transverse cross-uervures already noted. 
 
 Much less of the riylit n-uicj remains than of its fellow, but sufficient is present 
 to show that the neuration was not quite the same. The median ends in five 
 divisions in place of six, and the cubitns has seven final branches instead of six. 
 The greatest width of the wing is along a line drawn from the outer margin to the 
 middle of the cubital area on the inner margin, the width being 40 mm. The 
 absence of the apex in each wing is unfortunate, as it renders the outline of the 
 whole wing a little uncertain. The shape of the nodule indicates that very little 
 of the wings is missing, if they were wholly included in the nodule, as seems 
 probable. The somewhat semicircular inner wing-margin and the short wide 
 wings indicate a broadly rounded wing-apex. 
 
 Affinitie*. The, interstitial neuration is much like that of Hypermegetltes. The 
 great width of the wings is a character usually associated with the hind- wings of 
 members of the family Lithoinanticlse, although in this case the wings lack the 
 distal attenuation noticed in the hind-wings of that family. With Titanodicti/a 
 jvcnuda, Scudder, there is a close relationship, both in the general character of 
 the main veins, the interstitial mesh work in all areas other than intercostal and 
 subcosta-radial, and in the presence of the same oblique cross-nervures in the 
 intercostal area in these wings. 
 
 The true systematic position of the genus seems to lie between the Li thorn an - 
 tidse and the genus Titanodictya, and closest to the latter. Because of the greater 
 development of a mesliAvork neuration between the main veins, and the limitation of 
 cross-nervures to the intercostal area, I regard this genus as more primitive than 
 any of the Lithomantidse, but closely related thereto. 
 
 Family LITHOMANTIRE, Handlirsch. 
 1906. Handlirsch, Proc. U.S. National Mus., vol. xxix, p. 673 ; also Die Fossilen Insekten, p. 82. 
 
 This family is closely allied, by wing-structure, to the Dictyoneurida3. The 
 branching of the main veins has proceeded further than in the Dictyoneuridae, and 
 the body, where it has been preserved, shows striking differences. The family is 
 represented in the Coal Measures of Great Britain and North America and in the 
 Upper Carboniferous of continental Europe. 
 
 Genus LITHOMANTIS, Woodward. 
 1*70. Lifliomntitis, H. Woodward, Quart. Jouru. Geol. Soc., vol. xxxii, p. 60. 
 
 Generic Characters. Large insects with two pairs of flying wings ; the hinder 
 pair double the width of the anterior pair. Prothorax produced into a central
 
 LITHOMANTIS CARBONARIUS. 43 
 
 rostral-like process, acutely pointed, and expanded into two lateral bulbous lobes. 
 Thorax wide. The interstitial neuration consists usually of stout transverse 
 nervures, occasionally forking or in a loose meshwork. 
 
 Lithomantis carbonarius, Woodward. Plate II, fig. 4; Text-figure 9. 
 
 1876. Lithomantis carbonarius, Woodward, loc. cit., p, 60, pi. ix, fig. 1. 
 
 1893. Lithomantis carbonarius, Bronguiurt, Fanne En torn. Terr. Prim., p. 489, fig. 
 
 1906. Lithomantis carbonarius, Woodward, Geol. Mag. [5], vol. iii, p. 25, fig. 3 (noti fig. 1). 
 
 1906. Lithomantis carbonarius, Handlirsch, DieFossilen Iiisekten, p. 83. 
 
 Type. Portions of the fore- and hind-wings, with prothorax, and a left anterior 
 leg in nodule ; British Museum (no. I. 8118). 
 
 Horizon and Locality. Coal Measures; Scotland. 
 
 Specific Characters. Hind pair of wings double the width of the fore pair. 
 Outer wing-margin straight, costa and subcosta close together ; radius a weak 
 vein, giving off the radial sector far out. Median small. Cubitus a powerful 
 vein Avith numerous widely spaced branches. Anal veins six or more. 
 
 FIG. y.Liihomantis carlanarius, Woodward ; diagram of neuration of the left fore- and hind-wings, 
 natural size. Coal Measures ; Scotland. Brit. Mus. (no. I. 8118). 
 
 Description The type-specimen lies on the surface of one half of an ironstone 
 nodule, the counterpart being lost. It was obtained by Mr. Edward Charlesworth 
 from the Coal Measures of Scotland, but the locality and horizon are not known. 
 
 The remains consist of portions of two pairs of wings, those of the left side 
 being most nearly complete. In no case is the apex of the wing or the inner 
 margin preserved. Lying in front of the wings are a pair of very large convex 
 lobes which Dr. Woodward regards as part of the prothorax, and in front of these 
 is a roughly quadrangular structure prolonged forward in the middle line into a 
 styliform process or rostrum. Woodward describes the latter as " the small head
 
 44 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 with its eyes," but no definite trace of the latter are observable. The " head " is 
 small, not more than 7 mm. wide, and between 56 nun. long at the sides ; 
 medially it is prolonged into the rostral process, which is about 89 mm. long 
 and ending in a sharp point. I am unable to distinguish any dividing line between 
 the "head" and the prothorax ; on the other hand, the margin of the latter is 
 continuous with and inseparable from it. The marking which Woodward has 
 regarded as representing eyes is, I believe, the thickened margin. I am of opinion 
 that the whole structure is wholly prothorax, and that the head lies concealed 
 beneath. The main mass of the prothorax is 30 mm. wide, with a flattened margin, 
 best seen on the left side. Within the flat margins rise two low dome-shaped 
 lobes separated. by a wide hollow in front, their margins meeting in an obtuse 
 angle posteriorly. The median edge of each lobe dips sharply into a wide median 
 hollow, and from each of these edges arises a series of veins, which spread out to 
 the lateral margins of the lobes. The areas between the veins are occupied by a 
 fine mesh work of smaller veins. 
 
 A trace of the mesothorax is shown between the bases of the fore-wings as a 
 slight transverse bar, a small rounded tubercle lying in front and a little to the 
 left of the middle line. 
 
 The left fore- and hind-wings are the most nearly perfect, the hind-wing being 
 56 mm. long, with a greatest width of 30 mm. The fore-wing is a little shorter 
 and narrower. The outer margin appears to have been straight, and the costa 
 and subcosta closely approximated. Traces of both veins are present. The radius 
 is a straight thin vein not far removed from the subcosta along its whole length, 
 and giving off the radial sector beyond the middle of the wing. The radial sector 
 comes off at an acute angle, going out to the wing-apex and keeping closely 
 parallel with the radius. The median is somewhat inconspicuous owing to the 
 great length of the main stem, and the narrow areas which bound it between the 
 radius and the prominent cubitus. It gives off two outer branches before reaching 
 the broken edge of the wing. The median vein of the hind-wing has three 
 outer branches, the first arising nearer the base than the point of origin of the 
 radial sector, whereas in the fore-wing the first branch arises distally to the origin 
 of the radial sector. The median of the hind-wing is a more important vein than 
 its fellow in the fore- wing and occupies a greater area owing to its greater inward 
 curvature. The cubitus is a powerful vein with widely spaced divisions, the first 
 branch, both in fore- and hind-wings, coming off from the main stem on the outer 
 side, and low down near the base of the wing, and then passing in a bold convex 
 sweep down to the distal portion of the inner margin. On its inner side the 
 cubitus gives off five branches, the fourth forking in the middle of its length. 
 The branches arise at irregular distances, and the main stem reaches the margin 
 far out towards the wing-apex. The cubitus of the hind-wing gives off a large 
 outer braach which is almost equal in strength to the main stem. This arises
 
 LITHOSIALIS. 45 
 
 even nearer the base than its counterpart in the fore-wing, but its great length 
 is masked by the development of the median. Beyond the origin of the outer 
 branch the cubitus gives off three inward branches, the first forking. Owing to 
 the strong inward curvature of the cubitus and its branches, the succeeding anal 
 veins are directed almost straight inwards. The anal veins are six in number, the 
 first forking twice, and ending on the margin in three branches. 
 
 The remaining veins are undivided. The cubital and anal veins of the 
 fore-wing are more obliquely disposed than those of the hind-wing, and the 
 latter has an anal area much larger than that of the fore-wing. 
 
 The interstitial neuration consists of short stout cross-nervures, which 
 occasionally fork, and in the wider areas unite to form a meshwork. 
 
 A triangular area is marked off from the base of each wing by a deep furrow. 
 This Woodward correlates with a similar area in the wing of Gryllacris (Corydalis) 
 brongniarti, which Swinton (' Geol. Mag., [2], vol. i, 1874, p. 337, pi. xiv, fig. 3) 
 has described as a stridulating organ. As stated elsewhere, I am of opinion 
 that Swinton's conclusions in the case of G. brongniarti were founded on a 
 misinterpretation of the wing, and that no stridulating organ, or similar 
 structure, is present. In the case of //. carbonarius such a structure would 
 apparently be useless, as none of the wings could come into such close apposition 
 as would allow the structure to be used. I am unable to determine its purpose 
 or significance, unless it be a portion of the musculature attachment of the wing. 
 
 Traces of a fore-leg are present, projecting from beneath the left lobe of the 
 prothorax. Its structure is too indefinite for description. 
 
 The wings were probably one-fourth or one-third longer than the portion 
 preserved an estimate which would make each complete wing about 70 mm. long, 
 with a spread of about 140 mm., or 6 inches. 
 
 Genus LITHOSIALIS, Scudder. 
 1881. Litlwsialis, Scudder, Geol. Mag. [2], vol. viii, p. 299. 
 
 Generic Characters. Wings three to four times as long as wide. Outer and 
 inner margins almost parallel ; intercostal area wide basally, and diminishing to 
 extinction at apex of wing. Radius simple; median with two main branches, 
 and cubitus large ; anal veins few and oblique. 
 
 The wing on which this genus is founded is interesting as being the first 
 discovered in British Palaeozoic rocks, except the problematic examples' mentioned 
 by Lhuyd and not otherwise known. Supposed to be a plant, the wing .was 
 first sent by Man tell to Brongniart, who in turn referred it to Audouin. The 
 latter recognised its insect character, and brought it before the Entomological 
 Society of France, the Academy of Sciences, and the Assembly of German
 
 46 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 Naturalists at Bonn. Audouin described the wing as that of an unknown 
 Nenropteroos insect allied to Hemerobius, Semhlis and especially to Gorydalw. 
 Mantell (r Audouin) described it as closely resembling a species of living Cori/dalis 
 of Carolina. Swinton states that Mantell purchased the fossil at a sale of 
 Parkinson's collection, although Mantell (' Medals of Creation,' p. 554) says that 
 he " discovered " it in a nodule from Coalbrookdale. Possibly the nodule had 
 previously formed part of the Parkinson Collection. Both the figures by Mantell 
 and Murchison are badly drawn, and it was not until 1874 that a reliable drawing- 
 was published by Swinton. The latter author devoted considerable attention to 
 a " serrated vein " at the base of the wing, Avhich he regarded as a stridulating 
 organ. He therefore referred the wing to the Orthoptera, and to the genus 
 Gryflacris. Scudder threw considerable doubt on Swinton's conclusions, and 
 showed that such an organ so placed could not have been of any service. As 
 we shall see later, the supposed stridulating organ is merely a torn edge of the 
 base of the wing. A study of the wing-structure convinced Scudder that the 
 Aving was most closely related to the Litliomontis carboiuiri-us, Woodward. Being 
 generically distinct from Litkomantix, and from living types, he gave the insect 
 the generic name of Lithosialis . 
 
 Lithosialis brongniarti (Mantell). Plate III, fig. 1 ; Text-figure 10. 
 
 1833. Corydalis ?, Audouin, Ann. Soc. Eat. France, vol. ii, Bull., p. 7. 
 
 1836. Corydalis ?, Audouin, Buckland, Bridgewater Treatise, vol. ii, p. 77. 
 
 1844. Corydalis brongniarti, Mantell, Medals of Creation, ed. 1, vol. ii, p. 578, ligu. 124, fig. -2. 
 
 1854. " Sialidse," Pictet, Traitt- de Paleontologie, ed. 2, p. 377, pi. xl, fig. 1. 
 
 1867. Corydalis, "allied to," Murchison, Siluria, ed. 4, p. 300, woodcut 80. 
 
 1871. Corydalis brongniarti, Woodward, Geol. Mag., vol. viii, p. 387 (name only). 
 
 1874. Corydalis brongniarti, Swinton, Geol. Mag. [2], vol. i, p. 339, pi. xiv, fig. 3. 
 
 1875. Gryllacris (Corydalis) brongniarti, Woodward, G-eol. Mag. [2], vol. ii, p. 622. 
 
 1876. Corydalis brongniarti, Woodward, Quart. Jouvn. Geol. Soc., vol. xxxii, p. 62. 
 
 1880. Corydalis brongniarti, Nowak, Jahrb. k.k. Geol. Beichsanst., Wien, vol. xxx, p. 73, pi. ii, fig 4. 
 
 1881. Lithosialis brongniarti, Scudder, Geol. Mag. [2], vol. viii, p. 299. 
 
 1883. Lithosialis brongniarti, Scudder, Mem. Bost. Soc. Nat. Hist., vol. iii, p. 220, pi. xvii, figs. 1 , 2, 8, 9. 
 385. Protogryllacris brongniarti, Brongniart, Bull. Soc. Amis Sci. Nat. Rouen [3], aim. xxi, p. 59. 
 
 Lithomantis brongniarti, Brongniart, Faune Entom. Terr. Prim., p. 371, figs. 17, 18. 
 1906. Lithosialis brongniarti, Handlirsch, Die Fossilen Insekten, p. 84, pi. x, fig. 13. 
 
 Ti/p?.~A left fore-wing; British Museum (Mantell Coll., olim Parkinson Coll., 
 no. 11,619). 
 
 Horizon and Locality. Coal Measures; near Coalbrookdale, Shropshire. 
 
 Specific Characters. Radial sector with not more than three branches ; median 
 much branched, and occupying much of the distal inner wing-margin. Cubitus 
 with three divisions. Anal veins few, and very oblique. 
 
 '
 
 LITHOSIALIS BRONGNIARTI. 47 
 
 Description. The left fore-wing measures 61 mm. long and 18 mm. wide 
 across the distal third. The apex, the inner margin and a very small portion 
 of the base are missing. The wing appears to have been strap-shaped, with a 
 blunt apex, the inner margin curving most into the apex. 
 
 The outer margin is feeble, almost straight over the greater part of its length, 
 and slightly sloped backwards. B.isally it dips abruptly inwards to the point 
 of attachment. The subcosta is widely spaced from the outer or costal margin 
 proximally. the two gradually approaching each other until they meet at the 
 wing-apex. The radius passes straight out to the apex, giving off the radial sector 
 before the middle of the wing is reached ; the radial sector remains undivided up 
 to the distal third of the wing, beyond which it forks twice. The median vein 
 divides a little further out than the radial sector, forming two strongly divergent 
 branches, an outer large branch with two large forward twigs, and a small inner 
 branch which forks, the inner twig forking again. The median, therefore, ends 
 on the inner margin of the wing in six twigs. 
 
 Fia. 10. Lithosialis brongniarti (Mantell) ; diagram of neuration of left fore-wing, natural size. 
 Coal Measures; near ' Coal brookdale, Shropshire. Mantell Collection, olim Parkinson Collection, 
 Brit. Mus. (no. 11,619). 
 
 The cubitus divides into three branches, the first arising low down, and passing 
 in a bold sweep beyond the middle of the inner margin. The anal veins are five 
 in number, all but the first being undivided. The first anal forks near the base, 
 the outer branch forking again. 
 
 The whole wing is covered by a numerous series of strong transverse nervures, 
 rarely branching, and usually crossing at right-angles to the main veins. A few 
 are oblique or curved. 
 
 The "file" or "serrated vein," described at length by Swinton, and by him 
 correlated with that present in recent forms of Gryllacris, appears to be nothing 
 more than an irregular torn edge of the basal part of the wing, which is also lifted 
 up a little above the general level. The torn edge extends along the line of the 
 median vein for a short distance, and wholly lacks the symmetry and detail given 
 to it by Swinton. 
 
 Affinities. Although, closely allied to Lithomantis carbonarius, Woodw., this 
 wing differs in the great width basally of the intercostal area, and in the straighter 
 course of the cubitus and anal veins, due to the greater length of the wing. The 
 interstitial neuration is much the same.
 
 48 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 Genus PRUVOSTIA, novnni. 
 
 Generic Characters. Fore-wings two and a half times as long as wide ; outer 
 and inner wing-margins almost parallel ; wing-apex well rounded. Subcosta 
 widely removed from the costal margin basally, and apparently not connected with 
 the costa or radius. Radius straight with divergent radial sector. Radial sector, 
 median and cubitus all distally branched Anal veins few. Interstitial neuratkm 
 of straight cross-nervures. 
 
 Certain features of this wing are suggestive of the Protorthoptera, These are 
 the wide basal, intercostal area, the basal origin of the. radial sector, and the 
 remote branching of the radial sector, median and cubital veins. 
 
 It is, however, more nearly allied to Lithosialis than to Metri/in <mlis, Handl., 
 for example, among the Protorthopteroids. The great length before division of 
 the main stems of the radial sector, median and cubitus, their strong divergence, 
 and the many branches of the median, form an assemblage of characters not 
 elsewhere known, and certainly deserving of generic recognition. 
 
 It is with pleasure that I attach to this genus the name of Dr. P. Pruvost, of 
 Lille University, in recognition of his valuable work on the fossil insects of the 
 north of France. 
 
 Pruvostia spectabilis, sp. nov. Plate III, fig. 2; Text-figure 11. 
 
 Type. A left fore-wing; British Museum (Johnson Collection, no. I. 15,894). 
 
 Horizon and Locality. Middle Coal Measures (clay ironstone nodules in the 
 Binds between the "Brooch" and "Thick" coals); Coseley, near Dudley, 
 Staffordshire. 
 
 Fio. 11. Pruvostia spectabilis, sp. nov.; diagram of neuration of left fore-whig, natural size. Coal 
 Measures (clay ironstone nodules in the binds between the "Brooch" and "Thick" coals); 
 Coseley, near Dudley, Staffordshire. Johnson Collection, Brit. Mus. (no. I. 15,894). 
 
 Characters. Outer margin convex at base, and inclining inwards to 
 the apex. Subcosta straight, not reaching the apex of wing. Radius straight, 
 giving off radial sector near base ; radial sector twice branched, median divergent, 
 three times branched, the first two branches forking. Cubitus dividing basally 
 into two, the outer forking three times, the inner branch once. 
 
 Description. A left fore-wing 52 mm. long and 24 mm. wide, almost covering 
 the surface of one half of the median plane of a reddish-brown ironstone nodule. A 
 little of the base of the wing is missing, while a part of the wing-apex is concealed
 
 PRUVOSTIA SPECTABILIS. 49 
 
 by a film of ironstone which cannot be removed. Fortunately the slight film of 
 ironstone does not hide the outline of the wing, and it is evident that the outer and 
 inner wing-margins are almost parallel, while the apex is well rounded. The outer 
 margin merges by a well-rounded contour into the wing-apex. The subcosta is 
 well-marked, widely removed from the outer border proximall y, and lies in a shallow 
 groove formed of the intercostal and radial subcostal areas. These two areas are 
 reduced distally to less than half their proximal diameter owing to the backward 
 inclination of the outer margin. The subcosta seems to die out near the radius, 
 about 7 mm. from the wing-apex, and gives off a numerous series of irregularly 
 spaced and forwardly directed nervures, which are at first straight and then curved 
 towards the wing-apex. At their point of origin the cross-nervures are very 
 distinct, but they thin, and occasionally die out, before reaching the outer margin. 
 The radius is a strong vein, passing perfectly straight out from the wing-base to 
 the outer part of the wing-apex. The radial sector arises near the base of the 
 radius, gradually diverging from it up to the junction of the middle and outer 
 thirds, where a single branch is given off, the latter taking a position parallel with 
 the main stem of the radial sector, and entering the middle of the wing-apex. 
 Eleven mm. further out and within 7 mm. of the apex a second branch is 
 given off, which lies evenly between the main stem and the first branch. The 
 median vein arises so close to the radius as to appear united with it. Like the 
 radial sector, it does not divide until it reaches the junction of the middle and outer 
 thirds of the wing. Here the first inward branch arises, and at equal distances 
 further out are given off two more ; the first two each fork in the middle of their 
 length, the third remaining undivided ; the median ends therefore on the inner 
 wing-margin in six divisions. The cubitus consists of two, and possibly three, 
 main stems. Owing to the base of the wing being broken away these stems are 
 not seen to be in actual union. Little doubt can exist as to the union of the first 
 two, but if .the third vein is a branch of the cubitus, it can only join the other two 
 by a strong outward curvature. This third vein may be the first anal, although its 
 manner of division is similar to that of the cubitus, in this respect agreeing with 
 what is seen in the first anal of Lithosialis brongniarti. The next vein is parallel 
 with the one succeeding, which is undoubtedly anal, while it shows an increasing 
 basal divergence from the second cubital. For this reason I have regarded the 
 third vein as the first anal. The first branch of the cubitus diverges widely from 
 the median, passing obliquely to the inner margin of the wing, which is reached 
 just beyond the middle third'. Owing to the great divergence of the first cubital 
 branch from the median, the area between the two, at the point where they first 
 branch, is very wide almost twice the width of any other area. 
 
 The first cubital vein gives off three outward twigs. The second is parallel 
 to the first as far as its division into two equal twigs. The next vein is that to 
 which I have already alluded as a possible third branch. It is widely separated 
 7
 
 50 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 from the undoubted cubitus, and by reason of its two forward branches resembles 
 the cubital elements in front of it. Its wide separation from the second branch of 
 the cnbitns supports the view that it is the first anal vein. The undoubted anal 
 veins are two in number, undivided, and passing backwards very obliquely to the 
 wing-margin. More anal veins may have been present, but it is very doubtful. 
 The wing-surface is much wrinkled across both the length and breadth, and 
 traversed by numerous cross-nervures, which are most evident in the median area. 
 
 Affinities. The Aving is typically Palaaodictyopteroid, and in the development 
 of the radius, radial sector, and cubitus, shows an affinity to Lithosialis brongniarti. 
 It differs markedly from that species, however, in the great length of the main 
 stems before divisions arise. 
 
 Dr. Tillyard, to whom I have shown my enlarged drawings, is of opinion that 
 Prnrostia is allied to Pseudofouqnea rather than to the Lithomantidje. 
 
 Family BREYERIIDJ;, Handlirsch. 
 1906. Handlirsch, Die Fossilen Insekten, p. 95. 
 
 Wings markedly triangular, with broadened bases. Costa, subcosta and radius 
 brought closely together in the outer part of the wing, the median with few 
 divisions. Cubitus and anal veins directed almost straight inwards, at right 
 angles to the length of the wing. 
 
 Genus BREYERIA, Borre. 
 
 1875. Breyeria, Borre, Ann. Soc. Entom. Belg., vol. xviii, p. 40. 
 1908. StobMa, Handlirsch, Die Fossilen Insekten, p. 1348. 
 
 Generic Characters. Wings two arid a half times as long as wide. Outer wing- 
 margin straight; apex curved inwards. Inner wing-margin strongly convex. 
 Costa marginal, subcosta parallel and joining the radius near the wing-apex. 
 Radius giving off the radial sector near the base of the wing, and reaching the 
 apex undivided. Radial sector subparallel to radius. Median boldly curved inwards 
 almost at a right angle. Cubitus less curved than the median. Anal veins few. 
 Interstitial neuration of two kinds, that of the intercostal and radial sector areas of 
 short straight nervures, and that of the remaining areas of irregular thin nervures 
 which tend to anastomose into a loose meshwork or reticulate arrangement. 
 
 O 
 
 Breyeria woodwardiana (Handlirsch). Text-figures 12, 13. 
 
 1903. "Allied to Lithomantis carbonariw," Stobbs, Geol. Mag. [4], vol. x, p. 524. 
 1906. Palaeodictyopteron, sp., Handlirsch, Die Fossileu Insekten, p. 126,
 
 BREYERIA WOODWARDIANA. 
 
 1906. Lithonutntis carbonarius (/), Woodward, Greol. Mag. [5], vol. iii, p. 26, fig. 1. 
 1908. Stobbsia woodwardiatia, Handlirsch, Die Fossilen lusekteu, p. 1348, text-fig. 
 
 51 
 
 Type. Greater part of a left hind- wing ; Mr. J. T. Stobbs' Collection. 
 
 Horizon and Locality. Peacock marls overlying the Peacock Coal, and near the 
 top of the workable Coal Measures; Foley, near Longton, Staffordshire. 
 
 Specific Characters. Radial sector dividing in its distal half into fine inwardly 
 directed twigs which end on the distal third of inner margin. Median and cubitus 
 with few widely spaced branches, occupying greater part of inner margin. 
 Anal veins about three to four in number. 
 
 Description. The specimen has not been available for examination, and my 
 observations are based on the figures published by Woodward and Handlirsch. It 
 is probable that neither of these figures is wholly correct, Handlirsch pointing out 
 what appears to be an obvious error by Woodward in the character of the cubitus 
 
 FIG. 12. 
 
 PIG. 13. 
 
 Fia. 12. Breyeria woodwardiana (Handlirsch) ; left hind-wing as restored and figured by Dr. H. 
 Woodward under the name of Lithomantis carbonarius ?, natural size. Coal Measures (Peacock 
 marls overlying the Peacock Coal) : Foley, near Longton, Staffordshire. Mr. J. T. Stobbs' Collection. 
 
 FIG. 13. Breyeria woodu-ardiana (Handlirsch) ; same left hind-wing as restored and figured by Dr. A. 
 Handlirsch under the name of Stobbsia ivoodivardiana, nearly natural size. A = IX, anal ; Cu = 
 VII, cubitus ; M = V, median : R = III, radius ; Rs = IV, radial sector ; Sc = II, subcosta ; I, costa. 
 
 vein, and the position and mode of division of the radial sector being doubtful. 
 Notwithstanding this difficulty, I have by means of an old plaster cast been able 
 to satisfy myself upon the more essential details. 
 
 The fragment is 50 mm. long and 20 mm. wide, and consists of the greater 
 part of a left wing, of which the base and a portion of the basal third are stated 
 to be obscured by a pinnule of Neuropteris. The maximum width is across the 
 anal area, beyond which the wing rapidly narrows owing to the forward direction 
 of its inner margin. The costal vein is marginal and strong, passing in an almost 
 straight line distally until joined by the subcosta, after which it curves gradually 
 backwards into the wing-apex. The subcosta is parallel with the costal margin 
 over more than two-thirds of the wing, and then joins the costa. The base of 
 the radius is not shown in the figures, except a very small portion in front of the 
 point of origin of the radial sector, the latter arising in the basal third of the wing ; 
 beyond the radial sector the radius passes out to the wing-apex, keeping parallel 
 with the costal margin.
 
 52 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 The radial sector presents several difficulties. In neither of the two published 
 figures is this vein depicted as we might expect. In all other respects the wing 
 agrees remarkably closely with those of Breyerla and Meyaptiloide^ where the 
 radial sector sends off inwards a series of simple branches, and runs out fairly 
 parallel with the radius to the wing-apex. In place, however, of the radius 
 passing straight outwards, it is represented as dividing into two branches in the 
 distal fourth of the wing, the outer branch forking once and the inner forking 
 twice. The inner branch of the radial sector diverges widely from the outer, and 
 its three divisions go to the inner side of the wing-apex. A small branch is shown 
 by Woodward as joining the radial sector to the first branch of the median, while 
 the same branch is shown by Handlirsch as coming off from the radius immediately 
 in front of its division into two, and passing down towards the inner margin 
 between the inner branch of the radial sector and the first branch of the median, 
 but not uniting to the latter. The median vein arises near the radius and sweeps 
 out in a bold curve to the middle of the inner margin, giving off three outer 
 undivided branches. Dr. AVoodward, in his restoration of the base of the wing, 
 has inadvertently indicated the main stem of the cubitus as joining the median. 
 This is corrected in Handlirsch's drawing. The cubitus consists of a strongly 
 curved stem giving off two branches, but only the inner marginal portions of the 
 veins are present. The anal veins are three or four in number, and directed 
 backwards at right angles to the length of the wing. 
 
 The interstitial neuration consists of feeble transverse nervures, which either 
 pass irregularly across between the main veins or occasionally fork. 
 
 Affinities. Dr. Woodward doubtfully refers the wing to Lithomantis carlo- 
 narius, Woodw. Handlirsch, in the earlier part of his work, ' Die Fossilen 
 Insekten ' (p. 126), classed it as a " Palasodictyopteron " only, and afterwards, 
 owing to its supposed relationship to L. carbonarius, and its evident likeness to 
 Litlwsialis and Hadnmeura, established the genus Stobbxia for it, placing the species 
 in the Lithoinantidas. There are, however, certain features of the wing which 
 militate against his view. The close apposition of the costa, subcosta and radius 
 are in marked contrast to the condition in that family, where these veins are 
 widely spaced, and where there is also a very wide intercostal area. The radial 
 sector is also more complex. In those details in which the wing departs from the 
 Lithomantida3, it approaches the characters of the genera JJrvt/eria and Meyiijili- 
 loides. The resemblances to Meyaptiloides brodiei, Brong., and Breyeria borinensis, 
 Borre, are very close, so far as can be determined by the distal fragment of the 
 former wing and the more than three-fourths of the latter wing. The main 
 difference between this wing and those of Breyeria borinensis and It. laclilani is in 
 the character of the subcosta, which in the latter two species joins the radius. 
 Whether the published figures of the Staffordshire wing are correctly drawn in 
 this particular we do not know, and as we have already seen that these figures are 

 
 SPILAPTERA. 58 
 
 wrong in other details, we are compelled to assign the specimen to the family 
 Breyeriidas, and doubtfully to the genus Breyeria, with which it seems in agreement. 
 
 Family SPILAPTERIDJE (Brongniart), Handlirsch. 
 1906. Handlirsch, Die Fossilen Insekten, p. 101. 
 
 Radial sector more or less branched ; median divided into two main branches, 
 the outer much divided ; cubitus with an outer branch sending numerous twigs to 
 the inner margin. Intercostal area occupied by a series of straight cross-nervures. 
 
 The family Spilapteridse, founded by Brongniart in 1885, has been re-defined 
 by Handlirsch and Lameere. The. latter extended the group to include species 
 which Brongniart had placed partly in the family Platypteridse and partly in 
 Protephetneridae. All the forms thus brought together by Handlirsch agree in 
 the possession of a typical palaeodictyopteroid neuration and the general characters 
 'enumerated above. 
 
 Lameere ('Bull. Mus. Hist. Nat. Paris,' 1917, no. 1), who rejects Handlirsch's 
 views and classifications (see above, p. 15), has remodelled the family and given it 
 a new significance. He is of opinion that the genera Lamproptilia, Epitethe, 
 Becguerelia, PaltKoptilus, Compsonenra, Spiloptilus, Homaloneura, Graphiptilns and 
 Spilaptera form a natural family, the Spilapteridse, in which a progressive 
 evolution in the longitudinal venation can be observed. The family is regarded 
 as linked to that of Megasecopteridaa through the genus Becquerelia-, and to the 
 Protepherneridas through the genus Apopappu.s. The three families are then 
 grouped in his new order, Ephemeroptera. Such a classification is based on the 
 belief that a perfect evolutionary sequence can be made out. Unfortunately, in 
 presenting this classification, Lameere gives only a summary of his reasons and 
 evidence, and it is not possible to criticise his argument. It is, to say the least, 
 very doubtful if, in the present state of knowledge, we can judge relationships 
 always correctly, while the sequence of evolution is still more difficult. 
 
 Handlirsch has acknowledged that he is unable to undertake any division of 
 the family Spilapterida3, as he understands it, and is content to await the discovery 
 of the bodies of these insects for a fuller knowledge of the family. As the more 
 rational view, Handlirsch's definition of the family is adopted. 
 
 Genus SPILAPTERA, Brongniart. 
 1885. Spilaptera, Brougiiiart, Bull. Soc. Amis Sci. Nat. Boueii [3], ann. xxi, p. 63. 
 
 Generic Character*. Insects closely resembling Palseopt'dus ; fore-wings 
 narrower basally than the hind- wings ; body slender.
 
 54 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 Spilaptera sutcliffei, Bolton. Plate III, fig. 3 ; Text-figure 14. 
 
 1917. Spilaptera sutdi/ei, Boltoii, Quart, Jouru. Geol. Soc., vol. Ixxii, p. 53, pi. iv, fig. 1, and 
 text-fig. 
 
 Type. Basal third of a wing; Manchester Museum (no. L. 8197). 
 
 Horizon and Locality. Middle Coal Measures (grey -blue shales 135 180 feet 
 above the Royley or Arley Mine) ; Sparth Bottoms, Rochdale, Lancashire. 
 
 Specific Characters. Subcosta parallel with the costal margin ; median vein 
 dividing near the base into two branches, of which the outer forks just beyond the 
 point of origin of the radial sector. Cubitus a large and much-divided vein. 
 Anal veyis few in number. 
 
 Description. This specimen was formerly labelled " Stenodictt/a lolata" and is 
 one of three recorded by Sutcliffe, Baldwin, and others, in their papers on the 
 fossils found at Sparth Bottoms. The remaining two are described (p. 37) under 
 
 '**' f / / ' ! ' ' ' ' ' / ' ,'/">'/-/-// / / ' 
 "* ~'- - -' -/. J- i. _ /- -'- - - - '- -'- -'-- L /. / _ LJ- L -''- '" '* ' ' 
 
 FIG. 14. Spilaptera sutcliffei, Bolton ; restoration of left wing, natural size. Middle Coal Measures 
 (shales above the Royley or Arley Mine) ; Sparth Bottoms, Rochdale, Lancashire, Manchester 
 Museum (no. L. 8197). 
 
 Mecynoplem tuberculata, Bolton. The specimen consists of the basal third of a 
 wing lying on the median plane of a small irregular micaceous sandy nodule. The 
 finer structure of the wing has not been preserved, owing, no doubt, to the coarse 
 grain of the matrix. The chief veins of the wing are robust structures, and these 
 are fortunately well marked and clear. The wing-fragment is 35 mm. in length 
 along the outer margin, and 27 mrn. in greatest width, but as the inner margin is 
 broken away and lost, the total width of the wing exceeded this, and may have 
 been over 30 mm. It belongs to a left wing, and when complete must have been 
 at least 90 mm. in total length. The perfect insect must therefore have had a 
 spread of wing of nearly 200 mm., or about 8 inches. 
 
 The outer margin is feebly convex, the subcosta fairly parallel, sunk in a groove, 
 and gradually approaching the outer margin as it passes towards the wing-apex. 
 The rate of approach is so gradual that the junction must have been far out near 
 the wing-apex. The intercostal area is crossed by a series of straight nervures 
 which are oblique in their course outwards.
 
 SPILAPTERA STJTCLIFFEI. 55 
 
 The radius is strong, standing up above the 'surface, and almost parallel with 
 the subcosta. The radial sector arises from the radius at about 22 mm. from the 
 base of the wing. The median vein, owing to its inward direction away from 
 the radial sector, is better shown. It forks very low down into two equal 
 branches, both of which are widely spaced. The outer branch divides almost 
 opposite the point of origin of the radial sector, while the inner branch forks 
 much nearer the base of the wing, the innermost branch curving sharply 
 back from its fellow, so that the area between them is wide almost from the 
 commencement. 
 
 The general direction of the branches of the median is such that they would 
 reach the outer half of the inner margin of the wing. 
 
 The cubitus consists of two main stems, the basal union being missing ; the 
 outer branch passes, after twice forking, in a double curve down to the inner 
 margin of the wing, the inner branch being directed more directly backwards, 
 and forking three times in the basal third. Fragments of four anal veins are 
 distinguishable, the first dividing near the middle of its length into two equal 
 branches. The interstitial neuration seems to have consisted of comparatively few 
 straight and well-spaced nervures, some of which can be seen crossing the wide 
 area between the innermost twig of the median and its fellow. No others are 
 visible, except those in the intercostal area already mentioned. 
 
 Affinities. Notwithstanding the fact that only about one-third of the wing 
 is know r n, it is yet possible to determine the generic characters with a reasonable 
 degree of accuracy. The mode of division of the radius and median, the manifest 
 importance of the latter, the wide area between the median and cubitus, and the 
 few widely spaced cross-nervures, are all typical Spilapteroid characters. The 
 wing cannot be confused with that of Stenodictya, in which the median is much less 
 developed, and the interstitial neuration closely reticulated. The only other genus 
 to which it might be referred, that of Becquerelia, is distinguished by a union of 
 the anal veins, which in Spilaptera and in this specimen are distinct. 
 
 Spilaptera has hitherto comprised only three species. From 8. packardi, 
 Brong., the specimen differs by the subcosta being parallel with the outer margin ; 
 by the equal separation of the subcosta, radius and median in the basal third of 
 the wing ; and by the division of the median into two branches close to the base of 
 the wing. S. libelhiloides, Brong., of which the distal half of the wing only is 
 known, has a much feebler cubitus, which divides by forking further out than in 
 S. sutcli/ei. S. renusta, Brong., was established on a fragment much similar to that 
 of 8. libelluloides, in which the subcosta is a short vein, and the radial sector arises 
 nearer the middle of the wing.
 
 56 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 Family LAMPROPTILID/K (Bronguiart), Handlirsch. 
 
 1885. Brougniart, Bull. Soc. Amis Sci. Nat, Eouen [3], ami. xxi, p. 63. 
 1906. Handlirsch, Die Fossilen Insekten, p. 109. 
 
 Fore- and hind-wing strongly marked. Veins of the anal and cubital groups 
 numerous and directed obliquely backwards. Hind- wings short and broad. 
 
 Genus BOLTONIELLA, Handlirsch. 
 
 1885. Lamproptilia, Brougniart, Bull. Soc. Amis Sci. Nat. Eoueu [3], anu. xxi, p. 63. 
 1919. Boltoniella, Haudlirsch, Revision der Paliiozoischeu Insekteu, p. 21 . 
 
 Generic Characters. Fore-wings two and a half times as long as wide; hind- 
 wings as long as wide. Outer margin of former curved, of hind-wings almost 
 straight. Apex of wings rounded. The fore-wings are almost elliptical, and the 
 hind-wings rectangular in shape. The costa is marginal and the costal area some- 
 what narrow. The subcosta joins the costa far out. Radius simple, the radial 
 sector arising near the base of the wing and giving off three forked branches, 
 which pass obliquely to the inner half of the wing-apex. Median dividing into 
 two main branches, the inner twice as much divided as the outer. Cubitus with 
 two main branches, each breaking up into numerous twigs. Interstitial neuration 
 consisting of widely spaced straight cross-nervures. 
 
 Handlirsch, who has formed this genus, admits that it reminds one in mam r 
 ways of Lamproptilia, but by reason of the small size and closer spaced cross- 
 nervures he separates it from that genus. 
 
 Boltoniella tenuitegminata (Bolton). Plate III, fig. 4. 
 
 1911. Lamproptilia tenuitegminata, Bolton, Quart. Journ. Geol. Soc,, vol. Ixvii, p. 170, pi. x, fig. 6. 
 
 Type. A right hind-wing, in a small fragile block of brown mudstone or shale, 
 crowded with plant-remains ; Museum of Practical Geology, Jermyn Street, London 
 (no. 24509). 
 
 Horizon and Locality. Coal Measures (No. 2 Rhondda Seam, base of the Pennant 
 Series) ; 1} miles north-east of Resolven Station, Glamorganshire. 
 
 Specific Characters. Broad wings of great tenuity and blattoid-like in character. 
 Principal veins much divided, and fading out on the inner margin. Apex of wing 
 bluntly rounded. Anal veins very numerous. 
 
 Description. The type-specimen consists of a right hind-wing of considerable 
 tenuity, the underlying plant-remains being easily traceable through the texture 
 of the wing. The greatest length is 29 mm., and the greatest breadth 7 mm. The 
 neuration is so similar to that which I have observed in the hind- wings of blattotfs
 
 BOLTONIELLA TENUITEGMINATA. 57 
 
 from Commentary, France, that it is difficult to refrain from classing the wing as 
 blattoid. The neuration is extensively branched and the wing-boundaries are not 
 well marked, but the wing appears to have been somewhat quadrangular in outline, 
 with a sinuous inner margin, and the base much broader than is seen in the ordinary 
 form of blattoid hind-wing. The outer margin seems to have been straight, a 
 portion of it still remaining in the middle third. The two margins merge in a 
 well-rounded apex. The subcostal area is narrow, strap-shaped, and probably 
 extended over the whole length of the outer margin. No traces of cross-nervures 
 can be seen on it. The radius divides near the base, giving rise to a series of 
 branches which curve inwards as they approach the apex of the wing. The course 
 of the branches is irregular, the interspaces widening and narrowing, possibly 
 owing to the wing having crumpled during deposition. The median divides into 
 two branches low down, each of which is repeatedly forked, the final divisions 
 becoming attenuated and untraceable before the inner margin of the wing is 
 reached. The course of the cubitus is obscured by a reed-like plant, only two 
 basal portions being distinguishable. The anal area is filled by a broad series of 
 thread-like veins, which sweep obliquely inwards in a fan-shape, and occupy a large 
 part of the inner margin. 
 
 The inner margin of the wing to a third of its total length is quite filmy, the 
 veins crossing the area as faint shadowy lines. The distal two-thirds is more 
 strongly impressed, while in the broad base of attachment the stems of the 
 principal veins seem to have been more than usually robust. No trace of trans- 
 verse nervures or reticulation can be seen. 
 
 Affinities. The nearest analogues to this wing are, I believe, the forms described 
 by Brongniart as Lamproptilia grand' euryi and L. stirrupi (' Etudes sur le Terrain 
 Houiller de Commentry,' vol. iii [1893], pp. 46770, pi. xxxv [19], figs. 79). 
 It appears to be more closely related to L. stirrupi than to L. grand'euryi, but is 
 more quadrangular, and its costal area is broader. The anal portion of the wing 
 is of greater tenuity, and occupies fully half of the inner margin. 
 
 Family BRODIID^:, Handlirsch. 
 
 1906. Handlirsch, Die Fossilen Insekten, p. 113. 
 
 1919. Handlirsch, Revision der Paliiozoischen Insekten, p. 73. 
 
 Wings in which the anal area is much reduced and specialised, the radius 
 undivided, and the median, cubitus, and anal veins arched and directed back 
 towards the inner margin. 
 
 From a re-examination of the originals in the British Museum, Handlirsch has 
 concluded that the family does not belong to the Palaaodictyoptera, but to the 
 Order Megasecoptera. This view seems at variance with his own definition of the 
 8
 
 58 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 latter Order ('Die Fossilen Insekten,' p. 312), where lie states that the special 
 distinctive characters of the Megasecopteridae are the tendency to a reduction of the 
 anal area of the wing, and the partial fusion of the median and cubitns with 
 the base of the radius. Brocfia certainly agrees with the Megasecopteridae in the 
 reduction of cross-nervures, but the anal area is very long, and I have not seen 
 a single case among the twelve or thirteen specimens in the British Museum 
 where there is any clear sign of fusion between the median, cubitus, and 
 the base of the radius. The bases of the three veins are closely brought together 
 in the inner spatulate portion of the wing, but this seems a natural result of the 
 narrowing of the wing in this region. 
 
 The family Brodiidae is more fully represented by individuals in the British 
 Coal Measures than any other, thirteen specimens being known of" Brodia 
 priscotincta, Scd., alone. To this family also seem to belong a series of small 
 larval wings. The neuration in these is immature, and the wings themselves are 
 far too small and weak to have supported the body in flight. The abdomen seems 
 long and well segmented, the segments showing evidence of well-developed 
 pleura, and they may also have borne tubercles. The whole integumentary 
 structure is thin, and if at all chitinous, only feebly so, and but faintly outlined 
 on the surface of the nodules. The wing-shape is much like that of Brodia, and 
 the veins, so far as determinable, such as may reasonably be supposed to have 
 developed into the typical Brodia-type in the adult. 
 
 While general evidence and association point to these wings being those of 
 larval forms of Brodia, and possibly of B. priscotincta, Scd., it is more judicious 
 to retain them in a distinct group. The same difficulties occur with other larval 
 wings which cannot be allocated to known genera, and for which new generic 
 names are not advisable. I therefore class larval wings of unknown relationship 
 as a separate group under the name of " Pteronepionites" a name which has no 
 classificatory value, but merely serves to indicate their larval nature. When 
 evidence is forthcoming of the generic and specific relationship of any member 
 of the group, it can be removed without the necessity of reducing a generic term 
 which has passed into nomenclature to the rank of a synonym. 
 
 I would allot the term " Pteronepionites " to all larval insect- wings of all 
 geological periods. 
 
 Genus BRODIA, Scudder. 
 1881. Brodia, Scudder, Geol. Mag. [2], vol. viii, p. 293. 
 
 Generic Characters. Wings spatulate in shape, three times as long as wide. 
 Outer margin feebly convex. Costa and radius spinulose, radius undivided, radial 
 sector and median vein with few divisions. Cubitus a single vein. Anal veins 
 few and widely spaced.
 
 BRODIA PRISCOTINCTA. 59 
 
 Brodia priscotincta, Scudder. Plate III, figs. 5, 6 ; Plate IV, figs. 13 ; Text- 
 figures 15, 16. 
 
 1881. Brodia priscotincta, Scudder, Geol. Mag. [2], vol. viii, p. 293, text-fig. 
 
 1883. Brodia priscotincta, Scudder, Mem. Boston Soc. Nat. Hist., vol. iii, p. 213, pi. xvii, figs. 37. 
 
 1885. Brodia priscotincta, Bronguiart, Bull. Soc. Amis Sci. Nat. Rouen [3], aim. xxi, p. 63. 
 
 1893. Brodia priscotincta, Brongniart, Fauue Entoni. Terr. Prim., p. 528, pi. xl (24), fig. 4. 
 
 1906. Brodia priscotincta, Handlirsch, Die Fossilen lusekten, p. 113, pi. xii, fig. 13. 
 
 1917. Brodia priscotincta, Bolton, Proc. Birmingham Nat. Hist, and Phil. Soc., vol. xiv, pt. 2, p. 100, 
 
 pi. vii, figs. 34, text-figs. 23. 
 
 1919. Brodia priscotincta, Handlirsch, Revision der Palaozoischen Insekteu, p. 73, fig. 83. 
 
 1919. Brodia Scudderi, Handlirsch, op. cit., p. 74, fig. 84, Brit. Mus., no. I. 3879. 
 
 1919. Brodia petiolata, Haudlirsch, op. cit., p. 74, fig. 85, Brit, Mus., no. I. 2961. 
 
 1919. Brodia pictipennis, Haudlirsch, op. cit., p. 74, fig. 86, Brit. Mus., no. I. 2961. 
 
 1919. Brodia fasciata, Haudlirsch, op. cit., p. 75, fig. 87, Brit. Mus., no. I. 1557. 
 
 1919. Brodia nebulosa, Handlirsch, op. cit., p. 75, fig. 88, Brit. Mus., no. I. 2961. 
 
 Type. Incomplete wing; British Museum (Brodie Collection, no. I. 3896). 
 
 Horizon and Locality. Middle Coal Measures (clay ironstone nodule from the 
 binds between the " Brooch " and " Thick " coals) ; Dudley, Staffs. (Scudder gives 
 the locality as Tip ton). 
 
 FIG. 15. Brodia, priscotincta, Scudder; diagram of a complete left wing showing position of tubercles 
 and character of neuration, natural size. Middle Coal Measures ; Dudley, Staffordshire. 
 
 Specific Characters. Outer margin of wing spinulose, almost straight, or feebly 
 convex at the most. Subcosta remote from margin at base, and extending beyond 
 the middle of the wing, joining neither to the costa nor to the radius. Radius a 
 strong vein, and spinulose. Radial sector arising in the first half of the wing, and 
 giving off four branches. Median with two forward branches. Cubitus a single 
 vein. Anal veins two. Inner margin strongly convex. 
 
 Description. The Geological Department of the British Museum possesses no 
 less than eleven examples of this species in addition to the type, while two more 
 are in the Geological Collection of Birmingham University, one being in the Beale 
 Collection, and the other presented by Dr. Blake. The British Museum specimens 
 are registered as follows: I. 1557, I. 1567, I. 3879, I. 3896, In. 18431, all from 
 Coseley; I. 2961, In. 18433, In. 18434, all from Tipton; In. 18429 from Sedgeley; 
 and In. 18430, I. 5866, I. 2962, and In. 18432. The following description and 
 remarks are based on a study of all these specimens. 
 
 The longest wing I have seen measures 58 mm. in length, and we are not likely
 
 60 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 to be beyond the mark in estimating the total span of the insect as about 130 mm., 
 or over 5 inches. 
 
 The outer or costal margin is straight, or feebly convex at most, over the greater 
 part of its length, curving distally into the wing-apex. Close to the base it swells 
 out into a slight hump-like elevation which is seen in all cases where the wing has 
 been broken off close to the body. The whole of the outer margin bears a dense series 
 of minute, conical, sharply pointed spinules of a black colour. These are arranged 
 in two rows on the proximal half, with the points of the spinules directed towards 
 the wing-apex. The bases of the spinules are expanded, and give a doubly corded 
 or monilated appearance to the wing-margin. Tillyard regards these spinules as 
 modifications of large hairs which he has termed " macrotrichia," and I see no 
 reason to dissent from his view. 
 
 The subcosta at its origin is widely spaced from the outer margin, and passes 
 out beyond the middle of the wing, gradually approaching, but failing to reach it. 
 The subcosta is a strong, straight vein, rhe greater part elevated above the level 
 ' of the wing-membrane, and distally flattening into it and disappearing. 
 
 The radius is a strong vein, more convex than the outer or costal margin, and 
 therefore more widely separated from it at either end than in the middle. The 
 basal portion is parallel with the subcosta for the whole length of the latter ; it 
 then becomes parallel with the outer margin for a short distance, and afterwards 
 curves into the wing-apex. A single row of spinules can be distinguished along 
 the whole vein in some specimens, and gives a slight monilation to the vein-surface. 
 The radial sector is well marked, and comes off from the radius about the point of 
 origin of the first forward branch of the median. The two veins are close together 
 for some distance, but in the distal half of the wing they become parallel, the 
 interspace being equal in width to that between the radius and the outer margin. 
 The radial sector gives off four inward branches, which end on the inner side of 
 the wing-apex. The first branch arises just beyond the middle of the wing, at 
 an acute angle, and is separated from the second by an interval which is double 
 the length of that separating the second and third. The fourth branch is very 
 short, and so close to the margin as to be absent in some specimens. 
 
 The median vein for the first quarter of its course lies in the middle line of the 
 wing, and then bends inwards in a wide curve to the distal third of the inner 
 margin. It gives off two outward branches, both of which are larger and stronger 
 than those of the radial sector. Both branches have the same sweeping curve 
 possessed by the branches of the radial sector, and are parallel with the latter, 
 while the main stem becomes almost straight. 
 
 The cubitus is a single vein, not united at the base to the median, and passing 
 obliquely to the margin. Some wings have broken off so far out from the base 
 that the cubitus appears to join the wing-margin at the junction of the middle and 
 distal thirds. A striking feature of the cubitus is its isolated position upon the
 
 BRODIA PRISCOTINCTA. 61 
 
 margin, for while the interval between it and the stem of the median is much 
 wider than any other in the fore-part of the wing, the interval between the cubitus 
 and the next vein is nearly twice as wide. There is, in fact, a progressive 
 widening between the veins as they are traced from the apex to the base of the 
 wing, the areas between the median and its branches being wider than those 
 enclosed by the branches of the radial sector. 
 
 Two somewhat dissimilar anal veins are present. The first has a wide curve 
 to a point far out on the margin, sometimes giving off a short branch, while the 
 second vein is much shorter and joins the margin at an acute angle. In some 
 specimens this vein is seen to give off two, or even three, short oblique branches. 
 The shape of the wings is very similar to that of the wings of mosquitoes 
 (Anopheles), and they bear evidence of having been folded in a plicate or fan-like 
 fashion along their length. The first two folds are united at the wing-base, and 
 pass out along the radius and the first outward branch of the median. The third 
 fold lies along the line of the cubitus vein. The degree of plication which a wing 
 retained when silted up modifies considerably the apparent distance between the 
 several veins and their branches, and at times hides important junctions. In one 
 example (Brit. Mus., no. In. 18431) the wing was well flattened out before being 
 buried, and the origin of the veins and their true position can now easily be 
 determined. This specimen shows that the radial sector arises much nearer the 
 wing-base than the first outward branch of the median a feature not usually 
 shown in the remaining specimens. 
 
 The general build of the wing is such that the bases of the costal and subcostal 
 veins on the outer margin, and those of the anal veins on the inner, must have 
 served as the main support to the distal two-thirds of the wing, the latter consisting 
 mainly of the radial sector and its branches, the median, and the distal half of the 
 cubitus. Flight must have been mainly maintained by the action of this more 
 distal expanded area, while the strain of movement would fall across the narrow 
 neck-like base of the wing, and may ultimately have led to fracture and the loss of 
 the wings. It is quite possible, also, that these insects were capable of finding 
 food among the decaying vegetation of the coal forests, and thus prolonging life 
 for a considerable period after the wings were lost. Such a presumptive sequence 
 of events would account for the total absence of any trace of the bodies or legs, 
 although the wings of this species are more numerous than any others in the Coal 
 Measures and preserved in good condition. 
 
 Colour Bands. Scudder mentions this species as the most striking instance 
 among Palaeozoic insects of the preservation of " colour bands," and as he states, 
 some wings show three broad irregular belts of dull umber-brown colour across 
 the wings. Close examination of these " colour bands " in the type-specimen, and 
 in other examples, leads us to doubt the correctness of his view. In all cases 
 where the " colour bands " do not show on the wings, the areas appear to be
 
 62 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 totally destitute of any traces of the wing-membrane, and the course of the veins 
 only is shown across the matrix. It is much more likely that the " colour-band " 
 effect has been produced by conditions of preservation owing to the wing- 
 membrane being destroyed in those areas which do not show colour. 
 
 Affinities. Although Scudder founded both genus and species, he did not 
 attempt any diagnostic description of either, confining his efforts mainly to a 
 discussion of relationships. His figure is unusually poor, and adds nothing to the 
 text. Scudder's general conclusion was that the wing was neuropteroid in 
 character, but " refusing to affiliate closely with the restricted families of the 
 present day," A manuscript note in Brodie's handwriting placed with the type- 
 specimen would seem to show that Scudder's views acquired greater definition 
 later. Brodie writes under date, February, 1880: "I sent this wing to Mr. 
 Scudder, and he supposes it to belong to the white ants (Termitidse), or close to 
 the group of which Goldenberg's Dictyoneura is the best type." The first detailed 
 descriptive note of the species Avas published in 1893 by Brongniart, with an 
 
 FIG. 16. Brodia priscotincta, Scudder; immature wing, twice natural size. Middle Coal Measures 
 (clay ironstone nodule from binds between the " Brooch " and " Thick " coals) ; Coseley, 
 Staffordshire. Madeley Collection, Brit. Mas. (no. I. 2966). A, anal; C, costa; Cu, cubitus; 
 M, median ; R, radius ; Rs, radial sector ; Sc, subcosta. 
 
 excellent enlarged drawing. Brongniart very doubtfully assigned the wing to the 
 Protodonata, and " alongside the Campyloptera." 
 
 Immature Wings. The collection of insect-remains which I describe later 
 (p. 67) under the name of " Pteronepwnites" were found at the same horizon and 
 localities as Brodia priscotincta, and it is fairly certain that some of them are 
 immature forms of this species. It will be noted that these larval forms are broad- 
 bodied and well segmented, and with lateral outgrowths of a pleura-like character 
 upon the abdomen. They indicate that Brodia priscotincta went through a progres- 
 sive metamorphosis, the rudimentary wings gradually developing as the insects 
 lived as ground-feeders among dank and rotting vegetation. 
 
 One of these wings (PI. IV, fig. 3) in the Madeley Collection in the British 
 Museum (no. I. 2966) is an impression 18 mm. in length by 4 mm. in 
 maximum breadth, contained in a small grey ironstone nodule. As is usually 
 the case with these grey nodules, the details of the wing are much obscured 
 by the matrix, and the precise method of division of the veins is far from being 
 clear. 
 
 The outer margin is regularly convex and formed by the costa. It gradually 
 merges into the expanded and well-rounded wing-apex. The subcosta extends
 
 BRODIA PRISCOTINCTA. 63 
 
 the whole length of the wing, ending in the apex, and being parallel to the costal 
 margin. A wide area separates it from the next vein. 
 
 The course of the radius is similar to that of the subcosta. The point at 
 which the radial sector arises cannot be determined, but it lies in the base of 
 the wing. 
 
 The radial sector is parallel with the radius over the greater part of its length, 
 and gives off three inward branches in the distal third of the wing, all of which 
 end on the inner side of the wing-apex. 
 
 The course of the inner third of the median is indistinguishable. The main 
 stem reaches the middle of the wing before it gives off the first outer branch, 
 which bends round and becomes parallel with the inner branch of the radial sector. 
 The second branch arises a little further out, and at a more acute angle than the 
 first, passing down to the inner margin of the wing midway between the first 
 branch and the main stem. 
 
 The cubitus is a long undivided vein which passes, first in a curve, and then 
 in a straight line to the inner margin. 
 
 Lying inward to the cubitus are traces of another undivided vein, w r hich may 
 represent the anal. It bends inwards more rapidly than the cubitus, and the 
 interval between the two veins is very wide. The inner margin of the wing is 
 sigmoidal in outline. 
 
 The general characters of this immature wing are unmistakeably those of the 
 genus llrodia, and only the absence of spinules on the costal margin and radius, 
 and the lack of a fourth branch to the radial sector, distinguish the specimen from 
 adult wings of /?. priscotincta. These details are not of specific value, and may be 
 due to the immature condition of the wing, which is but one-third the length of 
 that of a normal B. priscotincta. 
 
 The general characters of the wing are much like those of Brodia priscotincta 
 (jnuenis), and the specimen may be a slightly older larva of that form. The 
 outline of the wing is much the same, but the branching of the principal veins is 
 better shown and their apical curvature less pronounced. 
 
 The wing also lends considerable support to the belief that these insects did 
 not possess a resting or pupal stage, but that the metamorphosis was regularly 
 progressive. 
 
 Brodia priscotincta, Scudder, forma juvenis, Bolton. Plate IV, figs. 1,2; Text- 
 figures 17, 18. 
 
 1919. Brodia nympha, Handlirsch, Revision der Paliiozoischen Insekten, p. 76, fig. 90. 
 
 Type. A pair of wings, one almost complete, the other showing the apical half 
 only; British Museum (Johnson Collection, no. I. 1563.).
 
 64 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 Horizon and Locality. Middle Coal Measures (binds between the " Brooch " 
 and " Thick " coals) ; Coseley, Staffs. 
 
 Description. The fossil is contained in a brown ironstone nodule, the whole 
 wing having a length of 21 mm., and a maximum diameter of 4 mm. It is 
 membranous, and oblanceolate in shape. The outer wing-margin, of which only 
 the distal half is preserved, is flatly convex and curves inwards, meeting the inner 
 margin in a rounded apical angle. The inner margin is feebly concave. The wing 
 has three longitudinal folds, and the extreme tenuity of the integument is accom- 
 panied by a corresponding thinness of the veins. The greater portion of the costa 
 and the whole of the subcosta are missing. The outer portion of the radius is present. 
 It is parallel with the outer margin, and curves round into the wing-apex. The 
 radial sector is parallel with the subcosta, and curving round into the apex is lost, 
 by reason of its tenuity. Lying inward to the main stem of the radial sector in 
 the distal half of the wing are two long veins, and traces of two others, all 
 following the same course, and curving to the inner margin. The two long veins 
 
 FIG. 17.- Brodia priscotincta, Scudder, forma juvenis, Bolton ; diagram of wing-neuration, two-and-a- 
 half times natural size. Middle Coal Measures (binds between the "Brooch" and "Thick" 
 coals) ; Coseley, Staffordshire. Johnson Collection, Brit. Mus. (no. I. 1563) Lettering as in 
 Text-figure 16, p. 62. 
 
 appear to arise from the radial sector, in which case the two of which traces only 
 are seen would do so also. The radial sector therefore seems to give origin to 
 four inwardly directed parallel branches. The whole course of the next vein is 
 not clearly determinable. It passes to just beyond the middle of the inner margin, 
 in an almost straight oblique course, and gives off a single forked outer branch 
 parallel with the fourth branch of the radial sector. This vein can only be the 
 median. The next vein is the cubitus. It is undivided, and goes to the middle of 
 the inner margin. Anal veins are only indicated by feeble traces of a single 
 undivided vein, which apparently reached the margin midway between the cubitus 
 and the base of the wing. 
 
 Another similar wing from the same horizon and locality in the British Museum 
 (no. I. 1564) is 17 mm. long and 5 mm. wide, and lies on the surface of a split 
 nodule of dark-brown ironstone. The wing-membrane is very thin, and forms a 
 slight glaze on the otherwise granulated surface a feature which has made the 
 details of structure difficult to determine. The costa is marginal, the outer margin 
 convex, and gradually curving into the rounded apex of the wing. The inner 
 margin is slightly convex distally, and straight proximally. 
 
 The subcosta is a feeble vein whose course cannot be traced with certainty
 
 BRODIA PEISCOTINCTA. 65 
 
 beyond the middle of the wing. It is close to the margin and parallel with it for 
 its whole length. The radius is well marked, sunken, and also parallel, with the 
 margin except distally, where it is more inwardly curved. It ends on the apex in 
 a short fork. The radial sector arises near the base, and is parallel with the radius. 
 It appears to give off three to four branches, the first arising very near the base of 
 the wing. The whole course of this branch can be traced, but of the middle two 
 only faint traces are left in the region of the wing-apex. The fourth branch is 
 very short, and corresponds in position with the last branch of the radial sector in 
 Brodia priscotincta. The median vein forks low down into two equal branches, 
 which reach the middle of the inner margin. 
 
 The cubitus is a long undivided vein passing out almost to the middle of the 
 inner margin, and separated from the next vein, which seems to be the first anal. 
 The course of the latter is more oblique to the inner margin than that of the 
 cubitus. 
 
 In its general character this wing is distinctly similar to that of B. priscotincta, 
 although it lacks a second branch to the median, and the first anal does not seem 
 to be forked. 
 
 FIG. 18. Brodia priscotincta, Seudder, forma juvenis, Bolton ; diagram of wing-neuration, twice natural 
 size. Middle Coal Measures (binds between the " Brooch " and " Thick " coals) ; Coseley, 
 Staffordshire. Madeley Collection, Brit. Mus. (no. I. 1564). Lettering as in Text-fig. 16, p. 62. 
 
 Affinities. Handlirsch regards these as nymphal wing-sheaths, but their 
 extreme tenuity militates against this view. He is probably wrong in referring 
 Brodia to the Megasecopteridse, but right in regarding these wings as those of 
 heterometabolous insects, and not holometabolous as Lameere supposed. 
 
 Immature though the wings undoubtedly are, the more nearly perfect example 
 possesses an assemblage of characters which I believe points to its relationship. 
 The shape of the wing, the character and course of the subcosta, the number and 
 position of the branches of the radius, the simple forking of the median and the 
 anal veins, are all characters pertaining to the genus Brodia. The differences in 
 detail between this wing and that of B. priscotincta are such as may be looked for 
 between the nymph-stage and the adult. Among previously described larval wings, 
 the only type which seems comparable is Lameereites cnrvipennis, Handlirsch, based 
 on four nymph-wings or wing-cases, as Handlirsch has them, found in the Coal 
 Measures (Pennsylvania!!) at Mazon Creek, Illinois, U.S.A. (1911, Handlirsch, 
 'Arner. Journ. SciV [4], vol. xxxi, p. 375). 
 
 In these wings, the outer or costal margin is more strongly curved, and the costa 
 and radius extend over the bluntly curved apex down to its junction with the 
 inner margin. The succeeding veins arise nearer the base of the radius, only one 
 9
 
 66 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 apparently being a branch of that vein, while the median and the cubital areas ar< 
 occupied by numerous veins whose origins are not indicated. 
 
 The differences between Lameereites curvipennis and this specimen are considei 
 able. The balance of evidence is greatly in favour of an affinity with Brodia pru 
 cotincta, and the specimen may represent a nymph or larval stage of that species. 
 
 It is undesirable to attach a specific name to immature wings agreeing 
 closely with a known species. I propose to regard it as B. prisc.otimta, fori) 
 juvenis. 
 
 Brodia furcata, Handlirsch. Plate III, figs. 7, 8; Text-figure 19. 
 1919. Brodia furcata, Handlirsch, Revision der Palaozoischen Insekteu, p. 75, fig. 89. 
 
 Type. A left wing and impression showing the under-surface ; British Museui 
 (no. I. 2962). 
 
 Horizon and Locality. Middle Coal Measures (above-, the "Brooch" Coal) 
 Dudley and Coseley, Staffs. 
 
 FIG. 19. Brodia furcata, Handlirsch ; showing forking of the second branch of the median into two 
 equal twigs, natural size. Coal Measures (clay ironstone nodule from binds between the " Brooch " 
 and "Thick" coals) ; Coseley, Staffordshire. Brit. Mus. (no. I. 2962). 
 
 Specific Characters. Radial sector reduced in area, and possibly with thret 
 branches. Median having the second branch dividing into two equal twigs, both 
 of which reach the margin. Median area enlarged. 
 
 Description. This wing differs so much from the type-form as to be worthy of 
 specific distinction. Its total length is 44 mm. and the greatest width 12'5 mm. 
 The base of the wing is much more nearly complete than usual, and very narrow 
 (5 mm.) for a short distance, beyond which it widens by the development of the 
 strong convex inner margin. The wing shows the usual plication, which fortu- 
 nately is not continued into its base, so that the course of the veins in the latter 
 is not obscured, as is so often the case in P>. priscotincta. The outer or costal 
 margin forms an almost straight line, and bears a double row of spinules; those at 
 the base of the wing pointing inwards. No basal hump, as in /?. priscotincta, is 
 shown. The subcosta is widely spaced from the outer margin proximally. The 
 radius and radial sector present no special features, and but one inward branch is 
 present. Whether four branches arose from the main stem, as in the type-species, 
 cannot be determined owing to the loss of the apical portion of the wing. It is 
 doubtful if such was the case, as the portion missing is not great. The outer three 
 branches of the radial sector are usually 10 12 mm. apart, so that if the wing
 
 " PTERONEPIONITES." G7 
 
 possessed the same number of brandies of the radial sector as in 1L priscotincta, 
 its total length would have been 70 mm. an unusual length. The median vein is 
 the most powerful of the whole series, and occupies a middle diagonal area equal 
 in extent to the combined costal and radial areas. The first branch arises in line 
 with the radial sector, and much nearer the base of the w r ing than in 13. priscotincta, 
 and also lies so much nearer the cubitus that the area separating the two is but 
 half the diameter of the area in the former species. This shortening of the main 
 stem and its movement inwards has been brought about by the division of the 
 second branch into two equal twigs, which pass out to the margin between the 
 first branch and the main stem. The median vein therefore takes a larger share 
 in the wing-structure than in B. priscotincta, and the radial and cubital areas are 
 correspondingly reduced. The cubitus presents no special features. Notwith- 
 standing the shortening up of the area separating it from the median, the area 
 between the cubitus and the anal veins is very large. The cubitus has suffered no 
 displacement by the increased division of the median. The first anal vein is very 
 long, passing well beyond the first third of the wing and dividing just before 
 reaching the wing-margin. The second anal vein is two-thirds the length of the 
 first, and bends inwards more gradually to the margin, giving off two short oblique 
 branches in its basal half. 
 
 " PTERONEPIONITES," 
 
 Many Iarva3 of fossil blattoids have been recorded, but very few of other 
 groups. These larvae may eventually reveal the changes undergone up to the 
 adult stage, and the development of the neuration of the wings. In addition, the 
 occurrence of larvae in deposits is more likely to be indicative of habitat than the 
 presence of adult wings, as the inability of Iarva3 to fly and their lesser power of 
 flotation would ensure inclusion in adjacent deposits. It is therefore necessary 
 that their occurrence, and as much as possible of their structural appearance, be 
 fully recorded. Any attempt to classify them under genera and species would 
 rather retard than accelerate progress, and it seems advisable to record them 
 under some term which will leave no doubt of their larval character. Handlirsch 
 ('Amer. Journ. Sdi.' [4], vol. xxxi, p. 375, 1911) has already described 
 larval "wing-cases" of a somewhat similar character under the generic name of 
 " Lameereites" and placed them under the order Megasecoptera, 
 
 Had not Handlirsch given a generic value to the name " Lameereites," it would 
 have been possible to extend the use of his term to all larval wings. Failing this 
 I would suggest the use of the word " Pteronepionites " for all larval wings which 
 cannot be referred to a known genus, adding a specific designation when any larval 
 wing presents features of a definitely recognisable character. Handlirsch restricts 
 the name " Lameereites " to the wings of larval Megasecopteridaj, but recognises the
 
 68 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 close similarity between them and Pteronepionites (' Revision der Palaozoischen 
 Insekten,' p. 76). 
 
 "Pteronepionites" johnsoni, sp. nov. Plate IV, fig. 4; Text-figure 20. 
 
 Type. Immature wing, 12 mm. long and 4 mm. wide; British Museum 
 (Madeley Collection, no. I. 2967). 
 
 Horizon and Locality. Middle Coal Measures (binds between the " Brooch " 
 and " Thick " coals) ; Coseley, Staffs. 
 
 Description. The wing, like all these immature structures, is of extreme 
 tenuity, and the finer details are masked by the coarse granular nature of the 
 matrix composing the nodule. The outer margin is straight, curving backwards 
 until it meets the inner margin in a bluntly-pointed apex. The inner margin is 
 strongly convex, a distal infolding indicating that the wing was apparently never 
 fully extended. Very feeble traces occur of a short straight subcosta, which 
 
 FIG. 20. "Pteronepionites" johnsoni, sp. nov.; diagram of wing-neuration, two-and-a-half times 
 natural size. Middle Coal Measures (binds between the "Brooch" and "Thick" coals) ; Coseley, 
 Staffordshire. Madeley Collection, Brit. Mus. (no. I. 2967). 
 
 reached the margin near the apex of the wing. The radius is stout and straight 
 and reaches the wing-apex, giving off a well-marked radial sector, which, diverging 
 from the radius, passes in a wide curve into the wing-apex. The median remains 
 undivided for nearly a third of its length, and then bifurcates into two equal 
 branches which assume a parallel position, and reach the outer third of the inner 
 margin. The cubitus is represented by two veins, the first having a sigmoidal 
 sweep and the second a simple curve. Both reach the middle third of the inner 
 margin. No trace of anal veins can be observed. 
 
 " Pteronepionites " ambigua, sp. nov. Plate IV, fig. 5. 
 
 '/'///">. A pair of larval wings still attached to the crushed and almost 
 obliterated body ; British Museum (Madeley Collection, no. I. 2968). 
 
 Horizon and Locality. Middle Coal Measures (binds between the "Brooch" 
 and " Thick " coals) ; Coseley, Staffs. 
 
 Description. The insect lies in a fragment of a light grey ironstone nodule, 
 and is not well preserved. One wing is whole, but more than half of the second 
 is missing, and its apex is obscured by the matrix. The whole wing has a length 
 of 6-5 mm. and a width of 2 mm. Each is much thickened over the basal half,
 
 "PTERONEPIONTTES" LEPUS. 69 
 
 and strongly ridged or furrowed along the line of a powerful vein or tracheal 
 trunk (the wing is so immature as to render the latter possible), occupying the 
 position of the radius and median veins. Traces of a third vein occupying the 
 position of the cubitus are present on the inner half of the wing. The body- 
 segments are numerous, 34 mm. in depth, and seem to have had pleura-like 
 expansions. The region in front of the thorax is bent backwards and below the 
 abdominal segments. The thoracic segments are larger and more robust than 
 those of the abdomen. 
 
 The thinness of the body, and the difficulty of determining boundaries with 
 satisfactory accuracy, render all attempts at a more precise determination impos- 
 sible. We can, however, say with confidence that the remains are those of an 
 insect possessing a long segmented body, an elongated head-region, and wings 
 carried upright over the back. 
 
 " Pteronepionites " lepus, sp. nov. Plate IV, fig. 6. 
 
 Type. Remains of a larval insect, having a segmented body and two immature 
 wings, in a flattened nodule of light-grey ironstone ; British Museum (Madeley 
 Collection, no. I. 2969). 
 
 Horizon and Locality. Middle Coal Measures (binds between the " Brooch " 
 and " Thick " coals) ; Coseley, Staffs. 
 
 Description. The impression of two wings is clearly discernible, the segments 
 of the abdomen are less so. With oblique lighting, six segments can be 
 made out behind the thorax. The wings are slightly unequal in size, and the 
 anterior is longer and thicker than the posterior wing; it is also somewhat 
 infolded at the base. The differences between the two wings, and their relative 
 positions, would seem to indicate that they are the fore- and hind-wings of the 
 one side. 
 
 The fore-wing has a length of 9 mm. and a width of 2'5 mm., while the hind- 
 wing is 8 mm. long and 2 mm. wide. The fore-wing has a strap-shaped 
 appearance, and the line of attachment to the body is its broadest part. A stout 
 ridge, swollen at its junction with the body, traverses the greater part of the 
 length of the wing, dying out before the apex is reached. The wing is too small 
 for definite determination of this swelling, but it appears due to incomplete 
 extension of the wing-membrane, rather than to the presence of a vein. The distal 
 fourth is flat and thin, and the apex well rounded. The outer and inner margins 
 are parallel and undulated. A few faint and irregular lines may be indications 
 of veins. 
 
 The hind-wing has undergone greater expansion than the fore-wing, and lies 
 almost flat on the nodule. Both outer and inner margins are gently convex, the 
 convexity of the inner margin being the greater. Feeble traces of a subcosta, and
 
 70 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 of a stout vein which forks before the middle of the wing is reached, are dis- 
 tinguishable. The latter vein occupies the position I should assign to the radius. 
 No further traces of veins are visible. 
 
 The two wings are separated by an interval of 3 mm. at their bases. No 
 definite details can be made out in the thoracic region. Behind the hind-wing 
 are faint impressions of a series of abdominal segments. These are about three 
 times as deep as wide, and appear to have borne lateral spiny processes. There 
 are also traces of tubercles. 
 
 The head-region is only indicated by faint discolorations. As in the case of 
 other examples of " Pteronepionites " we have met with, the remains are so filmy 
 in character, and merge so much into the matrix, that it is impossible to define the 
 outer limits of the various segments with absolute clearness, and no attempt can be 
 made at a systematic determination of characters. 
 
 The specimen is a larval form with wings not yet fitted for flight, but with a 
 degree of differentiation in the fore-wings which indicates that they were thicker 
 and less flexible than the hind-wings. 
 
 The abdomen is long, wide, and well segmented, the lateral expansions being 
 not unlike those of Eupholeriaferox. So closely does the abdomen resemble the 
 segmented body of a Diplopod, that in the absence of the wings we believe it 
 would be readily classed as belonging to that group, and as possessing nothing in 
 common with insects. 
 
 It is difficult to resist the belief that these larval insects were capable of 
 crawling about in decaying vegetation, and that their larval life was thus spent, 
 until by successive ecdysis the wings had acquired sufficient strength to lift the 
 body from the ground, and enable the insect to fly. 
 
 Family ^ENIGMATODIDTE, Handlirsch. 
 
 1906. Haudlirsch, Proc. U.S. National Mus., vol. xxix, p. 683. 
 1906. Handlirsch, Die Fossileu Iiisekten, p. 116. 
 
 Wing strongly arched, and broadly rounded at the apex. Anal area small, and 
 not marked off from the rest of the wing. Subcosta reaching almost to the wing- 
 apex; radius simple, and radial sector with three divisions. Median with four 
 branches. Costa represented by an oblique vein with a terminal fork, followed by 
 three simple, strongly curved anal veins. Interstitial neuration partly of regular 
 cross-nervures, and partly of a polygonal network. 
 
 Handlirsch founded this family on an incomplete wing in which the greater 
 part of the outer border, subcosta and radius is missing. From the upper Middle 
 Coal Measures of Mazon Creek, 111., U.S.A.
 
 ^ENIGMATODES (?) REGULARIS. 71 
 
 Genus ^ENIGMATODES, Handlirsch. 
 
 1906. JEnigmatodes, Handlirsch, Die Fossilen Insekten, p. 116. 
 
 Generic characters as above. 
 JEnigmatodes (?) regularis, sp. nov. Plate IV, fig. 7; Text-figure 21. 
 
 7 'ype. Fragmentary wing; British Museum (no. In. 18,604). 
 
 Horizon and Locality. Middle Coal Measures (over the Barnsley Thick Coal) ; 
 Monckton Main Colliery, Barnsley, Yorkshire. 
 
 Specific Characters. Radial sector, median, and cubital areas occupying 
 most of the wing, and all with well-spaced branches united by straight nervures, 
 except in the marginal area between the median and cubital veins where they 
 form a slight mesh work. Inner margin well rounded. 
 
 Description. Little more than half of this wing is preserved, and lies on a 
 fragment of hard grey bind associated with broken-up plant-remains. The 
 greatest length of the fragment is 42 mm., and the width 15 mm. The length 
 
 FIG. 2l.JEnigmatodes (?) regularis, sp. nov. ; diagram of neuration of wing-fragment, one-and-a-half 
 times natural size. Coal Measures (over the Barnsley " Thick " coal) ; Monckton Main Colliery, 
 Barnsley, Yorkshire. Brit, Mus. (no. In. 18,604). Lettering as in Text-figure 16, p. 62. 
 
 of the complete wing was probably 50 60 mm., and the width 18 20 mm. The 
 outer margin is missing, and of the subcosta and radius only portions of the basal 
 third are present. These are separated by a narrow area crossed by short trans- 
 verse nervures. 
 
 In the distal third of the wing is the innermost branch of the radial sector, 
 which ends on the margin in a small fork. The line of fracture of the wing has 
 closely followed the line of the radial sector, and a portion of another branch of 
 the radial sector may have been present along the line of the extreme broken edge 
 of the wing. The median vein sweeps in a convex curve to far out on the inner 
 margin, giving off two well-spaced branches. The next two veins, one of which 
 forks, are more oblique in their course, and may also belong to the median, but 
 the basal curve rather indicates that these veins are cubital, as is the succeeding 
 vein, of which only a small portion is left. A trace of an anal vein is present near 
 the wing-margin. The interstitial neuration consists of well-marked transverse 
 nervures, regularly spaced in the outer parts of the wing, and uniting in a loose
 
 72 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 mesh work in the wide area between the inner branch of the median and the 
 cubitus. 
 
 Affinities. The nearest approach to a wing of this character is that of 
 JEnigmatodes daniehi, Handlirsch, in which the greater part of the outer margin 
 is also missing. The Yorkshire specimen, which may be a hind-wing, is three 
 times as long as JE. daniehi, and the veins have a stronger inward curvature. 
 In the absence of more definite details, it seems best provisionally to refer the 
 specimen to the genus JEnigmatodes. 
 
 Genus PSEUDOFOUQUEA, Handlirsch. 
 
 1906. Pseudofouquea, Handlirsch, Die Fossilen Insekten, p. 125. 
 
 Wings three times as long as wide. Cubitus with inner and outer branches. 
 Anal veins, so far as known, not united. Interstitial neuration of feeble cross- 
 nervures except between cubitus and first anal, where it is irregularly reticulate. 
 
 Pseudofouquea cambrensis (Allen). Plate IV, fig. 8; Text-figure 22. 
 
 1901. Fouquea cambrensis, Allen, Geol. Mag. [4], vol. viii, p. 65, text-fig, on p. 66. 
 1906. Pseudofouquea cambrensis, Handlirsch. Die Fossilen Insekten, p. 125, pi. xiii, fig. 5. 
 1916. Pseudofouquea cambrensis, Bolton, Quart. Journ. Geol. Soc., vol. Ixxii, p. 59, pi. iv, figs. 4 5, 
 and text-fig. 
 
 Type. A. broken left fore-wing, of which the two parts are preserved on 
 fragments of black shale ; one fragment, bearing the basal part of the wing, in 
 the Museum of Practical Geology, Jermyn Street (no. 7272), the other, containing 
 the impression of the distal 28 mm. of the wing, in the Welsh National Museum, 
 Cardiff (no. 13,120). 
 
 Hwizon and Locality. Lower Coal Measures (top of the Four Foot Seam) ; 
 Llanbradach Colliery, near Cardiff. 
 
 Specific Characters. Wings stout and obtusely pointed. Costa marginal and 
 flatly convex. Subcosta reaching margin in the outer third. Radius parallel with 
 subcosta, and ending in apex of wing. Radial sector diverging widely from radius, 
 and ending on the wing-apex in five divisions. Comstock regards it as a typical 
 dichotomous radial sector, with an accessory vein on the second branch (' Wings 
 of Insects,' 1918). Radius and radial sector occupying all the wing-apex. Median 
 dichotomously branched, with an accessory vein in the fourth branch. Cubitus 
 widely divergent from median, and giving off alternate twigs from its outer and 
 inner sides, those of the inner side being much the feeblest. Anal veins four in 
 number. Interstitial neuration of feeble cross-nervures, except between the base 
 of the first anal vein and the cubitus, where it is irregularly reticulate. 
 
 Description. The total length of the wing is now 32 mm., 9 mm. having been
 
 PSEUDOFOUQUEA CAMBRENSIS. 73 
 
 lost from its tip since it was measured by Allen, who gives the total length as 
 41 mm. The greatest width is 15 mm. 
 
 The subcosta is widely separated from the outer or costal margin in the base of 
 the wing, and gradually approaches and unites with it beyond the middle. The 
 radius is parallel with the subcosta throughout its length, and gives origin to the 
 radial sector basally. The radial sector diverges from the radius over the whole 
 of its course. It now shows but one inwardly directed branch, which forks at the 
 broken edge of the wing. Allen's figure indicates that two more branches were 
 given off, both being undivided. The radius and radial sector occupied almost the 
 whole of the apex of the wing. 
 
 The median vein forks in the basal fourth, the outer branch again forking 
 before the middle of the wing is reached. The inner branch of the median diverges 
 almost in a straight line from the outer branch, and also forks, and bears an 
 accessory twig upon the fourth branch. A feeble accessory twig appears to be 
 o-iven off near the middle of the wing, but dies out in the integument. The cubitus 
 
 FIG. 22. Pseudofouquea cambrensis (Allen) ; restoration of whole wing, natural size. Lower Coal 
 Measures (top of the Four Foot Seam) ; Llanbradach Colliery, near Cardiff. Basal portion of 
 wing in Mus. Pract. Geol. (no. 7272). Impression of apical portion of wing in the Welsh National 
 Museum (no. 13,120). 
 
 is a remarkable vein, unlike that of any other fossil wing (Comstock, ' Wings of 
 Insects,' 1918, p. 106). For nearly half its length it passes in a broad curve to 
 the inner margin, giving off a series of alternate twigs upon its outer and inner 
 sides, those of the inner being weaker than those of the outer side. The two 
 outer twigs are strongly developed, while those on the inner side of the cubitus, 
 four in number, are weaker and shorter. The feeble continuation of the main stem 
 reaches the margin between the two sets of branches. Four anal veins are 
 distinguishable. The inner two arise from a common base, the outer two not 
 uniting. This is unlike the condition in Fonquea, where the anal veins branch off 
 regularly from a single stem. 
 
 The area lying between the first anal vein and the main stem of the cubitus is 
 very wide much wider, indeed, than any other area. 
 
 The interstitial neuration consists of weak cross -nervures, except between the 
 base of the first anal vein and the main stem of the cubitus, where it is irregularly 
 reticulate. 
 
 Affinities. The characters of the cubital and anal veins definitely remove the 
 
 species from the genus Fouqnea, and the cubitus, with its strong, anteriorly 
 
 directed twigs, and its feebler inner series, is wholly unlike that of any other 
 
 insect, and would alone suffice to justify the generic rank given by Handlirsch. 
 
 10
 
 74 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 So far I am in agreement with Handlirsch, but I regard the enlarged areas 
 between the inner divisions of the radial sector and the cubitus, and between the 
 cubitus and the anal veins, as more suggestive of the Protorthoptera, notably 
 Thoronysis ingbertensis, Ammon. More than this cannot be said, and Pseudo- 
 fonquea cambrensis must be regarded provisionally as Palseodictyopterid, with a 
 possibility of Protorthopterid or even Orthopterid affinities. 
 
 INCERT^E SEDIS. 
 Genus ARCHJEOPTILUS, Scudder. 
 1881. Archxoptilus, Scudder, G-eol. Mag. [2], vol. viii, p. 295. 
 
 A wing of unusually robust type. Only one is known, consisting of not more 
 than the basal fifth of a whole wing whose total length may have been 25 '4 cm. to 
 35-5 cm. The fragment is too small for a correct determination of its systematic 
 position, and has been referred to widely separated families by various workers. 
 
 Archseoptilus ingens, Scudder. Plate IV, fig. 9 ; Text-figure 23. 
 
 1881. Archxoptilus ingens, Scudder, Geol. Mag. [2], vol. viii, pp. 295, 300. 
 
 1883. Archeeoptilm ingens, Scudder, Mem. Bost. Soc. Nat. Hist., vol. iii, pp. 217, 223, pi. xvii, 
 
 figs. 1012. 
 
 1885. Archxoptilus ingens, Brongniart, Bull. Soc. Amis Sci. Nat. Rouen [3], ann. xxi, p. 60. 
 
 1885. Archieoptilm ingens, Scudder, Zittel's Haudbuch der Palseontologie, vol. ii, p. 757. 
 
 1893. Archseoptilus ingens, Brongniart, Faune Entom. Temps Prim., p. 498, pi. xxxvii, fig. 6. 
 
 1906. Archteoptilus ingens, Handlirsch, Die Fossilen Insekteu, p. 117, pi. xii, fig. 18. 
 
 Type. Basal fifth of wing, in counterpart; British Museum (no. I. 3997). 
 
 Horizon and Locality. Middle Upper Coal Measures; between Shelton and 
 Clay Lane, near Chesterfield, Derbyshire. 
 
 Specific Characters. Wings very large ; costa, subcosta, and radius broad 
 and robust. The costal border spiny. Interstitial neuration of stout transverse 
 nervures. 
 
 Description. Only the basal part of the wing and its counterpart are 
 preserved, having a total length of 43 mm., and a greatest breadth of 33 mm. 
 Scudder's estimate of the length of the whole wing as 35'5 cm. is probably 
 excessive. 
 
 Scudder (he. cit., 1881) thus describes the specimen : "All the principal veins 
 are a millimetre or more thick, and the cross-veins of the upper interspaces are 
 tolerably distant, stout, prominent, and generally simple. The marginal (costa) 
 vein, forming the front (outer) border of the wing is studded with short oblique 
 spines (Pmacrotrichia). The other veins lie at very different levels on the stone,
 
 ARCH^OPTILUS INGENS. 75 
 
 and below the interspaces mentioned, seem rather closely crowded, and much 
 more curved, sweeping downward, while the upper veins show little tendency to 
 turn from a longitudinal course." 
 
 The great apparent width of the costa is caused by the formation of an 
 expanded chitinous bar along its outer margin, the free edge bearing the spines 
 described by Scudder. The costa, with its chitinous bar, the subcosta and 
 the radius are so broad as to appear strap-like, are widely separated, and the 
 intervening areas are crossed by equally strong nervures. The costa can be 
 distinguished on the cast from the frontal bar and appears as a narrow 
 rounded vein. 
 
 The subcosta is a very broad vein, crossed by oblique striae which are directed 
 outward from the upper edge of the vein. A broad interval (9 mm.) separates 
 it from the costa, the area being crossed by stout, slightly oblique, transverse 
 nervures. The general direction of the subcosta is such that it must have reached 
 the margin of the wing near the apex. At the base it is much enlarged, the 
 
 FIG. 23. Archaeoptilus ingens, Scudder; diagram of whole wing from Scudder's original restoration, one-quarter 
 r'natural size. Upper Coal Measures ; near Chesterfield, Derbyshire. Brit. Mus. (no. I. 3997). 
 
 enlargement probably indicating the attachment to the body of the insect. The 
 expanded inner portion of the base is fused with the equally expanded base 
 of the radius. 
 
 The radius diverges from the subcosta in its outward course, and is even more 
 enlarged than the subcosta. It is connected with the latter by a series of thirteen 
 transverse nervures in the wing-fragment. Most of these are slightly convex, 
 and two are united by lateral branching. 
 
 The median vein was apparently closely apposed to, or united with the base 
 of the radius, and is much less robust than the latter, the intervening area being 
 narrow, and crossed by short, thin, straight, transverse nervures which do not 
 appear to continue into the base of 'the wing. The cubitus is not so readily 
 distinguishable, and diverges sharply inwards towards the inner margin. The 
 anal veins are 4 5 in number, and are strongly curved inwards, occupying 
 not more than one-fifth of the inner margin of the wing. The first three may 
 have been united at their base. The wing is marked by a series of folds along 
 the lines of the principal veins. 
 
 Ani nities. This remarkable wing-fragment has caused considerable conjecture 
 as to its true character and relationship. Scudder in his second note (loc. cit.,
 
 76 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 1883) somewhat vaguely placed it, " with a strong degree of probability, in the 
 same general group as some other Paleozoic wings." This reference can only be 
 to the Palseodictyoptera. Later he published a restoration of the wing, and classed 
 it with the Protophasmidae. Brongniart at one time regarded it as a Dictyoneuron, 
 and later as a " Neurorthopteron " of the group Sthenaropterida. Still later 
 he placed it as a Neuropteron in the group Platypteridae. Handlirsch considers 
 that the costa is marginal, and that this character, with the sharp inward curve 
 of the anal veins, justifies its inclusion in the Palaeodictyoptera, although he does 
 not attempt to indicate its allies. 
 
 A second species was described by Brongniart as A. lu.casi (' Bull. Soc. Amis 
 Sci. Nat. Rouen' [3], aim. xxi, p. 60, 1885), but this throws no light on the 
 genus, nor does it appear to be generically referable to Archseoptilus. 
 
 The only details preserved which can be used in the determination of relation- 
 ship are the spiny outer margin, the great width of the principal veins, the 
 well-developed cross-nervures, and the strongly curved and numerous anal veins. 
 Even these are too fragmentary for safe conclusions to be drawn in the absence 
 of other material. 
 
 The general structure of the wing-fragment is to me more suggestive of the 
 Protodonata than of the Palaeodictyoptera, but the presence of well-marked anal 
 veins discounts this view, unless we are prepared to accept the specimen as an 
 early and archaic prototype of the Protodonata. 
 
 Order MIXOTERMITOIDEA, Handlirsch. 
 
 1906. Haudlirsch, Proc. U.S. National Museum, vol. xxix, p. 695, and Die Fossilen Insekten, p. 126. 
 1919. Handlirsch, Revision dev Paliiozoischen Insekteu, p. 26. 
 
 Subcosta much shortened ; radial sector arising close to the base, with 23 
 branches, of which only one forks. Median long, four-branched and suggestive of 
 the Pateodictyoptera. Cubitus with 23 inward branches. Anal vein 3, simple. 
 Cross-nervures strong, wide-spread and regular. 
 
 This is a provisional order established by Handlirsch to include two forms 
 onlyMixotermes luganensis, Sterzel, from the Coal Measures of Saxony, and 
 Geroneura wilsoni, Matthew, from the Carboniferous of St. John, New Brunswick, 
 North America. Both wings show clearly their Palseodictyopteroid ancestry, but 
 Handlirsch is uncertain whether they should be brought near to the Protorthoptera 
 or to the Perlidae. 
 
 Genus GERONEURA, Matthew. 
 1889. Geroneura, Matthew, Trans. Roy. Soc. Canada, vol. vi, sect, iv, p. 57. 
 
 Generic Characters. Wing three times as long as wide; apex obtusely rounded;
 
 GERONEURA (?) OVATA. 77 
 
 subcosta short and joining outer margin before middle of wing. Radius and 
 radial sector occupying almost the whole of the wing-apex. Median, cubitus, and 
 anal divisions few. Interstitial neuration of stout cross-nervures at wide intervals. 
 
 Geroneura (?) ovata, sp. nov. Plate V, fig. 1. 
 
 Type. Portion of left wing; British Museum (Madeley Collection, no. I. 2965). 
 
 Horizon and Locality. Middle Coal Measures (binds between the "Brooch" 
 and " Thick " coals) ; Coseley, near Dudley, Staffs. 
 
 Specific Characters. Radial sector rising in outer third of wing, with three 
 divisions. Median vein large, the two outer branches forking in line with origin 
 of the radial sector, the third branch undivided. Cubital veins few, undivided, and 
 reaching the distal part of the inner margin of the wing. 
 
 Description. The specimen consists of the impression of the upper surface 
 of the distal portion of a left wing, having a length of 32 mm., and a breadth of 
 20 mm. The impression lies on the surface of a thin flattened half-nodule of fine 
 sandy grit, and is but faintly indicated. The total length of the wing was 
 probably from twice to three times the length of the portion preserved, and its 
 breadth may have been a little more than 20 mm. The outer margin is gently 
 convex, and curves into the broadly rounded apex. Very little is left of the inner 
 margin, which also seems to have been convex. The distal portion of the costal 
 margin is present for a length of 20 mm. There is no trace of the subcosta, so 
 that this vein did not extend much, if at all, beyond the middle of the wing. The 
 radius gives off the radial sector about the distal third of the wing, the two veins 
 remaining almost parallel with the wing-apex. The radial sector gives off a single 
 inward forked vein. The next three veins seem to be divisions of the median. 
 The first two each give off an outer branch in line with the division of the radius 
 and radial sector, and the outer of the two also forks before reaching the edge of 
 the wing. The third vein is single for its whole length, but evidently united with 
 the second a short distance outside the line of fracture of the nodule. The 
 remaining four veins appear to belong to the cubitus. No anal veins are 
 distinguishable. All the veins, with the exception of the small forks of the 
 radial sector and first median, are parallel and widely spaced. They are united by 
 a series of strong, straight cross-nervures placed widely apart. Notwithstanding 
 the strength of the veins and of the cross-nervures, the smooth impression of the 
 wing-fragment seems to indicate that the veins were not sunk below the general 
 surface of the wing, as is usually the case. 
 
 Affinities. The determination of the relationship of so small a wing-fragment 
 would be difficult were it not for the unusual direction of the main veins, their 
 mode of branching, and the character of the cross-nervures. These characters are 
 a special feature of the order Mixotermitoidea, Handl.
 
 78 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 The wing-fragment must be referred to this provisional order in the absence of 
 knowledge of the whole wing-structure. The much-divided median vein is more 
 comparable with that of Geroneura wilsoni, Matthew, than with that of Mixotermes 
 luganensis, Sterzel, and is also correlated with a shorter subcostal vein, although in 
 G. wilsoni that vein extends beyond the point at which the radial sector arises from 
 the radius. An open series of cross-nervures is present in both genera, as in this 
 specimen, and both have the same well-rounded apex. The wing-fragment is 
 suggestive of Hemeristia occidentalis, Dana, but has a less branched radial sector. 
 I provisionally refer it to Geroneura with the specific name of ovata. 
 
 Order PROTORTHOPTERA, Handlirsch. 
 
 1906. Handlirsch, Proc. U.S. National Museum, vol. xxix, p. 695, and Die Fossilen Insekten, p. 123. 
 1919. Haudlirsch, Eevision der Paliiozoischen Insekten, p. 28. 
 
 Head large, with strong mouth-parts, and bearing long slender antennae ; 
 prothorax large and elongated, and the body strongly built. Legs either uniform 
 in character and fitted for running, or the hind-legs modified for leaping. Wings 
 more specialised than those of the Palaeodictyoptera, and capable of folding on the 
 abdomen when at rest, with the enlarged anal areas of the hind-wings doubled 
 under, owing to the formation of a fold between the anal area and the rest of the 
 wing. The principal veins and their subdivisions not so strongly curved inwardly 
 as in the Palseodictyoptera. 
 
 Handlirsch established this order to include a series of insects intermediate in 
 character between true Orthoptera and Palaeodictyoptera, to which Scudder had 
 previously given the name of Pala3odictyoptera Neuropteroidea. 
 
 Genus JED(EOPHASMA, Scudder. 
 1885. jEdceophasma, Scudder, Geol. Mag. [3], vol. ii, p. 265. 
 
 Generic Characters. Large wings two-and-a-half times as long as wide ; inner 
 margin more convex than outer margin, and curving distally into the latter. 
 Principal veins broad and flat in the basal third, and diminishing in size distally. 
 Subcosta and radius reaching the wing-apex. Median vein with two main 
 branches, the outer with most subdivisions. Cubitus with two main branches, each 
 much subdivided. Anal veins numerous. Interstitial neuration of irregular 
 nervures, and a loose meshwork in the wider areas. 
 
 -Edceophasma anglica, Scudder. Plate V, fig. 2 ; Text-figure 24. 
 
 1885. ^Edoeophasma anglica, Scudder, Geol. Mag. [3], vol. ii, p. 265, and in Zittel's Handbuch der 
 Palseontologie, vol. ii, p. 758, fig. 941.
 
 vEDCEOPHASMA ANGLIC A. 79 
 
 1906. Mdcewphasma anglica, Handlirscli, Die Fossilen Insekten, p. 125, pi. xiii, fig. 4. 
 1916. jEdoeophasma anglica, Bolton, Quart. Journ. Geol. Soc., vol. Ixxii, p. 43, pis. iii, iv, and 
 text-figure. 
 
 Type. Greater part of a left wing in an ironstone nodule ; Liverpool Museum 
 (presented by Major Chambers in 1858). 
 
 Horizon and Locality. Middle Coal Measures; South Lancashire (locality 
 unknown, but the nodule so similar to those derived from the Ravenhead Railway 
 Cutting that it may be from that section). 
 
 Specific Characters. As generic characters. 
 
 Description. The specimen was partly described and named by Scudder in 1885, 
 and re-examined and figured by the present writer in 1916. 
 
 The wing lies in counterpart in a fine-grained ironstone nodule, and its total 
 length as now exposed is 87 mm., its greatest breadth (across the middle) 40 mm. 
 When whole, the wing was probably 100 mm. long. 
 
 FIG. 24. JEdmoplnasma, anglica, Scudder ; restoration of whole wing, showing the general character of 
 the venation, natural size. Middle Coal Measures ; South Lancashire. Liverpool Museum. 
 
 The outer or costal margin is gently convex. The subcosta is a broad flat 
 vein, gradually diminishing in width towards the wing-apex, which it just fails 
 to reach. 
 
 The radius is an even broader vein than the subcosta, is also flattened in its 
 basal third, and reaches the outer angle of the wing-tip, keeping parallel with 
 the subcosta. 
 
 The median vein divides low down into two equal branches, the outer giving 
 off four inwardly directed twigs. The first of these remains undivided ; the second 
 forks twice, and the outer and inner divisions of the second bifurcation again 
 divide, so that the vein ends on the apical margin in six divisions. The remaining 
 two branches are undivided. The divisions of the outer branch of the median 
 occupy the greater part of the wing-apex. 
 
 The inner branch of the median does not divide until it has reached the apical 
 fourth of the wing, where it gives off four twigs which pass inwards to the 
 junction of the inner margin with the apex. Only the first of these twigs forks. 
 
 The cubitus has the same broad flattened basal portion which is so character- 
 istic of the veins we have already dealt with. The main stem lies somewhat near
 
 80 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 the first branch of the median, and remains parallel with it over almost the whole 
 of its length. Inwardly it is separated somewhat widely from a slighter vein 
 which, I believe, joined it near the base, and formed the first inward branch. The 
 main stem sends off at the middle of the wing a strongly curved branch which 
 bends first inwards and then outwards towards the apex, breaking up into five 
 twigs before reaching the inner margin. The second of these twigs forks. A 
 second undivided branch comes off a little further out, and a third very small one 
 almost on the margin. The next two veins were probably united a little way out 
 from the base, and their direction is such that the single stem from which they 
 arose may have arisen, as suggested above, as the first inward branch of the 
 cubitus. The outer of the two veins is undivided, and reaches the inner margin 
 beyond the middle of the wing. The innermost vein runs fairly parallel with the 
 first along its whole length, giving off, as it does so, four inwardly directed twigs, 
 of which the first and fourth fork. The whole vein ends on the margin in six 
 twigs. Four anal veins are shown, one only forking. 
 
 The interstitial neuration of the radial and median areas consists of straight 
 or slightly curved nervures, placed at nearly equal distances. The very wide 
 cubital and cubi to-anal areas are filled by a loose mesh work, and a few irregular 
 wavy nervures. The anal area is crossed by simple straight nervures. 
 
 Affinities. Scudder was originally of opinion that this wing was related to 
 Meganeura (Dictyoneura) mont/i, Brong., representing a member of the group 
 Protophasmidse. Handlirsch removed the genus to the group of Palseodictyoptera 
 incertae sedis. 
 
 Scudder was undoubtedly mistaken in referring the wing to the Protophasmida?, 
 as a glance at the figure of Protophasma dumasii, Brong., will at once show (' Die 
 Fossilen Insekten,' pi. xvi, figs. 12). Handlirsch did not see the specimen, and 
 had to base his determination on a sketch of the wing which he considered " con- 
 fusedly drawn." The latter probably accounts for the interpretation which he 
 placed on the various principal veins. More recently I have been able to expose 
 more of the structure, and diagnosed the wing accordingly. If Handlirsch 's view 
 were correct, the radial sector would be of enormous proportions, and occupy all 
 the wing-apex. The base of the radius, so far as shown, is widely divergent from 
 the base of the median more so, in fact, than at any other part of the whole 
 course of the radius and supposed radial sector. These veins, therefore, have 
 come into union only at the actual point of origin of the wing. This may have 
 been the case, but in my opinion, the radius is wholly simple and undivided, and 
 no radial sector is present. The median and cubitus are large, much divided, 
 and take up the greater part of the wing-area, while the anal veins are few. 
 
 If this view be correct, the wing is a very primitive example of the Protor- 
 thoptera, still retaining evidence in the costa, subcosta and radius, of its 
 Palasodictyopteroid origin.
 
 COSELIA PALMIFORMIS. 81 
 
 Family COSELIID^E, nova. 
 
 A group of large- winged insects in which- the principal veins are openly 
 spaced ; the radial sector joined by an outer branch of the median, and the median 
 sending a strong inward commissure to the cubitus. 
 
 The family shows relationships with the (Edischiidae, Omaliidae, and Cacurgidaa. 
 
 Genus COSELIA, novum. 
 
 Generic Characters. Costa, and subcosta feeble, intercostal area very wide ; 
 radius straight ; radial sector arising near middle of wing. Cubitus S-shaped, 
 its branches occupying most of the inner margin of the wing. Interstitial 
 neuration of transverse nervures uniting in some areas to form an open 
 meshwork. 
 
 Coselia palmiformis, sp. nov. Plate V, fig. 3 ; Text-figure 25. 
 
 Type. Basal half of a left wing in counterpart in an ironstone nodule ; British 
 Museum (Johnson Collection, no. I. 15893). 
 
 Horizon and Locality. Middle Coal Measures (binds between the "Brooch" 
 and " Thick " coals) ; Coseley, near Dudley, Staffs. 
 
 ^l>cific Characters. Subcosta united to radius basally, giving off numerous 
 branches, united by irregular cross-nervures, to the outer margin. Radius strong, 
 divergent from subcosta. Radial sector diverging from the radius. Median 
 steadily divergent from radius and radial sector, and giving off an inner branch in 
 the first third of the wing to the main stem of the cubitus. Beyond the middle of 
 the wing, the median sending off several outer branches, the first uniting with the 
 radial sector. Cubitus with its divisions extending over the greater portion of the 
 inner margin of the wing. First anal vein strongly curved, the next thrice forked. 
 
 Description, This species is represented by a little over the basal half of 
 a left wing, 54 mm. long and 26 mm. wide, showing the under surface only. The 
 outer or costal margin is convex and very thin. The costal area very wide at its 
 base (6 mm.) and diminishing towards the wing-apex. 
 
 The subcosta is a feeble vein, apparently united basally to the radius, and 
 passing outwards in a straight line towards the distal end of the outer margin, 
 or into the outer part of the wing-apex. It gives off numerous branches to the 
 outer margin, the branches being disposed regularly, and inclining apically in 
 their course. They are joined by a few irregular cross-nervures, most of which 
 are at right-angles to the branches of the subcosta. 
 
 The radius arises with an inward curve near the point at which the subcosta 
 is given off, diverging from it and following a straight line to the wing-apex. 
 11
 
 82 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 The radial sector arises near the middle of the wing, and at a distance of 
 38 mm. from the wing-base. The angle which it makes with the radius is 
 fairly large, and the enclosed area must have been very wide in the outer half 
 of the wing. 
 
 The median arises close to the radius, and then bends inwards as it passes 
 to the inner half of the wing-apex. Its divergence from the radius is more than 
 double that of the radius from the subcosta, Portions of two outwardly directed 
 branches are shown, the first joining the radial sector; only the base of the 
 second is preserved; it follows a course parallel with the first branch. About 
 21 mm. from the base the median gives off a strong inward branch passing 
 obliquely to the cubitus and fusing with it. The cubitus is S-shaped, being 
 convex outwardly to the junction with the commissural vein from the median, 
 and concave inwardly afterwards. Owing to this inward inflexion, that part 
 
 FIG. 25. Coselia palmiformis, gen. et sp. nov. ; diagram of basal portion of left wing, showing the 
 general character of the venation, three-halves natural size. Middle Coal Measures ; Coseley, 
 Staffordshire. Johnson Collection, Brit. Mus. (no. I. 15893). 
 
 of the cubital area lying beyond the junction with the median is extremely wide. 
 The cubitus gives off four inwardly directed branches at wide intervals, the first 
 branch having a wide forking in the first third of its length. The branches 
 of the median must have occupied the greater part of the inner margin of 
 the wing. 
 
 The cubitus gives off six inward branches, the first arising very low down, 
 and the remaining five at regular intervals, the first being given off a little way 
 beyond the junction with the commissural vein. The first and fourth of these 
 branches fork into two widely separated twigs. 
 
 The first anal vein is simple, and has a broad convex outward sweep down to 
 the inner margin ; the second anal vein gives off three branches on its inner side. 
 Other anal veins may have been present, but the wing-structure is missing at 
 this point. 
 
 With the exception of the subcosta and radius, all the main veins are widely 
 spaced, and no branches are given off, except the first median, in the basal third 
 of the wing. 
 
 The interstitial neuration is well marked, and of unusual character. That
 
 COSELIA PALMIFORMIS. 83 
 
 of the intercostal area has been already described; that between the subcosta 
 and radius, between the radius and radial sector, and between the radial sector 
 and the median, consists of a few irregular cross-nervures, one or two of which 
 unite. In the median-cubital area the cross-nervures are joined up by longitudinal 
 branches following a zig-zag course. A similar neuration is seen between the 
 inner branches of the cubitus, while the anal area is crossed by irregular 
 cross-nervures occasionally uniting, as in the fore-part of the wing. 
 
 Affinities. The general characters agree most closely with those of the 
 Cacurgidas, a family of insects which Handlirsch has described from ironstone 
 nodules found in and around Mazon Creek, Illinois, U.S.A. It comprises insects 
 which possess well-marked characters, but which, as a whole, Handlirsch is 
 unable to assign with certainty to the Protorthoptera, or to the Protoblattoidea 
 (Handlirsch, 'Amer. Journ. Sci.' [4], vol. xxxi, 1911, p. 323). 
 
 The wing of Gacurgus spilopterus, Handlirsch, has the same wide costal area; 
 the radial sector arises far out, and the median gives off a commissural vein 
 to the cubitus. The central portions of the median and cubital areas are also 
 very wide, and the anal veins have the same strong inward curve. 
 
 The interstitial neuration is much similar, but the formation of a mesh work 
 has not proceeded so far in the British specimen as in Gacurgus spilopterus, 'where 
 it is present between all the main veins, except the costal and subcostal. In the 
 British specimen the mesh work is chiefly developed in the median and cubital 
 areas. Areolse are present in the wings of both species. 
 
 The differences between the wings of the two species are, however, important. 
 In Gacurgus spilopterus, the first branch of the median does not unite with the 
 radial sector, nor does it approach it very closely, whereas in the British wing, 
 the first outer branch must have united with the radial sector. The median 
 also branches again further out, the median in 0. spilopterus having one outer 
 branch only. Lithosialis brongniarti (Mantell) shows less affinity to the wing, 
 but possesses the wide costal area, the same irregular and ramified interstitial 
 neuration, but a more complex median and simpler cubitus. The commissural 
 vein is also wanting. 
 
 One other form with which a comparison needs to be made is Omalia 
 macroptem, Van Beneden and Coemans. This species was obtained from the 
 Westphalian (Middle Upper Coal Measures) of Hainault, Belgium. In its wing 
 the costal area is wide, and crossed by oblique branches from the subcosta; the 
 median sends an outer branch to join the radial sector, but there is no union 
 of the median and cubitus. The interstitial neuration is not mesh-like. 
 
 These wings are so similar that there seems no doubt they are closely 
 related, but the relationship of the British specimen is closest with the genus 
 Gacurgus.
 
 84 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 Family (EDISCHIIDJE, Handlirsch. 
 
 1906. Handlirsch, Proc, U.S. National Museum, vol. xxix, p. 700, and Die Fussileu Insekten, p. 142. 
 1919. Haudlirsch, Revision der Palaozoischen Insekten, p. 39. 
 
 Radial sector coalescing with the main stem of the median, and branching as if 
 a part of the latter vein. 
 
 Handlirsch regards the family as closely related to the Sthenaropodidae. 
 
 Genus GENENTOMUM, Scudder. 
 
 1885. Genentomum, Scudder, Mem. Bost. Soc. Nat. Hist., vol, iii, p. 329. 
 1893. (Edischia, Brongniart, Faune Eiitom. Terr. Prim., p. 559. 
 
 Generic Characters. Wings about three times as long as wide ; principal veins 
 and their branches well spaced, and united by a series of straight and well- 
 defined nervures. Apex of wing obtuse. 
 
 Considerable doubt exists as to the true relationship of this genus, Scudder 
 referring it to the Homothetidse, Brauer considering it to have affinity with the 
 Sialida?, while Brongniart placed it with the (Edischiidse. 
 
 Genentomum (?) subacutum, Bolton. Plate V, fig. 4. 
 
 1911. Genentomum subacutum, Bolton, Quart. Jouru. Greol. Soc., vol. Ixvii, p. 334, pi. xxvii, 
 figs. 18, 19. 
 
 Type. Two small wing-fragments, 9 mm. in length, and 6 mm. in breadth; 
 Bristol Museum (no. C. 972). 
 
 Horizon and Locality. Lower Coal Measures (637 feet below the Bedminster 
 Great Vein, and 137 feet above the Ashton Great Vein) ; South Liberty Colliery, 
 Bristol. 
 
 Description. One wing is partly superposed on the other, and the lower is also 
 partially concealed by shale. The upper wing is represented by a portion of the 
 distal inner portion, and by the wing-apex which is bluntly rounded. The outer 
 margin is broken away, and the original size and shape of the wings cannot be 
 determined. 
 
 The manner in which the wing-fragments are exposed on the shale renders 
 systematic analysis of the neuration impossible. All the veins are narrow and sunk 
 in the wing-membrane, while the interstitial neuration consists of transverse 
 nervures arranged at right-angles, and dividing up the various areas into quad- 
 rangles. Where forking of the principal veins takes place, the resultant branches 
 first diverge rapidly, and then pass outwards in parallel lines and at right-angles 
 to the margin.
 
 XEROPTERA OBTUSATA. 85 
 
 The lower wing-fragment shows a well-defined outer margin, evidently bounded 
 by the costal vein. This margin is regularly and broadly convex, and shows signs 
 of passing proximally into a straight or incurved line. The apex of the Aving is 
 narrow and bluntly rounded. The principal veins exposed appear to be portions 
 of the radius, radial sector, and the distal branches of the median. 
 
 Affinities. Fragmentary though these wings are, they nevertheless present 
 features of unusual interest, as they are wholly unlike the blattoid wings found in 
 the South Wales Coalfield, and are as yet the only insect-wings found in the 
 Lower Coal Measures of the Bristol district. 
 
 The resemblance in venation and structure to locustid wings is remarkably 
 close. There are the thin, sharply defined branch veins, and the division of the 
 interspaces into rectangular areas by straight transverse nervures ; and the 
 texture is quite filmy, apart from the incised veins, as in Locustidse. 
 
 G-enus XEROPTERA, novum. 
 
 Generic Characters. Wings more than three times as long as wide. Outer 
 margin feebly convex. Subcosta ending far out. Radius reaching almost or to 
 the wing-apex. Radial sector united by commissure to median. Median large, 
 forming two main, equal-sized branches. Cubitus simple. Interstitial neuration 
 of numerous straight cross-nervures. 
 
 Xeroptera obtusata, sp. nov. Plate V, fig. 5 ; Text-figure 20. 
 
 Type. A right fore- wing in counterpart in nodule ; British Museum (Johnson 
 Collection, no. I. 1558). 
 
 FIG. 26. Xeroptera obtusata, gen. et sp. nov. ; diagram of venation of wing, three-halves natural 
 size. Middle Coal Measures; Coseley, Staffordshire. Johnson Collection, Brit. Mus. (no. I. 1558). 
 Lettering as in Text-figure 16, p. 62. 
 
 Horizon and Locality. Middle Coal Measures (binds between the " Brooch " 
 and " Thick " coals) ; Coseley, near Dudley, Staffs. 
 
 Description. The wing is not complete, the whole of the outer margin being 
 missing, or hidden by the matrix ; the apex is raggedly torn, and much of the
 
 86 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 inner margin is missing. So far as preserved it is 41 mm. long, and 10 nun. in 
 greatest breadth. 
 
 The fore- wing lies on a hind-wing which is partly traceable, and both have 
 been crumpled together, some of the veins of the hind-wing showing through 
 the fore-wing. This renders the unravelling of the venation difficult, while 
 the fragmentary condition of the stems of the main veins in the basal part 
 of the wing hinders the definite determination of their course, and their relation 
 to one another. 
 
 A small portion of the distally outer or costal margin has been uncovered 
 sufficient to show that it was slightly convex, ('lose to the margin is a trace of 
 the subcostal vein, proving that it continued well towards the wing-apex, if it did 
 not reach the latter. 
 
 The outer part of the wing has been torn away along the area separating the 
 subcosta and radius, and the latter vein is seen to traverse the whole length of the 
 wing, but whether it joined the outer margin distally or came out on the wing- 
 apex is not discernible. 
 
 The basal half of the radius is confused with the basal portions of the median 
 and cubitus veins, owing to the crumpling which the wing has undergone. The 
 radial sector arises in the basal half of the wing and diverges from the radius, a 
 wide area separating the two. Soon after its origin it receives a branch from the 
 median, and immediately afterwards gives off the first of three inwardly directed 
 branches, the middle one soon dividing into two equal twigs, while the first branch 
 forks on the margin. These divisions of the radius and the radial sector occupy 
 the whole of the outer part of the wing-apex. The first branch of the radial 
 sector probably represents the union of two twigs of the radius and median, and 
 the small marginal fork may also represent their separation. 
 
 The median vein is incomplete basally, and appears to be large and much 
 divided. Apparently it separates early into two main branches, the first dividing 
 into two almost equal twigs, which again fork, the outer branch of each fork going 
 forwards as an oblique commissure and joining the vein next in front. This is a 
 somewhat unusual feature. The course of the outer twig after its juncture with 
 the stem of the radial sector Ave have already traced. The outer twig of the 
 second branch of the median, after fusing with the inner twig of the first branch, 
 is probably continued by the inner of the two later divisions formed by that twig. 
 The third branch of the median passes obliquely inwards, giving off an outer twig 
 which forks twice, and, beyond the middle of the wing, gives off outer and inner 
 branches from one point, afterwards forking twice before it reaches the inner 
 margin of the wing. The whole structure ends in fourteen divisions on the distal 
 half of the inner margin, and occupies a considerable area of the whole wing. It 
 is possible that this complex of branching veins represents the median and cubitus 
 combined, but I do not think it likely. If such be the case, the first branch given
 
 XEROPTERA OBTUSATA. 87 
 
 off from the main stem is the median with four ultimate divisions, plus a branch to 
 the radial sector, all the rest forming a much-divided cubitus with outer branches. 
 
 If this interpretation is correct, the next veins in the series are anal. Unfor- 
 tunately, these are far too broken and uncertain in character to supply any 
 evidence on the point. The first vein has a bold sweep from the base of the 
 wing 1 well outwards towards the middle of the inner margin, and seems to divide 
 into a wide fork before reaching it. The second vein is simple, and may also fork, 
 but low down, although it is impossible to say whether or no the forked appear- 
 ance is due to a fold of the crumpled wing. If the two veins present are anal, 
 then the anal area is very large, and the divisions of the cubitus are directed so as 
 to lie almost wholly in the distal half of the wing. This appears unlikely. The 
 divisions of the cubitus are usually simple, or but once forked, and usually directed 
 inwards, ending normally on the middle of the wing-margin. The vein is united 
 to the undoubted median stem and has a similar mode of branching, while its 
 position on the margin is in the distal half. I see no reason, therefore, to regard 
 this vein as a cubitus. The next vein, with its bold semicircular sweep to the 
 margin, has the character of a cubitus, and so I regard it. 
 
 The second more incomplete vein may also be a cubital vein or the first anal. 
 The interstitial neuration consists of a numerous series of straight cross-nervures. 
 
 The wing is thin and membranous. 
 
 Affinities. The salient features of this wing include the union of the median 
 with the radial sector and of one branch of the median with another, the con- 
 siderable branching of the median and its extension on to the inner margin, the 
 reduced cubitus, a small anal area and the long spathulate shape of the wing. 
 
 These general characters belong in varying degrees to a group of families, 
 among which may be instanced the (Edischiidae, Sthenaropodidae, Spanioderidae, 
 and Greraridae. The genus* Becquerelia has the same development of the median, 
 and in B. superba, a fusion of an outer branch of the median with the radial 
 sector; but the wing is widest across the anal region, and the cubitus has a 
 correspondingly increased development. 
 
 With Sclnich&rtiella and Gerarus the relationship is perhaps closer, as in these 
 genera the median is large and much divided, and the. cubitus correspondingly 
 reduced. Material differences debar a reference of the specimen to any of these 
 families. J would assign it a position close to (Edischiidae, in which the outer 
 margin is nearly straight, the wing-apex rounded, and the median and radial 
 sector united. Dr. Tilly ard has suggested to me that the wing be cpmpared with 
 his archetype wing of the Panorpoid complex. There certainly appears to be a 
 relationship, but this wing is specialised beyond the archetype, and too fragmentary 
 for definite conclusions to be drawn from it. It cannot be referred to (Edischiidae, 
 however, because in that genus the subcosta is longer, the cubitus shorter, and 
 with fewer twio-s. I have therefore formed for it the new genus Xeroptera.
 
 88 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 Genus SCAL^OPTERA, novum. 
 
 Large wings, 60 mm. or more in length. Costa marginal and slightly curved. 
 Subcosta widely separated from margin and giving off straight, oblique and widely- 
 separated branches to the margin. Radius parallel with the subcosta, and giving 
 off the radial sector in the basal half of the wing. Median with few divisions. 
 Cubitus long, with few divisions. Wing feebly plicate. Interstitial nenration 
 forming a fine, close reticulation. 
 
 Scalseoptera recta, sp. nov. Plate VI, fig. 1 ; Text-figure 27. 
 
 2' ?/ _^. Basal half of left wing in counterpart, having a length of 29 mm., and 
 a width of 14 mm. ; British Museum (no. I. 13878). 
 
 Horizon and Locality. ~ Middle Coal Measures (binds between the "Brooch" 
 and " Thick " coals) ; Coseley, near Dudley, Staffs. 
 
 Desertion. The inner margin of the wing is lost or concealed in the matrix, 
 
 FIG. 27. Scalxoptera recta, gen. et sp. nov. ; diagram of the basal half of whig, twice natural size. 
 Middle Coal Measures ; Coseley, Staffordshire. Brit. Mus. (no. I. 13878). 
 
 while the anal area, and possibly a part of the cubital area, is missing. A portion 
 of the nodule containing the distal half of the wing is lost. The total length 
 of the wing could not have been less than 60 mm., and judging from its apparent 
 relationships, may have been more. 
 
 The surface of the wing is slightly plicated, two low ridges bearing the radius 
 and principal branch of the cubitus, while the subcosta and the median are deeply 
 sunk. Its membranous portion is strongly coriaceous, with stout cross-nervinvs 
 at wide intervals. The outer margin and the principal veins are feebly spinulose, 
 as best seen on the radius and radial sector. The outer margin is feebly convex, 
 the fairly straight course of the margin, and of all the principal veins, pointing 
 to the wing being much longer than wide ; a view also borne out, as we shall see 
 later, by its possible relationships. 
 
 The subcosta is widely removed from the margin, parallel with it, and giving 
 off a series of stout branchevS passing obliquely outwards to the margin. Of these 
 divisions, one only is clearly forked. 
 
 Two portions of the radius are present, a short* basal portion, and a second 
 piece further out. The vein is separated from the subcosta by an interval less
 
 SCAL^OPTERA RECTA. 89 
 
 than half that separating the subcosta and the margin. It lies also on a ridge 
 which increases in strength distally. The radius and snbcosta are parallel. The 
 actual point of origin of the radial sector is not shown, but can be determined 
 within narrow limits. It arises almost in the basal fourth of the wing, and so far 
 as its course can be traced, diverges from the radius. 
 
 The median diverges along its whole course from the radius, and gives off 
 on its outer side two branches, which remain undivided to the broken edge of 
 the nodule. The first branch arises in line with the origin of the radial sector, 
 and at an acute angle which soon enlarges, so that the interval separating it from 
 the radial sector is less than half that separating them at their origin. 
 
 The cubitus divides into two nearly equal branches very low down in the base 
 of the wing, the second branch sending off a short division against the broken 
 edge of the nodule. 
 
 The stem of the median, and the two branches of the cubitus, have a parallel 
 course, like the outer margin, subcosta and radius, the widening interval between 
 the two groups of veins being occupied by the radial sector, and the two outer 
 branches of the median. The direction of the subcosta, radius, radial sector, and 
 first branch of the median, indicates that they traversed almost the whole length 
 of the wing and ended on the wing-apex. The main stem of the median and 
 the branches of the cubitus would reach the distal half of the inner margin. 
 There are no traces of anal veins. 
 
 Affinities. The extreme width of the intercostal area, the marked divergence 
 of the costa, subcosta and radius from the median and cubitus, are features not 
 readily recognisable among other known fossil insects. The few widely-spaced 
 cross-nervures are not wholly peculiar to the specimen. Similarly, the spinulose 
 principal veins are paralleled by those of Brodia priscotincta and Archxoptihis 
 ingens. I have not seen so marked a coriaceous texture in any British fossil 
 insect. The length of the wing-fragment before branching of the principal veins 
 occurs, implies a long and relatively narrow wing. Such a wing-form is seen in 
 the Pala?odictyoptera and in the Protorthoptera. The wing differs from that of 
 Brodia priscotincta in its much greater size, in the presence of a well-developed 
 series of cross-nervures, and in the coriaceous surface of the wing-membrane. 
 The direction of the radial sector and the divisions of the median vein are unlike 
 those in Brodia, where these veins curve steadily from their points of origin 
 inwards to the wing-margin. 
 
 Two groups of insects described by Handlirsch ('Amer. Journ. Sci.' [4], 
 vol. xxxi, 1911), from the Pennsylvanian Series of the Carboniferous of Mazon 
 Creek, 111., under the family names of Spanioderida3 and Geraridae, appear to 
 resemble most nearly the wings we are considering. The wings in these families 
 are long and narrow, and obtusely rounded at the apex, and the principal veins, 
 especially in the Spanioderidse, pass straight out for the whole, or the greater 
 12
 
 90 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 part, of their length. Their costal area is also broader than the subcostal-radial 
 area, and in both families the interstitial neuration consists of straight cross- 
 nervures, except in the costal area, where they are oblique, as in this specimen. 
 
 These resemblances, however, are not sufficient to justify the inclusion of the 
 new wing among the Geraridae, because they are accompanied by equally or more 
 important differences, such as the spinulose character of the principal veins, the 
 great development of the intercostal area, the marked divergence of the stems 
 of the radius and the median, the coriaceous wing-membrane and the much fewer 
 cross-nervures. The wing, too, possesses quite a leathery texture, while the 
 cross-nervures are broad and flat, forming conspicuous details of the surface. 
 
 With our present knowledge, it is only possible to infer that the wing is 
 related to the Geraridas and Spanioderida3, that it is a Protorthopteroid, and that 
 it still retains traces of its Palasodictyopteroid ancestry. 
 
 Order PROTOBLATTOIDEA, Handlirsch. 
 
 The Protoblattoids have a well-rounded head, a prothorax showing little or no 
 expansion, and the wings intermediate in type between those of Palseodictyopteroids 
 and Blattoids. The anal areas of the fore-wings or tegmina 1 are well marked out, 
 and crossed by arcuate or oblique veins reaching the inner margin, while those of 
 the hind-wings are enlarged and marked off from the rest of the wing by an anal 
 fold. The body is more slender than in the majority of Blattoids. 
 
 The members of this order form a connecting link between the Palasodictyoptera 
 and the Blattseformes. In many respects there is a close resemblance between the 
 members of the order and those of the Protorthoptera, and the two may have 
 arisen from nearly related stocks. 
 
 Genus PTENODERA, novum. 
 
 Generic Characters. Subcosta reaching almost to the wing-apex. Radial 
 sector twice furcate ; median well divided, with its minor divisions crowded on the 
 inner half of the wing-apex. 
 
 I have formed this genus to include a wing-fragment of unusual character in 
 which the principal veins are long, and directed towards the wing-apex with few 
 divisions, and the interstitial neuration consists of straight cross-nervures. The 
 specimen shows that the branching of the principal veins is much similar to that of 
 Polyetes among the Protoblattoidea, and to that of Spaniodera schucherti, Handl., 
 among the Protorthoptera, in this respect supporting Pruvost, who has established 
 
 1 The thickened ri gl d fore-wings of Blattoids are usually termed " tegmina,"
 
 - PTENODERA DUBIUS. 91 
 
 a sub-order, Archiblattides, to receive many of the insects classed as Protorthoptera 
 by Handlirsch. 
 
 The wing is apparently related also to Scalasoptera recta, Bolton, and as the 
 latter shows no trace of the formation of an anal lobe, I class this genus with the 
 Protorthoptera rather than with the Protoblattoidea. The Protoblattoidea 
 probably arose from a Protorthopteroid stock. 
 
 Ptenodera dubius, sp. nov. Plate VI, fig. 2 ; Text-figure 28. 
 
 Type. Distal half of a left wing, on the split surface of one half of an ironstone 
 nodule; British Museum (Johnson Collection, no. I. 1559). 
 
 Horizon and Locality. --Middle Coal Measures (binds between the "Brooch" 
 and " Thick " coals) ; Coseley, near Dudley, Staffs. 
 
 FIG. 28. Ptenodera dubius, gen. et sp. nov.; diagram of apical portion of wing, twice natural size. 
 Middle Coal Measures ; Coseley, Staffordshire. Johnson Collection, Brit. Mus. (no. I. 1559). 
 
 Description. The ironstone nodule has been broken across, and that portion 
 which contained the basal half of the wing has been lost. The distal portion of 
 wing remaining is 27 mm. long, with a width of 16 mm. across the widest part. 
 The width of the basal half of the wing was probably greater. 
 
 The distal half of the wing is thin and membranous, and a little wrinkled. 
 The principal veins are well shown, but the interstitial nervures are only clearly 
 visible when the nodule is immersed in water. 
 
 The costa is marginal and gradually curves into a well-rounded wing-apex. 
 The subcosta is widely spaced from the costal margin basally, and gradually 
 approaches it as it passes out towards the wing-apex, which it does not reach. It 
 gives off to the wing-margin a series of oblique divisions, several of which fork. 
 
 The radius is simple, and passes straight to the wing-apex. Somewhat widely 
 separated from the radius is the radial sector, which must therefore have been 
 given off low down in the base of the wing. It divides into two equal branches, 
 each of which is again equally divided, so that it ends on the middle of the wing- 
 apex in four divisions. 
 
 The median vein is large and well branched. The nodule has been broken
 
 92 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 across just beyond the point of division of the two main branches. The outer 
 branch remains undivided until near the margin, where it forks twice, ending on 
 the inner side of the wing-apex in three divisions. The inner branch forks into 
 two equal twigs, the outer breaking up twice by equal forking into four divisions, 
 while the inner twig forks once. Most of the divisions of the inner branch of the 
 median end on the distal portion of the inner wing-margin. 
 
 Two incomplete branches of the cubitus are present, but the character of the 
 vein cannot be determined. No traces of the anal veins are shown. 
 
 The principal veins and their main branches are well spaced, and follow the 
 same general direction without being truly parallel. The interstitial neuration 
 consists of fine straight cross-nervures. 
 
 Affinities. The general character of the veins other than the subcosta is seen 
 in several Palaeodictyopteroid genera such as Eubleptus, but in none that I know 
 are these characters accompanied by a similar type of subcosta. In the oblique, 
 forked divisions of the latter, and the manner in which they are given off, as well 
 as in the course of the subcosta itself, the wing most resembles that of the Proto- 
 blattoids, and to this group I would assign it. 
 
 INCERT^ SEDIS. 
 
 Genus PLESIOIDISCHIA, Haridlirsch. 
 1906. Plesioidischia, Handlirsch, Die Fossilen lusekteu, p. 346. 
 
 Plesioidischia sp. Plate VI, fig. 3 ; Text-figure 29. 
 
 Type. A fragment of a right wing, 40 mm. long and 10 mm. wide ; 
 Manchester Museum (no. L. 4905). 
 
 Horizon and Locality. Middle Coal Measures (binds between "Brooch" and 
 " Thick " coals) ; Tipton, Staffs. 
 
 FIG. 29.-Plesioidischia sp ; diagram of venation of wincf-fragment, three-halves natural size.-Middle 
 Coal Measures ; lipton, Staffordshire. Manchester Museum (no. L. 4905). 
 
 Description. The wing-fragment lies on the irregular surface of a small 
 ironstone nodule, the outer costal margin and costal area on one half of 
 the nodule, and the middle portion of the wing on the other. The base
 
 BLATTOIDEA. 93 
 
 and apex of the wing are missing, or still concealed in the matrix. Few definite 
 factors are presented for determination, and the following description is the 
 best we can give. 
 
 The outer margin of the wing is formed by the costa, and seems to be 
 curved basally and straight distally. The costal area is widest at the base, 
 and gradually narrows by the approach of the subcosta, which doubtless reached 
 the outer margin far out. The costal area is crossed by a few stout straight 
 branches of the subcosta. The subcosta passes straight from the base to the 
 distal portion of the wing-margin. 
 
 The radius is close to and parallel with the subcosta. It gives off two 
 inward diverging branches, the first uniting with the median, but giving off a 
 secondary branch before it reaches the latter. The median gradually diverges 
 from the radius. It gives off two well-marked inner branches, and then bends 
 forwards to join the first branch of the radius. The point of union of the two 
 branches gives origin to a backward twig parallel with the secondary branch 
 of the radius, and with the two inner branches of the median. 
 
 The cubitus is represented by a long oblique vein going down to the inner 
 margin. It gives off near its base an outer branch, which curves inwards 
 parallel with the main stem, and reaches the wing-margin much beyond it. 
 The wing appears to have broken along the level of a succeeding vein, but 
 whether this was also a cubital or an anal vein it is impossible to determine. 
 
 The interstitial neuration consists of a series of stout strong cross-nervures. 
 The texture of the wing seems to have been membranous. 
 
 Affinities. The chief features of the wing are the strong parallelism of the 
 veins, the numerous and strong cross-nervures, and the unusual mode of union 
 of the radius and median. This assemblage of characters serves to place the 
 specimen in the Protorthoptera, but it is not referable to any known genus, and 
 is too incomplete to justify a new genus for its reception. The nearest approach 
 appears to be Progenentomum carbonis, Handl., in which the subcosta reaches 
 the middle of the wing, the radial sector unites with the median, and the interstitial 
 neuration is of straight cross-nervures. 
 
 Order BLATTOIDEA. 
 
 The most numerous and varied insects found in the Coal Measures are the 
 Blattoids, the recorded forms being classified in several families, embracing nearly 
 100 genera and 400 species. The early history of their study is dealt with by 
 S. H. Scudder ('Mem. Bost, Soc. Nat, Hist.,' vol. iii, pt. 1, no. 3, 1879), and need 
 not be repeated here. 
 
 Notwithstanding the abundance of specimens and forms known, they are 
 insufficient for a satisfactory classification of the group, and do not provide the
 
 94 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 stages in the life-history of any species. Handlirsch has recorded the occurrence 
 of several egg-cases of Blattoids from the Carboniferous (' Proc. U.S. National 
 Museum, 1 vol. xxix, p. 716, 1906). Larval forms have been described by 
 Scudder, Handlirsch, Woodward, and especially by Sellards ('Amer. Journ. 
 Sci.' [4], vol. xviii, p. 113, 1904), but in all cases the remains are too incomplete 
 for satisfactory study. 
 
 The Coal Measure Blattoids show unmistakably that the race had a much 
 earlier ancestry, as they depart widely from the Palasodictyopteroid type, and 
 have attained a high degree of specialisation. The eggs were enclosed in capsules 
 (Ootheca) much as in modern cockroaches, and the development was by a 
 progressive metamorphosis, in which the rudimentary wings were formed at 
 a relatively early stage, and increased in complexity of structure and size 
 at successive ecdyses. It would also appear that the rudimentary wings were 
 attached to the thorax by broad bases, and that the formation of an articular 
 joint^was not developed until the adult stage was reached. The elements of the 
 thorax were as well developed in Coal Measure times as they are to-day, both in 
 structure and function, while the legs, as seen in examples from Commentry 
 (Allier), France, were long, spiny, or covered with stiff hairs, and well adapted 
 for walking or running. 
 
 The wings display a remarkable diversity of neuration. The fore-wings are 
 invariably modified by the formation of chitin into stout, horny structures, usually 
 termed " tegmina," which served to protect the more delicate membranous hind- 
 wings concealed beneath them. The costa is always marginal. 
 
 The hind-wings are not well known. When found, they are thin, membranous, 
 larger than the tegmina, and folded beneath them. The enlargement of 
 the wing has taken place over the inner half, the costal and radial areas being 
 reduced, and the anal area not marked off from the rest of the wing by a furrow, 
 as is seen in the fore-wings or tegmina. The Avide variation in the neuration 
 of the tegmina furnishes the only satisfactory data for classification. Assuming, 
 as we must, that the Blattoids were derived from the Palseodictyoptera, the 
 simplest form of Blattoid must be that in which the wing-neuratiou most nearly 
 approximates to the latter type. This principle is the basis of Handlirsch's 
 classification. 
 
 Modification of the wing-membrane for flight in the case of the hind- wings 
 has brought about, or been accompanied by, a narrowing of the costal and 
 radial areas, a reduction in the strength and extent of these veins, and an 
 increased development of the median and cubital areas, coupled with an increased 
 growth of the wing-membrane inwardly. The hind-wing is, therefore, a much 
 more asymmetrical structure than the fore-wing or tegmen. 
 
 The abdomen is broad, somewhat flattened and well-segmented, but no positive 
 evidence is known of an invagination of the terminal segments to carry the 

 
 BLATTOIDEA. 95 
 
 egg-capsule. Cerci are present, and Sellards has identified long ovipositors, 
 though this identification is not accepted by Handlirsch. 
 
 The habits of Carboniferous Blattoids can only be inferred from those of living 
 forms. I have previously (p. 11) drawn attention to the occurrence of Blattoid 
 wings among the leaves of Gordaites bearing numerous shells of the small annelid, 
 Spirorbis piisillvs, and suggested that the Blattoids frequented decaying vegetation 
 to feed on the Spirorbis and similar organisms. 
 
 The tegmina of Phylomylacris mantidioides occur among masses of the spat 
 ("Ancylus vintii") of Anthracomya phillipsii in coal shales of the Durham Coalfield. 
 Most of the other Blattoid remains from the Coal Measures of this country have 
 been found in sedimentary deposits in which Ostracods also occur. Apart from 
 these facts, the general faunal associations of fossil insects to which I drew attention 
 on pp. 18 24 are, I think, of great importance, as furnishing definite clues to habits. 
 The same Arthropod-association still persists, and the lagunal and marshy phases 
 of the Coal Measures would furnish a fitting and desirable environment for the 
 whole series. 
 
 Classification. The classification of the Carboniferous Blattoids is difficult, 
 owing to lack of material. The known British forms are few in number, and 
 have been found in several coalfields at horizons which cannot be correlated with 
 each other or placed in a regular sequence. 
 
 Scndder's classification does not sufficiently take into account the relationship 
 which undoubtedly exists between the Blattoids and the Palasodictyoptera, and it 
 is based mainly on a knowledge of living forms. For this reason, probably, the 
 Mylacridae are placed as the first tribe in his group " Palaeoblattariae," in 
 preference to the Archimylacridas, in which a Palseodictyopteroid relationship is 
 more clearly evident. 
 
 Handlirsch's classification gives greater attention to the ancestral type, and is 
 also based on an examination of more numerous specimens than were accessible to 
 Scudder. It is far from satisfactory, and will certainly be much modified when 
 the collection of several thousand Blattoid remains from Commentry is fully 
 worked out by Prof. Meunier, whose results are not yet published. Lameere's 
 general classification ('Bull. Mus. Hist. Naturelle/ 1917, no. 1) is too uncertain 
 to justify adoption. 
 
 Dr. P. Pruvost, who has given several years to a close study of the fossil 
 Blattoids of the French and Belgian Coalfields, has modified Handlirsch's classifica- 
 tion, and in his latest memoir (1920) he classifies the Blattoidea of Northern 
 France as follows : 
 
 BLATTOIDEA. 
 
 Family ARCHIMYLACKID.S;. 
 
 Genera ActinoUatta, Manoblatta, ArcUmylacris, Asemollatta, PhyloUatta, Grypoblattina, 
 Arcluvotiphe, Barroisiblatta, and Mesitoblatta.
 
 96 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 Family MYLACRIDJE. 
 
 Genera Hemimylacris, Phylomylacris, TrilopJiomylacris, Soomylacrls, LitJiontylacrin, Ortlionujlacris, 
 Stenomylarr'i*. 
 Family POROBLATTINID^E. 
 
 Grenus Premnoblatta. 
 
 This classification seems to accord most nearly with recent discoveries, and is 
 therefore adopted here. 
 
 Family ARCHIMYLACRID.E, Handlirsch. 
 
 Body slender in primitive forms, broadened out in more specialised types. 
 Cerci well developed and jointed. Legs slender and armed with spines. Antennae 
 slender. Pronotura snbcircular. Subcosta distinct, with pectinate or equally 
 spaced branches. Radius well branched, with or without radial sector. Median 
 of one or two main branches. Radius, radial sector and median with outward 
 branches only. Cubitus inwardly curved, and with inward branches. Anal 
 furrow present. Anal area large, extending beyond the basal third of the wing. 
 Interstitial neuration of straight nervures, or an irregular reticulation, or the t 1 
 combined. 
 
 The greater number of Palaeozoic Blattoids belong to this family, which i* 
 united to the Palaeodictyoptera by transitional forms. 
 
 Genus APHTHOROBLATTINA, Handlirsch. 
 
 1906. Aphthoroblatfina, Handlirsch, Proc. U.S. National Museum, vol. xxix, p. 719, and Die Fossil* 
 Insekten, p. 183. 
 
 Generic Characters. Tegmina two-and-a-half times as long as wide, witl 
 convex outer margin. Costal area narrow and strap-shaped. Subcosta extending 
 over two-thirds the wing-length. Radius giving off radial sector near middle 
 wing, and sending a few simple branches to the apex. Radial sector forking inl 
 four or six branches. Median small, with few branches directed inwardly 
 Cubitus giving off 7 8 branches along its whole length, and extending well on 
 towards the wing-apex. Anal area small. 
 
 Aphthoroblattina johnsoni (Woodward). Plate VI, figs. 4, 5 ; Text-figures 30, 3 1 
 
 1887. Etollattina johnsoni, Woodward, Greol. Mag. [3], vol. iv, p. 53, pi. ii, figs. 1 o, 1 b. 
 1906. Aplithoroblattina, Handlirsch, Die Fossilen Insekten, p. 183. 
 
 Type. The pronotum and tegmina in an inverted position with traces of hind- 
 wings on a small ironstone nodule, in counterpart; British Museum (Johnsoi 
 Collection, no. I. 1067).
 
 APHTHOROBLATTINA JOHNSON!. 
 
 97 
 
 Horizon and Locality. Middle Coal Measures (binds between the " Brooch " 
 and " Thick " coals) ; Coseleyj Staffs. 
 
 Specific Characters.- Tegmina slightly convex and expanded towards the apex, 
 about two-and-a-quarter times as long as wide. Apex broadly rounded. Costal 
 area occupying three-fourths of the outer margin of the wing. Subcostal vein 
 weak. Radius almost straight and giving off the radial sector about the middle of 
 its length. Radial sector doubly furcate. Median vein widely spaced across the 
 middle of the wing. Cubital vein reaching the inner angle of the wing-apex, 
 giving off outer and inner branches. Anal area large, and crossed by a few small 
 anal veins. Hind- wings thin, larger than the tegrnina, and with strong veins. 
 
 Description. The elucidation of the neuration of the wings of the type 
 specimen is difficult, owing to the venation of the hind-wings being impressed 
 
 FIG. 30. Fm. 31. 
 
 PIG. 30. Aphthoroblattina johnsoni (Woodward) ; diagram of venation of the two fore-wings of the 
 
 type-specimen, twice natural size. Middle Coal Measures (binds between " Brooch " and " Thick " 
 
 coals) ; Coseley, Staffordshire. Johnson Collection, Brit. Mus. (no. I. 1067). 
 FIG. 31. Aphthoroblattina johnsoni (Woodward) ; diagram of venation of the fore-wings of a normal 
 
 example of the type-series, twice natural size. Same horizon and locality as Fig. 30. 
 
 W. Egginton's Collection, " no. 2." 
 
 on that of the tegmina. The veins of the hind-wings are stronger than the 
 distal portions of those veins of the tegmina which they overlie. It is probably 
 owing to these factors that the original figures of the type are incorrect. 
 
 In both tegmina, as indicated by Dr. Woodward, the median and cubitus are 
 shown as united, whereas this union takes place in the left wing only. Neither 
 of the figures gives an adequate indication of the presence of the hind-wings. 
 In the larger figure the radius and cubitus are represented by a single vein, 
 which is also incorrect. Allusion has already been made to the wings lying on 
 the smaller portion of the nodule in an inverted position. This is demonstrated 
 by the pronotum, which dips under the mesonotum into the matrix, and by the 
 deep impressions of the veins of the hind-wings crossing over, and not under, 
 the veins of the tegmina. The veins of the tegmina are also in relief, instead 
 of being sunken as they normally are on the dorsal surface. 
 13
 
 98 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 The pronotum has a diameter of 13 mm., and 8 mm. of its length is exposed. 
 The sides curve evenly backwards, and show no sign of flattening, so that its 
 general shape is semicircular. The hinder border is hidden, and disposed at 
 an angle to the surface-plane of the wings. The presence of a sunken, central, 
 pyriform area on the inner surface of the pronotum probably marks the position 
 occupied by the head of the insect. The downward inclination of the front 
 margin of the pronotum, and the elevation of the hinder border, is a frequent 
 feature in the Coal Measure Blattoids. 
 
 During life a small gap interposed between the pro- and mesonotum, and 
 this became filled up by sediment on burial in sufficient quantity to cause the 
 pronotum to be lifted from the body of the insect by that half of the nodule 
 which carries the wing-impressions. This inclined position of the pronotum is so 
 common that it appears to have been normal. 
 
 The mesonotal area is covered by the anal areas of the tegmina, and its 
 character cannot be determined. 
 
 The venation of the right tegmen differs considerably from that of the left. 
 The under surface of the latter is the more nearly perfect, and the venation is more 
 distinct. The costal margins of the tegmina form two broad arcs, sharply bent 
 inwards at the base, and distally merging into the wing-apex. The left subcosta 
 is widely separated from the margin, giving off three oblique branches which 
 fork, the first twice, before reaching the margin. Beyond these branches, 
 the left costa curves inwards and unites with the main stem of the radius at the 
 distal third of the wing. The right costa is more normal, and does not unite with 
 the radius. It gives off six outward branches, of which only the first is forked. 
 
 The left radius, after a slight basal curve, passes straight out to the apex, 
 giving off four forward branches, the first forked. This first forked branch 
 appears to be the continuation of the subcosta. The radius of the right tegmen 
 gives off two simple outer branches only. 
 
 There is a wide difference between the radial sectors of the two tegmina. 
 That of the left divides in the middle of its length into two equal branches, which 
 in turn fork, the outermost twig forking again, so that the radial sector ends on 
 the margin in five divisions. The right radial sector forks once only beyond the 
 middle of its length, and ends on the margin in only two divisions. 
 
 The median in each tegmen is a comparatively simple vein, that of the left 
 being united with the cubitus in the basal .fourth, and separating from it at an 
 acute angle. Beyond the middle of the wing it gives off a single forked branch 
 on its outer stfe. These divisions of the median, with the first branch of the 
 radial sector, occupy the inner half of the apex. The median of the right tegmen 
 is well separated from the cubitus along its whole length, and passes in a bold 
 curve to the inner half of the apex, giving off four simple outer branches. 
 
 Apart from its union with the stem of the median, the cubitus of the left
 
 APHTHOROBLATTINA JOHNSONI. 99 
 
 tegmen agrees fairly well with its fellow. Both veins curve inwards, and reach 
 the junction of the inner margin with the apex, and both give off 6 7 branches 
 inwardly, the first branch in each case forking. 
 
 Which forward branches were given off on the outward side of the cubitus in 
 the left tegmen cannot be made out, but faint furrows on the right tegmen seem 
 to indicate that three simple outer veins were present. 
 
 The anal area is long and acutely ovate, extending over nearly a third of the 
 inner margin. Six anal veins are present, the distal one with a strong fork. 
 
 The interstitial neuration consists of stout, straight nervures, not always well 
 shown. The inner margin is nearly straight. 
 
 The veins of the hind-wings are very fragmentary. They are much thinner 
 than those of the tegmina, and have left a much slighter impress. They appear 
 to consist of a straight costa, separated by a very narrow area from an equally 
 straight subcosta, below which can be made out a portion of the radius, radial 
 sector, and median. 
 
 All the veins are widely spaced, and the breadth of the hind-wings must have 
 been abont double that of the tegmina. 
 
 The dissimilarity between the neuration of the left and right tegmina in the 
 type-specimen suggests a wide varietal range among Coal Measure Blattoids, a 
 feature which needs to be taken into account in the definition of species. 
 
 The presence of these abnormalities in the type-specimen somewhat militates 
 against its value for reference, and I therefore add other details from a second 
 example which I received from Mr. W. Egginton. The specific characters given 
 above have been drawn up from this specimen, which lies in a small ironstone 
 nodule, marked " No. 2," having a length of 43 mm. and a breadth of 38 mm. 
 The Blattoid has a total length of 33'5 mm., and a maximum breadth across the 
 tegmina of 24 mm. The inner margins of the tegmina overlap, evidently in their 
 position of rest during life. 
 
 The pronotum, apparently in natural position, lies a little out of the horizontal, 
 the front margin dipping downwards, and the hinder margin upwards and a little 
 forwards, so that a slight gap occurs between it and the wings. This feature we 
 have already alluded to, and in this case the pronotum has been carried away as 
 usual, but the wings remain, so that the wing-impressions with the pronotum are 
 on the upper surface of one half of the nodule, the other half carrying the 
 pronotal impression and the wings. 
 
 The pronotum is 12 mm. wide and broadly rounded, only a little more than 
 half being visible. It shows a central raised area, oval in outline, and defined 
 from the margins by lateral grooves. The meso- and meta-notal segments show as 
 a low, flat, elongated, heart-shaped region overlain by the anal areas of the tegmina. 
 
 Both tegmina are still attached high up on the sides of the body. So far as 
 can be determined through the substance of the tegmina, the hind- wings are
 
 100 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 attached nearer the middle line, and almost on the upper surface. All four wings 
 are present, the tegmen of the right side 'being perfect, while that on the left has 
 lost a portion of the inner part of the apex. The right tegmen is 25 mm. long, and 
 12mm. wide at its broadest part. 
 
 The costal margin is broadly arcuate, and merges into a well-rounded apex. 
 The subcosta is feeble, and gives off 7 10 very oblique branches, each breaking up 
 into numerous twigs before reaching the margin. The intercostal area is wide and 
 strap-shaped, and occupies the greater part of the outer margin. 
 
 The radius is well separated from the subcosta at its base, and diverges a little 
 from it along its course. It gives off the radial sector before the middle of the 
 wing is reached, and then passes forward undivided until near the end of the sub- 
 costa, where it gives off four or more short oblique branches forward. 
 
 The radial sector diverges strongly from the radius", and, like the latter, remains 
 undivided for the greater part of its length. In the last third it gives off two, 
 possibly three, outer twigs, which fork before reaching the margin. The inter- 
 radial area at the point where the first branch arises is very wide. 
 
 The median arises close to the radius, but soon bends strongly towards the 
 inner side of the apex. It gives off three outer parallel branches, the first of 
 which forks. 
 
 The cubitus is a powerful and large vein, well apart from the median at its 
 origin, and passing above the middle of the wing in its proximal third, beyond 
 which it curves inwards to the end of the inner margin. It gives off six simple 
 branches and ends in a feeble fork, while two strong' branches arise on the outer 
 side and pass to the apex. 
 
 The anal area is large, occupying one-third of the margin, and containing six 
 anal veins, the first, second and third forking. 
 
 The inner margin is almost straight. The interstitial neuration consists of 
 numerous raised cross-nervures, best seen in the anal and cubital areas. In the 
 radial and median areas the cross-nervures unite laterally and occasionally fork. 
 
 Portions of the hind-wings are present, but only the distal branching of the 
 radial sector, median and cubitus is shown beyond the broken end of the left 
 tegmen. The course of the radius and median is traceable across the surface of 
 the tegmen of each side. 
 
 The wing-membrane is so thin as to be scarcely discernible. 
 
 Aphthoroblattina eggintoni, sp. nov. Plate VI, fig. ; Text-figure 32. 
 
 Type. Pronotum, tegmina and portions of hind-wings lying on the surfaces of 
 a split ironstone nodule; " No. 1 " in the Collection of Mr. W. Egginton. 
 
 Horizon and Locality. Middle Coal Measures (binds between the " Brooch " 
 and " Thick " coals) ; Coseley, near Dudley, Staffs.
 
 APHTHOROBLATTTXA EGGINTONI. 
 
 101 
 
 Specific Characters. Costal margin convex. Cubitus long, strap-shaped, and 
 extending almost the whole length of the outer margin. Radial sector with few 
 simple branches. Median vein forking three times, and ending in middle and 
 inner side of the wing-apex. Cubitus extending the whole length of the inner 
 margin, with no evident outer branches, and feeble forking. 
 
 Description. The specimen is contained in a small greyish-brown nodule, 
 48 mm. long and 35 mm. wide, and lies with the dorsal surface of the wings and 
 the impression of the pronotum on one half of the nodule, and the wing-impressions 
 and pronotum on the other half. The remains are in good condition, and show a 
 small portion of the head, the pronotum and mesonotum, almost the whole of the 
 left tegmen, and the basal two-thirds of the right tegmen, while a portion of 
 the left hind-wing is disclosed over that area from which the fragment of the 
 
 FIG. 32. AphthoroUattina eggintoni, sp. nov. ; diagram of venation of fore- wings and portion of left hind- 
 wing, twice natural size. Middle Coal Measures (binds between ' Brooch " and "Thick" coals) ; 
 Coseley, Staffordshire. W. Egginton's Collection, "no. 1." 
 
 tegmen is missing. That both hind-wings are present is indicated by their 
 principal veins showing in feeble fashion through the tegmina. 
 
 The head is small, not more than 3 mm. of its length being visible, and not all 
 of this would be seen were the front margin of the pronotum intact. The latter 
 has been broken away in the middle line, and allows a portion of the head to 
 become visible. The hinder part of the head is well rounded and narrowed, the 
 front being wide, and having a forward and downward slope. It is finely 
 tuberculated. A small aperture in the middle of the head may be due to a loss 
 of matrix. 
 
 The pronotum is attached to that half of the nodule which contains the 
 wing-impressions, and the hinder border is still covered with the matrix. Its 
 whole contour cannot be seen, and only the inner surface is exposed. It is semi- 
 circular, with a well-defined margin, 12 mm. wide, and 8 mm. long from front to
 
 102 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 back. The central area has a deep median furrow, and is marked off from the 
 lateral halves by two ridges (representing the furrows on the dorsal surface). 
 These ridges converge anteriorly and almost meet. The inner surface is slightly 
 concave, and indented by three faint furrows on each side, parallel with which are 
 traces of fine striations. 
 
 The mesonotum is 4'5 mm. long, and much wider anteriorly than posteriorly. 
 It is convex, and the hinder border has a raised rounded edge. It is strongly 
 chitinous, much more so than the metanotum, which is crushed down beneath the 
 anal areas of the tegmina, only a little of the hinder, portion of the left side 
 showing at one point where the anal areas of the wings have been broken away. 
 The right tegmen lies in the position of rest, its inner margin being in line witli 
 the axial line of the body. The left tegmen was rotated outwards to a slight 
 degree before entombment a feature which, combined with the breaking away of 
 its distal third, has enabled the left hind-wing to be better shown. The right 
 tegmen has a length of 33 mm. and a breadth of 13 mm. 
 
 The costal margin is moderately thickened, convex, and merges gradually into 
 the wing-apex, the latter bluntly rounded. 
 
 The subcostal vein is thin, and encloses a long strap-shaped area, crossed by 
 9 10 oblique branches, most of them forking twice before reaching the margin. 
 
 The radius is a strong vein, nearly parallel with the subcosta, and ending on 
 the outer margin in four small branches, the second only forking. The radial 
 sector arises just before the middle of the wing, diverging but slightly from the 
 radius. It gives off three short outward branches to the apex. 
 
 The main stem of the median passes along the middle line of the wing for some 
 distance, and curves inwards to the distal end of the inner margin. The first 
 outward branch arises a little beyond the middle of the wing, and forks into two 
 equal-sized twigs ; the remaining two branches remain undivided to the broken 
 edge of the wing. The median and its subdivisions occupy the centre and inner 
 half of the wing-apex. 
 
 In the left tegmen the second branch of the median forks. The cubitus is 
 strongly convex in the proximal two-thirds, and bends forwards in the distal 
 third and inwards again to the inner margin. It gives off six inward branches, 
 the fourth and fifth forking before reaching the margin. The anal veins are eight 
 in number, three being very small and short. The first forks twice, and the 
 second once. The inner margin is almost straight. The interstitial neuration is 
 made up of straight nervures. The surface of the tegmina is chitinous. 
 
 The hind-wings appear not only to have been covered by the tegmina, when 
 they were in a position of rest on the body, but to have stretched a little beyond 
 them like a fringe. 
 
 The left hind-wing is thin and membranous, and marked by numerous veins, 
 the bases still hidden under the tegmen. A little of the distal portion of the
 
 ARCHIMYLACRIS HASTATA. 103 
 
 costal margin is shown, passing into a very broad and expanded apex. The 
 subcosta reaches nearly to the end of the costal margin, and gives off numerous 
 oblique and forked branches Immediately behind the subcosta is a stout vein 
 with two outer branches, both forking. This vein seems to be the radius. ~Tts 
 basal extension under the tegrnen can be traced by a raised line. Behind the 
 radius are two stout veins, each of which forks into equal-sized twigs some 
 distance from the margin. Their position justifies the inference that they unite 
 proximally and join the radius, in which case they must belong to the radial 
 sector. The median is a doubly forked vein. The remainder of the wing is 
 either bent under, or folded upon itself, and further details cannot be made out. 
 
 Affinities. The species differs from AphtJioroblattinajohnsonim several important 
 details. The radial sector arises further out, and instead of forking twice, gives 
 off three simple forward veins. The median forks three times, the proximal 
 branch forking again before reaching the wing-apex. In A. johnsoni there are two 
 branches, both of which fork. The cubitus vein differs considerably owing to the 
 fact that the main stem divides at the outer third into two branches of equal 
 strength, the outer immediately forking again into two equal twigs, while the 
 inner gives off a small inner twig. No definite outer branches are distinguishable. 
 A strong anal furrow is present, and the first anal vein on both sides divides into 
 three twigs, the remaining 5 6 anal veins being undivided. 
 
 Genus ARCHIMYLACRIS, Scudder. 
 1868. Archimylacris, Scudder, in Dawson's Acadian Geology, ed. 2, p. 388. 
 
 Generic Characters. Tegmina twice as long as wide, with convex outer margin. 
 Inner margin concave. Radial sector arising in basal half of wing. Anal veins 
 few, anal area large. Neuration of abundant stout straight cross-nervures. 
 
 Archimylacris hastata, Bolton. Plate VII, fig. 1 ; Text-figure 33. 
 
 1911. Archimylacris (EtoUaUina) hastata, Bolton, Quart. Journ. Geol. Soc., vol. Ixvii, p. 160, pi. ix, 
 figs. 13. 
 
 TIJI><>. The greater part of a left tegmen and its counterpart, showing the 
 upper surface; Museum of Practical Geology, Jermyn Street (nos. 24501 and 24502). 
 
 Horizon and Locality. Upper Coal Measures (Gellideg Level of the Mynyd- 
 dislwyn Vein) ; near Maes-y-cwmmer, Monmouthshire. 
 
 Specific Characters. Wing broadly elliptical. Costal margin convex ; costal 
 area broad at base and stretching beyond the middle of the wing. Subcosta 
 with numerous branches, all oblique. Radius forking twice, and ending near
 
 104 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 apex of wing. Median branching further out than the radius. Cubitus 
 reaching margin in distal third of wing. Anal veins numerous. Inner margin 
 almost straight. 
 
 Description. A portion of the base of the wing and a considerable part of the 
 apical and inner margin are missing, but the length preserved is 33'4 mm. and its 
 breadth is 16' 5 mm. 
 
 The wing is strongly outlined, especially along the outer margin, partly owing 
 to a broad concave depression which runs almost the whole length of the subcostal 
 and radial areas, causing the anterior margin to be reflexed dorsally. The inner 
 margin is preserved only along a portion of the anal border, and the inner two- 
 thirds of the wing-apex are also missing. Sufficient is left of the apex to show 
 that it must have been bluntly rounded. 
 
 Fio. 33. Archimylacris hastata, Bolton ; diagram of venation of left fore-wing, three times natural 
 size. Upper Coal Measures (Gellideg Level of the Mynyddislwyn Vein) ; near Maes-y-cwmmer, 
 Monmouthshire. Mus. Pract. Geol. (nos. 24501, 24502). Lettering of veins. I. costa ; II. sub- 
 costa ; III. radius ; IV. median ; V. cubitns ; VI. anal. 
 
 The subcostal vein passes out obliquely from the point of attachment and 
 reaches into the distal third of the wing. It gives off 9 10 very oblique branches, 
 the basal three forking twice, the next two undivided, and the last branch but one 
 forked. The subcostal area is broad basally, and terminates in an acute angle 
 distally. 
 
 The actual origin of the principal veins is lost, and the radius appears at its 
 origin therefore to be widely removed from the snbcosta. It follows a parallel 
 course to the origin of the first anterior branch, and afterwards diverges, so that 
 it finally ends on the apex of the wing near the middle line. The first branch is 
 important, forking three times in its course and ending in six divisions. The 
 second branch forks only once. 
 
 The subcosta and radius together occupy a little less than half the whole wing- 
 area. Dr. Pruvost supposes that a commissural branch unites the radial sector 
 and the median in this species (1919, 'Memoires pour servir a 1'Explication de la 
 Carte Ge"ologique Detaillee de la France,' p. 151), but I cannot perceive this. 
 
 The median vein arises in actual contact with the stem of the radius, or is
 
 ARCHIMYLACRIS HASTATA. 105 
 
 united with it, rapidly diverging to the distal part of the inner margin. Four 
 branches are given off on the outer side, the first forking before the broken edge 
 of the wing is reached. The remaining three probably forked also on the missing 
 part of the wing-apex. The subdivisions of the median occupy the inner half of 
 the wing-apex, and the apex itself. 
 
 The regularity of arrangement of the minor veins is a marked feature of this 
 wing, and lends colour to our supposition. 
 
 The cubitus gradually diverges from the median along the whole of its course. 
 Ten branches are given off on the inner side, the first only forking. 
 
 The anal area is sharply marked off from the rest of the wing by a deep anal 
 groove, the anal area being elevated and somewhat convex in outline. Ten to 
 eleven anal veins can be distinguished, the marginal ones being very short and 
 feeble. 
 
 The interstitial neuration consists of close, strong, transverse nervures so 
 numerous as to give the wing-surface a finely corrugated appearance. 
 
 The tegminawere apparently of great strength, the unusual development of the 
 transverse nervures adding to the rigidity, while the presence of an oblique ridge 
 served to support and strengthen the radius, median and cubitus at their base, and 
 a similar ridge gave support to the whole of the anal area. 
 
 Affinities. In 1911 (Joe. cit.) I gave reasons for regarding this species as an 
 Archimylacrid, comparing it with A: spectabilis, Goldfuss, and A. venusta. More 
 recently, Dr. Pruvost ('Ann. Soc. Geol. Nord,' vol. xli, p. 335, pi. ix, figs. 4 4 a, 
 1912) has expressed the opinion that it is so nearly like A. belgica,, Handlirsch, and 
 a specimen he has himself described from the roof of the Alfred Vein, Lievin, 
 Northern France, as to be referable to that species. An examination of the figures 
 published by Handlirsch and Pruvost does not support this view. Archimylacris 
 belgica was founded by Handlirsch (' Mem. Mus. Roy. Hist. Nat. Belg.,' vol. iii, 
 p. 12, pi. iv, figs. 1718, 1906) on a wing in the Brussels Museum, from the West- 
 phalian or Middle Upper Carboniferous of Jemappes, Belgium. It was afterwards 
 transferred by him to a new genus, Parelthoblatta (' Die Fossilen Insekten,' p. 184, 
 pi. xviii, fig. 54, 1906). This new genus is not accepted by Pruvost, who considers 
 that it is founded chiefly on the pectinate character of the first branch of the radius. 
 He therefore replaces the species in the genus Archimylacris, and refers his ow r n 
 specimen to the same genus and species. With this conclusion I cannot agree. 
 Handlirsch's species is characterised not only by the pectinate character of the 
 first branch of the radius, but (and much more important) by the presence of a 
 meshwork of interstitial neuration between the radius, median and cubitus. This 
 latter feature clearly separates not only the French specimen from Handlirsch's 
 genus Parelthoblatta, and therefore from P. belgica, but the British specimen also ; 
 for in both the interstitial neuration consists wholly of a close series of straight 
 cross-nervures, with not the slightest trace of the development of a network. 
 14
 
 106 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 A. liastata is closely allied to the French species, but the divisions of the main 
 veins are fewer and of a more simple nature. I see no reason to remove the 
 British species from the genus Archimi/lacris, and would refer the French example 
 also to the same genus. This view is now accepted by Pruvost (loc. cit., p. 158). 
 
 Archimylacris woodward!, Bolton. Plate VII, fig. 2 ; Text-figure 34. 
 
 1910. Archimylacris (Etoblattina) woodwardi, Bolton, Greol. Mag. [5], vol. vii, pp. 147 151, pi. xv, 
 figs. 1 1 a . 
 
 Type. A left tegmen or fore- wing, 18 mm. long and 10 mm. wide; collection 
 of Mr. D. Davies, F.G.S. 
 
 Horizon and Locality. Coal Measures (a ten-foot shale overlying the No. 2 
 Rhondda Seam, base of the Pennant Series) ; Clydach Vale, South Wales. 
 
 Specific Characters. Wing short and broad. Apex well rounded and merging 
 
 Fia. 34. Archimylacris woodwardi, Bolton ; diagram of wing- venation of left fore-wino- three times 
 natural size.-Base of Pennant Series (roof of ten-foot shale overlying the No. 2 Khondda Seam) ; 
 Clydach Vale, South Wales. David Davies Collection. 
 
 into costal border. Anal area convex. Interstitial neuration of abundant stout 
 cross-nervures, occasionally uniting laterally. 
 
 Description. -The tegmen, which is short, broad and strongly chitinous, lies 
 with the convex dorsal surface uppermost. The well-rounded outer margin of the 
 wing merges into the equally well-rounded apex. The inner margin is nearly 
 straight, a slight hollowing only of the middle portion being shown. Notwitli- 
 * standing its small size, the details of the wing are very clear, the veins and the 
 interstitial neuration being well shown. The basal portions of the main veins are 
 elevated above the surface, the rest of their length and also their divisions being 
 sunk below the general level. 
 
 The subcostal vein is weak, and sunk along its whole course. It is widely 
 separated from the costal margin, following a parallel course to beyond the middle 
 of the wing, where it bends outward and joins the margin. The subcosta gives 
 off a numerous series of forked and simple divisions to the costal margin. 
 
 The radius arises close to the subcosta and median, diverging from the former 
 in the middle of its length, and approaching it again by the first of the four outer
 
 ARCHIMYLACRIS INCISA. 107 
 
 branches which it gives off. The first two branches of the radius are forked, 
 the rest undivided. The radius with its divisions occupies the distal third of the 
 wing-margin. At the junction of the basal and middle thirds of the wing, 
 the radius gives off a strong radial sector which diverges from it, giving off two 
 outer branches, the first forking twice into four twigs. The radial sector occupies 
 the outer half of the wing-apex. 
 
 The median curves regularly inwards along its whole course, reaching the inner 
 end of the wing-apex, and giving off three outer branches, of which the first is 
 forked. With its subdivisions it occupies a much less area than any other main 
 vein. 
 
 The cubitus follows a course nearly parallel with the median, and gives off nine 
 inward branches. With its subdivisions, it occupies almost the whole of the distal 
 two-thirds of the inner margin. 
 
 The anal area is strongly convex, and crossed by at least five anal veins, the 
 third being forked. 
 
 The interstitial neuration is well developed in the neighbourhood of the 
 subcosta, and consists of a close-set series of straight cross-nervures. These are 
 in relief, and in places seem to unite laterally, and to cross the main veins. 
 
 Affinities. The wing is typically Archimylacrid. At the time of its discovery 
 in 1910 I was unable to show that it possessed a close relation to any known 
 species, but more recently Dr. Pruvost has recorded a very similar example, A. 
 atrebatica, Pruv., from the Coal Measures of Bruay, Northern France. 
 
 The presence of a well-defined radial sector is a Palaeodictyopteroid feature 
 which most Blattoids have lost, and stamps the species as primitive. It is 
 therefore interesting to find that Pruvost draws special attention to the fact that 
 A. atrebatica, Pruv., is the oldest Archimylacrid of the Northern French Coal 
 Measures, being found in the lower portion of the Formation of Bruay, over the 
 vein Ernestine at Lens, in a shale containing seeds and drifted plant-remains. 
 
 Archimylacris incisa, sp. nov. Plate VII, fig. 3. 
 
 Type. Pronotum, mesonotum, and the basal portions of the tegmina in an 
 ironstone nodule; British Museum (Johnson Collection, no. I. 15900). 
 
 Horizon and Locality. Middle Coal Measures (binds between the "Brooch" 
 and " Thick " coals) ; Coseley, near Dudley, Staffs. 
 
 Specific Characters. Wing robust. Principal veins widely spaced. Interstitial 
 neuration of stout cross-nervures, which unite in a meshwork in the wide areas. 
 
 Description These Blattoid remains are 21 mm. long. The pronotum lies 
 with its underside uppermost on that portion of the nodule which also bears the 
 impressions of the under surfaces of the tegmina, while the impression of the 
 underside of the pronotum and the fragments of the tegmina upside down occur
 
 108 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 on the other half. The hinder margin of the pronotum is still hidden in the 
 matrix, and the impressions of the under-surf aces of the wings would need to be 
 destroyed to expose it. 
 
 The pronotum is semicircular in outline, so far as can be determined. The 
 inner surface is marked out into two lateral areas which join in front of, and 
 behind, the central area, the latter being hollow, probably to lodge the head of the 
 insect during life. The central hollow of the pronotum is crossed in front and 
 behind by close-set parallel wrinkles, while a suture-like crack traverses the whole 
 structure in the middle line. Posteriorly to the central hollow, the lateral halves 
 have separated and become wrinkled up against each other. 
 
 The condition of the pronotum shows that it was convex, the central hollow of 
 the inner surface marking off an area which was more convex than the sides, and 
 separated from them by grooves which bent round and joined behind the central 
 more elevated area. The central area is thinner than the sides and infolded. 
 
 Lying behind, and partially overhung by the hind border of the pronotum, is a 
 heart-shaped structure, irregularly indented, owing its superficial appearance to 
 the overlap of the anal areas of the wings on it. On one side of this structure 
 can be seen what appears to be part of a stout segment, having a thickened anterior 
 border, and a convex surface bearing a few low tubercles. The central indented 
 structure occupies the position of the mesonotum, and the lateral stouter segment 
 may be a part of it, or of the metanotum which has been pushed forward. 
 
 The base of the left tegmen is in actual articulation with what I suppose to be 
 the mesonotum. As the tegmina lie with their under surface uppermost, the 
 impressions on the opposite half of the nodule are also of the same surface. 
 
 The fragment of the left tegmen is largest, and in the best condition, but not 
 more than a third of it is present. The tissue is thickly chitinous, much more so 
 than that of the pronotum. 
 
 The costal margin is convex, and the costal area strap-shaped and wide. Seen 
 from the underside it appears smooth, a few diagonal wrinkles alone crossing the 
 surface. We may therefore infer that the divisions of the subcosta are weakly 
 incised on the upper surface. 
 
 The subcostal vein is more convex in direction than the margin, the costal area 
 being widest at the base. 
 
 The radius arises close to the subcosta, and gradually diverges from it. It 
 shows no divisions in the basal portions preserved. 
 
 The median vein arises even closer to the radius than does the latter to the sub- 
 costa, and bends inwards in a convex curve for some distance, afterwards flattening 
 and becoming parallel with the radius. 
 
 The cubitus arises closer to the anal furrow than to the median, but bends out- 
 wards in a bold sweep away from it, becoming almost parallel with the median. 
 It gives off three inward branches in the part preserved, the first low down near
 
 ARCHIMYLACRIS (SCHIZOBLATTA) OBOVATA. 109 
 
 the base, and the remaining two at wide intervals further out. The course of the 
 main stem is such as to indicate that it reached the distal portion of the inner 
 margin, so that the wing-apex was wholly occupied by the divisions of the radius 
 and the median. 
 
 The anal veins seen are five in number, the first forking twice and the rest 
 single. They probably occupied the proximal third of the inner wing-margin. 
 
 The articulation of the tegmen to the mesonotum seems to have been strengthened 
 by the formation of three ridges, two formed by the deepening of the bases of the 
 subcosta and radius, and a third by a similar thickening just inside the anal furrow, 
 which in this specimen coincides with the line of the first branch of the cubitus. 
 
 The basal part of the subcostal area shows a deep hollow ; this feature is seen 
 in several of the British Blattoids, and favours the assumption that while the point 
 of attachment lay immediately behind, the frontal part of the mesonotum developed 
 an area which served as a fulcrum for the tegmina. 
 
 The right tegmen is represented only by the base of the costal border and sub- 
 costa, short portions of the radius, median, a part of the cubitus with two proximal 
 branches, and the inner anal vein dividing into three. 
 
 The interstitial neuration consists of strong cross-nervures, frequently uniting 
 laterally, and on the broader areas forming a slight open meshwork. 
 
 Affinities. Fragmentary though the wing is, it yet shows many of the features 
 of A. venusta, Scd., having the same Avide costal area, the same relation of subcosta, 
 radius and median so far as these can be traced, and the cubitus beginning to branch 
 low down before the radius and median show any trace of branching. The inter- 
 stitial neuration in both consists of straight cross-nervures occasionally uniting 
 laterally. 
 
 This wing is more robust than that of A. venusta, Scd., and the anal veins are 
 more oblique in direction. The whole of the main veins and those of the anal area 
 are more widely spaced. In its robust character the wing approaches A. acadica, 
 Scd., but in that species the interstitial neuration does not unite laterally. 
 
 Archimylacris (Schizoblatta) obovata, Bolton. Plate VII, fig. 4; Text-figure 35. 
 
 1911. Archimylacris (Schizoblatta) obovata, Boltou, Quart. Jouru. G-eol. Soc., vol. Ixvii, p. 157, pi. vii, 
 figs. 4-6. 
 
 Type. The distal two-thirds of a left tegmen, having a length of 23 mm. and a 
 greatest breadth of 10mm.; Museum of Practical Geology, Jermyn Street (nos. 
 24506 and 24507). 
 
 Horizon and Locality. Base of the Upper Coal Measures (Grwernau Level of 
 the Mynyddislwyn Vein) ; Maes-y-cwmmer, Monmouthshire. 
 
 Specific Characters. Costal border moderately convex, and passing into a
 
 110 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 subacute apex. Costal area narrow and strap-shaped. Subcosta extending just 
 beyond the middle of the wing. Radius large, much divided, and occupying the 
 distal portion of the outer margin, and the outer half of the apical margin and 
 apex. Median a small vein with few divisions ending on the inner apical margin. 
 Cubitus occupying the whole of the inner margin beyond the anal veins. Surface 
 covered with a close series of oblique wrinkles. 
 
 Description. When I first described this specimen in 1911, I was of opinion 
 that the length of the complete wing was not more than 25 mm. AVith a larger 
 knowledge of the structure and form of the Blattoid tegmen I should now 
 estimate the length at nearer 35 mm., the complete structure being a little over 
 two-and-a-half times as long as broad. 
 
 The costal border is broadly convex, more so in the distal than in the proximal 
 half. 
 
 The subcostal area is narrow, the subcosta being a delicate vein, sending a 
 number of forked and simple branches obliquely to the costal margin. Probably 
 half of the subcosta is missing, though three branches of it are present. One 
 
 FIG. 35. Archimylacris (Schizoblatta) obovata, Bolton ; diagram of venation of left fore-wing, three 
 times natural size. Upper Coal Measures (Gwernau Level of the Mynyddislwyn Vein) ; Maes-y- 
 cwmmer, Monmouthshire. Mus. Pract. Geol. (nos. 24506, 24507). Numbering of veins as in 
 Text-figure 33, p. 104. 
 
 remains single, the next gives off a simple basal twig, and forks just before 
 reaching the margin, while the outer branch is undivided. 
 
 The radius is much the largest of the principal veins, and is widely separated 
 from the subcosta over the middle portion of its length. Before reaching the 
 middle of the wing it divides into two diverging branches, each of which forks, 
 and afterwards gives off a series of smaller veins, eleven in number, which occupy 
 the distal outer margin, and the outer half of the wing-apex. 
 
 The median vein is not complete, and seems to consist of a main stem, giving 
 off a long branch in the basal third of the wing, a second branch in the middle, 
 and a final branch in the outer third. The first branch only divides, so that the 
 median ends on the inner apical margin in five divisions. 
 
 The remaining marginal veins are probably wholly cubital. They are five in 
 number, one only forking.
 
 ARCHIMYLACRIS (SCHIZOBLATTA) OBOVATA. Ill 
 
 There is no trace of the anal portion of the wing, or of the remainder of the 
 cubitus. 
 
 The wing is thick, coriaceous in texture, and convex dorsally. The whole 
 surface is covered by a dense series of irregular oblique wrinkles. In some places 
 a close-set series starts out from the sides of a vein, and dies out in the interspace. 
 In other places sets of wrinkles are interrupted by smooth interspaces, this 
 arrangement occurring at haphazard. There are a few cases in which the 
 wrinkles unite. 
 
 Affinities. There can be no doubt as to this specimen representing an 
 Archimylacrid. Dr. Handlirsch, who makes ArcMmylacris the type of a family, 
 Archimylacridae (' Proc. U.S. National Museum,' vol. xxix, p. 722, 1906), has also 
 founded a new genus, Schizoblatta, and with the type-species of this genus the 
 specimen here described is in close agreement. 
 
 The points to which I attach importance are the following: In both, the 
 subcostal area extends for a short distance beyond the middle line, the veins in 
 each case passing out obliquely to the margin. A very wide interval separates the 
 stern of the subcosta from that of the radius in the middle of their length, and this 
 area is narrowed distally in each case by the approach of the marginal veins. 
 The radius is a large and much branched vein, and separates into two main 
 divisions, which fork at the same level and reach the apical point of the wing 
 in this specimen just beyond it. The median is relatively small, while the 
 cubitus has few branches, passing obliquely out, like those of the subcosta, to the 
 margin. 
 
 The anal area in the type, Schizoblatta alutacea, is long, attaining nearly half 
 the length of the wing. In the specimen here described this part is missing, as is 
 also a part of the cubitus. The missing portion of the inner margin extends 
 beyond the middle of its length, and knowing how frequently the anal vein 
 determines the line of fracture, this extended broken area becomes significant. 
 
 Dr. Handlirsch's definition of the genus Schizoblatta is as follows: "Front wing 
 elliptical, about two and two-fifth times as long as broad. Costal area extending 
 about three-fifths the length of the wing, with about nine or ten normal veins; not 
 expanded at the base. Radius divided into two principal stems, the superior of 
 which separates into six branches and the inferior into eight, the majority of the 
 latter ending in the apical border. The median likewise divides into two main 
 stems, the anterior of which forms five branches, and the posterior four, all of which 
 fuse in the apical margin. The eight branches of the gently vaulted cubitus take 
 up the entire inner border. The anal area attains nearly half the length of the 
 wing. Cross-veins area not to be distinguished, but instead there is a fine-grained 
 leathery structure " (loc. cit.). 
 
 If the genus is to be maintained apart from that of Arcliimylacris, I would base 
 the characters on the obliquity of the marginal veins of the subcosta and cubitus,
 
 112 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 the presence of a wide interval between the former and the radius, and the wide 
 area occupied by the latter. 
 
 The division of the radius into two unequal branches, with its symmetrical 
 double bifurcation, is also, I venture to think, a feature of primary importance. 
 
 In the wide divergence of the radius and median, this species agrees with 
 Scudder's genus SpiloblaUina ; but in this case the veins do not converge again to 
 enclose an elongated or oval area, 
 
 Archimylacris, sp. indet. Plate VII, fig. 5. 
 
 1911. Archimylacris, sp. indet., Bolton, Quart. Journ. Geol. Soc., vol. Ixvii, p. 163, pi. x, fig. 3. 
 
 Type. A fragmentary wing, lacking the apex and the base ; Museum of 
 Practical Geology, Jermyn Street (no. 24503). 
 
 Horizon and Locality. Base of Upper Coal Measures (Gellideg Level of the 
 Mynyddislwyn Vein) ; near Maes-y-cwmmer, Monmouthshire. 
 
 Description. The specimen is much too fragmentary for any attempt* at specific 
 determination. A portion of the distal margin is present, with three branches of 
 the subcosta. These are succeeded by eight straight veins, two at least forking, 
 which belong to the radius and median series. These are followed by an equal 
 number of veins partially hidden by the surface of the integument, which is much 
 wrinkled over this area, These, I assume, are parts of the cubitus vein. The 
 wrinkling of the integument over the cubital area is very marked. 
 
 The special interest of this wing-fragment is in its association with portions of 
 a leaf of Cordaites. Scudder and others have commented on the general association 
 of the wings of Blattoids with leaves of Cordaites, but have not, so far as I am 
 aware, drawn attention to a feature which is well shown by this leaf, namely, pits 
 on its surface. I have repeatedly found such depressions on the leaves of Cordaites 
 in the Lancashire Coalfield, and in many cases seen the hollows occupied by the 
 shells of Spirorbis pusillus. The shallow pits on the Cordaites-\eai associated with 
 this fragmentary wing show faint traces of a spiral, similar to the impression 
 of Spirorbis, and such shells were, I think, once attached to it. While the 
 Carboniferous Blattoids may have been wholly phytophagous, we are led by our 
 knowledge of the living Blattids, especially by the common cockroach (Periplnnrta 
 orientalis), to assume the contrary, and to regard them as more likely to have been 
 omnivorous, in which case the association of Blattoid remains with the leaves of 
 Cordaites bearing the sedentary Spirorbis is easily understood. The association 
 also lends support to the belief that the Blattoids were semi-aquatic in habit, or 
 lived in marshes and swamps in which decayed vegetation formed a home for 
 Spirorbis.
 
 PHYLOBLATTA SULCATA. 113 
 
 Archimylacris, sp. indet. Plate VII, fig. 6. 
 1911. Archimylacris, sp. indet., Bolton, Quart. Journ. Geol. Soc., vol. Ixvii, p. 152, pi. vii, fig. 2. 
 
 Type Impression of basal portion of left tegmen ; Museum of Practical 
 Geology, Jermyn Street (no. 24508). 
 
 Horizon and Locality. Base of Upper Coal Measures (Mynyddislwyn Vein, 
 Gellideg Level) ; near Maes-y-cwmmer, Monmouthshire. 
 
 Genus PHYLOBLATTA, Handlirsch. 
 1906. PhyloUatta, Handlirsch, Bull. U.S. Nat. Mus., vol. xxix, p. 738. 
 
 Generic Characters. Tegmina elliptical, t\vo-and-a-quarter to two-and-a-half 
 times as long as wide ; costal area strap-shaped, rarely wide, and extending usually 
 to three-fifths or two-thirds the length of the wing ; not expanded basally. Radius 
 with numerous branches, all of which reach the outer margin. Median with outer 
 branches only, reaching wing-apex. Cubitus large, much branched, and occupying 
 distal two-thirds of inner margin. Anal area large with numerous veins. Inter- 
 stitial neuration rugose leathery or much cross-wrinkled. 
 
 Phyloblatta sulcata (Bolton). Plate VIII, fig. 1; Text-figure 36. 
 
 1911. GeraUattina (Aphthoroblattina) sulcata, Bolton, Quai't. Journ. G-eol. Soc., vol. Ixvii, p. 165, 
 pi. viii, figs. 1 3. 
 
 Type. Greater part of tegmen and its counterpart impression showing the 
 under surface only ; Museum of Practical Geology, Jermyn Street (nos. 24504 
 and 24505). 
 
 Horizon and Locality. Base of the Upper Coal Measures (Gwernau Level of 
 the Mynyddislwyn Vein) ; near Maes-y-cwmmer, Monmouthshire. 
 
 Specific Characters. Costal area broad ; subcosta strap-shaped, and extending 
 far out. Radius with six branches, occupying outer half of wing-apex. Median 
 with few divisions and occupying a small area of the wing. Cubitus united with 
 median at the base. Anal furrow well developed. Anal veins 5 8. 
 
 Description. The specimen comprises the greater part of a right tegmen, 
 showing the under surface, the counterpart impression being on a second piece of 
 black shale. An irregular narrow fringe of the wing-membrane has been lost 
 along the outer and inner margins, and the inner half of the wing-apex is also 
 missing. The length of the fragment is 35 mm., and the greatest width 16 mm., 
 the perfect wing being probably about 45 mm. long, and 2223 mm. wide. If this 
 estimate is correct, the wing is short. Tegmina are usually at least two-and-a-half 
 15
 
 114 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 times as long as wide. The structure is well preserved, and the principal veins 
 and their subdivisions are in relief. 
 
 The costal margin is thickened, strongly convex, and well rounded at the base. 
 The costal area is unusually broad and strap-shaped, extending over two- thirds of 
 the costal margin. The basal portion of the costal area is smooth, and not crossed 
 by branches of the subcosta. 
 
 The subcosta is thin, elevated basally, and somewhat crenulated, doubtless 
 owing to post-mortem change or pressure. The subcosta is parallel with the 
 costal margin over more than one-half the length of the wing, and gives off six 
 oblique branches to the margin. Three of these branches are forked. 
 
 The radius passes out to the apex of the wing, giving off six outer branches, 
 the first forking once, and the second twice before reaching the margin. The 
 
 PIG. 36. Phyloblatta sulcata (Bolton) ; diagram of the venation, and restored outline of fore-wing, two- 
 and-a-half times natural size. Upper Coal Measures (Gwernau Level of the Mynyddislwyn Vein) ; 
 near Maes-y-cwmmer, Monmouthshire. Mus. Pract. Geol. (nos. 24504, 24505). Numbering of 
 veins as in Text-figure 33, p. 104. 
 
 remaining branches are undivided. The radius and subcosta together occupy a 
 little less than half the total wing-area. 
 
 The median vein is convex outwards in its basal half, and then bends inwards 
 to the junction of the inner and apical margins. It gives off three long, outward, 
 undivided branches, all of which are parallel with the main stem of the radius. 
 Basally, the median curves sharply inwards and unites with the cubitus, the two 
 veins having a very short common stem. 
 
 The cubitus is the largest and most important vein in the wing, although but 
 faintly outlined at the base, where its union with the median is nevertheless quite 
 clear. It gives off seven inwardly directed branches, the third only forking. On 
 its outward side two branches are given off. 
 
 The anal furrow is represented by a strong ridge, and is therefore a well- 
 marked groove on the dorsal surface of the wing. The anal area is large, 
 extending at least one-third the length of the inner margin, and crossed by five
 
 PHYLOBLATTA TRANSVERSALTS. 115 
 
 anal veins, the first and second only being forked. The whole of the anal area is 
 not shown, and 3 4 veins may be missing. 
 
 The interstitial neuration consists of a close series of transverse nervures or 
 wrinkles, it is impossible to say which. They are best seen in the anal and 
 cubital areas. 
 
 Affinities. The union of the stems of the median and cubitus veins, the 
 occupation of the greater part of the inner half of the wing by the median, whose 
 branches take up the whole of the inner margin outside the anal area, and the 
 strong parallelism of the branches of the principal veins, are well-marked features 
 which in the main agree most, I think, with Phi/loblatta. Its resemblance to 
 Gemllattina is also marked, and it is well to remember that Handlirsch has 
 referred several species originally placed in the latter genus to Phyloblatta. This 
 question of the generic relationship is a good example of the difficulty of allocating 
 species to genera whose characters are not wholly known. 
 
 Phyloblatta transversalis, Bolton. Plate VIII, fig. 2 ; Text-figure 37. 
 
 1917. PhylloUatta transversalis, Bolton, Proc. Birmingham Nat. Hist. Phil. Soc., vol. xiv, pp. 100 
 103, pi. vii, figs. 1, 2. 
 
 Type. Remains of two Blattoids, consisting of the tegmina, two pronota, and 
 portions of the hind- wings, in a split nodule of ironstone, 60 mm. long, and 45 mm. 
 broad ; Geological Museum, University of Birmingham. 
 
 Horizon and Locality. Coal Measures; Staffordshire. 
 
 The nodules were collected by Dr. Blake and agree in all respects with those 
 found at Tipton and Coseley. Staffordshire, in the binds between the " Brooch " 
 and " Thick " coals of the Middle Coal Measures. 
 
 Specific Characters. Radius relatively simple with five branches, radial sector 
 large and evenly forked ; median vein small, with three outer branches ; cubitus 
 vein long with seven inward branches ; anal area small with few anal veins. 
 
 Description. The remains of three insects are shown on the split surfaces of 
 two small ironstone nodules. The larger nodule contains the remains of two 
 Blattoid tegmina, two pronota, and fragments of hind-wings. One tegmen is 
 almost complete, while of the other more than half is shown, the distal end having 
 been lost in splitting the nodule. 
 
 The tegmina lie with the under surfaces uppermost, the impression of the 
 underside being preserved on the larger portion of the nodule. As both are from 
 the left side, it follows that they belong to two insects a conclusion confirmed by 
 the presence of a pronotum in close association with each. In the case of the 
 more nearly complete tegmen, the pronotum lies a little apart, and upside down, 
 like the tegmen, while the pronotum of the more incomplete wing lies on the basal
 
 116 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 portion of the tegmen itself, and has the dorsal surface uppermost. Traces of 
 hind-wings are present, lying on a slightly lower plane than the tegmina, and 
 showing only the underside. 
 
 As the two tegmina are specifically identical, the more nearly complete example 
 has been taken as the type and is here described in detail. 
 
 The tegmen has a length of 32 mm., and, as a small part of the apex is missing, 
 its total length must have been about 35 mm. The width is uniform over the 
 basal half of the wing, and averages 11 '5 mm. 
 
 The costal margin forms a strong convex curve, in this respect contrasting with 
 the inner margin, which is nearly straight. The wing-apex is directed backwards 
 owing to the great convexity of the costal margin. The whole wing has a broad 
 semilunate appearance. 
 
 The subcosta is a feeble vein, not easily discernible; its outwardly -directed 
 twigs pass out obliquely, and are twice or thrice forked. Like the main stem, they 
 are faintly impressed, and cannot be traced up to the costal margin. Eight twig 
 can be determined. The subcostal area is strap-shaped, and extends beyond the 
 middle of the wing. 
 
 FIG. '67.Phyloblatta transversalis, Bolton ; diagram of venation of left fore-wing of type-specimen, 
 twice natural size. Middle Coal Measures ; Staffordshire. Geol. Mus. Univ. Birmingham. 
 
 The radius is a strong vein, and shows an inner sub-division which has all the 
 characters of a radial sector. Basally, it seems joined to the median, but it is an 
 apparent junction only, the wing-base being pressed in upon itself, and bringing 
 the two close to one another. After giving off the inner (radial sector) branch 
 the radius gives off five outer branches which reach the distal third of the outei 
 margin. The first of these branches seems to break up into three twigs, but the 
 structure of the wing is not clear at this point. The radial sector, which arises in 
 the basal half of the wing, forks at the distal third into two equal branches, the 
 outer forking twice into four divisions, and the inner forking once. The radius 
 and radial sector occupy the whole of the outer half of the wing-apex. 
 
 The median vein diverges from the radius along its whole course, and readies 
 the distal end of the inner margin, giving off three outer branches in the distal 
 half of the wing. These pass straight to the apex, the first forking twice, and 
 the second once, the third remaining undivided to the broken edge of the wing. 
 The branches of the median occupy the whole of the inner half of the wing- 
 apex.
 
 PHYLOBLATTA TRANSVERSALIS. 117 
 
 The cubitus remains above the middle line of the wing in the basal third, 
 curving inwards and flattening in the outer two-thirds of the wing, and giving off 
 seven inward branches, the second alone forking low down near the margin. 
 
 The anal area is small, and crossed by six oblique anal veins, the first and third 
 of which fork. 
 
 The interstitial neuration consists of a close-set series of straight cross-nervures 
 which unite laterally as they cross the wider areas. 
 
 The pronotum in close apposition with this wing has its inner surface upper- 
 most. This sho\vs a central shallow concave area, bounded by two lateral ridges, 
 which die away as they approach the hinder border. The general outline is 
 broadly circular. 
 
 The second tegmen is much less exposed than the first, and has its apex buried 
 under the second pronotum. So far as its structure can be determined, it agrees 
 with the wing already described. 
 
 The second pronotum has been crushed, the anterior border being turned round 
 upon the base of the wing, and partly broken away. The posterior margin is 
 almost straight, and that of the anterior Avell rounded. 
 
 Traces of two hind-wings are shown, one a little in advance of the more nearly 
 complete wing, and partly underlying it, and the other underlying the costal and 
 radial areas of the second incomplete wing. The first hind-wing shows a portion 
 of the distal extremity, with a series of incomplete veins, probably belonging to 
 the radius, median, and cubitus; the remains of the second hind-wing probably 
 consist of the distal radial twigs only. 
 
 Affinities. The distinguishing features of the tegmina are essentially those of 
 Phyloblatta, unless we except the interstitial neuration. This is composed of 
 straight cross-nervures, which unite laterally across the wide areas between the 
 main stems of the radius, median, and cubitus, but not producing anywhere a 
 meshwork such as is usually seen in the Phyloblattids. 
 
 Handlirsch doubts the presence of a cross-neuration in the genus Phyloblatta 
 (' Proc. U.S. National Museum,' vol. xxix, p. 731, 1906), but there seems no reason 
 Avhy it should not be present in the more archaic members of the genus, as they 
 all have an Archimylacrid ancestry, in which a cross-neuration is a dominant 
 feature. 
 
 The species is closely allied to a form figured as Gemblattina sp. by Brongniart 
 (' Insectes Fossiles des Temps Primaires,' pi. xlvi, fig. 7, 1893), and repeated by 
 Handlirsch (' Die Fossilen Insekten,' pi. xxx, fig. 35, p. 295, 1 906) under the name 
 of Blattoidea, sp. Handlirsch regards Brongniart's specimen as belonging either to 
 the Spiloblattinidae or to the Archimylacridae. Unfortunately the interstitial 
 neuration is not shown. If, as in these specimens, Brongniart's species possesses a 
 cross-neuration, or one in which the nervures unite laterally, it must be referred 
 to Phyloblatta.
 
 118 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 (Archimylacridse) kirkbyi (Woodward). Plate VIII, fig. 3. 
 
 1887. Lithomylacris kirkbyi, Woodward, Greol. Mag. [3], vol. iv, p. 55, pi. ii, figs. 4 a 4&. 
 1887. / Hermatoblattina kirkbyi, Scudder, Proc. Boat. Soc. Nat. Hist., vol. xxiii, p. 357. 
 1906. (Archimylacridffi) kirkbyi, Handlirsch, Die Fossilen Insekten, p. 238, pi. xxiv, fig. 37. 
 
 Type. Left wing (tegmen) ; formerly in the collection of the late Mr. James 
 Kirkby. 
 
 Horizon and Locality. Upper Coal Measures (bed No. 33) ; near Meithil, coast 
 of Fifeshire. 
 
 Description. Our knowledge of this wing depends on the description and 
 figures published by Dr. H. Woodward. His description is as follows : " Outline 
 of wing pointed-ovate, slightly flattened on its superior border ; length of wing 
 15 mm., breadth 8 mm. The 'mediastinal' [subcostal] vein occupies rather more 
 than a fourth of the entire area of the wing ; it extends to about two-thirds of the 
 length, where it unites with the superior margin of the wing ; it branches six 
 times ; one of these branches has three forks. The ' scapular ' [radius] vein 
 extends nearly to the extremity of the wing; it remains single for over one-fourth 
 of its length, and then branches into three veins, the middle one of which is again 
 forked. The * externo-median ' [median] vein continues parallel to the scapular 
 vein for a slightly longer distance before it branches at the extremity of the wing 
 into three inferior veins, two of which are again forked. The 'interno-median' 
 [cubitus] vein occupies about one-fourth the entire 'area of the wing ; it gives off 
 three almost equidistant branches, none of which appear to be forked. The anal 
 vein is nearly straight, and has three other almost parallel oblique simple veins 
 occupying the anal area." 
 
 Affinities. Neither Woodward's figures nor the later modified drawing 
 published by Handlirsch are very helpful in enabling us to identify the genus. 
 Handlirsch has reversed the figure given by Woodward, and shows three veins 
 which may belong to the subcosta, a radius with three forward branches, and a 
 well-divided median ending on the wing-apex in nine outer branches. Woodward's 
 " mediastinal " becomes the cubitus, and no anal veins are shown. This seems a 
 more reasonable interpretation of the wing. 
 
 Woodward's comparison of the wing with Lithomylacris pittstonianum, Scd., is 
 unfortunate, as in that species the radius is a large, much-branched vein, occupying 
 nearly half the outer margin, while the radius (scapular) in A. kirkbyi, Woochv., has 
 three divisions only as shown by him, although it is a more branched structure 
 when the figure is reversed and the veins reconsidered, as by Handlirsch. 
 
 In the absence of the type-specimen, it is not possible to do more than refer 
 the species to the family Archimylacridge.
 
 BLATTOIDS INCERT.E SEDIS. 119 
 
 BLATTOIDS INCERT^E SEDIS. 
 
 Remains of Blattoids have been found at four horizons and in four localities in 
 the Kent Coalfield. They were obtained from small cores, the edges of which had 
 cut through the wings, and rendered them so fragmentary as to be useless for the 
 determination of genera or species. They serve, however, to indicate that the Kent 
 Coal Measures may eventually prove to be as rich in fossil insect-remains as the Coal 
 Measures of the Pas-de-Calais. The depths of the cores given with the original 
 description of these fossils were found afterwards to have been reckoned from a 
 private datum, while the depths now given have been corrected to Ordnance 
 datum. 
 
 Phyloblatta (?), sp. Plate VIII, fig. 4. 
 
 1912. Phylloblatta (?), sp., Boltoii, Quart. Journ. Geol. Soc,, vol. Ixviii, p. 321, pi. xxxiii, figs. 3, 4, 5. 
 1915. Phylloblatta (?), sp., Bolton, Trans. Inst. Min. Eng., vol. xlix, p. 45. 
 
 Specimen. Impression of a Blattoid wing, 5 mm. long and 2*5 mm. wide; 
 Museum of the Kent Coal Concessions Company, Dover. 
 
 The impression is that of the middle part of the margin of a tegmen. This 
 may be a part of the outer margin, as I formerly supposed, but I now think it 
 more likely to belong to the inner margin, and to show branches of the median and 
 cubitus veins. 
 
 Horizon and Locality. Shales at a depth of 1967 ft. in the Maydensole boring, 
 Kent Coalfield. 
 
 Phyloblatta (?), sp. 
 
 1912. Phylloblatta (?), sp., Bolton, Quart. Journ. Geol. Soc,, vol. Ixviii, p. 321, pi. xxxiii, figs. 8, 9. 
 1915. Phylloblatta (?), sp., Bolton, Trans. Inst. Min. Eng., vol. xlix, p. 45. 
 
 Specimen. Impression of part of a Blattoid wing, 6 mm. long and 3 mm. wide ; 
 Museum of the Kent Coal Concessions Company, Dover. 
 
 The fragment appears to belong to the proximal part of the wing, and shows a 
 long vein giving off six branches. 
 
 Lying more inward are the remains of a short forked vein. 
 
 Horizon and Locality. Shales at a depth of 954 feet in the Ripple boring, 
 Kent Coalfield. 
 
 BLATTOID WING-FRAGMENT. Plate VIII, fig. 5. 
 
 1912. " Blattoid," Bolton, Quart, Journ. Geol. Soc., vol. Ixviii, p. 321, pi. xxxiii, figs. 6, 7. 
 1915. " Blattoid," Bolton, Trans. Inst. Min. Eng., vol. xlix, p. 45. 
 
 Specimen Impression of a wing-fragment 3mm. long, and 2'25 mm. wide; 
 Museum of the Kent Coal Concessions Company, Dover.
 
 120 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 The fragment shows a little of a well-curved margin and three veins, one with 
 a small fork on the wing-margin. The curvature indicates that the fragment 
 belongs to the apical end of the wing. 
 
 Horizon and Locality. Black shale at a depth of 2180 feet in the Barfreston 
 boring, Kent Coalfield. 
 
 BLATTOID WJNG-FRAGMENT. Plate VIII, fig. 6. 
 
 1915. " Blattoid," Bolton, Trans. Inst. Min. Eng., vol. xlix, pp. 45,- 46, pi. ix, fig. 24. 
 
 Specimen. Distal portion of a tegmen, including the whole of the wing-apex; 
 Museum of the Kent Coal Concessions Company, Dover. 
 
 Portions of the subcosta, radius, and median veins can be determined. The 
 subcosta is represented by a little of the main stem which sends off four simple 
 branches to the outer margin. The radius divides low down into two branches, 
 the outer dividing again into two nearly equal twigs, which pass out to the outer 
 margin of the wing. The inner branch divides into four twigs, all reaching the 
 apical margin. The cubitus is represented by a long vein, giving off a series of 
 eight to nine inward branches. 
 
 Horizon and Locality. Shales at a depth of 2424 feet in the Stonehall boring, 
 Kent Coalfield. 
 
 Family MYLACRIOE, Scudder. 
 
 1879. Scudder, Mem. Best, Soc. Nat. Hist., vol. iii, pt, 1, no. 3, p. 40. 
 1906. Handlirsch, Die Fossilen Insekten, p. 258. 
 
 Fore-wings variable in shape, but generally broad and short, nearly always 
 widest at the base. Costal area more or less triangular. Subcosta with branches 
 arising in a radial manner. Radius sending numerous branches outwardly, or 
 dividing into two widely dividing branches. The median either gives off its 
 branches serially, or forms two compound main branches. Cubitus with a variably 
 number of inward branches. Anal area large, with the anal veins usually ending 
 on the inner margin (Handlirsch). 
 
 Scudder established this family for a numerous series of North American 
 Blattoids, all of which are characterised by the subcostal area being widened out 
 basally, and the subcostal vein and its branches reduced from the strap-shaped 
 form seen in Archimylacrids to a shorter structure, in which the branches arise 
 mainly from the root of the vein, and radiate in fan-fashion into the subcostal area. 
 
 The humeral portion of the subcostal area is usually smooth, and destitute of 
 subcostal branches, or where these are present, they are short and do not reach the 
 margin. Owing to the enlargement of the subcostal area, the point of attachment
 
 HEMIMYLACRIS. 121 
 
 of the wing becomes central, instead of being in front of the wing-axis as in 
 the Archimylacridae. The principal veins and their branches show a greater 
 tendency to pass straight outwards to the wing-margin than in the Archimylacridse, 
 while the pronotum has become broader and shorter, and assumes more of a 
 reniform appearance. 
 
 The wing-surface is leathery and coriaceous, and often cross-wrinkled. Pruvost 
 states that the interstitial neuration is composed of a fine network of nervures, 
 which is more correct, the thickened wrinkled condition being a later development. 
 The body is broad and flat. 
 
 Handlirsch regards the Mylacrida3 as an early, extremely developed, lateral 
 branch of the Blattoid series, and thus still retaining rather primitive characters, 
 best seen in the median vein. Handlirsch adds (1906, ' Proc. U.S. Nat. Mus.,' 
 vol. xxix, p. 766) : " Perhaps they owe their origin to an adaptation to their 
 environment, for it is remarkable how similar many of them are to certain leaves 
 of ferns, with which they are generally found (a fact to which Scudder had already 
 drawn attention). Probably they lived under deciduous fern-fronds, and by their 
 similarity to the pinnae were protected from their enemies." 
 
 Pruvost (1920) considers that the Mylacrida3 may constitute a well individualised 
 phyletic series which sprang late from the Archimylacrid stock. He divides the 
 family Mylacrida3 into two divisions, Hemimylacridian and Mylacridian, the former, 
 as typified in Pliylomylacris and Hemimylacris, being really intermediate forms 
 lying between the true Archimylacrids and the true Mylacrids (Mylacridians), and 
 more nearly descended from the first. Pruvost also points out that Handlirsch has 
 placed Apophthegma (the Hemimylacris of Saxony) with the Archimylacrids, and 
 the American members of the Hemimylacris among the MylacridaB. He is some- 
 what uncertain whether to place the Hemimylacridians with the Arehiraylucridse 
 or with the Mylacridse, but decides in favour of the latter, suggesting that both the 
 Hemimylacridians and the Mylacridians are of polyphyletic origin. 
 
 Genus HEMIMYLACRIS, Handlirsch. 
 1906. Hemimylacris, Handlirsch, Proc. U.S. Nat. Mus., vol. xxix, p. 767. 
 
 Generic Characters. Wings twice as long as wide. Costal area broad, in one 
 species almost triangular, in another somewhat strap-shaped. Radius % with four 
 outer branches, the first dividing into two or three twigs. Median with three 
 branches directed inwards. Cubitus with 45 branches which do not occupy the 
 whole of the free inner margin. Anal area two-fifths the length of the wing, and 
 more than twice as long as wide. 
 
 Handlirsch remarks that this genus may be classed almost as well with the 
 Archimylacridse as with the Mylacridas, thus agreeing with Pruvost,
 
 122 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 Hemimylacris obtusa, Bolton. Plate VIII, fig. 7 ; Text-figure 38. 
 
 1911. Hemimylacris obhtsa, Bolton, Quart, Journ. Geol. Soc., vol. Ixvii, p. 154, pi. x, figs. 4, 5. 
 
 Type. A stout right tegmen, lying on a surface of fireclay full of Stigmarian 
 rootlets, and distorted by pressure ; Museum of Practical Geology, Jermyn Street 
 (no. 24510). 
 
 Horizon and Locality. Upper Coal Measures (Four- foot Seam of Swansea) ; 
 Gladys Colliery, one mile east-south-east of Penller-gaer Church, Glam. 
 
 Specific Characters. Costal margin convex ; subcostal vein dividing by 
 repeated forking into five twigs; costal area triangular. Radius with a radial 
 sector, the latter much divided. Median with a few forked inward branches. 
 Cubitus large, with four branches, the second twice forked. Anal area wide, and 
 crossed by numerous veins, the first giving off a simple branch, and then dividing 
 twice by equal forking. Inner margin almost straight. 
 
 ff 
 
 I 
 
 FIG. 38. Hemimylacris obtusa, Bolton ; diagram of right fore-wing with apex restored, three times 
 natural size. Upper Coal Measures (four-foot Seam of Swansea) ; Gladys Colliery, near Penller- 
 gaer Church, Glamorganshire. Mus. Pract. Geol. (no. 24510). Numbering of veins as in Text- 
 figure 33, p. 104. 
 
 Description. A short rounded right tegmen, 23 mm. long and 14 mm. wide, 
 broken across the basal third along the anal furrow, and the apex missing. 
 
 The subcostal area is broadly triangular, the subcostal vein sunken, and 
 passing out obliquely, giving off a basal branch which forks into two equal twigs, 
 the most distal twig forking again. A single undivided branch is given off near 
 the end of the subcosta. 
 
 The radius gives off a strong radial sector, and afterwards sends outwardly 
 three simple branches. The radial sector sends four branches to the margin, the 
 first forking twice, and the fourth once, the second and third being simple. 
 
 The median vein appears to give off the stem of the cubitus near its base, but 
 this appearance is probably due to the crumpling of the tegmen. The first of the 
 three inward branches of the median arises further out than any of the divisions 
 of the radius or cubitus. The first branch divides into three twigs, and the second 
 forks on the broken edge of the wing. All the branches pass straight out towards 
 the wing-apex.
 
 HEMTMYLACRIS CONVEXA. 123 
 
 The cnbitus is convex in its basal half, and straightens out distally, giving off 
 four inward branches, the second dividing into three, and the fourth forking. 
 The divisions of the cubitus occupy the distal half of the inner margin. 
 
 The anal area is large, and marked off from the rest of the wing by a deep 
 anal furrow. The first anal vein breaks up into five divisions, by a triple forking, 
 and the second forks twice. The third anal vein forks near its base, and the 
 remaining 3 4 veins appear simple. The anal veins end on the margin in at least 
 fourteen twigs. 
 
 The interstitial neuration cannot be determined. 
 
 Affinities. The mode of branching of the subcostal vein, the presence of a 
 decided radial sector and the character of the first anal vein, are features which 
 would justify the inclusion of this species in the genus Soomylacris, were it not for 
 the fact that the median is not divided into two equal branches. There is a close 
 correspondence between this species and H. ramificata, Handl., and I therefore 
 retain it in the genus Hemimylacris^ although, as I have stated elsewhere, I am 
 inclined to merge the two genera into one, as marking a transitional group between 
 the Archimylacridae and the Mylacridas. 
 
 Hemimylacris convexa, Bolton. Plate VIII, fig. 8. 
 
 1911. Hemimylacris convexa, Boltoii, Quart. Journ. Geol. Soc., vol. Ixvii, p. 156, pi. vii, fig. 3. 
 
 Type. Proximal half of a tegmen lying on an irregular surface of black shale; 
 Museum of Practical Geology, Jermyn Street (no. 24512). 
 
 Horizon and Locality. Pennant Series (Shales associated with the Graigola 
 Seam) ; Clydach Merthyr Colliery, Clydach Valley (Swansea Vale), Glam. 
 
 Specific Characters. Costal area triangular. Radius with a well-defined radial 
 sector, the two occupying most of the outer margin and the outer half of the 
 wing-apex. Median small. Cubitus arcuated and ending on the middle of the 
 inner margin, and outer third of the apex. Anal veins few. 
 
 Description. Only the proximal half of the tegmen is shown, and this does not 
 exceed 10 mm. in length. The subcostal area is much crushed and broken, and 
 but two branches of the subcostal vein can be seen on it. 
 
 The radius arises in the middle of the base, and at once divides, evidently 
 giving off a radial sector. The radius shows two marginal veins, while the radial 
 sector sends off a long branch towards the wing-apex, the branch forking near the 
 broken edge of the wing. 
 
 The median vein is represented by a portion of the main stem, which f, 
 
 twice. 
 
 The cubitus has an arcuated stem bifurcating twice into four branches. 
 
 anal veins can be distinguished.
 
 124 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 Affinities. The fragmentary and crumpled condition of this wing makes its 
 elucidation difficult and unsatisfactory. The chief features distinguishable are the 
 almost equal costal and anal areas, the presence of a radial sector, and the origin 
 of several anal veins arising from one stem. Few though these characters be, they 
 are sufficient to refer the specimen to the Mylacridas, and to the genus Hemimylacns, 
 
 Genus PHYLOMYLACRIS, Pruvost. 
 
 1919. Phylomylacris, Pruvost, Faune Continent. Terr. Houill. N. France (Mem. Explic. Carte G-col. 
 France), p. 199. 
 
 Generic Characters. Tegmina semi-ovate, with rounded apex. Surface coria- 
 ceous, with a fine close meshvvork of interstitial nervures giving a shagreen-like 
 appearance. Wing-attachment in middle of wing. Costal area triangular. Sub- 
 costal branching, partly pectinated and partly radial, extending beyond the middle 
 of the wing. Radius well developed. Median with numerous branches. Cubitus 
 large and well branched. Anal area large, convex, and crossed by numerous 
 anal veins. 
 
 Pruvost has formed this genus to include certain Blattoids found in recent 
 years at Lens and Lievin in northern France. They possess a superficial 
 resemblance to the type-species of Necijmylacris, N. heros, Scd., but their 
 differences are nevertheless so marked as to justify the formation of a new genus. 
 Pruvost in the work in question placed Goldenberg's Blattidium mantidioides in 
 the genus Archasotiphe, having at the time only the original drawings of Kirkby 
 to rely on. After borrowing the type-specimen, I came to the conclusion that 
 Goldenberg's species must be referred to a new genus, in which I also placed 
 Necymiilacris villeti, Pruv., N. lofittei, Pruv., and N. godoni, Pruv. To this new 
 genus I proposed to attach the name of my French friend, Dr. P. Pruvost, and 
 intimated my intention to him, at the same time sending my -manuscript. I received 
 a reply and the proof-sheets of his new work, in which it appeared that he also had 
 recognised that the three species previously referred to Necymylacris must be 
 placed in a new genus, to which he had already given the name of Phylomylacds. 
 Our conclusions were identical as to the generic value of his species, and as his 
 work antedates my own, I am unable to attach his name to the new genus, but 
 must adopt the generic name of Phi/lomylacris for the British species. 
 
 Had Dr. Pruvost received my photographs and drawings earlier, he would not 
 have referred B. mantidioides to the genus ArcJiseotipke, but to Phylomijlaens, as 
 he based the characters of the latter on the same details of structure as I had 
 determined for the genus in which B. mantidioides should be placed. The fact 
 that we arrived at the same conclusion, and by the selection of the same structures, 
 has been a satisfaction to both.
 
 PHYLOMYLACRIS MANTIDIOIDES. 125 
 
 Phylomylacris mantidioides (Goldenberg). Plate IX, fig. 1 ; Text-figure 39. 
 
 1867. Blatta or Blattina, Kirkby, Greol. Mag., vol. iv, p. 388, pi. xvii, figs. 6, 7. 
 
 1877. Blattidium mantidioides, Goldeiiberg, Faima Sarsepout. Foss., vol. ii, p. 20. 
 
 1879. Etoblattina mantidioides, Scudder, Mem. Bost. Soc. Nat. Hist., vol. iii, p. 72, pi. iii, fig. 6. 
 
 1906. (Archimylacridie) mantidioides, Handlirsch, Die Fossilen Iiisekten, p. 237, pi. xxiv, fig. 27. 
 
 1919. Arclueotiphe mantidioides, Pruvost, Faune Coutiuent. Terr. Houill. N. France, p. 170. 
 
 Ti/pe. Basal half of left tegmen in nodule; Kirkby Collection, Hancock 
 Museum, Newcastle-on-Tyne. 
 
 Horizon and Locality. Upper Coal Measures (zone of Atithmcomya phillipm) ; 
 South Hylton, opposite Claxheugh on the Wear, Durham. 
 
 Specific Characters. Surface of wing coriaceous. Costal margin convex ; 
 subcosta feeble, with few simple branches passing obliquely to the margin. Costal 
 area acutely pyriform. Radius much divided, and occupying outer half of wing- 
 apex Median with numerous divisions, and passing to inner half of wing-apex. 
 
 FIG. 39. Phylomylacris mantidioides (Goldenberg) ; diagram of venation of type, a left fore-wing, 
 three-and-a-half times natural size. Upper Coal Measures (zone of Anlhracomya phillipsii) ; South 
 Hylton, opposite Claxheugh on the Wear, Durham. Hancock Mus., Newcastle on-Tyne (Kirkby 
 Coll.). 
 
 Cubitus large, extending to the distal end of the inner margin, and giving off five 
 or more inner branches, the first and third dividing. Anal area tumid, anal furrow 
 well defined. Anal veins 9 10 in number, the first forking twice, and also sending 
 three weak branches forward. Inner margin convex, but less so than costal margin. 
 
 Description. The surface of the nodule surrounding the wing-fragment is 
 covered with the " spat " or young fry of a species of mollusc to which Kirkby 
 gave the name of Ancijlus vinti, but which I have since shown to be the young fry 
 of Antliracomya pUillipsii ('Trans. Inst. Mining Engin.,' vol. xlix, p. 675, 1915); 
 also Trechmann and Woolacott (< Geol. Mag.' [6], vol. vi, p. 207, 1919). 
 
 The wing-fragment, measuring 13'5mm. long and 8 mm. wide, is in excellent 
 condition, and lies with the dorsal surface uppermost. The whole of the venation 
 is well marked. Kirkby's original figure is not correct, and his errors have been 
 repeated by Scudder and Goldenberg. Goldenberg did not describe the species he 
 named, and Scudder, to whom we owe the first detailed account, was hindered by 
 the faulty drawing, and does not seem to have seen the specimen. Handlirsch 
 removed the species from Etoblattina, regarding the specimen as an indeterminable
 
 126 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 form of the Archimylacridse. Re-examination of the type-specimen shows errors 
 in the earlier descriptions and figure, and I now re-describe and re-figure the type, 
 placing it in the new genus established by Pruvost. 
 
 The wing is a stout structure, with a coriaceous surface which has enabled it to 
 be perfectly preserved. The costal margin is well rounded inwardly at the base, 
 and flattens over the middle of the wing. Assuming that it agreed with wings of 
 a Phylomylacrid type, the margin would gradually merge distally into a broadly 
 rounded apex. 
 
 The subcostal vein is the weakest of the whole series, and passes obliquely 
 outwards, reaching the margin about the middle of the wing. It gives off five 
 undivided branches which are oblique in direction, and fail to reach the margin. 
 
 The costal area is broad basally, and ends in an acute angle against the margin. 
 It may be best described as pyriform. 
 
 The radius is a powerful vein, giving origin, in the inner third of the wing, to 
 what seems to be a well-marked radial sector, and dividing into two equal branches 
 a little further out. The outer of the two distal branches divides by twice forking 
 into three twigs, which may have reached the middle of the outer margin. The 
 inner of the two branches forks once only. 
 
 A wide area separates the radius from the subcosta over the greater part of its 
 length, the widest interval being at the point where the radius divides into two 
 main branches. The branch (? radial sector), of which but few divisions are 
 shown on the wing-fragment, diverges inwardly from the main stem of the radius 
 into the middle of the wing, giving off two branches, the first forking close to the 
 broken edge of the nodule. From its position the ultimate divisions must have 
 ended upon the outer half of the wing-apex, and the distal part of the outer 
 wing-margin. 
 
 The various divisions of the radius are so divergent that they enclose a con- 
 siderable portion of the whole wing, while they are equally well spaced out from 
 the subcosta in front, and the median behind. The base of the radius stands up 
 in relief, and owing to a little enfolding of the wing, overhangs the bases of the 
 median and cubitus, but does not unite with them. 
 
 The median vein arises close to the radius, and like the latter, bends inwards 
 at the base in a strong curve, which is continued until the direction is obliquely 
 inward, when it becomes straightened. 
 
 At the summit of the basal curve the median sends off a short oblique vein 
 inwards to the cubitus. This vein is a distinct commissure, and not an enlarged 
 interstitial nervure. The significance of this commissural vein is not yet fully 
 understood. Pruvost has recorded a similar structure in species of the genus 
 Archimylacris (op. cit. t p. 151), and mentions that Scudder and Handlirsch have 
 also indicated its occurrence in other forms. It is not, however, present in 
 Archimylacris woodwartU, Bolton, and A. hantata, Bolton.
 
 PHYLOMYLACRIS MANTJDIOIDES. 127 
 
 The median remains undivided in the proximal third of the wing, and then 
 forks, the inner of the resultant veins again dividing on the broken edge. The 
 areas on either side of the main stem of the median are, like those of the radius, 
 well spaced. 
 
 The ciibitus vein is less curved basally than the median and radius, well 
 separated from the latter, and passing obliquely towards the distal end of the 
 inner margin of the wing. It gives off six inward branches, the first arising near 
 the base, and margins the anal furrow. The second branch gives off two irregular 
 twigs which do not reach the margin, and the fourth is forked. The rest are 
 undivided. 
 
 The whole of the anal area is remarkably well developed. It stands out as a 
 tumid mass, marked off from the rest of the wing by the first branch of the 
 cubitus, and with all the anal veins in high relief. 
 
 A broad area interposes between the first anal vein and the anal furrow, and 
 upon this run out a few small and wavy veins which die out on the membrane. 
 The first well-defined vein has a short stem dividing at once into two equal 
 branches, which continue to the margin. The inner of these two branches either 
 forks again, or has received the larger and distal portion of the next vein, the 
 basal part being joined to the third vein, and forming an enclosed area. The 
 next vein forks, and also the sixth, the remainder being closely packed together, 
 and rapidly diminishing in strength and length as they crowd down upon the 
 margin. 
 
 The interstitial neuration consists of a well-defined meshwork which shows a 
 tendency to a longitudinal arrangement in the direction of the wing-margin. 
 
 Affinities. The incorrect figure published by Kirkby has been a source of 
 confusion, and probably accounts for Goldenberg's reference of the species to 
 Blattidium, Scudder. At the time when he so referred it, Scudder had become 
 familiar with the distinctive characters of wings of this type, and had created a 
 new genus Neci/mylacris to receive them ('Mem. Boston Soc. Nat. Hist.,' vol. iii, 
 p. 52, 1879). We can only account therefore for his reference of the wing to 
 Etoblattina on the score of the faulty drawing, as he does not seem to have seen 
 the wing. The later drawing by Handlirsch is wrong in almost every detail, and 
 his reference of the wing to the Archimylacridre was the only one possible under 
 the circumstances. 
 
 Scudder's genus Necymi/lacris was founded, not upon N. lacoann, as Hand- 
 lirsch assumes (a fragment only of that wing being known), but upon N. heros. 
 Handlirsch's later name of Eumorphoblatta ('Die Fossilen Insekten,' p. 195, 1906) 
 is rightly reduced by Pruvost ('Ann. Soc. Geol. Nord,' vol. xli, p. 350, 1912) to 
 the position of a synonym. 
 
 The figures of the three new species published by Pruvost in that paper are 
 admirably reproduced, and as they show a complex of raised nervures in nearly
 
 128 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 all tlie areas of the wings, such as is seen in the British specimen, and as the 
 species agree among themselves while differing from the type of Scudder's genus 
 Necymylacris, I considered them to represent a new genus, as already mentioned. 
 In N. heros Scudder plainly shows an interstitial neuration of closely-packed 
 transverse nervures, which he describes as " a minute tracery of nearly straight, 
 very closely approximated, excessively delicate, scarcely impressed, cross-lines." 
 
 Apart from this difference, there is a close relationship between Necymylacris 
 and Pruvost's Phi/lomylacris. Regarding, as I do, the character of the interstitial 
 neuration, especially when well developed, as one of more than specific importance, 
 I consider that Phylomylacris is a valid genus. 
 
 Genus SOOMYLACRIS, Handlirsch. 
 1906. Soomylacris, Handlirsch, Die Fossilen lusekten, p. 260. 
 
 Generic Characters. Wings short and wide, being two to two-and-a-half times 
 as long as wide. Costal area broadly triangular, the subdivisions of the subcostal 
 arising from a central point, the inner branches only reaching the costal margin 
 after a very oblique course. Radius with distinct radial sector. Median dividing 
 low down into two main branches, while the cubitus is long, with simple or forked 
 branches. Anal area large, nearly as long as the subcostal area, strongly convex, 
 and crossed by numerous anal veins, the first usually thrice divided, both the 
 subcostal and anal areas extending nearly to the middle of the wing. 
 
 Soomylacris deanensis (Scudder). Plate IX, fig. 2; Text-figure 40. 
 
 1895. EtoUattina deanensis. Scudder (in part), Bull. U.S. Geol. Surv., no. 124, p. 34, pi. xii, fig. 1. 
 
 1896. EtoUattina deanensis, Scudder, Geol. Mag. [4], vol. iii, p. 12, fig. 1. 
 
 1906. Soomylacris deanensis, Handlirsch, Die Fossilen Insekten, p. 260, pi. xxvii, fig. 15. 
 
 Type. Right tegmen, lower view, lacking apical portion; U.S. National 
 Museum, Washington (Lacoe Collection, no. H. 21326; Nat. Mus. no. 38090). 
 
 Horizon and Locality. Coal Measures; Foxe's Bridge, Forest of Dean, 
 Gloucestershire. 
 
 Description. Scudder described and figured two specimens from the Forest of 
 Dean under the name of EtoUattina deanensis, but the second is dealt with in this 
 monograph under the new specific name of Soomylacris stocld (see p. 130). 
 
 The type tegmen of 8. deanensis has a length of 25 mm., and a maximum 
 breadth of 13 mm. across the anal area. Scudder gives the length as 38 mm., but 
 the wing has lost a second portion of the apical end since he figured the specimen. 
 The anal area is broken, and has become slightly displaced along the line of the 
 anal furrow, but to so slight an extent as not to hinder measurement.
 
 SOOMYLACRIS DEANENSIS. 129 
 
 The costal margin is strongly convex basally, becoming almost straight over 
 the middle of the wing, in this respect differing from 8. stocki, which is increasingly 
 convex in the middle of the wing a fact noted by Scudder. The inner margin is 
 more nearly straight, and the wing narrows as the apex is reached, the maximum 
 width being across the anal area. 
 
 The subcosta is made up of three main branches, the first forking into two equal 
 twigs, the second forking twice into three twigs, and the third forking three times 
 into four twigs. The subcostal area is very wide basally, and terminates in an 
 acuminate angle beyond the junction of the first and second thirds of the wing. 
 
 The radius is a powerful vein dividing low down into two main branches, the 
 outer breaking up into four twigs by a trifid forking, while the inner branch forks 
 into two, and each of these into two again, the two twigs of the inner secondary 
 fork each dividing into two. The radius thus ends on the broken edge of the wing 
 in ten twigs. 
 
 rtJ& 
 
 FIG. 40.-SoomyZoms deanensis (Scudder) ; diagram of venation of right tegmen (the diagram reversed), 
 twice natural size.-Coal Measures ; Foxe's Bridge, Forest of Dean, Glos. U.S. National Museum, 
 Washington (Lacoe Coll., no. H. 2132 b; Nat. Mus. no. 38090). 
 
 The median vein forks into two main branches at about the end of the proximal 
 third of the wing. The outer branch divides into two equal twigs, the inner 
 forking again further out. The inner branch forks once only on that portion of 
 the wing which is now preserved, although Scudder has figured it as breaking up 
 into six ultimate twigs by repeated forking of the innermost of the two branches 
 now seen. Scudder shows twelve twigs of the median ending on the inner half of 
 the wing-apex, while the actual specimen now shows only five. Scudder is wrong 
 in showing a twig coming off inwardly from the first main division of the median, 
 and dying out in the wing-membrane. The appearance is due to a slight furrow 
 on the impression. This furrow can be traced across the branches of the adjoining 
 
 cub it us. 
 
 The cubitus, like the median, is sigmoidally curved, and gives off five inward 
 branches, and ends upon the distal end of the inner margin in a small fork, 
 second and third branches of the cubitus fork twice, and end on the : 
 three twigs each. 
 
 Unlike the rest of the wing, which is flatly convex, the anal area i 
 rounded, and must have projected somewhat above the general surface. 
 17
 
 130 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 crossed by six anal veins, the first being a large structure breaking up into four 
 branches, and more widely separated from the anal furrow proximally than is its 
 outer branch distally. The second vein is simple. The third forks twice, and ends 
 in three twigs. The fourth and fifth fork once each. Traces of two more anal 
 veins are seen on the margin. 
 
 The interstitial neuration can only be distinguished clearly in the anal area, 
 and consists of irregular cross-nervures which cross the main veins and unite 
 laterally to form a meshwork, which is not, however, of a strictly reticulate 
 character. 
 
 Soomylacris stocki, sp. nov. Plate IX, fig. 3; Text-figure 41. 
 
 1895. Etoblatti/ia deanensis, Scudder (in part), Bull. TJ.S. Geol. Surv., no. 124, p. 34, pi. xii, fig. 3. 
 
 1896. Etoblattina deanensis. Scudder, Greol. Mag. [4], vol. iii, p. 12, fig. 2. 
 
 1906. Soomylacris deanensis, Handlirsch, Die Fossileu Insekten, p. 260, pi. xxvii, fig. 16. 
 
 Horizon and Locality. Coal Measures; Crump Meadow, Forest of Dean, 
 Gloucestershire. 
 
 Type. Fragments of two tegmina; U.S. National Museum, Washington (Lacoe 
 Collection, no. H. 2132 c; Nat. Mus. no. 38090). 
 
 Description. The fragment of a left tegmen has lost much of its base and apex, 
 while the greater part of the anal area is obscured by the anal area of the 
 presumably opposite right tegmen. It lies with the under-surface uppermost, the 
 anal area of the second tegmen being right side uppermost. Feeble traces of what 
 appear to be a hind-wing can be distinguished through the fragment of tegmen, 
 and project a little beyond its margin at one point. The specimens are on a dark 
 grey shale in which occur numerous remains of Neuropteris, Annularia, etc. 
 
 The outer margin is seen on the displaced shoulder of the wing, and on the 
 portion of the outer half of the wing that remains. It is more convex than in 
 8. deanensis, and the shoulder bends inwards almost at a right-angle, that of 
 8. deanensis being regularly rounded. 
 
 The subcosta is represented by the bases of three stout veins which spring 
 from the same point, all showing forking, and by two straight veins running out 
 on the costal margin on the fragment. These two seem to be the terminals of the 
 innermost forking seen on the shoulder of the wing. The intercostal area is 
 broader than that of 8. deanensis. The base of the radius is missing, and the 
 union of the two twigs into which the outer branch divides is missing. Each of 
 these twigs forks again half-way between its origin and the margin. 
 
 The inner branch of the radius divides by twice forking, and the inner of the 
 resultant twigs forks again, so that the radius ends on the margin in at least nine 
 twigs.
 
 SOOMYLACRIS BURRI. 131 
 
 The median forks equally twice, the inner of the four twigs again forking twice 
 into three. 
 
 The cubitus is a simpler structure than that of 8. deanensis. It consists of a 
 sigmoidally curved main stem giving off five inward branches, the first and third 
 only forking. The cubitus, therefore, ends on the inner margin in eight twigs, 
 that of 8. deanensis in eleven. 
 
 The anal area of the right tegmen is superficially much like that of S. deanensis, 
 but the first anal vein gives off three branches in place of two, the innermost 
 forking. In 8. deanensis the middle branch forks. The second vein forks twice, 
 that of 8. deanensis being undivided ; the third forks once into two twigs, that of 
 S. deanensis having three ; the fourth forks once, the fifth is undivided, and the 
 sixth forks twice into three twigs. Traces of a seventh vein are present. Portions 
 of the anal area are shown near the inner margin, and above the outer edge of the 
 anal area of the right wing, but they are too fragmentary to determine the course 
 of the veins. 
 
 YIQ. 41 Soomylacris stocki, sp. nov. ; diagram of venation of two tegmina, twice natural size. Coal 
 Measures ; Crump Meadow, Forest of Dean, Glos. U.S. National Museum, Washington (.Lacoe 
 Coll., no. H. 2132 c ; Nat. Mus. no. 38090). 
 
 The inner margin is almost straight, and owing to the greater convexity of the 
 outer margin, the wing appears to have been narrower apically than S. deanensis. 
 
 The interstitial neuration is similar to that of 8. deanensis. Traces of possibly 
 one of the hind- wings can be seen underneath the costal margin and under the 
 outer twigs of the radius. The wing is closely related to that of 8. deanensis, but 
 the differences, especially in the cubitus, are sufficient to distinguish the two. 
 
 Soomylacris burri, Bolton. Plate IX, fig. 4 ; Text-figure 42. 
 
 1912. Soomylacris (Etoblattina) burri, Bolton, Quart. Journ. Geol. Soc., vol. Ixviii, p. 318, pi. xxxm, 
 figs. 1, 2. 
 
 Tt/pe.Lett tegmen ; Museum of the Kent Coal Concessions Co., Dover. 
 
 Horizon and Locality. Coal Measures (dark shale from a depth of 1208 feet) ; 
 Barfreston Boring, Kent. 
 
 Specific Characters. Subcosta with few divisions. Radius with well-marked 
 radial sector, which is well developed and much divided. Median with two main
 
 132 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 divisions. Cubitus large, with six simple branches. Anal area very wide basally, 
 and with 46 veins. Tegmen tapering along both margins to the apex. 
 
 Description. The tegmen lies with the under surface uppermost, lacking only 
 a small portion of the wing-apex. It is short and broad, measuring 14mm. long 
 and 8mm. wide. The dorsal surface is convex, the exposed under surface being 
 concave. 
 
 The costal margin is rounded, especially in the basal portion of the subcostal 
 area. Distally the margin curves into the wing-apex, which is bluntly rounded. 
 The inner margin curves outwards in its distal half,, so that the wing is widest 
 basally, and narrows into the wing-apex. 
 
 The subcosta is a strong vein with few branches, the first two being simple, 
 and passing out obliquely. A forked branch arises further out, and near the 
 margin of the wing a single simple branch. The costal area is broad basally, and 
 well rounded. 
 
 The radius is a large, much-divided vein, occupying, with the subcosta, the 
 outer half of the wing. It is short, and gives off a radial sector in the basal 
 
 FIG. 42. Soomylacris burri, Bolton ; diagram of venation of left tegmen, four times natural size. 
 Coal Measures (dark shale from a depth of 1208 ft.) ; Barfreston Boring, Kent. Mus, Kent Coal 
 Concessions Co., Dover. 
 
 fourth, afterwards passing out obliquely to the outer margin, and giving off four 
 outward branches, the first irregular in its course, and with two feeble twigs 
 which do not reach the margin, the second and third forking well out, and the 
 fourth small and undivided. The radial sector passes straight to the wing-apex, 
 giving off six strong branches, the first, second, and fourth forking, the rest 
 remaining undivided. 
 
 The median is convexly curved basally, and bent inwards to the inner margin. 
 It gives off a strong forward triple-divided branch in the basal fourth, and further 
 out a second branch, which forks three times into four branches. The eight 
 divisions of the median end on the distal third of the inner margin and part of the 
 apex. 
 
 The cubitus describes a regular sigmoidal curve, and gives off at regular 
 intervals a series of six inwardly directed branches, all of which are undivided. 
 
 The anal area is very wide basally, and crossed by 4 6 veins, the first forking 
 twice, the second once, the third undivided, and the fourth forked. Traces of two
 
 ORTHOMYLACRIS LANCBOLATA. 133 
 
 very small anal veins are also seen in the curve of the wing-margin. The anal 
 area is strongly convex between the anal furrow and the first anal vein. 
 
 The interstitial neuration of the anal area appears to consist of straight cross- 
 nervures. Over the rest of the wing the interstitial neuration cannot be made out. 
 
 Affinities. When first describing this species, I regarded it as closely allied to 
 forms which Handlirsch has placed in his genus Hemimylacris. There can be no 
 doubt, from what has been already said, that a close relationship exists. Pruvost 
 (op. cit., pp. 222 4) has recorded two species of Soomylacris from the Lens and 
 Lievin Coalfields of Northern France, 8. lievinensis, Pruv., and 8. aff. deanensis. 
 The occurrence of these British species may serve to indicate the Westphalian 
 affinities of the Kent and Forest of Dean Coalfields. 
 
 Genus ORTHOMYLACRIS, Handlirsch. 
 1906. Orthomylacris, Handlirsch, Proc. U.S. Nat. Mus., vol. xxix, p. 768. 
 
 Generic Characters. Tegmina two to two-and-a-half times as long as wide, 
 with a sub-cordate outline. Costal area extending from one-half to two-thirds the 
 length of the tegmen. Radius extending to the tegmen-apex, and giving off a 
 large number of outward branches. Median with few veins, directed towards the 
 apex and the inner margin. Cubitus never reaching the apical margin, and with 
 few branches. Anal area at least twice as long as high. Structure leathery, and 
 with cross-wrinkles. 
 
 Orthomylacris lanceolata, Bolton. Plate IX, fig. 5 ; Text-figure 43. 
 
 1911. Orthomylacris lanceolata, Bolton, Quart. Journ. Geol. Soc., vol. Ixvii, p. 167, pi. x, figs. 1, 2. 
 
 Type. Left tegmen; Museum of Practical Geology, Jermyn Street (no. 24511). 
 
 Horizon and Locality. Coal Measures (shales associated with the Graigola Seam, 
 Pennant Series) ; Clydach Merthyr Colliery, Clydach Valley, Swansea Vale, Grlam. 
 
 Specific Characters. Tegmen long, and uniformly tapering to the apex. 
 Upper surface regularly convex. Subcostal area broadly triangular, with oblique 
 branches. Radius much branched. Median unbranched in basal half, with three 
 outer branches. Cubitus divided into two main branches, the outer forking into 
 two equal twigs, and the inner giving off three inward twigs, the first only forking. 
 Outer and inner margins convex. 
 
 Description. The tegmen, which measures 23 mm. long and 10 mm. wide, lies 
 with the dorsal surface uppermost, and is gently rounded along its length, a slight 
 flattening only being visible over the middle of the outer margin. The uniform 
 tapering of the two margins to the apex produces an elongate form of tegmen of 
 an unusual type among Blattoids. The base of the tegmen is lost, and the costal 
 area appears therefore to occupy nearly half the outer margin.
 
 134 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 The subcosta is thin, sunk in the tegminal structure, and gives off three forked 
 branches which pass in straight oblique lines to the margin. 
 
 The radius is a large vein, with four branches, the first three doubly-forked 
 and the last undivided. The main stem of the radius is convex in the basal part 
 of its length, and concave in the outer half. The branches pass out very obliquely 
 and extend on to the apical margin. 
 
 The median divides about the middle of the tegmen, on a level with the third 
 branch of the radius. It gives off three branches, the first forking twice into four 
 equal fcwigs, the second of the series forking again near the margin. The inner 
 pair of twigs unite in the middle of their length, and separate again further out, 
 so that a lenticular " cell " is produced. The second branch is undivided. The 
 third branch divides into a long outer twig which bends forwards towards the 
 apex, and a smaller and weaker twig which goes straight out to the inner margin. 
 There is, as a result of this separation, a wide part of the inner margin destitute 
 of veins. 
 
 nr 
 
 FIG. 43. Or thorny lacris lanceolata, Bolton ; diagram of venation of left tegmen, three times natural 
 size. Coal Measures (shales associated with the Graigola Seam, Pennant Series) ; Clydach 
 Merthyr Colliery, Clydach Valley, Swansea Vale, Glam. Mus. Pract. Geol. (no. 24511). Number- 
 ing of veins as in Text-figure 33, p. 104. 
 
 The cubitus arises close to the median, and at once divides into two main 
 branches, the outer being forked once only, and the inner giving off a forked 
 branch succeeded by two which are undivided. 
 
 The anal area has broken away along the line of the anal furrow. It is shorter 
 than the subcostal area. 
 
 The interstitial neuration cannot be determined with certainty. The surface 
 near the apex is marked by cross-wrinkles, but I cannot affirm that this is a part 
 of the neuration. 
 
 LARVAL BLATTOIDS. 
 (Blattoidea) peachi (Woodward). Plate IX, fig. 6. 
 
 1887. EtoUattina peachii, Woodward, Geol. Mag. [3], vol. iv, p. 433, pi. xii, fig. 1. 
 1906. (Blattoidea) peacUi, Handlirsch, Die Fossilen Insekteu, p. 178, pi. xviii, fig. 26. 
 
 Type. Upper surface of head, pronotum, rudimentary wings, and broad seg- 
 mented abdomen, in nodule of fine grey sandstone ; Kilmarnock Museum.
 
 (BLATTOIDEA) PEAOHI. 135 
 
 Horizon and Locality. Coal Measures ("grey sandy shale with nodules of 
 impure clay and ironstone" at 91 ft. 6 in. below the surface); Greenhill Pit, 
 Kilmarnock. 
 
 Description. The length of the insect is 23 mm. and its width across the fore- 
 wings 15 mm. The pronotum is 12 mm. wide, and 5 mm. long. The abdomen is 
 12 mm. long and diminishes in width from before backwards. The coarse 
 character of the stone has obliterated or failed to preserve all the finer detail of 
 the specimen, but the insect still retains the gently convex dorsal surface which it 
 doubtless had during life. 
 
 The very small head, not well defined, is apparently divided into two small 
 anterior and two larger posterior areas by shallow longitudinal and transverse 
 grooves. No appendages are visible. A raised V-shaped portion of the matrix in 
 front of the head may indicate that it formerly extended beyond the line of the 
 pronotum. The two anterior plates covering the head are notched in a small v on 
 the middle outer edge. 
 
 The pronotum is nearly two-and-a-half times as wide as long, with rounded 
 latero-posterior angles and almost straight posterior border. The front border 
 forms a semicircle, recessed in the middle, and enclosing the head-shield termed by 
 Dr. Henry Woodward an " epicranial plate." 
 
 Two pairs of rudimentary wings are present, the first pair articulating well 
 forward, as if attached to the mesonotum under the hinder margin of the pronotum ; 
 the hinder pair being joined to the metanotum nearer the middle line. The wings 
 are about 10 mm. long and 5 mm. wide, with blunt apical angles. The front pair 
 of wings have begun to take on the character of tegmina, being stouter than the 
 inner pair. 
 
 The venation of the wings is but faintly indicated. Woodward has stated 
 that the " mediastinal " vein and the veins of the anal and intermedian area are 
 seen in all four wings. In present nomenclature this means that the subcosta 
 and the cubitus are present, with some traces of the anal veins. 
 
 This seems hardly to be the case. The left hind-wing has the veins best 
 marked, and all that can be said is, that there are indications of a few short, thick 
 veins passing from near the attachment of the wing towards the inner margin, and 
 crossing the middle of the wing. The condition is much like that figured by 
 Comstock and Needham (< Amer. Nat/ [4], vol. xxxii, p. 773, fig. 56, 1898) in the 
 hind-wing of the nymph of a cockroach, except that the veins in the Kilmarnock 
 specimen spread fanwise, instead of keeping in close order down to the wing-apex. 
 The veins present will therefore agree best with the radius, median and cubitus. 
 The vein which I consider the radius is nearly parallel with the outer margin and 
 better defined than the rest. 
 
 The length of the mesonotum and metanotum together is 6 mm. Neither is 
 well seen owing to the overlap of the wing-bases.
 
 136 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 The abdomen is in excellent preservation, and has several features of interest. 
 It is 12 mm. in length, diminishing from a breadth of 10 mm. across the body and 
 expanded epimera to 3'5 mm. across the ninth segment. The central axis of the 
 abdomen is more convex than the lateral epimera, and the middle line is slightly 
 ridged. The abdomen forms about one-third of the total breadth, being 4 mm. 
 wide in the first segment, diminishing to 1-5 mm. on the ninth segment. The tenth 
 segment is missing. 
 
 The epimera are broad, with slightly thickened posterior edges. From the 
 posterior dorsal margins of the first to fourth segments arise thin plate-like 
 expansions. These pass back over the succeeding two segments, showing ragged 
 edges, as if they had a greater extension during life, and had since been partially 
 torn away. The precise relation of these structures to the segments is not quite 
 clear. On the right side of the first, second and third segments, these processes 
 seem to emerge from beneath the hinder edges of the terga. If they do not, but 
 are continuous with the hinder edge, they are yet distinct from the latter, as the 
 suture-line between the adjacent terga can be traced outwards along the front edge 
 of the epimera, and if the two are united this furrow may have functioned as a 
 joint. 
 
 The undeveloped wings, scarcely wider than the pronotum, and the broad 
 epimera of the abdomen, clearly indicate the larval condition of the specimen. 
 It must be more advanced towards the adult stage than Leptoblattiva exilis, 
 Woodw., from the Coal Measures of Coseley, Staffordshire, as it differs considerably 
 in the greater breadth of the abdomen and the development of the epimera. 
 
 In size (Blattoidea) peachi is two-thirds the length of such an adult form as 
 Apldlioroblattma johnsoni (Woodw.), and as size is fairly well correlated with 
 development in Blattoids, we may assume that the insect had not fully reached 
 the nymph stage. This conclusion seems to be also confirmed by comparison with 
 the wing-length of L. exilis, where the wings are double the length of the 
 pronotum. 
 
 The absence of determinable details in the wings prevents any successful 
 attempt to assign the specimen to a position in any accepted classification. The 
 most that can be said is. that the great breadth of the pronotum, as contrasted 
 with the length, would seem to indicate a relationship with the Mylacrida?. 
 
 Leptoblattina exilis, Woodward. Plate IX, figs. 7, 8. 
 
 1887. Leptoblattina exilis, Woodward, Geol. Mag. [3], vol. iv, p. 56, pi. ii, figs. 2, 3. 
 1906. (Blattoidea) exilis, Handlirsch, Die Fossilen Insekten, p. 173, pi. xvii, figs. 16, 17. 
 
 Type. Nearly perfect larval blattoid in an ironstone nodule ; British Museum, 
 Johnson Collection (no. 1065).
 
 LEPTOBLATTINA EXILIS. 137 
 
 Horizon and Locality. Middle Coal Measures (binds between the ".Brooch " 
 and " Thick " coals) ; Coseley, Staffs. 
 
 Description. Vhe type-specimen and a second (no. 1066), described at the same 
 time, show an almost complete pronotum, two pairs of rudimentary wings, and a 
 long jointed abdomen. Dr. Henry Woodward recognises a portion of the head 
 projecting in front of the pronotum in each, but of this I am doubtful. He also 
 describes the head as follows : " The head is very small, and somewhat bluntly 
 pyramidal in form, and measures 2 mm. in breadth at its base, where it disappears 
 beneath the pronotum, and is 2 mm. in length. There is a suture visible down the 
 centre which divides the two epicranial plates, at the sides of which the eyes 
 would be seen ; in front of the epicranium a small projection no doubt represents 
 the clypeus with the labrum at its extremity." 
 
 In the type-specimen (PL IX, fig. 7) the pronotum, 7*5 mm. long and 9 mm. 
 wide, is well rounded in front and on the sides, the margin of the latter passing 
 by blunt rounded angles into an almost straight hinder border. The surface is 
 marked out into the usual central raised area, bordered by slight grooves which 
 deepen backwards, and curve inwards. The lateral portions are flattened. 
 
 The dorsal part of the mesonotum is partially exposed, its anterior edge being 
 hidden beneath the pronotum. The front pair of wings is attached to the 
 mesonotum so far forward as to touch the hinder edge of the pronotum. The 
 surface of the mesonotum was originally rounded, and judging from its present 
 condition, somewhat thin. It is now slightly puckered by folding. 
 
 The metanotum is similar in character to the mesonotum, but a little more 
 robust, and has the second pair of wings still articulated near the front margin. 
 
 The abdomen is long, the segments immediately following the body being 
 flattened by pressure, and their boundaries not clearly discernible. Behind follow 
 at least five well-marked segments which decrease in diameter backwards. The 
 last three show lateral epimera with the points turned backwards. Traces of 
 what appear to be cerci follow the last segment. These seem to be curved 
 inwards at their tips, and widely spaced, but their definition is unsatisfactory. 
 The length of the a-bdomen behind the metanotum is 14mm., and the last segment 
 is not more than 2 mm. wide, the first of the six segments being double that 
 diameter. 
 
 The wings are short, stout structures, those of the right side being perfect, and 
 those of the left side incomplete. The right tegmen has a length of 11 mm., and 
 the hind-wing a length of 10 mm. The venation, as in all larval wings, is very 
 obscure, and can be best studied on the impression of the wings. It consists of a 
 stout vein passing from the point of articulation of the wing outwards towards the 
 wing-apex, and keeping close and parallel with the outer margin. From this vein 
 a series of fine branches spreads fan-wise towards the inner margin of the wing, 
 and seems to reach it. Woodward has recognised a mediastinal (subcostal) vein 
 18
 
 138 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 and an anal vein. An examination of a good cast of the specimen by oblique light 
 shows that there are a strong outer vein parallel with the outer margin, and at 
 least two others with short, thick sterns breaking up into a fan-like series of 
 oblique veins which spread out towards the inner margin. From the researches 
 of Messrs. Comstock and Needham (1899, 'American Naturalist' [4], vol. xxxiii, 
 pp. 573-582, fig. 74), these veins seem recognisable as the radius, median and 
 anal. I am unable to distinguish any detached limb such as that mentioned by 
 Woodward. 
 
 The second example (PI. IX, fig. 8) differs from the first in several small 
 details. It lies on an inclined surface, and the abdomen has been dislocated and 
 twisted over to the right, so that it is in a different plane from the rest of the insect. 
 The small " head " described by Woodward can be seen projecting from the 
 middle of the frontal margin of the pronotum, which is a little broken away. It 
 consists of a wedge-shaped structure, which narrows forwards, broad at its base 
 and slightly swollen. It is divided into an anterior centrally placed plate of 
 quadrangular outline, the hinder angle joining a sunken line (suture ?) dividing the 
 two basal plates, which Woodward terms " epicranial." Too little is seen of the 
 anterior central plate for description, or for its recognition as a labrum. 
 
 The pronotum is raised in the centre and hollowed on the sides, while the 
 hinder margin is partially broken away. It overhangs the bases of the first pair 
 of wings. A ridge runs down the median line of the pronotum, the sides of the 
 latter being less rotund than in the first specimen, and the whole pronotum is 
 more sub-pyramidal in form. 
 
 The first pair of wings is still in place, and about 11 mm. in length, allowing 
 for the basal part hidden under the hinder margin of the pronotum. The wings 
 stand out at a low angle from the body, leaving the mesonotum and metanotum 
 exposed. Both are partially crushed and crumpled, and the characters cannot be 
 made out. The left wing is the better marked of the two. 
 
 The outer margin of the wing of the first pair is stout, and raised above the 
 general level. Two stout ridges rise from the middle of the base and pass along 
 the wing, the second ridge being the strongest, and traceable over two-thirds of 
 the length, while the first ridge does not extend beyond the basal fourth. The 
 first ridge occupies the place of the subcostal vein, and dies out in the direction of 
 the outer margin, but with no evident branches. The second and larger ridge has 
 all the appearance of a radius, is parallel with the outer margin, and can be seen 
 to send off four branches to the middle of the wing-apex, and the inner half of it. 
 The anal area is more slightly ridged, but with no discernible divisions. 
 
 The hind-wings have a straighter outer margin than the fore-wings, and are 
 more membranous. Very little is seen of the left hind-wing. The outer margin 
 of the right hind-wing seems to be folded back on itself, and the radius vein is the 
 first distinguishable. It is more branched than its fellow of the fore-wing, giving
 
 MEGANEURID^E. 139 
 
 off at least five branches, the fourth. being forked. These branches reach a portion 
 of the outer side, and apex of the wing, and the whole of the inner side of the 
 apical margin. More inward is seen the distal half of another vein which may be 
 the median. It separates into two branches which reach the distal part of the 
 inner margin. The anal area is hidden by being folded in against the body. 
 
 The wings appear too small and of too rigid a type to have been serviceable 
 for flight, and the elongated abdomen could not have been supported by such 
 rudimentary wings. The size and condition of the hind-wings shows that the 
 great increase in width over that of the tegmina in the adult form is a com- 
 paratively late development, and probably synchronised with the shortening up of 
 the abdomen. 
 
 The proximal segments of the abdomen are obscure, but beyond the dislocation 
 can be seen four well-rounded segments, followed by the crushed terminal segment 
 and indications of one of the cerci. 
 
 Affinities. The generic name attached to these specimens by Dr. Woodward is 
 of doubtful value, since in the present state of our knowledge it is impossible to 
 assign larvas to the adult forms of which they are the immature representatives. 
 The only adult forms known from the same beds are Archimylacris johnsoni, 
 A. eggintoni and A. incisa. 
 
 Order PROTODONATA (Brongniart), Handlirsch. 
 1906. Handlirsch, Die Fossilen Insekten, p. 304. 
 
 Large insects with slender bodies similar to those of the Odonata. Wings 
 large, outspread in the condition of rest, and only capable of an up and down 
 movement in one place. Wing-neuration specialised by the union of several 
 longitudinal veins into accessory or interpolated sectors. Cross-nervures well 
 developed in a regular order. Head large, with large eyes and powerful jaws; 
 thorax much as in the Odonata, Legs strong and similar. Antennae short. 
 Abdomen long. 
 
 Family MKGANEURID^E, Handlirsch. 
 1906. Handlirsch, Die Fossileu Insekten, p. 306. 
 
 Wings Protodonate in character, with a pre-costal marginal area destitute of 
 veins. Costa almost straight; subcosta simple, and uniting with the costa far o 
 Radius simple, and followed by a series of accessory veins which represent 
 radial sector, these all curving towards the distal inner margin. Median
 
 140 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 apparently united to the radius at the base, and dividing about the middle of the 
 wing into two widely divergent branches, between which are developed a numerous 
 series of accessory veins. Cubitus (in the type species) consisting of a single 
 undivided vein on the outer side, and of an inner branch which is much subdivided. 
 Anal vein reaching middle of inner margin, and giving off many strongly recurved 
 branches. Inner margin broadly convex. Interstitial neuration consisting of 
 numerous series of straight cross-nervures. Wings membranous. 
 
 This family was formed for a small group of insects of enormous size, discovered 
 at Commentry and described by Brongniart. 
 
 Genus MEGANEURA, Brongniart. 
 1885. Meganeura, Brongniart, Bull. Soc. Amis Sci. Nat. Rouen [3], ann. xxi, p. 60. 
 
 Generic Characters. Gigantic insects with long narrow fore-wings and broader 
 hind-wings. Precostal area extending to the middle of the outer margin. Costa 
 almost straight, powerful, and joined far out by the subcosta. Radius a single 
 strong vein, followed by a radial sector arising from the radius in the fore-wing, 
 and in the hind-wing from the outer branch of the median. Median united with 
 the radius at the base, having a simple outer branch and a much divided inner 
 branch. Many of the branches of the latter accessory. Cubitus with outer and 
 inner branches dividing at the base of the wing, and curved in an S-shape. The 
 inner branch well divided, the outer feebly so. Anal veins numerous, strongly 
 curved to the inner margin. 
 
 Genus BOLTONITES, Handlirsch. 
 1919. Bolto'nites, Handlirsch, Revision der Paliiozoischen Insekten, p. 61. 
 
 Generic Characters. Pre-costal field very small ; cubitus and anal veins much 
 as in Gilsottia titana, Meunier. " The bridge clearly preserved." By this I 
 understand that Handlirsch alludes to the transverse vein which I described as an 
 "oblique inward branch of the cubitus joining on to the anal." Cubitus and anal 
 with few branches. 
 
 Boltonites radstockensis (Bolton). Plate X, fig. 1 ; Text-figure 44. 
 
 1914. Meganeura radstockensis, Bolton, Quart. Journ. Geol. Soc., vol. Ixx, p. 119, pis. xviii, xix. 
 1919. Boltonites radstockcnsis, Handlirsch, Revision der Paluozoischen Insekten, p. 61. 
 
 Specific Characters. Outer wing-margin thickened, coriaceous and tubercular. 
 Outer and inner wing-margins spinulose. Cubital-anal vein present. Areolae
 
 BOLTONITES RADSTOCKENSIS. 141 
 
 (P aborted spiracles) in anal area. Length of wing when complete about 190 mm. 
 or seven and a half inches. 
 
 Type- Proximal third of wing, in dark shale with plant-remains ; Sedgwick 
 Museum, Cambridge. 
 
 Horizon and Locality. Upper Coal Measures; Tyning Colliery, Radstock, 
 Somersetshire. Precise horizon uncertain. 
 
 Description. The fragment consists of the proximal portion only of a wing, 
 64 mm. long and 40 mm. in greatest width, and appears to have formed about 
 one-third of the whole. The wing has also been broken along the middle, and a 
 portion lost, due apparently to the shale breaking irregularly, and leaving an 
 uneven surface. The outer and inner parts of the wing-fragment still retain their 
 normal position relatively to each other, but a portion of the inner part of the 
 base is missing. 
 
 The outer part of the specimen consists of two veins, the costa and sub-costa, 
 The inner portion shows the cubital and anal veins. The veins missing are there- 
 fore the radius and median. 
 
 FIG. -44. Boltonites radstockensis (Bolton) ; restoration of right wing, based on that of Meganeura 
 monyi, Brongt., one-half natural size. Upper Coal Measures (precise horizon uncertain) ; Tyning 
 Colliery, Badstock, Somerset. Sedgwick Museum, Cambridge. 
 
 The outer wing-margin consists, not of a free alar expansion, as is usual in the 
 Protodonata, but of a coriaceous and tuberculated mass, which is well marked off 
 from the costa along its whole length. This thickened mass apparently represents 
 the free alar development seen in other members of the group. 
 
 That other veins existed in the middle area of the wing is proved by the 
 remains of three short portions of a vein, lying in the middle of the interspace. 
 Not until these fragmentary vein-structures had been perceived, and the type of 
 M^ijaneura monyi studied in the Museum at the Jardin des Plantes, Paris, could its 
 relationship with the Protodonata be confirmed. The direct comparison of the 
 two wings also showed that the Radstock specimen had the same parallelism of 
 the principal veins, the co-existence of similar cross-veins, and the same distinctive 
 character of the anal area as M. monyi. 
 
 The basal portion of the outer margin of the Radstock wing is swollen into an 
 elongated mass, which thins out distally, and probably did not extend beyond the 
 proximal third of the wing. It can be traced for a length of 20 mm., and near 
 the articulation is covered with numerous low, smooth-topped tubercles, arranged
 
 142 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 irregularly. The more median part of the costal margin appears as a flat, straight, 
 knife-like edge, crossed by a series of fine striations. 
 
 The thickened pre-costal portion of the outer margin forms a rigid bar, 
 enormously strengthening the wing for flight. The costa is distinct, and passes 
 directly to the wing-apex, as a broad, strap-shaped vein, broadest at its point of 
 origin, and slowly diminishing in width to the tip of the wing. 
 
 The subcosta has an apparent union at its base with the costa, but this is due 
 to a slight backward displacement of the latter. Some distance beyond the 
 thickened margin, the costa bears, at regular intervals, a series of low spines, 
 projecting freely forwards, and gradually inclining towards the wing-apex. A 
 few scattered tubercles, and a faint ridge, are seen along the middle of the vein, 
 and eventually die out. The distal part of the costa is straight, and is joined to 
 the subcosta by a series of transverse branches which are well marked. Of these 
 branches, eleven are still whole, and portions of nine others can be made out. It 
 would therefore seem that the whole of the costal area was covered by a parallel 
 system of straight cross-branches with no intervening network. 
 
 The subcosta arises in close contact with the costa, diverging rtipidly until the 
 two are about 5 6 mm. apart. Once this distance has been attained, they remain 
 nearly parallel for some distance. There are indications that the two come 
 together some distance beyond the broken edge of the wing-fragment. 
 
 The costa and subcosta are nowhere more than 5 6 mm. apart, in the main 
 but 4 mm., the interval narrowing to the broken edge of the wing. While the 
 subcosta has the same flat, strap-shaped character as the costa, it is more delicate, 
 and a narrower vein. A few tubercles are disposed along the median line of its 
 length. 
 
 The radius and median veins being lost, except for the trivial pieces already 
 noted, they cannot be compared with those of Meganeura. In the French species 
 of this genus a complex of parallel veins arises from one or two roots between the 
 subcosta and the cubitus. In the case of M. monyi, this complex arises from a 
 strong radius and a weak median. In M. selysii it would appear that the radius 
 and median are united at their base. In the restoration of the Radstock wing it 
 will be seen that the vein-fragments appear as parts of the median vein, the radius 
 being wholly lost. 
 
 The cubitus is separated from the outer part of the wing by the interval 
 formerly occupied by the radius and median. This interval is variable in width, 
 owing to the strong curvature of the cubitus. This is a large flattened vein, 
 strongly flexuous, and finally curving inwards to reach the middle of the inner 
 margin. The base of the vein has been broken away and lost, but its course may 
 be indicated by a faint groove which curves forward towards the base of the 
 subcosta. The outer inward curve of the cubitus is the greater, so that the 
 interval between the radius and the subcosta becomes increasingly wide, thus
 
 BOLTONITES RADSTOCKENSIS. 143 
 
 agreeing exactly with what obtains in the same part of the wing in Meganeura 
 niont/i. It is an indication of the rapid inward curvature which takes place a little 
 further out, and of the presence in the complete wing of the wide distal area 
 occupied by the many branches of the radius and the median. 
 
 Close to the basal broken end of the cubitus a strong oblique inward vein is 
 given off, which reaches the anal vein, and fuses Avith it. At the time when I first 
 described this wing I wrote as follows : " It has the appearance of an important 
 commissure between the cubitus and the anal, or of a posterior branch of the 
 former which has fused with the latter." Since then, Dr. R. J. Tillyard has 
 published the results of his studies on recent dragon-flies (' The Biology of 
 Dragon-flies,' Cambridge University Press, 1917), in which he notes our discovery 
 of this oblique vein. He recognises it as an anal-cubital vein which he had 
 previously (' Proc. Linn. Soc., N.S.W.,' vol. xxxix, pp. 163 216,' 1914) 
 described as indicating the point where the true anal vein diverges from the 
 cubitus. He concludes (op. cit., p. 305) : " Hence it would appear that Cu (cubitus) 
 and A (anal) were fused basally as in all recent forms. Thus the gap between 
 Protodonata and Odonata is being gradually lessened until to-day we may almost 
 certainly see in the Meganeuridaa the giant relatives of the direct ancestors of 
 some at least of our recent families." 
 
 Beyond this anal-cubitus vein the cubitus is joined to the anal by a system of 
 parallel slightly curved branches, similar in character to those which unite the 
 sub-costa and the costa. No fewer than twenty-five of these branches can be 
 distinguished. Basally to the anal-cubital vein are two transverse branches, a little 
 more curved than the rest. As in the costa and sub-costa, a median line of 
 tubercular ornament is present. 
 
 The anal vein is strongly marked, and its course is much similar to that of the 
 cubitus, but the second inward flexure is less marked, so that the two veins are 
 closer together in the middle of their length, and more widely separated basally. 
 The anal gives origin, along the whole length of its hinder margin, to a series of 
 branches, which arise at slightly increasing intervals, being closest basally, and 
 most widely separated between the twelfth and thirteenth branches, beyond which 
 the interspaces narrow again to the fifteenth branch. The basal branches pass in 
 straight or slightly oblique lines inwards to the wing-margin. Further out they 
 become curved, with the convexity outwards. The twelfth branch is a strong and 
 important vein, sweeping in a powerful double curve distally, and inwardly to the 
 wing-margin. It corresponds in position to Brongniart's " vein X." Beyond it 
 are the remains of three feebler branches, which bend in simple curves to the 
 margin. 
 
 The spaces between the anal branches are divided up into a series of quad- 
 rangular areas or cellules, by a great series of secondary branches, arising at 
 right-angles. The twelfth dorsal branch stands out from the rest by reason of its
 
 144 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 robust character and sweeping curve, and gives off, on its inner side, a series of 
 twigs, which divide and separate so widely that first a double and then a treble 
 series of cellules are enclosed between them. On its distal side it intercepts five 
 branches arising from the inner border of the main stem of the anal, the enclosed 
 quadrangular areas being increasingly elongated antero-posteriorly up to the 
 eighteenth branch, the course of that branch to the margin being uninterrupted. 
 The integument in many of the quadrangular areas or cellules is marked by a 
 slight central elevation, which, under high magnification, presents the appearance 
 of a circular thickened lip, with a central depression or perforation. These 
 structures have no regularity of arrangement, but are most numerous on each side 
 of the twelfth branch of the anal. The integument within the cellules is, in some 
 instances, obliquely wrinkled. These structures appear to resemble aborted 
 spiracles. 
 
 The inner margin of the wing is sharply sloped forwards from the broken edge 
 onwards to the point of attachment, and is well-defined and spinous along the 
 whole or the greater part of its length. The spines are low, directed distally, and 
 in character somewhat like those on the outer margin of the wing. They are not 
 so clearly shown, however, and they may not be truly marginal, as it is a matter 
 of doubt whether the integument stretches a little outside them. I incline to the 
 latter view. The ventral surface of the wing seems to be uppermost, in which 
 case the wing-fragment is the proximal portion of a right wing. 
 
 Affinities. The general characters of the wing-fragment are those typical of 
 the genus Meganeura, and it cannot be confused with the genus Paralogus. It is, 
 however, unlike M. monyi, in which the development of marginal spines has not 
 been recorded, and the tuberculation so characteristic of the Radstock wing is not 
 shown in that species. M. monyi is a wider and longer wing, and does not possess 
 the areolse seen in the quadrangular cellules of this species. 
 
 Considered as a Meganeurid wing, the Radstock specimen is no primitive 
 structure, but highly specialised, an anal cubitus connection is present, the costal 
 border has become thickened and tuberculated, and a secondary development of 
 spinules and tuberculations has arisen. 
 
 INCERT^E SEDIS. 
 
 Genus TILLYARDIA, novum. 
 
 Generic Characters. Wings elongated, five times as long as wide. Subcosta 
 short; radius simple, radial sector forming an accessory vein with several divisions. 
 Median vein with few branches. Cubitus with two main branches, the inner much
 
 TILLYARDIA MULTIPLICATA. 145 
 
 divided. Accessory sectors present. Interstitial neuration forming a regular 
 series of straight cross-nervures. 
 
 I refer to this genus a highly specialised wing from the Scottish Coal Measures, 
 which is unlike any other Carboniferous wing that I know. It is at once suggestive 
 of the Palaeodictyoptera and of the Protodonata, but belongs, I believe, to the 
 latter Order. Where shown, the neuration of the wing is clear, and the absence 
 of the outer part of the base, and the central portion of the wing, renders it 
 difficult to determine the actual course of certain veins and their true nature. 
 
 I have much pleasure in naming this genus after Dr. R. J. Tillyard, whose 
 researches on the venation of the wings of nymphs of recent insects have helped 
 to interpret the older fossil forms. 
 
 Tillyardia multiplicata, sp. nov. Plate X, fig. 2 ; Text-figure 45. 
 
 Type. Greater part of the impression of the under surface of a right wing ; 
 Museum of Geological Survey of Scotland, Edinburgh (no. T. 40981). 
 
 FIG. 45. Tillyardia multiplicata, sp. nov.; diagram of venation of right wing, enlarged 
 one-and-a-half times. Upper Coal Measures; Barony Pit, Auchinleck, Ayrshire. Museum of 
 Geological Survey of Scotland, Edinburgh (no. T. 4098 6). 
 
 Horizon and Locality. Upper Coal Measures; Barony Pit, Auchinleck, Ayrshire. 
 
 Specific Characters. As above. 
 
 Description. The wing lies on the irregular surface of a soft purple sandstone, 
 is imperfect, and has been badly rubbed, so that the junctions of some of the veins 
 have been obscured. The whole of the base of the wing is missing, and the 
 middle portion of the impression of the wing has also been destroyed. It is 
 67 mm. long and 13 mm. wide. 
 
 The outer margin appears to have been almost straight over the greater part 
 of its length, inclining gently into the wing-apex, the latter joining the inner 
 margin in a blunt right-angle. By analogy with the Protodonata, the outer 
 margin was extended outwardly in its basal half, and the subcostal area was 
 widened out. This feature is seen in Paralogus seschiwides, Scd., with which the 
 wing has much in common. No trace of an undoubted subcostal vein is dis- 
 tinguishable. It must therefore have been short, and joined to the outer margin 
 in the basal half. The radius is a long vein, parallel with the outer margin, and 
 19
 
 146 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
 reaching the wing-apex, curving inwards as it does so. The junctions of the 
 branches of the radial sector are destroyed. It would seem to have begun between 
 the radius and the median as an accessory vein, rather than as an offshoot of the 
 radius. Its branches occupy the outer half of the wing-apex. The median is a 
 large and important vein, dividing near the base into two branches. The outer 
 branch is undivided until close to the distal end of the outer margin, where it gives 
 off an outer and two inner twigs. The inner branch of the median gives off five 
 simple inner twigs, and an outer one which forks close to the margin. The cubitus 
 is represented by two veins, an outer vein which remains undivided and an inner 
 which gives off at least five twigs. It is possible that the vein which I thus describe 
 as the cubitus is really part of a cubito-anal, but as the basal part is missing the 
 exact conditions cannot be determined. The inner margin of the wing is nearly 
 straight. 
 
 Affinities. The wing-structure, so far as it can be made out, is much like that 
 of Paralogus aeschnoides, Scd., and Protagrion audouini, Brong., and of these it 
 more nearly accords with the latter in its great length as contrasted with the 
 breadth. The plication of the wing is unusually well developed, and is a feature 
 best seen in P. a&schnoides. The presence of accessory sectors is again a feature 
 seen in Protagrion. The general wing-structure, therefore, has certain characters 
 of both genera, and where it departs from the one it approaches the other. 
 
 ADDENDUM. 
 Archimylacris pringlei, Bolton. Plate X, fig. 3. 
 
 1921. Archimylacris pringlei, Bolton, Quart. Journ. G-eol. Soc., vol. Ixxvii, pp. 23 29, pi. i, figs. 1 2, 
 
 text-fig. 1. 
 
 Type. The basal two-thirds of a fore- wing, and counterpart impression, partly 
 obscured by plant remains ; Museum of Practical Geology, Jermyn Street (nos. 
 30725, 30726). 
 
 Horizon and Locality. Keele Group, Upper Division of the Coal Measures; 
 from rocks between the surface level and 97 feet in borehole at Slang Lane, 
 Wellington, Shropshire. 
 
 Specific Characters. Wing twice to two-and-a-half times as long as wide, 
 costal margin flattened ; subcostal area strap-shaped and very wide, crossed by 
 numerous parallel branches of the subcosta. Radius dividing beyond middle of 
 wing. Median parallel with radius. Cubitus curving to inner margin, with 
 widely separated inward branches. Anal area long. 
 
 Description. The wing-fragment has a length of 18 mm. The costal margin
 
 ADDENDUM. 
 
 147 
 
 is flatly convex over the greater part of its length, and well rounded at the base 
 of the wing. The subcosta is parallel to and widely spaced from the outer margin, 
 to which it sends eight or nine branches, two of which are forked, and one, the 
 most distal, is not fully developed. The radius is not complete, and has but a 
 slight divergence from the subcosta. It gives off the radial sector at a point 
 between the seventh and eighth branches of the subcosta. The radial sector goes 
 out to the wing-apex, sending its branches forward to the outer margin. The 
 median diverges a little more from the radius than the latter from the subcosta. 
 Portions of three inner branches are present. The cubitus is a convex vein 
 passing well out to the distal end of the inner margin of the wing, and giving off 
 a series of widely separated inward branches, of which four are present on the 
 wing-fragment preserved. Owing to the forward displacement of the anal portion 
 of the wing, the first branch of the cubitus passes under the first anal vein. The 
 second shows a simple fork, and the remaining two veins end undivided on the 
 line of fracture. Portions of six anal veins are present, the first obscure and 
 the rest parallel. The interstitial neuration consists of a compact series of close- 
 set transverse nervures, which in the basal areas between the radius, median, and 
 cubitus unite laterally and irregularly until they form a coarse irregular meshwork. 
 
 Affinities. The widely spaced subcosta and the character of the interstitial 
 neuration of the wing show a close relationship to that of two forms of Archimy- 
 lacris (A. desaillyl and A. lerichei) recorded from the Coal Measures of Lievin, 
 Northern France, but, as the accompanying tabulated comparison will show, the 
 wing is nevertheless specifically distinct from either. 
 
 Observations. The presence of this fossil Blattoid in the Keele Group of the 
 Upper Division of the Coal Measures of Shropshire, and its specific relation to 
 forms known only from the Westphalian Series of Lievin, Northern France, is a 
 matter of considerable interest, especially as Dr. Pruvost, of Lille, had previously 
 drawn attention to the fact that the fauna at the top of the Coal Measures in 
 Great Britain (Keele Group, Newcastle-under-Lyme Group, Etruria Group) does 
 not notably differ from the fauna at the top of the Westphalian in Northern 
 France. 
 
 A. pringlei, Bolton. 
 Costal Area : 
 
 Strap -shaped, widely spaced 
 from wing-margin. 
 
 Subcostal Vein : 
 
 Numerous divisions, mostly 
 forking. 
 
 19 
 
 TABULATED COMPARISON. 
 A. desaillyi, Leriche. 
 
 A. lerichei, Pruvost. 
 
 Strap-shaped, widely spaced Outer third oblique to wing- 
 
 from wing-margin. 
 
 Numerous divisions, mostly 
 forking. 
 
 margin. 
 
 Few divisions, much branched.
 
 148 FOSSIL INSECTS OF THE BRITISH C 
 
 A. pringlei, Boltou. A. desaillyi, Leriche. 
 
 Radius Vein: 
 
 Few branches. Radial sector Seven branches. Radial 
 
 arising opposite outer fourth of sector arising opposite outer 
 subcosta. 
 
 Radial Sector : 
 
 Branching not known. 
 
 Median Vein : 
 
 Branches beyond origin of 
 radial sector. 
 
 Cubitus Vein : 
 
 First branch simple, second 
 forked. 
 
 Anal Veins : 
 Undivided. 
 
 third of subcosta. 
 Eight branches. 
 
 Branches opposite first fork 
 of radial sector. 
 
 First branch simple, second 
 and third forked. 
 
 First vein only forked. 
 
 Interstitial neuration : 
 
 Transverse nervures, except Transverse nervures, except 
 
 in the median basal part of the in the median basal part of the 
 
 wing, where it is reticulate. wing, where it is reticulate. 
 
 OAL MEASURES. 
 
 A. lericJiei, Pruvost. 
 
 Twelve branches. Radial 
 sector arising opposite middle 
 of subcosta. 
 
 Seven to eight branches. 
 
 Branches much beyond origin 
 of radial sector. 
 
 First branch dividing into 
 four, second simple, third and 
 fourth branched. 
 
 First anal forking twice. 
 
 Transverse nervures, except in 
 the median basal part of the 
 wing, where it is reticulate. 
 
 BIBLIOGRAPHY. 
 
 AUDOUIN, V. " Sur une Empreinte d'Aile d'un Insecte ne'vroptere inconnu, trouve 
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 1833. 
 
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 ibid., vol. Ixviii, pp. 310323, pis. xxxi xxxiii, 1912.
 
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 pp. 661820, 1906. 
 
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 MAXTELL, G. A. 'The Wonders of Geology,' 1st American from 3rd London 
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 1867. (Brief reference to figure of an insect " allied to Corydalis" from 
 
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 152 FOSSIL INSECTS OF THE BRITISH COAL MEASURES. 
 
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 " La Faune Contineiitale du Terrain Houiller du Nord de la France. Intro- 
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 SCUDDER, S. H. " On an English Fossil Insect described as Lepidopterous," ' Proc. 
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 " Two New British Carboniferous Insects, with Remarks on those already 
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 Univ. Bull.,' no. ii, p. 175, 1881. 
 
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 ferous Formation," ' Mem. Boston Soc. Nat. Hist,,' vol. iii, pp. 143 152, 
 1883. 
 
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 747831, Mimchen und Leipzig, 1881 1885. 
 
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 vol. ii, pp. 748833, Paris, 1887. 
 
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 1904. 
 
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 BIBLIOGRAPHY. 153 
 
 TILLYARD, R. J. " The Panorpoid Complex, A Study of the Phytogeny of the Holo- 
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 " The Panorpoid Complex, Part 2. The Wing-tracheation and its Relation- 
 
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 pis. Ixvii Ixix, 1918. 
 
 " The Panorpoid Complex, Part 3. The Wing Venation," ibid., vol. xliv, 
 
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 Paleozoologie, Article " Insecta," by S'. H. Scudder, pp. 746833, Paris, 
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 682691', London, 1900.
 
 154 
 
 INDEX. 
 
 NOTE. The romaii and arable numerals following a semicolon refer to the plates and figures 
 
 illustrating those species which are recognised and described. 
 
 
 PAGE 
 
 
 PAGE 
 
 ./EdcBophasma 
 
 78 
 
 Boltonites 
 
 140 
 
 
 
 
 
 auglica . 
 
 78; V, 2 
 
 radstockensis 
 
 140; X, 1 
 
 jEnigmatodes 
 
 71 
 
 Breyeria 
 
 50 
 
 danielsi 
 
 39, 72 
 
 borineusis . . . 
 
 52 
 
 regularis 
 
 . ... 71; IV, 7 
 
 lachlani 
 
 52 
 
 jEnigmatodidse 
 
 70 
 
 woodwardiana 
 
 50 
 
 . , , , . ,, 
 
 96 
 
 Breyeriids8 
 
 50 
 
 Apnt'liorouiaT'tina ... 
 
 
 
 
 eggiutoni . 
 
 ... 100; VI, 6 
 
 Brodia ... ... 
 
 58 
 
 johnsoni . 
 
 96 ; VI, 4, 5 
 
 f asciata 
 
 59 
 
 sulcata 
 
 113 
 
 - f uivata 
 
 66 ; III, 7, 8 
 
 Apopappus 
 
 53 
 
 - iiebulosa 
 
 59 
 
 Archseoptilus ... ... - 
 
 74 
 
 nympha 
 
 63 
 
 ingens ... 
 
 ... 74; IV, 9 
 
 petiolata 
 
 59 
 
 lucasi 
 
 76 
 
 pictipennis 
 
 59 
 
 Archseotiphe mantidioides .. 
 
 125 
 
 priscotincta . . . 
 
 59 ; III, 5, 6 ; IV, 1-3 
 
 Archiblattides 
 
 91 
 
 - scudderi 
 
 59 
 
 Archimylacridse 
 
 96 
 
 Brodiidse 
 
 57 
 
 (Archimylacridse) kirk by i . 
 
 118; VIII, 3 
 
 
 
 Archimylacris 
 
 103 
 
 Cacurgus spilopterus 
 
 83 
 
 acadica 
 
 109 
 
 Classification 
 
 11 
 
 atrebatica 
 
 107 
 
 Colour bands 
 
 61 
 
 belgica 
 
 105 
 
 Compsoneura 
 
 53 
 
 hastate 
 
 ... 103; VII, 1 
 
 Cordaites 
 
 11,112 
 
 incisa ... 
 
 ... 107; VII, 3 
 
 Corydalis brongniarti 
 
 45,46 
 
 obovata 
 
 ... 109; VII, 4 
 
 Coselia 
 
 81 
 
 priuglei 
 
 ... 146; X, 3 
 
 palmiformis . . . 
 
 81; V, 3 
 
 spectabilis 
 
 105 
 
 Coseliidse 
 
 81 
 
 venusta 
 
 105 
 
 Cryptovenia ... 
 
 
 woodwardi 
 
 ... 106; VII, 2 
 
 moyseyi 
 
 35; II, 1 
 
 spp 
 
 112, 113 ; VII, 5, 6 
 
 Crypto veniidse 
 
 
 Athymodictya 
 
 34 
 
 
 
 parva ... 
 
 35 
 
 
 25 
 
 , . r 
 
 
 higginsii 
 
 25; I, 1 
 
 Becquerelia 
 
 53, 55 
 
 libelluloides 
 
 27 
 
 superba ... 
 
 87 
 
 Dictyoneuridse 
 
 
 Blattidium mantidioides 
 
 125 
 
 
 
 Blattoid wing-fragments 
 
 119, 120; VIII, 5, 6 
 
 
 53 
 
 Blattoidea 
 
 93 
 
 Epitethe 
 
 
 (Blattoidea) exilis 
 
 136 
 
 Etoblattina deanensis 
 
 128, 130 
 
 peachi 
 
 134; IX, 6 
 
 1 in ^t ;i 1 LI 
 
 103 
 
 Boltoniella 
 
 56 
 
 johnsoni 
 
 
 tenuitegminata . 
 
 56 ; III, 4 
 
 mantidioides 
 
 125
 
 INDEX. 
 
 155 
 
 
 PAGE 
 
 PAGE 
 
 Etoblattina peachii ... 
 
 ... 134 Lithomylacris kirkbyi 
 
 118 
 
 woodward! 
 
 106 
 
 pittstonianum 
 
 118 
 
 Eubleptus 
 
 92 
 
 Lithosialis 
 
 45 
 
 
 
 brongniarti 
 
 ... 46; III, 1 
 
 Faunal Association ... 
 
 18 
 
 
 Food of Coal Measure Insects 
 
 -^Q Mecynoptera ... 
 
 ' 36 
 
 Fouquea 
 
 73 
 
 splendida 
 
 39 
 
 cambrensis ... 
 
 72 tuberculata . . . 
 
 ... 37; II, 2 
 
 
 Mecyuopteridse 
 
 36 
 
 Genentomum 
 
 84 
 
 Meganeura ... 
 
 ... 140 
 
 subacutum 
 
 84; V, 4 m y [ 
 
 141 
 
 Gerablattina sulcata 
 
 113 ; radstockensis 
 
 140 
 
 Oerarus 
 
 g7 ! selysii 
 
 142 
 
 G-eroneura ... 
 
 j-g Meganeuridae... 
 
 .. 139 
 
 ovata 
 
 77 V 1 Megaptiloides 
 
 ... 52 
 
 ., - wilsoni 
 
 7g ; brodiei 
 
 52 
 
 Graphiptilus ... 
 
 53 Megasecopteridae 
 
 '... ... 53 
 
 Gryllacris bronguiarti 
 
 45,46 Metryia aualis 
 
 48 
 
 
 i Mixotermes luganensis 
 
 76 
 
 Habits of Fossil Insects 
 
 ^ Mixotermitoidea 
 
 76 
 
 Hadroneura ... 
 
 ^9 Mode of Occurrence ... 
 
 5 
 
 Hemeristia occidentals 
 
 70 Mylacridae 
 
 120 
 
 Hemimylacris 
 
 121 
 
 
 convexa 
 
 123- VIII 8 Necymylaeris 
 
 127 
 
 obtusa ... 
 
 122; VIII, 7 
 
 godoni 
 
 124 
 
 raniificata 
 
 123 
 
 heros ... 
 
 127 
 
 Hermatoblattina kirkbyi 
 
 118 
 
 lacoana 
 
 127 
 
 Homaloneura... 
 
 53 
 
 lafittei 
 
 124 
 
 Hypermegethes 
 
 32 
 
 villeti ... 
 
 124 
 
 
 
 northumbrisa 
 
 32; 1,4 
 
 
 
 schuclierti . . . 
 
 33 
 
 (Edischia 
 
 84 
 
 Hypermegethidse 
 
 31 
 
 (Edischiidaa 
 
 84 
 
 
 
 
 
 Omalia macroptera ... 
 
 83 
 
 Introduction- ... 
 
 1 
 
 Oi'thocosta 
 
 27 
 
 
 
 splendens 
 
 27; 1,2 
 
 Larval Blattoids 
 
 134 
 
 Orthocostidse ... 
 
 27 
 
 Larval Insect Life ... 
 
 7 
 
 Orthomylacris 
 
 133 
 
 Lameereites 
 
 67 
 
 lanceolata 
 
 ... 133; IX, 5 
 
 curvipennis 
 
 65 
 
 
 
 3jtHllpyOT)tlliclj 
 
 53 56 Palseodictyoptera 
 
 24 
 
 grandeuryi 
 
 57 Palseodictyopteron ... 
 
 25 
 
 stirrupi 
 
 57 
 
 higginsii 
 
 25 
 
 tenuitegniinata 
 
 56 
 
 Palaeomantis 
 
 40 
 
 Lamproptilidae 
 
 56 
 
 macroptera 
 
 ... 40; 11,3 
 
 Leptoblattina exilis 
 
 136 ; IX, 7, 8 
 
 Palaeoptilus 
 
 ... 53 
 
 Lithomautidse 
 
 42 
 
 Paralogus .*.. 
 
 144 
 
 Lithomantis ... 
 
 42 
 
 Parelthoblatta 
 
 105 
 
 cai-bonarius 
 
 ... 43; 11,4 
 
 belgica 
 
 105
 
 156 
 
 INDEX. 
 
 Phvloblatta 
 
 PAGE 
 
 113 
 
 Schizoblatta alutacea 
 
 PAGE 
 
 111 
 
 sulcata ... 
 transversalis 
 spp. 
 
 113 ; VIII, 1 
 115; VJII, 2 
 119; VIII, 4 
 
 obovata ... 
 Schuchertiella 
 Sherborniella ... 
 
 109 
 87 
 27 
 
 Phylouiylacris 
 
 124 
 
 higginsii 
 
 25 
 
 mantidioides 
 
 ... 125; IX, 1 
 
 Sooinylacris ... 
 
 128 
 
 Plesioidischia 
 
 
 92 
 
 burri 
 
 ... 131; IX, 4 
 
 sp. 
 
 ... 92; VI, 3 
 
 deanensis 
 
 ... 128; IX, 2 
 
 Polycreagra elegans ... 
 
 
 27 
 
 lieviuensis 
 
 133 
 
 Progenentomum carbonis 
 
 ... 
 
 93 
 
 stock! 
 
 ... 130; IX, 3 
 
 Protepliemeridse 
 
 
 53 
 
 Spilaptera 
 
 53 
 
 Protoblattoidea 
 
 
 90 
 
 libelluloides 
 
 55 
 
 Protodonata ... 
 
 139 
 
 packardi ... 
 
 55 
 
 Protogryllacris brongniarti 
 
 
 46 
 
 sutcliffei ... 
 
 ... 54 ; III, 3 
 
 Protophasma dumasii 
 
 
 80 
 
 venusta ... 
 
 55 
 
 Protorthoptera 
 
 
 78 
 
 Spilapteridse ... 
 
 53 
 
 Pruvostia 
 
 
 48 
 
 Spiloblattina ... 
 
 112 
 
 spectabilis 
 
 ... 48; III 
 
 ( 2 
 
 Spiloptvlus ... 
 
 53 
 
 Pseud of ouquea 
 
 
 72 
 
 Spirorbis pusillus 
 
 11. 112 
 
 cambrensis . . . 
 
 ... 72; IV 
 
 ,8 
 
 Stenodictya lobata 
 
 37, 54 
 
 Ptenodera 
 
 
 90 
 
 Stobbsia 
 
 50 
 
 dubius 
 
 ... 91; VI 
 
 ,2 
 
 woodwardiana 
 
 ... ... 50 
 
 Pteronepionites 
 
 ... 
 
 67 
 
 
 
 ambigua 
 
 ... 68; IV 
 
 ,5 
 
 
 
 johnsoni 
 
 ... 68; IV 
 
 . 4 
 
 Tillvardia 
 
 144 
 
 lepus 
 
 ... 69; IV, 6 
 
 multiplicata 
 
 ... 145; X, 2 
 
 Pteronidia 
 
 
 30 
 
 Titanodictya 
 
 42 
 
 plicatula ... 
 
 30; I 
 
 , 3 
 
 jucunda 
 
 42 
 
 Pteronidiidae 
 
 
 29 
 
 
 
 Scalaeoptera ... 
 
 
 88 
 
 Xeroptera 
 
 85 
 
 recta 
 
 ... 88; VI 
 
 , 1 
 
 obtusata ... 
 
 85 ; V, 5 
 
 ADLARD AND SON AND WEST NEWMAN, LTD., 1MPB., LONDON AND DORKING.
 
 PLATE I. 
 
 FIG. PAGE. 
 
 la. Dictyoneura higginsii (Handlirsch) ; basal portion of left wing, x 1 J. 
 Middle Coal Measures (horizon not known) ; Ravenhead Railway 
 Cutting, nr. St. Helens, Lancashire. Liverpool Museum. 25. 
 
 Ib. Ditto; impression of the wing. X 1J- 25. 
 
 2rt. Orthocosta splendens, Bolton; left wing. Natural size. Middle Coal 
 Measures (below the Top Hard Coal) ; Shipley Manor Claypit, 
 Ilkeston, Derbyshire. Mus. Pract. Geol. (Moysey Coll.), no. 30222. 27. 
 
 2b. Ditto ; impression of the wing. Natural size. Mus. Pract. Geol. (Moysey 
 
 Coll.), no. 30223. 27. 
 
 3ft. Pteronidia plicatula, Bolton; inner half of left wing, x 1^. The 
 straight inner margin and the plication are well shown. Middle 
 Coal Measures (below the Top Hard Coal) ; Shipley Manor Claypit, 
 Ilkeston, Derbyshire. Mus. Pract. Geol. (Moysey Coll.), no. 30224. 30. 
 
 36. Ditto; impression of portion of the wing, showing the oblique cross- 
 
 nervures. x 1 J. Mus. Pract. Geol. (Moysey Coll.), no. 30225. 30. 
 
 4a. Hypermegethes nortkumbrix, Bolton ; basal half of a left wing. X 1^. 
 Coal Measures (shale above the Crow Coal) ; Phoenix Brickworks, 
 Crawcrook, Durham. British Museum, no. In. 18524. 32. 
 
 46. Ditto ; counterpart of portion of basal half of the wing, the interstitial 
 neuration having been rendered visible by immersion of the nodule 
 in water during photography. X 1^. 32.
 
 PAL/EONTOGRAPHICAL SOCIETYJ9I9 
 
 Bolton, Insects of Coal Measures. 
 
 PLATE I. 
 
 l.DICTYONEURA. 2 .ORTHOCOSTA. 3.PTEROMDIA. 
 4.HYPERMEGETHES. 
 
 J.W. Tutcher, photo. London Stereoscopic Co. ir
 
 PLATE II. 
 
 FIG. PAGE. 
 
 1 a. Cryptovenia moyseyi, Bolton ; greater part of left wing, dorsal surface 
 
 uppermost. X 3-J. Middle Coal Measures (below Top Hard Coal) ; 
 Shipley Manor Clay pit, Ilkeston, Derbyshire. Mus. Pract. Geol. 
 (Moysey Coll.), no. 30226. 35. 
 
 16. Ditto; impression of dorsal surface of left wing showing interstitial 
 
 neuration. X 3|. 35. 
 
 2 a. Mecynoptera tubercidata, sp. nov. ; portions of two fore-wings and cubito- 
 
 anal portion of hind-wing superposed, x 1^. Middle Coal Measures 
 (shales above the Royley or Arley Mine) ; Sparth Bottoms, Rochdale, 
 Lancashire. British Museum, no. In. 18576. 37. 
 
 2&. Ditto; impression of the two fore-wings showing interstitial neuration. 
 
 x li. 37. 
 
 3 a. Palxomantis macroptera, Bolton ; fragment of left wing, dorsal aspect, 
 
 the underside of the right wing showing beneath. Natural size. 
 Middle Coal Measures ; Eavenhead railway cutting near St. Helens, 
 Lancashire. Liverpool Museum. 40. 
 
 3&. Ditto ; impression of greater part of dorsal surface of left wing. Natural 
 
 size. 40. 
 
 4. Lithomantis carbonarius, Woodward ; portions of fore- and hind-wing, 
 the prothorax with domed lateral lobes, and the anterior styliform 
 process. (The latter ends in a sharp point not shown in the photo- 
 graph. A little of the mesothorax is seen between the bases of the 
 fore-wings.) Natural size. Coal Measures ; Scotland. British 
 Museum, no. I. 8118. 43.
 
 PAL/EONTOGRAPH ICAL SOCIETY, 1919. 
 
 Bolton . Insects of Coal Measures. 
 
 PLATE H. 
 
 l.CRYPTOVENIA. 2. MECYNOPTERA. 3. PAL.EOMANTIS. 
 4.LITHOMANTIS.
 
 PLATE III. 
 
 Fia. PAGE. 
 
 la. I/ithosialis brongniarti (Mantell) ; left fore-wing. Natural size. Coal 
 Measures; near Coalbrookdale, Shropshire. British Museum 
 (Mantell Coll. dim Parkinson Coll.), no. 11619. 46. 
 
 Ib. Ditto ; impression of greater part of left fore-wing. Natural size. 46. 
 
 2. Pruvostia spectabilis, gen. et sp. nov. ; left fore-wing, x 1J. Middle 
 Coal Measures (ironstone nodules in the binds between the 
 " Brooch " and " Thick " coals) ; Coseley, near Dudley, Stafford- 
 shire. British Museum (Johnson Coll.), no. I. 15894. 48. 
 
 2b. Ditto; impression of same. X ]|. 48. 
 
 3. Spilaptera sutcliffei, Bolton ; basal part of left wing. X 1J. Middle 
 
 Coal Measures (shales above the Royley or Arley Mine) ; Sparth 
 Bottoms, Rochdale, Lancashire. Manchester Museum, no. L. 8197. 54. 
 
 4. Boltoniella temiitegminata (Bolton) ; right hind-wing. X 2. Coal 
 
 Measures (No. 2 Rhondda Seam, base of Pennant Series) ; If miles 
 north-east of Resolven Station, Glamorganshire. Mus. Pract. Geol., 
 no. 24509. 56. 
 
 5. Brodia priscotincta, Scudder; almost complete left wing, showing the 
 
 attenuated base. X 1^. Middle Coal Measures (clay ironstone 
 nodule from binds between the " Brooch " and " Thick " Coals) ; 
 Dudley, Staffordshire. British Museum, no. I. 2961. 59. 
 
 6. Ditto; portion of right hind-wing, under surface uppermost. (The 
 
 original furrows of the wing have been flattened out, the veins more 
 widely separated.) x If. Middle Coal Measures (clay ironstone 
 nodule from binds between "Brooch" and "Thick" Coals) ; Coseley, 
 Staffordshire. British Museum, no. In. 18431. 59 
 
 7. Brodia furcata, Handlirsch ; wing lacking about one-sixth of the apical 
 
 end. X 2. Middle Coal Measures (above the " Brooch " Coal) ; 
 Dudley, Staffordshire. British Museum, no. I. 2962. 66. 
 
 8. Ditto ; impression of same, showing the narrow base of wing. X 1 k- 66.
 
 PAL/EONTOGRAPHICAL SOCIETY, 1919. 
 
 Bolton. Insects of Coal Measures. 
 
 PLATE 
 
 W. Tutcher , photo 
 
 l.LITHOSIALIS. 2.PRUVOSTIA. 3. SPILAPTERA. 
 4.BOLTONIELLA. 5-8.BRODIA. 
 
 London Stereoscopic Co. imp.
 
 PLATE IV. 
 
 FIG. PAGE. 
 
 1. Brodia, priscotincta, Scudder (juvenis) ; wing. X 1^. Middle Coal 
 
 Measures (binds between the " Brooch " and " Thick " Coals) ; 
 Coseley, Staffordshire. British Museum, no. I. 1563. 63. 
 
 2. Ditto; slightly older wing with a broad base. X 3. Middle Coal 
 
 Measures (binds between " Brooch " and " Thick " Coals) ; Coseley, 
 Staffordshire. British Museum, no. I. 1564. 63. 
 
 3. Brodia priscotincta, Scudder ; immature wing. X 1^. Middle Coal 
 
 Measures (binds between " Brooch " and " Thick " Coals) ; Coseley, 
 Staffordshire. British Museum (Madeley Coll.), no. I. 2966. 59. 
 
 4a. " Pteronepionites " johnsoni, sp. nov. ; very immature wing. x 2. 
 Middle Coal Measures (binds between " Brooch " and " Thick " 
 Coals) ; Coseley. Staffordshire. British Museum (Madeley Coll.), no. 
 I. 2967. 68. 
 
 46. Ditto ; impression of same showing undeveloped apex of wing. X 2. 68. 
 
 5. " Pteronepionites " ambigua, sp. nov. ; crushed larval insect Avith partially 
 
 developed wings, x 4. Middle Coal Measures (binds between 
 "Brooch" and "Thick" Coals); Coseley, Staffordshire. British 
 Museum (Madeley Coll.), no. I. 2968. 68. 
 
 6. " Pteronepionites " lepus, sp. nov. ; remains of larval insect with immature 
 
 wings and long segmented abdomen, the photograph accidentally 
 placed upside down. X 2. (Obscure traces of what may have been 
 pleural spines can be distinguished.) Middle Coal Measures (binds 
 between "Brooch" and "Thick" Coals); Coseley, Staffordshire. 
 British Museum (Madeley Coll.), no. I. 2969. 69. 
 
 7. JEnigmatodes (?) regularis, sp. nov. ; portion of inner half of wing, x 2. 
 
 Middle Coal Measures (over the Barnsley Thick Coal) ; Monckton 
 Main Colliery, Barnsley, Yorkshire. British Museum, no. In. 18604. 71. 
 
 8 a. Pseudofouquea cambrensis (Allen) ; greater part of left wing. x 2. 
 
 Lower Coal Measures (top of the Four Foot Seam) ; Llanbradach 
 Colliery. Mus. Pract. Geol., no. 7272. 72. 
 
 8b. Ditto; impression of missing apex of same wing. X 2. 72. 
 
 9 a. Arckxoptilus ingens, Scudder; basal fifth of (? right) wing. Natural size. 
 
 Middle Upper Coal Measures ; between Shelton and Clay Lane, near 
 Chesterfield, Derbyshire. British Museum, no. I. 3997. 74. 
 
 96. Ditto; impression of same. X 1J. British Museum, no. I. 3997. 74.
 
 Bolt on. Insects of Coal Measures. 
 
 PLATE IV. 
 
 Life 
 
 3*2 
 
 ': 
 
 mW 
 
 1-3.BRODIA. 4-6.PTERONEPIONITES. 7. ^ENIGMATODES. 
 8.PSEUDOFOUQUEA. 9. ARCH^EOPTILUS.
 
 PLATE V. 
 
 FIG. PAGE. 
 
 ] . Geroneura (?) ovata, sp. nov. ; impression of upper surface of apex of 
 left wing. X 1^. Middle Coal Measures (binds between " Brooch " 
 and " Thick " Coals) ; British Museum (Madeley Coll.), no. I. 2965. 77. 
 
 2 a. jffldoeophasma anglica, Scudder ; almost complete left wing, x 1. 
 Middle Coal Measures ; Ravenhead Railway Cutting, near St. Helens, 
 Lancashire. Liverpool Museum. 78. 
 
 2 b. Ditto; impression of same. X 1. 78. 
 Sa. Coselia palm if or mis, gen. et sp. nov.; basal half of left wing, x 1^. 
 
 Middle Coal Measures (binds between "Brooch" and "Thick" 
 Coals) ; Coseley, Staffordshire. British Museum (Johnson Coll.), 
 no. I. 15893. 81. 
 
 3 b. Ditto ; impression of same showing the intercostal area, x 1J. 81. 
 
 4 a. Genentomum (?) subaciUum, Bolton ; apical parts of two wings. X 3 J. 
 
 Lower Coal Measures (637 feet below the Bedminster Great Vein) ; 
 South Liberty Colliery, Bristol, Somerset. Bristol Museum, no. 
 C. 972. 84. 
 
 46. Ditto; impression of same. X 3J. 84. 
 
 5 a. Xeroptera obtusata, gen. et sp. nov. ; greater part of right fore- wing 
 
 lacking the outer margin, x 2. Middle Coal Measures (binds 
 between the " Brooch " and "Thick" Coals) ; Coseley, Staffordshire. 
 British Museum (Johnson Coll.), no I. 1558. 85. 
 
 5 b. Ditto ; impression of same. X 2. 85.
 
 PAL/EONTOGRAPHICAL SOCIETY, I92O 
 
 Bolton. Insects of Coal Measures. 
 
 PLATE V. 
 
 l. 3 /2 
 
 l.GERONEURA? 2.^1DCEOPHASMA. 3.COSELIA. 
 4.GENENTOMUM. 5.XEROPTERA. 
 
 xJ.W. Tu.tch.er, photo. London Stereoscopic Co. imp.
 
 PLATE VI. 
 
 FIG. PAGE. 
 
 la. Scalseoptera recta, sp. nov. ; basal half of left wing. X 2. Middle Coal 
 
 Measures (binds between the "Brooch" and "Thick" Coals); 
 
 Coseley, nr. Dudley, Staffordshire. British Museum, no. I. 13878. 88. 
 1 b. Ditto ; impression of the same wing-fragment. X 2. 88. 
 
 2. Pteiwdera dubins, sp. nov. ; distal half of left wing, x 3. Middle Coal 
 
 Measures (binds between the " Brooch " and " Thick " Coals) ; 
 
 Coseley, nr. Dudley, Staffordshire. British Museum (Johnson Coll.), 
 
 no. I. 1559. 91. 
 
 3. Plesioidisckia, sp. ; fragment of right wing. X 1^. Middle Coal 
 
 Measures (binds between the " Brooch " and " Thick " Coals) ; 
 
 Tipton, Staffordshire. Manchester Museum, no. L. 4905. 92. 
 
 3 b. Portion of costal border of same wing borne on opposite half of nodule. 
 
 X H. 92. 
 
 4. Aphthoroblattina johnsoni (Woodward); type specimen, x 1 1. Middle 
 
 Coal Measures (binds between the " Brooch " and " Thick " Coals) ; 
 
 Coseley, Staffordshire. British Museum (Johnson Coll.), no. I. 1067. 96. 
 
 5 a. Ditto, x 2. Middle Coal Measures (binds between the " Brooch " 
 
 and " Thick " Coals) ; Coseley, Staffordshire. Mr. W. Egginton's 
 Collection, no. 2. 99. 
 
 . 5 b. Ditto ; impression, x 2. 99. 
 
 6 a. Aphthoroblattina engintoni,[sp. nov.; pronotum, portions of tegmina, and 
 
 the distal portion of left hind-wing showing beyond broken edge of 
 tegmen. x 2. Middle Coal Measures (binds between " Brooch " 
 and " Thick " Coals) ; Coseley, Staffordshire. Mr. W. Egginton's 100. 
 Collection, no. 1. 
 66. Ditto; counterpart, x 2. 100.
 
 PAL/EONTOGRAPHICAL SOCIETY, I92O 
 
 Bolton. Insects of Coal Measures. 
 
 PLATE VI. 
 
 IcA 
 
 l.SCAL^OPTERA. 2. PTENODERA. 3. PLESIOIDISCHIA. 
 4-6. APHTHOROBLATTINA.
 
 PLATE VII. 
 
 FIG. PAGE. 
 
 1 a. Arcliimylacrls hastata, Bolton ; greater part of left fore-wing. X 2J. 
 Base of Upper Coal Measures (Gellicleg Level of Mynyddislwyn 
 Vein) ; near Maes-y-cwmmer, Monmouthshire. Mus. Pract. Geol., 
 no. 24501. 103. 
 
 1 b. Ditto ; counterpart of left wing, x 2f . Mus. Pract. Geol., no. 24502. 103. 
 
 2 Archimylacris ivoodivardi, Bolton; left fore-wing. X 3J. Base of 
 
 Pennant Series (10 feet shale overlying the No. 2 Rhondda seam) ; 
 Clydach Vale, South Wales. Mr. D. Davies' Coll. 106. 
 
 3 a. Arcliimylacris incisa, sp. nov. ; impression of the under surface of the 
 
 pronotum, and of the remnants of the fore-wings, x 3. Middle 
 Coal Measures (binds between the " Brooch " and " Thick " Coals) ; 
 Coseley, nr. Dudley, Staffordshire. British Museum (Johnson 
 Coll.), no. I. 15900. 107. 
 
 36. Ditto; counterpart. X 3. 107. 
 
 4 a. Archimylaerut obovata, Bolton ; portion of left fore-wing, x 3. Upper 
 
 Coal Measures (Gwernau Level of the Mynyddislwyn Vein) ; Maes-y- 
 cwmmer, Monmouthshire. Mus. Pract. Geol., no. 24506. 109. 
 
 4 b. Ditto ; impression less incomplete than the broken wing, x 3. Mus. 
 
 Pract, Geol., no. 24507. 109. 
 
 5. Arehimylaeris, sp. indet. ; fragmentary fore- wing, showing the middle 
 
 third only, the rest being concealed under a leaf of Cordaites, the 
 latter bearing pits in which Spirorbis pusillus (Martin) had 
 established themselves. Much enlarged. Base of Upper Coal 
 Measures (Gellideg Level of Mynyddislwyn Vein) ; near Maes-y- 
 cwmmer, Monmouthshire. Mus. Pract. Geol., no. 24503. ] 12. 
 
 6. Arcliimylacris, sp. indet. ; impression of the basal portion of a left 
 
 fore-wing, and of a broken pronotum. Much enlarged. Base of 
 Upper Coal Measures (Gellideg Level of Mynyddislwyn Vein) ; near 
 Maes-y-cwmmer, Monmouthshire. Mus. Pract. Geol., no. 24508. 113.
 
 PAL/EONTOGRAPHICAL SOCIETY, I92O. 
 
 Bolton . Insects of Coal Measures. 
 
 PLATE Vll. 
 
 1-6. ARCHIMYLACRIS. 
 
 J.W. Tut cher, photo. 
 
 London Stereoscopic Co. imp.
 
 PLATE VIII. 
 
 FIG. PAGE. 
 
 1 . Phyloblatta sulcata (Bolton) ; greater part of a right fore- wing, x 3. 
 
 Upper Coal Measures (Gwernau Level of the Mynyddislwyn Vein) ; 
 
 near Maes-y-cwmmer, Monmouthshire. Mus.Pract. Geol., no. 24504. 113. 
 1 />. Ditto ; counterpart. X 3. 113. 
 
 2. Phyloblatta transversalis, Bolton ; remains of two Blattoids, consisting 
 
 of the tegmina, pronota, and portions of the hind-wings. Natural 
 size. Coal Measures; Staffordshire. Geological Museum, University, 
 Birmingham. 115. 
 
 3. (Archimylacrida?) kirkbyi (Woodward) ; reproduction of Woodward's 
 
 figure of a single left wing. X 5. Upper Coal Measures (bed 
 no. 33) ; Meithil, coast of Fifeshire. Present whereabouts of 
 specimen unknown ; formerly in the possession of Mr. James 
 Kirkby. 118. 
 
 4 a. Phyloblatta (?) sp. ; portion of Blattoid wing, x 4. Coal Measures 
 
 (shales at a depth of 1967 feet) ; Maydensole boring, Kent. 
 Museum of the Kent Coals Concession Co., Dover. 119. 
 
 4>b. Ditto; impression. X 4^. 119, 
 
 5 a. Phyloblatta (?) sp. ; portion of Blattoid wing. X 4^. Coal Measures 
 
 (at depth of 2180 feet) ; Barfreston boring, Kent. Museum of the 
 Kent Coal Concessions Co., Dover. ] 19. 
 
 5fc. Ditto; outline of wing-fragment, x 4J. 119. 
 
 6. Blattoid wing-fragment. X 4. Found in a core from a depth of 
 
 2424 feet at the Stonehall boring, Kent. Museum of the Kent Coal 
 Concessions Co., Dover. 120. 
 
 7. Hemimylacris obtusa, Bolton ; right fore-wing, x 2. Base of Upper 
 
 Coal Measures (Four-foot Seam of Swansea) ; Gladys Colliery, one 
 mile E.S.E. of Penller-gaer Church, Glamorganshire. Mus. Pract. 
 Geol., no. 24510. 122. 
 
 8. Hemimylacris convexa, Bolton ; basal half of a fore-wing, x 3J. Coal 
 
 Measures (shales associated with the Graigola Seam, Pennant 
 Series) ; Clydach Merthyr Colliery, Clydach Valley, Swansea Vale, 
 Glamorganshire. ] 23.
 
 PAL-CONTOGRAPHICAL SOCIETY, I92O. 
 
 Bolton. Insects of Coal Measures. 
 
 PLATE Vlll. 
 
 2.M 
 
 1.2.4^.5. PHYLOBLATTA. 3. ARCHIMYLACRIDAE . 
 6.BLATTOID WING FRAGMENT. 7. 8. HEMIMYLACRIS. 
 
 J.W.TutcKer, photo. London Stereoscopic Go.il
 
 PLATE IX. 
 
 PIG. PAGE. 
 
 1 . Phylomylacris mantidioides (Goldenberg) ; basal portion of a left fore- 
 
 wing, x 4J. Upper Coal Measures (zone of Anthracomya 
 phillipsii); South Hylton, opposite Claxheugh on the Wear, Durham. 
 Hancock Museum, Newcastle-on-Tyne (Kirkby Coll.). 125. 
 
 2. Soomylacris deanensis (Scudder) ; a right fore-wing, showing the under 
 
 surface. X 1J. Found by Mr. F. Stock in Coal Measures at 
 Foxe's Bridge, Forest of Dean, Gloucestershire. U.S. National 
 Museum, Washington, D.C. (Lacoe Coll., no. H. 2132 6; Nat. Mus., 
 no. 38090). 128. 
 
 3. Soomylacris stocki, sp. nov. ; fragmentary remains of a fore-wing, partly 
 
 overlaid by the remains of a second. x 2J. Found by Mr. F. 
 Stock in Coal Measures at Crump Meadow, Forest of Dean, 
 Gloucestershire. U.S. National Museum, Washington, D.C. (Lacoe 
 Coll., no. H. 2132 c; Nat. Mus., no. 38090). 130. 
 
 4. Soomylacris bum, Bolton ; almost complete fore-wing. X 4|-. Coal 
 
 Measures (in dark shale from a depth of 1208 feet) ; Barfreston 
 Boring, Kent. Museum of the Kent Coal Concessions Co., Dover. 131. 
 
 5. Orthomylacris lanceolata, Bolton ; left fore- wing, with the base missing. 
 
 X 2^. Coal measures (shales associated with the Graigola Seam, 
 Pennant Series) ; Clydach Merthyr Colliery, Clydach Valley, 
 Swansea Vale, Glamorganshire. Mus. Pract. Geol., no. 24511. 133. 
 
 6. (Blattoidea) peachi (Woodward) ; complete upper surface of insect, 
 
 showing the head, pronotum, larval wings, and the segments of the 
 abdomen. X 2. Coal Measures (grey sandy shale with nodules 
 of impure clay and ironstone, at 91 feet 6 inches below the surface); 
 Greenhill Pit, Kilmarnock. Kilmarnock Museum. 1 34. 
 
 7. Leptoblattina exilis, Woodward ; co-type of almost complete insect, seen 
 
 from the upper surface. X 2. Middle Coal Measures (binds 
 between the " Brooch " and " Thick " Coals) ; Coseley, Stafford- 
 shire. British Museum (Johnson Coll.), no. 1065. 136. 
 
 8. Ditto; a second co-type, with the enlarged epirnera well seen on the 
 
 left side of the abdomen. X 2. British Museum (Johnson Coll.), 
 
 no. 1066. 136.
 
 PALyEONTOGRAPHICAL SOCIETY, I92O . 
 
 Bolton . Insects of Coal Measures. 
 
 PLATE IX. 
 
 7.Vi 
 
 1. PHYLOMYLACRIS. 2-4. S OOMYLACRIS . 5. ORTHOMYLACRI S. 
 6. ( BLATTOIDEA.) 7. 8. LEPTOBLATTINA.
 
 PLATE X. 
 
 FIG. PAGE. 
 
 la. Boltonites radstockensis (Bolton) ; basal portion of the wing, x 2. 
 Upper Coal Measures ; Tyning Colliery, Radstock, Somersetshire. 
 Sedgwick Museum, Cambridge. 140. 
 
 16. Ditto ; proximal portion of costa and subcosta, showing the tubercular 
 anterior edge, and the backward slipping of the costa over the sub- 
 costa. X 4. 140. 
 
 Ic. Ditto; distal portion of the costa and subcosta, showing the spinous 
 anterior border of the former, the cross-branches uniting the two 
 veins, and a median line of tubercular ornament on the subcosta. 
 X 4. 140. 
 
 Id. Ditto; proximal portion of the inner margin, showing the development 
 
 of stout submarginal spines, x 4. 140. 
 
 1 e. Ditto ; more distal portion of the inner margin, showing the submarginal 
 
 spines, x 4. , 140. 
 
 2. TUlyardia multiplicata, gen. et sp. nov. ; impression of the under 
 
 surface of right wing. X 1J. Upper Coal Measures; Barony Pit, 
 Auchinleck, Ayrshire. Mus. Geol. Surv. Scot., no. T. 4098 b. 145. 
 
 3. Archimylacris pringlei, Bolton ; basal two-thirds of fore-wing. X 4. 
 
 Upper Coal Measures (Keele Group) ; Slang Lane, "Wellington, 
 Shropshire. Mus. Pract. Geol., no. 30725. 146.
 
 PALXEONTOGRAPHICAL SOCIETY, I92O 
 
 ol ton, Insects of Coal Measures. 
 
 PLATE X. 
 
 la. 
 
 l.a,e.BOLTONITES. 2. TILLYARDIA. 
 3. ARCHIMYLACRIS. 
 
 -o'ho>.n
 
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