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PALHONTOGRAPHICAL SOCIETY. 


VOLUME XXIII. 


CONTAINING 


SUPPLEMENT TO THE FOSSIL CORALS. Part II.—No. 2 (Creracrous). By Dr. Duncan. Six 
Plates. 

THE CRETACEOUS ECHINODERMATA. Vol. I—Part III. By Dr. Wricur. Ten Plates. 

THE BELEMNITIDA. Part V (Oxrorp Cray, &c., Betemnites). By Prof. Puitirrs. Nine Plates. 

THE FISHES OF THE OLD RED SANDSTONE. Part I (concluded). By Messrs. J. Powrre and FE. 
Ray Lanxester. Nine Plates. 

THE REPTILIA OF THE LIASSIC FORMATIONS. Part Il. By Prof. Own. Four Plates. 

THE CRAG CETACEA. No.1 (Zirntus). By Prof. Owzn. Five Plates. 


ISSUED FOR 1869. 


JANUARY, 1870. 


Council and Officers elected 9th April, 1869. 


resident, 
J. S. BOWERBANK, LL.D., F.R.S., GS. 


Vite-residents. 


T. Davrpson, Esa., F.R.S., G.S. 
Str R. I. Murcutson, Bart., F.R.S., G.S. 


Pror. Owen, M.D., F.R.S., G.S. 
Pror. Puriuuips, F.R.S., G.S. 


Council. 
Pror. AnstepD, M.A., F.R.S., G.S. J. Gwyn Jerrreys, Esa., F.R.S., G.S. 
Anronto Brapy, Esq., F.G.S. H. Les, Esa, F.L.S., G.S. 
W. Boyp Dawkins, Esq., F.R.S., G.S. W. H. Letenton, Esa., F.G.S. 
P. M. Duncan, M.B., F.R.S., Sec. G.S. J. PrcknrinG, Bsq. 
R. Erueripes, Hsq,, F.G.S. J. Prestwicu, Ese., F.R.S., G.S. 
J. W. Frower, Esq., F.G.S. Pror, TENNANT, F.G.S., Z.S. 
Pror. Huxtry, LL.D., F.R.S., G.S. C. Tyter, Esq., F.L.S., G.S. 
J. W. Itortt, Esq. H. Woopwarp, Esq,, F.G.S., Z.8. 
Trensurer. 


Srartes Woop, Esa., F.G.S., Brentwood, Essex. 


Honorary Secretary, 
Rev. T. Wittsuire, M.A., F.G.S., 13, Granville Park, Lewisham, Kent, S.E. 


Hocul Secretaries. 


Alton—Wmo. Curtis, Jun., Esa. 
Bath—Wnmn. Watton, Esq. 
Berlin—Mussrs. FRIEDLANDER & Son. 
Birkenhead—Wm. Frrouson, Esa. 
Birmingham—W. Matuews, Jun., Esq., F.G.S. 
Blackburn—Wm. Harrison, Esq,, F.G.S. 
Bristol—W. SanveErs, Esq., F.R.S., G.S. 
Cambridge—Jamus Carter, Esa. 
Cheltenham—T. Wricut, M.D., F.G.S. 
Cirencester—J. BravuenvER, Hsa., F.G.S. 
Colchester—C. R. Bren, M.D. 


R.G.S. 
Devizes—Wm. CuUNNINGTON, Esq., F.GS. 
Dublin—W. HE. Stnets, M.D., F.R.D.S. 
Edinburgk—Pror. Batrour, M.D., F.R.S8., LS. 


Guildford—R. A. C. Gopwin-AustEN, Esq., F.R.S., 
G.S. 

Kendal—Tuomas Govueu, Esq. 

Leeds—Tuos. NuNNELEY, Esa. 

Leicester —Jamus Puant, Esa. 

Liverpool—G. H. Morton, Esq., F.G.S. 

New York.—B. Watnrnouse Hawkins, Esq., 
F.L.S., GS. 

Norfolk—Rrv. J. Gunn, M.A., F.GS. 

Oxford—Pror. J. Pures, F.R.S., GS. 


| Paris—M. F. Savy, 24, Rue Hautefeuiile. 
Deddington—Cuas. FauLKner, Esq., F.S.A., G.S., | 


Plymouth—J. H. Fuce, Esq. 

Richmond, Yorkshire—EDWarpd Woop, Esq, F.G.S. 
Scarborough—Joun LecKENBY, Esq., F.G.S. 
Tonbridge Wells—J. Suarv, Esq., F.G.S. 
Torquay—Wmn. PrncE._ey, Esa., F.R.S., G.S. 


Wolverhampton—Hy. Bucxert, Bsq., F.G.S. 


LIST OF MEMBERS, 


JANUARY, 1870. 


Her Most Gracious Masrsty THE QUEEN. 


Adams, William, Esq., F.G.S., Cardiff, Glamorganshire. 

Adlard, J. E., Esg., Bartholomew Close, E.C. 

Albert Memorial Museum, Queen Street, Exeter. 

Aitken, John, Esq., Lane End, Bacup, Manchester. 

Allport, S., Esq., Snow Hall, Birmingham. . 

Angelin, Professor, Stockholm. 

Ansted, Professor D. T., M.A., F.R.S., G.S., &c., Atheneum Club; and 33, Brunswick 
Square, W.C. 

Arbuthnot, Capt. W., 25, Hyde Park Gardens, W. 

Asher and Co., Messrs., 18, Bedford Street, Covent Garden, W.C. 

Atheneum Library, Liverpool. 

Auerbach, Professor, Moscow. 

Austen, Rev. J. H., M.A., Ensbury House, Wimborne. 

Austen, Miss Helena E., 19, Henrietta Street, Bath. 

Aveline, W. T., Esq., F.G.S., Museum of Practical Geology, Jermyn Street, S.W. 


Backhouse, Edward, Esq., Ashburne House, Sunderland. 

Bair, M., Frankfort. 

Balfour, Professor, M.D., F.R.S., L.S., &c., Local Secretary, 27, Inverleith Row, Edinburgh. 
Balme, E. B. Wheatley, Esq., Loughrigg, Ambleside. 

Barclay, Joseph G., Esq., 54, Lombard Street, E.C. 

Barlow, H. C., M.D., F.G.S., 11, Church Yard Row, Newington Butts, Surrey, 5. 
Barr, W. R., Esq., Park Mills, Stockport. 

Barthes and Lowell, Messrs., 14, Great Marlborough Street, W. 

Bath Royal Literary and Scientific Institution. 

Bathurst, Rev. W. H., Lydney Park, Gloucester. 

Battersby, R., M.D., Torquay. 

Beaufoy, George, Esq. (Executor of the late), South Lambeth, S. 

Becker, M. Edvald, Breslau, Silesia. 

Beckett, Henry, Esq. F.G.S., Local Secretary, Darlington Street, Wolverhampton. 


* The Members are requested to inform the Secretary of any errors or omissions in this list, and of any delay 1) 
the transmission of the Yearly Volumes. 


Bell, Professor T., F.R.S., L.8., G.S., &c., The Wakes, Selborne, Alton, Hants. 

Benson, Starling, Esq., F.G.S., Swansea. 

Bernard, Ralph M., Esq., 5, Victoria Square, Clifton. 

Bewley, John, Esq., 4, Brown’s Buildings, Liverpool. 

Bigsby, J. J., M.D., F.R.S., G.S., 89, Gloucester Place, Portman Square, W. 

Bilke, Edward, Esq., F.G.S., &c., 1, Chester Square, Pimlico, 8.W. 

Binney, E. W., Esq., F.R.S., G.S., &c., 40, Cross Street, Manchester. 

Birmingham Free Library, Radcliffe Place, Birmingham. 

Birmingham Old Library, Union Street, Birmingham. 

Blackburne, Major John George, F.G.S., Oldham. 

Blackmore, Humphrey P., M.D., Salisbury. 

Bladon, James, Esq., Pontypool. 

Blake, W., Esq., Bridge House, South Petherton. 

Blanford, H. F., Esq., F.G.S., 21, Bouverie Street, Whitefriars, E.C. 

Boase, H. S., M.D., F.R.S., G.S., &c., Claverhouse, near Dundee. 

Bohn, Henry G., Esq., 18, Henrietta Street, Covent Garden, W.C. 

Bonney, Rev. George, F.G.S., St. John’s College, Cambridge. 

Bordeaux, La Faculté des Sciences de. 

Borradaile, Charles, Esq., Hast Hothly, Hurst Green, Sussex. 

Bosquet, Mons. J., For. Cor. G.S., Pharmacien, Maestricht. 

Boston Society of Natural History, Boston, U.S.A. 

Bower, Rev. E., Closworth, near Yeovil. 

Bowerbank, J. 8., LL.D., F.R.S., &c., President, 2, Hast Ascent, St. Leonard’s-on-Sea. 

Bradford Philosophical Society, 31, Manor Row, Bradford. 

Brady, Autonio, Usq., F.G.S., Stratford, Essex; and Admiralty Office, Somerset House, W.C. 

Brady, Sir M., Bart., 26, Upper Pembroke Street, Dublin. 

Braikenridge, Rev. G. W., M.A., F.S.A., G.S., &c., Clevedon, near Bristol. 

Brassey, Thomas, Esq., jun., 56, Lowndes Square, S.W. 

Bravender, John, Esq., F.G.S., Local Secretary, Coxwell Street, Cirencester. 

Bree, C. R., M.D., Local Secretary, Fast Hill, Colchester. 

Briggs, General John, F.R.S., G.S., &c., Oriental Club, Hanover Square, W.; and Bridge 
Lodge, Hurstpierpont. 

Briggs, Miss Ellen, 55, Lincoln’s Inn Fields. 

Brighton and Sussex Natural History Society, 7, Pavilion Parade, Brighton. 

British Museum, Printed Book Department, W.C. 

British Museum, Departmental Mineralogical and Geological Library, W.C. 

Broome, C. E., Esq., M.A., &c., Elmhurst, Batheaston, Bath. 

Brown, Edwin, Esq., F.R.G.S., G.S., Burton-on-Trent. 

Brown, Isaac, Esq., Ackworth, Pontifract. 

Brown, T. C., Esq., Cirencester. 

Browne, Wm. Meredith, Esq., Westminster Fire Office, King Street, Covent Garden, W.C. 

Buckman, Professor James, F.G.S., &c., Bradford Abbas, Sherborne, Dorset. 

Busk, George, Esq., F.R.S., L.S., &c., 32, Harley Street, Cavendish Square, W. 


Cambridge University Library. 

Carpenter, Alfred, M.D., Croydon, S. 

Carpenter, W. B., M.D., F.R.S., &c., 56, Regent’s Park Road, N.W. 
Carter, James, Esq., Local Secretary, 30, Petty Cury, Cambridge. 


Cassini, H. T., Esq., Victoria Cottage, Archway Road, Highgate, N. 
Cavell, Edmund, Esq., F.G.S., Saxmundham. 

Chamberlin, Rev. a ea 

Chambers, Robert, Esq., F.R.S.E., G.S., &e., St. Andrew’s, 
Champernowne, Arthur, Esq., Darlington Hall, Totness, Devonshire. 
Chapman, Thomas, Esq., 14, Cockspur Street, Charing Cross, S.W. 
Cheltenham Permanent Library, 18, Clarence Street, Cheltenham. 
Cherbullier, Mons., Paris. 

Christ’s College, Cambridge, Library of. 

Clabon, J. M., Esq., 21, Great George Street, S.W. 

Clark, Dr. W., F.R.S., &c., Cambridge. 

Clarke, Rev. W. B., F.G.S., &c., St. Leonards, near Sydney, New South Wales. 
Clayton, Rev. J. H., Liphook, Hants. 

Cobbold, Rev. R. H., Rectory, Broseley, Salop. 

Cocchi, Sig. Q., Professor of Geology, llorence. 

Colchester, W., Esq., F.G.S., Grundesburgh Hall, Ipswich. 
Collings, Rev. W. T., M.A., F.L.8., G.S., Lord of Sark, Channel Islands. 
Collingwood, F. J. W., Esq., Glanton Pyke, Glanton, Northumberland. 
Colvin, Col. John, C.B., Leintwardine, near Ludlow. 

Compton, Rev. John, Minesteed Parsonage, Lyndhurst. 

Cooper, Charles J., Esq., Bridgenorth, Salop. 

Cornalia, Prof. Emilio, Milan. 

Cornthwaite, Rev. T. M., M.A., Walthamstow. 

Corporation of London, Library Committee of, Guildhall, E.C. 
Corrie, Adam J., Esq., M.A., Quintella, Torquay. 

Cotteau, Mons., Paris. 

Cranage, Dr. J. E., The Old Hall, Wellington, Shropshire. 

Crickitt, E., Esq., 4, Belvedere Villas, Bath. 

Crisp, F., Esq., 131, Adelaide Road, N.W. 

Cross, Rev. J. E., Appleby Vicarage, Brigg, Lincolnshire. 

Cross, R. A., Esq., Hill Cliff, Warrington. 

Crowley, Alfred, Esq., Bramley Oaks, Croydon, 8S. 

Cubitt, George, Esq., M.P., F.G.S., 17, Princes Gate, W. 

Cull, R., Esq., 18, Tavistock Street, Bedford Square, W.C. 
Cunnington, W., Esq., F.G.S8., Local Secretary, Devizes, Wilts. 
Curtis, W., Esq., Local Secretary, Alton, Hants. 

Currey, Elliot 8., Esq., Erlwood, Bagshot. 

Cust, Lady Elizabeth, 13, Eccleston Square, S.W. 


Darwin, Charles, Esq., M.A., F.R.S., G.S., &c., Down, Bromley, Kent. 

Davidson, Thomas, Esq., F.R.S., G.S., Mem. Geol. Soc. France, &c., Vice-President, 22 
Crescent, Brighton. 

Daw, E., Esq., Mynfield, Amwell, Ware. 


Park 


Dawkins, W. Boyd, Esq., F.R.S., G.S., Birch View, Norman Road, Rusholme, Manchester. 


Dawson, Principal J. W., LL.D., F.R.S., G.S., &c., MeGill’s University, Montreal. 
Day, Rev. Hen. Geo., M.A., Sedbergh, Kendal. 
Deane, H., Esq., F.L.S., Clapham Common, Surrey, S.W. 


De Gray and Ripon, Earl, 1, Belgrave Square, S.W. 

Denison, Col. Sir William, R.E., K.C.B., F.R.S., R.A.S., Observatory, East, Sheen, S.W. 

Deshayes, Mons. G. P., F.M.G.S., Paris. 

Devonshire, Duke of, Chancellor of the University of London, F.R.S., G.S., &c., Devonshire 
House, Piccadilly, W. 

Devon and Exeter Institution, Exeter (by Ed. Parfitt, Esq.) 

Dewilde, G. R., Esqg., 14, Peak Hill Avenue, Sydenham. 

Dickinson, F. H., Esq., King Weston, Somerton. 

Dickinson, W., Esq., Thorncroft, Workington. 

Dickinson, W., Esq., 6, Princes Street, Lothbury, E.C. 

Digby, Lady, Minterne, near Dorchester. 

Dilke, Sir C. Wentworth, Bart., '.G.S., L.S., &c., 76, Sloane Street, S.W. 

Dollfus, Mons., Paris. 

Dorset County Museum Library, Dorchester. 

Douglas, Rev. Robert, Manaton Rectory, Moreton-Hampstead, Exeter. 

Dover Proprietary Library. 

Doyen, Mons. J. M. 

Ducie, the Earl of, F.R.S., G.S., &c., Belgrave Square, S.W. 

Dudley and Midland Geological and Scientific Society and Field-Club. 

Dumortier, Mons. E., Lyons (by Mons. F. Savy, Paris). 

Duncan, P. M., M.B., F.R.S., Sec.G.S., 40, Blessington Road, Lee, S.E. 

Durham, the Dean and Chapter of (by Samuel Rowlandson, Esq., the College, Durham). 

Dyer, Prof. W. T., Royal Agricultural College, Cirencester. 


Eccles, James, Esq., Springwell House, Blackburn. 

Edgell, Rev. E. Wyatt, 2, Lansdowne Terrace, Ladbroke Square, Notting Hill, W. 

Edinburgh Geological Society, 5, St. Andrew Square, Edinburgh. 

Edinburgh Museum of Science and Art, Argyle Square, Edinburgh. 

Edwards, F. E., Esq., F.G.8., 22, Woburn Square, W.C. 

Egerton, Sir Philip de Malpas Grey, Bart., M.P., Trustee Brit. Museum, F.R.S., G.S., &c., 
Oulton Park, Cheshire; and 288, Albemarle Street, W. 

Elhott, John, Esq., Kingsbridge, Devon. 

Elliot, Sir Walter, K.S.I., F.L.8., Wolfelee, Hawick, N.B. 

Enniskillen, Wilham Willoughby, Earl of, D.C.L., F.R.S., G.S., &e., Florence Court, 
Enniskillen; and 65, Eaton Place. 

Eskrigge, R. A., Esq., 3, Batavia Buildings, Liverpool. 

Etheridge, R., Esq., F.G.S., &c., Museum of Practical Geology, Jermyn Street, S.W. 

Evans, John, Esq., F.R.S., Sec.G.S., Nash Mills, Hemel Hempstead. 

Evans, Thomas, M.D., Gloucester. 

Eyton, Thomas C., Esq., F.L.S., G.S., &c., Eyton, near Wellington, Salop. 


Falconer, Thomas, Esq., F.G.S., Usk, Monmouthshire. 

Falkner, Frederick, Esq., Somersetshire Bank, Bath. 

Faulkner, Charles, Esq., F.S.A., G.S., R.G.S., Local Secretary, Museum, Deddington, Oxon. 
Favre, Mons. Alph., Professor of Geology, Academy, Geneva. 


9 


Ferguson, William, Esq., F.L.S., G.S., R.G.S., &e., Local Secretary, Kinmundy, Mintlaw, 
Aberdeenshire; and 2, St. Aidan’s Terrace, Birkenhead. 

Fielding, Edward, Esq., 142, St. Paul’s Road, Camden Square, N.W. 

Fisher, Rev. Osmond, M.A., F.G.S., Harlston Rectory, Cambridge. 

Fletcher, T. W., Esq., M.A., F.R.S., G.S., S.A., Lawneswood House, Stourbridge. 

Flower, J. W., Esq., F.G.S., Park Hill, Croydon, S. 

Forbes, John Edward, F.G.S., 3, Faulkner Street, Manchester. 

Fotherby, H. J., M.D., 40, Trinity Square, Tower, E.C. 

Fox, Rev. W. Darwin, Delamere Rectory, near Chester. 

Fraser, John, M.D., Wolverhampton. 

Friedlander, Messrs., Local Secretaries, Berlin. 

Froggat, John, Esq., Church Gate, Stockport. 

Fryer, A., Esq., Chatteris, Cambridgeshire. 

Fuge, J. H., Esq., F.R.C.S.E., 16, Frankfort Street, Plymouth, 


Galton, Captain Douglas, R.E., F.R.S., G.S., &c., 12, Chester Street, Grosvenor Place, S.W. 

Gardner, J. S., Esq., F.G.S., Park House, St. John’s Wood Park, N.W. 

Gassiot, I. P., Esq., F.R.S., &c., Clapham, S. 

Gatty, Charles, Esq., F.G.S., Felbridge Park, East Grinstead. 

Geological and Polytechnic Society of the West Riding of Yorkshire, Leeds. 

Geological Society of Manchester. 

Geological Survey of Great Britain, Paleontological Department, Jermyn Street, S.W. 

Geologists’ Association, University College, W.C. 

Gibson, G. S., Esq., Saffron Walden. 

Gibson, Thomas F., Esq., F.G.S., &c., Broadwater Down, Tunbridge Wells. 

Gilbertson, Henry, Esq., Hertford. 

Gilchrist, J., M.D., Crichton Royal Institution, Dumfries. 

Glasgow Geological Society, Andersonian University, Glasgow. 

Gloucester Literary Society, Gloucester (by Dr. B. Wagbourn). 

Godlee, Rickman, Esq., Lillies, Upton, Essex. 

Godwin-Austen, R. A. C., Esq., F.R.S., G.S., &e., Local Secretary, Chilworth Manor, Guildford, 
Surrey. 

Gordon, P. L., Esq., Craigmyle. 

Gough, Capt. the Hon. George S., F.G.S., L.S., &c., Lough Cutra Castle, Gort, Galway, Ireland. 

Gough, Thomas, Esq., Local Secretary, Kendal. 

Gould, John, Esq., F.R.S., L.S., Z.S., &c., 26, Charlotte Street, Bedford Square, W.C. 

Gray, John, Esq., Lyttleton Terrace, Hagley, near Stourbridge. 

Greenwell, G. C., Esq., Poynton, Stockport. 

Gregory, J. R., Esq., 15, Russell Street, Covent Garden, W.C. 

Griffith, Sir Richard, Bart., LL.D., F.R.S.E., G.S., &c., 2, Fitzwilliam Place, Dublin. 

Grindrod, R. B., M.D., F.G.S., Townshend House, Great Malvern. 

Grundy, Thomas, Esq., Beatlands, Sidmouth, Devon, 

Guise, Sir W. V., Bart., F.G.S., &c., Elmore Court, near Gloucester. 

Gunn, Rev. J., M.A., Local Secretary, Irstead Rectory, Norwich. 

Guppy, R. Lechmere, Esq., F.G.S., Government House, Trinidad. 


Hall, Hugh F., Esq., F.G.S., 17, Dale Street, Liverpool. 


co) 


10 


Hall, Townshend M., Esq., F.G.S., Pilton Parsonage, Barnstaple. 

Harford, Frederick, Esq., Ocean Marine Insurance Company, 2, Old Broad Street, E.C. 

Harkness, Professor Robert, F.R.S., G.S., Queen’s College, Cork. 

Harris, W., Esq., F.G.S., Charing, Ashford, Kent. 

Harrison, William, Esq., F.G.S., S.A., R.G.S., R.S. Ant., &c., Local Secretary, Samlesbury 
Hall, Preston, Lancashire; and Conservative Club, St. James’s Street, S.W. 

Hartley Institution, Southampton (by Dr. F. T. Bond). 

Haughton, Rev. Professor 8., M.D., F.R.S., G.S., Fellow of Trinity College, Dublin. 

Hawkes, Rev. Henry, B.A., F.L.8., &c., 5, Elm Grove, Southsea, Portsmouth. 

Hawkins, B. Waterhouse, Esq., F.L.8., G.S., Local Secretary, New York. 

Hawkins, Rev. H. S., Forest Lodge, Liphook, Hants. 

Hawkshaw, J. Clarke, Esq., 25, Cornwall Gardens, South Kensington, W. 

Haythornthwaite, William, Esq., Kirkby Lonsdale. 

Heywood, James, Esq., F.R.S., G.S., &c., Athenzeum Club, S.W. 

Hicks, Henry, Esq., St. David’s, Pembrokeshire. 

Higgins, E. T., Esq., 122, King Henry’s Road, Haverstock Hill, N.W. 

Hindson, Isaac, Esq., Kirkby Lonsdale. 

Hirst, John, jun., Esq., Dobcross, Saddleworth, near Manchester. 

Hodgson, Miss E., Cavendish Street, Ulverstone. 

Holden, Adam, Esq., 48, Church Street, Liverpool. 

Hony, Rev. W. E., F.G.S., &c., Archdeacon of Sarum, Beverstock, near Salisbury. 

Hopgood, James, Esq., Clapham Common, 8.W. 

Horen, Dr. F. Van, St. Trond, Belgium. 

Horner, Rev. John §., Mells Park, near Frome. 

Howitt, Thomas, Esq., Queen Square, Lancaster. 

Huddersfield Literary and Scientific Society. 

Hughes, T. M‘K., Esq., B.A., F.G.S., Museum of Practical Geology, Jermyn Street, S.W. 

Hudson, Robert, Esq., F.R.S., G.S., Clapham, S.W. 

Hull, W. D., Esq., F.G.S., &c., 36, Queen’s-Gate Terrace, South Kensington. 

Hunter, J. R. 8., Esq., Braidwood, Carluke, N.B. 

Hutton, R., Esq., F.G.S., M.R.1.A., &c., Putney Park, 8.W. 

Huxley, Professor T. H., LL.D., Ph.D., F.R.S., L.S., G.S., Museum of Practical Geclogy 
Jermyn Street, S.W. 


Illingworth, R. S., Esq., 9, Norfolk Crescent, W. 
Ilott, James William, Esq., Bromley, Kent. 
Ipswich Museum, Ipswich. 


J’Anson, E., Esq., 7a, Laurence Pountney Hill, E.C. 

James, Colonel Sir H., R.E., F.R.S., G.S., &e., Ordnance Survey, Southampton. 
Jarvis, Richard H., Esq., F.G.S., Beckenham, Kent, S.E. 

Jeffreys, J. Gwyn, Esq., F.R.S., L.S., G.S., 25, Devonshire Place, Portland Place, W. 
Johnes, J., Esq., F.G.8., Dolancothy, Llandilo, Wales. 

Johnson, William, Esq., Eton College. 

Jones, John, Esq., Newport Road, Middlesborough, 

Jordan, Swifen, Esq., Cherith Lodge, Clifton Park, Bristol. 


ll 


Jukes, J. Beete, Esq., M.A., F.R.S., F.G.S., &c., Geological Survey of Ireland, 51, Stephen’s 
Green, Dublin. 
Judd, J. W., Esq., F.G.S., Museum of Practical Geology, Jermyn Street, S.W. 


Kenyon, the Hon. Mrs. Thomas, Pradoe, near Shrewsbury. 
King, W. P., Esq., Avonside, Clifton Down, Bristol. 

Kinnaird, Lord, Rossie Priory, Inchture, N.B. 

Kingston, G. 8., Esq., Grote Street, Adelaide, South Australia. 
Knight, Captain E., Hornacott Manor, Launceston. 

Krantz, Herr, Bonn. 


Lawrance, John, Esq., F.G.S., Elton, Oundle. 

Leckenby, John, Esq., F.G.S., Local Secretary, Scarborough. 

Lee, Henry, Esq., F.L.S., G.S., The Waldrons, Croydon, 8S. 

Lee, John Edward, Esq., F.G.S., The Priory, Caerleon, Monmouthshire. 
Leeds, E. C., Esq., B.A., Eyebury, Peterboro’. 

Leeds Library, Leeds, Yorkshire. 

Leicester Literary and Philosophical Society, Town Museum, Leicester. 
Leighton, W. H., Esq., 2, Merton Place, Turnham Green, W. 

Leonard, Edward J., Esq., Engineer’s Office, West India Docks, E. 
Lightbody, Robert, Esq., Ludlow. 

Lindsay, Charles, Esq., Ridge Park, Lanark, N.B. 

Lingard, John R., Esq., 10, Booth Street, Piccadilly, Manchester. 

Linnean Society, Burlington House, Piccadilly, W. 

Lister, J. J., Esq., F.R.S., &c., Upton, Essex. 

Literary and Philosophical Society of Manchester. 

Literary and Philosophical Society of Newcastle, Westgate Street, Newcastle-on-Tyne. 
Literary and Philosophical Society of Sheffield (by J. Holland, Esq., Music Hall, Sheffield). 
Liveing, Professor G. D., M.A., St. John’s College, Cambridge. 

Liverpool Free Public Library. 

Liverpool Natural History Society. 

Lloyd, J. H., Esq., 10, Lancaster Gate, W. 

Lobley, J. L., Esq., 50, Lansdowne Road, Kensington Park, W. 

London Institution, Finsbury Circus, E.C. 

Lovén, Professor 8., Stockholm. 

Lubbock, Sir John W., Bart., M.A., F.R.S., L.S., 15, Lombard Street, E.C. 
Lucas, Joseph, jun., Esq., Upper Tooting, Surrey. 

Lucas, John F., Esq., Middleton, Yolgrave, Bakewell, Derbyshire. 

Ludlow Natural History Society. 

Lyall, George, Esq:, F.G.S., 38, Winchester Street, South Shields. 

Lyell, Sir C., Bart., M.A., F.R.S., L.8., G.S., &c., 72, Harley Street, Cavendish Square, W. 
Lyon, Bibhothéque de la Ville de. 


McAndrew, R., Esq., F.R.S., Bond Street Chambers, Walbrook, E.C. 
McCollough, D.M., M.D., Larchfield, Abergavenny. 


12 


Mackeson, Henry B., Esq., F.G.S., &c., Hythe, Kent. 

Mackey, Colonel, Fairhill, near Exeter. 

Maclean, William C., Esq., 5, Camperdown Terrace, Great Yarmouth. 

Maemillan, Messrs., Cambridge. 

McMorran, A., Esq., Cheapside, E.C. 

Macredie, P. B. M., Eisq., Perceton House, Irvine, N.B. 

Madras Government Museum (per Messrs. Williams and Norgate). 

Major, Charles, Esq., Red Lion Wharf, Upper Thames Street, E.C. 

Mann, C. S., Esq., F.G.S., Eltham, Kent, 8.E. 

Mansel, Ton Esq., F.G. S. ee CRBs Blandford, Dorset. 

Manzoni, Dr. Angelo, Havenia: 

Marés, Mons. P., Paris. 

Marshall, James G, Esq., F.G.S., Headingley, near Leeds. 

Marshall, Matthew, Esq,, Bank of England, E.C. 

Marshall, Reginald D., Esq., Cookridge Hall, Leeds. 

Marsham, Hon. Robert, F.G.S., 5, Chesterfield Street, Mayfair, W. 

Martin, Miss, Bredon’s Norton, near Tewkesbury. 

Mathews, W., jun., Esq., F.G.S., Local Secretary, 5, Carpenter Road, Edgbaston, Birmingham. 

Maton, Professor B. 

Matthieson, James, Esq., 22, Belitha Villas, Barnsbury Park, N. 

Maw, G., Esq., F.S.A., L.S., G.S., Benthall Hall, Broseley, Salop. 

Meade, Rev. R. J., Castle Car, 

Merian, Professor Dr. Pierre, '.M.G.S., Directeur du Muséum, Basle. 

Meryon, Edward, M.D., F.G.S., 14, Clarges Street, W. 

Millar, John, M.D., F.L.8S., G.S., Bethnall House, Cambridge Road, N.K. 

Milne-Edwards, Dr. Henry, F.M.G.S., Paris. 

Mitchell, F. J., Esq., Llanbrechba Grange, Newport, Monmouthshire. 

Mitchell, W. S. , Esq., LL.B., F.L.S., G.S., New University Club, St. James’s Street, S.W. 

Mitchenson, Hen John, D.C. ie Keane: s School Canterbury. 

Moller, Captain Valerian, Tagen des Mines, St. Petersburg. 

Monk, James, Esq., Aden Cottage, Durham. 

Moore, J. Carrick, Esq., M.A., F.R.S., G.S., &c., 118, Eaton Square, S.W. 

Moore, Charles, Esq., F.G.S., 6, Cambridge Place, Widcome Hill, Bath. 

Moore, Joseph, Esq., Grasmere Lodge, Lower Tulse Hill, S. 

Morgan, William, Esq., Bryn Nant, Swansea. 

Morton, George eetnela) Esq., F.G.S., Local Secretary, 5, London Road, Liverpool. 

Mosley, Sir Oswald, Bart., D.C.L., F.L. 8., Gis. ccs Rollecton Hall, Burton-on-Trent. 

Murchison, Sir R. L., Bae K.C.B,, G.C.StS., D.C.b., MA. ERIS: G.S.) ish) ace 
Vice President, 16, Belgrave Square, S.W. ° 

Murdock, James Barclay, Esq., 33, Lynedoch Street, Glasgow. 

Museum of Practical Geology, Jermyn Street, S.W. 

Mushet, Robert, Esq., F.G.S., Royal Mint, Little Tower Hill, E. 


Nantes, Musée d’ Histoire Naturelle de. 

Neale, Edward Vansittart, Esq., West Wickham, Kent. 

Nicholl, W. A., Esq., Ham Court, Glamorganshire. 

Norfolk and Norwich Literary Institution, Norwich. 

Nottingham Literary and Philosophical Society, School of Art, Nottingham. 


13 


Nunneley, Thomas, Esq., Local Secretary, 22, Park Place, Leeds. 
Nutt, D., Esq.. Strand, W.C. 


Oldham, Mrs., Hyde House, South Littleton, Eversham, Worcestershire. 

Onate, Countess of, Madrid. 

Oswestry Naturalists’ Field Club, Oswestry. 

Ormerod, G. W., Esq., M.A., F.G.S., &e., Brookbank, Teignmouth. 

Owen, Professor R., M.D., LL.D., F.R.S., L.S., G.S., &c., Vice-President, British Museum, W.C. 


Paine, Mrs. J. M., Farnham, Surrey. 

Papillon, Rev. J., Rectory, Lexden, Colchester. 

Parker, J., Esq., F.G.S., Turl Street, Oxford. 

Parry, Thomas G., Esq., F.G.S., Highnam Court, near Gloucester. 

Pattison, 8. R., Esq., F.G.S., 50, Lombard Street, E.C. 

Paynter, Rev. Samuel, Stoke Hill, Guildford, Surrey. 

Peabody Institution, America. 

Pearson, Sir Edwin, K.H., M.A., F.R.S., &c., Wimbledon, S.W 

Pease, Thomas, Esq., F.G.S., Cote Bank, Westbury-on-Trym, Bristol. 

Peckover, Algernon, Esq., F.L.S., Wisbeach. 

Pengelly, William, Esq., F.R.S., G.S., Local Secretary, Torquay. 

Penny, Rev. James, M.A., &c., Blandford. 

Penruddocke, Charles, Esq., Compton Park, near Salisbury. 

Perceval, Spencer George, Esq., Severn House, Henbury, Bristol. 

Perkins, Rev. R. B., Wootton-Underedge, Gloucestershire. 

Phillips, John, M.A., LL.D., F.R.S., G.S., &c., Professor of Geology in the University of 
Oxford, Vice-President, Museum, Oxford. 

Philosophical Society of Glasgow. 

Phear, Rev. George, F.G.S., Emmanuel College, Cambridge. 

Phené, John 8., Esq., 34, Oakley Street, Chelsea, S.W. 

Pickering, John, Esq., 29, Loraine Road, Upper Holloway, N. 

Pictet, Mons. F. J., Professor of Zoology, Academy of Geneva. 

Pidgeon, Jonathan S., Esq., Pembridge Villa, Bayswater, W. 

Plant, James, Esq., Local Secretary, 40, West Terrace, West Street, Leicester. 

Pollock, Lady, Clapham Common, S.W. 

Portal, Wyndham S8., Esq., Malshanger House, Basingstoke. 

Portman, Hon, Miss, 5, Princes Gate, Hyde Park, S.W. 

Poynton, Rev. Francis, Rectory, Kelston, Bath. 

Prestwich, Joseph, Esq., F.R.S., G.S., Shoreham, Sevenoaks. 

Prout, Rev. E., F.G.S., &c., Fairfield, Torquay. 

Powrie, James, Esq., F'.G.S., Reswallie, Forfar. 

Provis, William A., Esq., F.G.S., The Grange, Ellesmere, Salop. 

Puxty, C., Esq., Maldon, Essex. 


Quaritch, B., Esq., Piccadilly, W. 
Queen’s College, Cork (by Messrs. Hodges and Smith). 


Queen’s College, Galway. 
Queen’s College, Oxford. 


Raban, Capt. R. B., Shirehampton, Bristol. 

Radcliffe Library, Oxford. 

Ramsay, Mrs. Wm., Rannagubzion, Blairgowrie. 

Ramsay, Professor A. C., LL.D., F.R.S., G.S., &e., Museum Pract. Geology, Jermyn Street, S.W. 
Ransome, Robert Charles, Esq., Ipswich. 

Regnés, Mons., Paris. 

Renevier, Mons. E., Professor of Geology, Academy of Lausanne, Switzerland. 
Richardson, James, Esq., Glaurafon, Swansea. 

Richardson, William, Esq., High Field, Southouram, near Halifax. 
Ricketts, Charles, M.D., F.G.S., 22, Argyle Street, Birkenhead. 
Ripon Scientific Society. 

Risley, Rev. William Cotton, M.A., Deddington, Oxfordshire. 
Robertson, E. H., Fair Ville, Rose Mount, Oxton, Cheshire. 
Robbins, George, Esq., F.G.S., 9, Royal Crescent, Bath. 

Rofe, John, Esq., F.G.S., &c., 7, Queen Street, Lancaster. 

Roper, F. C. 8., Esg., F.G.S., L.S., 157, Maida Vale, W. 

Rose, C. B., Esq., F.G.S., &c., 28, King Street, Great Yarmouth. 
Rothery, H. C., Esq., M.A., F.L.S., 94, Gloucester Terrace, Hyde Park W. 
Rothery, Charles W., Esq., Greta Hall, Keswick. 

Royal Artillery Institution, Woolwich, 8.E. 

Royal College of Science for Ireland, Stephen’s Green, Dublin. 
Royal College of Surgeons, Lincoln’s Inn Fields, W.C. 

Royal Dublin Society, Dublin. 

Royal Geological Society of Cornwall, Penzance. 

Royal Institution of Cornwall, Truro. 

Royal Institution of Great Britain, Albemarle Street, W. 

Royal Institution, Liverpool. 

Royal Institution of South Wales, Swansea. 

Royal Inmish Academy, 19, Dawson Street, Dublin. 

Royal Society of Edinburgh. 

Royal Society of London, Burlington House, W. 

Rudd, Rev. Leonard H., M.A., Kempsey, Worcester. 

Rutter, John, Esq., UJminster. 

Rylands, T. G., Esq., F.L.8., G.S., Heath House, Warrington. 


Sabine, General, R.A., F.R.S., L.8., &c., 13, Ashley Place, Westminster, S.W. 

Salford Borough Royal Museum and Library, Peel Park. 

Salt, 8., Esq., Ulverstone. 

Sanders, Gilbert, Esq., M.R.I.A., &c., 53, Claverton Street, S.W. 

Sanders, W., Esq., F.R.S., G.S., &c., Local Secretary, 21, Richmond Terrace, Clifton, Bris tol. 
Sanford, W. A., Esq., F.G.S., Nynehead Court, Wellington, Somerset. 

Saul, G. T., Esq., F.Z.S., Bow Lodge, Bow Road, E. 

Saunders, James Ebenezer, Esq., F.U.8., G.S., 9, Finsbury Circus, E.C. 


Saunders, W. Wilson, Esq., F.R.S., L.S., &c., Lloyd’s, E.C. 

Savy, Mons. F., Local Secretary, 24, Rue Hautefeuille, Paris. 

Scarborough Philosophical Society. 

Sedgwick, Rev. Adam, M.A., F.R.S., G.S., &c., Cambridge. 

Severs, J., Esq., Airthwaite, Kendal. 

Sharman, George, Esq., 9, St. George’s Road, Kilburn. 

Sharp, John, Esq., F.G.S8., Local Secretary, Culverden Hill, Tunbridge Wells, Kent. 
Sharp, Samuel, Esq., F.G.S., S.A., Dallington Hall, Northampton. 

Shaw, John, M.D., F.G.S., &c., Hop House, Boston, Lincolnshire. 

Sheppard, T. Byard, Esq., Selwood Cottage, Frome. 

Sheppard, Alfred B., Esq., Torquay. 

Sidney Sussex College Library, Cambridge. 

Simpson, J. B., Esq., Hedgefield House, Blaydon-on-Tyne. 

Skaife, John, Esq., 6, Union Street, Blackburn. 

Sloper, G. E., Esq., Devizes. 

Smith, Rev. Charles Lesingham, Little Canfield Rectory, near Chelmsford, Essex. 
Smith, Captain Robert, Frankford Avenue, Rathgar, Dublin. 

Smith, Rev. Urban, Stoney Middleton. 

Smithe, J. D., Esq., C.E., F.G.S., Madhopoor, Punjib (by Messrs. Smith, Elder, and Co.). 
St. Peter’s College, Cambridge. 
Somersetshire Archeological and Natural History Society, Museum, Taunton. 
Sopwith, T., Esq., F.R.S., G.S., 103, Victoria Street, 5.W. 

South Shields Geological Club. 

Sowerby, James de Carle, Esq., F.L.S., &c., Royal Botanic Gardens, Regent’s Park, N.W 
Spicer, Henry, Esq., jun., 19, New Bridge Street, Blackfriars, H.C. 

Spink, J., Esq., Drax, Selby. 

Spragge, W. Kennaway, Esq., the Quarry, Paignton, South Devon. 

Stebbing, Rev. T. R. R., M.A., Tor Crest Hall, Torquay. 

Steedman, John, Esq., Charlestown by Dunfermline. 

Stewart, Mrs. John, 7, Grosvenor Crescent, Edinburgh. . 

Stobart, W. C., Esq., Etherley House, Darlington. 

Stoddart, W. W., Esq., F.G.S., 9, North Street, Bristol. 

Stubbs, Charles E., Esq., Henley-on-Thames. 

Studer, Herr B., For.M.G.S., Professor of Geology, Berne. 

Sunderland Corporation Museum. 

Sunderland Geological Society. 

Sunderland Subscription Library. 

Swayne, H. J. F., Esq., The Island, Wilton, Wilts. 


Pate, Geo., Esq., F.G.S., Alnwick, N.B. 
Tawney, E. B., Esq., F.G.S., Woodville, Ilfracombe, N. Devon. 
Taylor, S. Watson, Esq., Erlestoke Park, Devizes. 
Tennant, Professor Jas., F.G.S., &c., 149, Strand, W.C. (Two Copies.) 
Teuchner, Professor. 
The General Administration of Mines and Salt-works in Munich (by Messrs. Williams and 
; Norgate). 
Thomas, Capt. F. W. L., R.N., H.M.S. “ Woodlark,” Trinity, near Edinburgh. 
Thompson, Miss S., Stamford. 


16 


Thompson, Thomas, Esq., Hull. 

Thomson, Alexander, Esq., F.R.S.E., Banchory House, Aberdeen. 

Thomson, Professor Wyville, LL.D., F.G.S., Queen’s College, Belfast. 

Tomes, John, Esq., 37, Cavendish Square, W. 

Torquay Natural History Society. 

Traquair, Professor R. H., M.D., Royal Agricultural College, Cirencester. 

Trautschold, Dr., Moscow. 

Trevelyan, Sir W. C., Bart., F.G.S., Wellington, Newcastle-on-Tyne ; and Athenzeum Club, S.W. 
Trinity College, Cambridge. 

Turner, E. J., Esq., Fairlight Villas, Clapham, 8. 

Twamley, Charles, Esq., F.G.S., 6, Queen’s Road, Gloucester Gate, Regent’s Park, N.W. 
Tyler, Chas., Esq., F.L.8., G.S., 24, Holloway Place, Holloway, N. 

Tylor, Alfred, Esq., F.L.8., G.S., Warwick Lane, Newgate Street, E.C. 


University of Edinburgh. 
University of Glasgow. 
University Library, Aberdeen. 
University Library, Leipzig. 
University Library, St. Andrew’s. 


Verneuil, Mons. Edouard de, Mem. de I’Instit., F.M.G.S., 57, Rue de la Madeleine, Paris. 
Vernon Park Museum, Stockport. 
Vicary, William, Esq., F.G.S., The Priory, Colleton Crescent, Exeter. 


Wall, Geo. P., Esq., F.G.S., Pitmoor, near Sheffield. 

Walmstedt, Dr. L. P., Professor of Mineralogy, Upsala. 

Walton, William, Esq., 11, Paragon, Blackheath, S.E. 

Walton, William, Esq., Local Secretary, 17, Grosvenor Place, Bath. 

Ward, Henry, Esq., F.G.S., Oaklands, Wolverhampton. 

Wardle, Thos., Esq., F.G.S., St. Edward Street, Leek. 

Waring, Samuel Long, Esq., F.G.S., The Oaks, Norwood, Surrey, S. 

Warrington Museum and Library. 

Warwickshire Natural History Society, Warwick. 

Watson, Rev. R. B., F.G.S., 4, Bruntsfield Place, Edinburgh. 

West, G. Herbert, Esq., B.A., F.G.S., 61, Wimpole Street, W. 

Wetherell, N. T., Esq., F.G.S., &c., Highgate, N. . 

White, Alfred, Esq., F.L.S., &c., Castle Street, Cow Cross, E.C. 

Willaume, T. B. T., Esq., jun., 24, Chester Terrace, Regent’s Park. 

Willcock, J. W., Esq., 6, Stone Buildings, Lincoln’s Inn, W.C. 

Williams and Norgate, Messrs., Henrietta Street, Covent Garden, W.C. 

Willis and Sotheran, Messrs., Strand, W.C. 

Wilson, J. M., Esq., B.A., F.G.S., Rugby. 

Wilson, Rev. Dr. W., Southampton. 

Wiltshire, Rev. Thomas, M.A., F.L.S., G.S., &c., Honorary Secretary, 13, Granville Park, 
Lewisham, Kent, S.E. 


17 


Winstone, Benjamin, M.D., 7, Ely Place, Holborn, E.C. 

Witts, Rev. E. F., F.G.S., Rectory, Upper Slaughter, near Stow-on-the- Wold. 
Winwood, Rev. Henry H., F.G.S., 4, Cavendish Crescent, Bath. 

Wolley-Dod, Rev. Charles, Eton College. 

Wood, Edward, Esq., F.G.S., R.S.L., &c., Local Secretary, Richmond, Yorkshire. 
Wood, Henry, Esq., 10, Cleveland Square, Bayswater, W. 

Wood, Rev. Henry H., F.G.S., Holwell Rectory, Sherborne, Dorset. 

Wood, Rev. J. E. Tenison, F.G.S., Penola, South Australia. 

Wood, S. V., Esq., F.G.S., &c., Treasurer, Brentwood, Essex. 

Woodall, Major J. W., M.A., F.G.S., &c., St. Nicholas House, Scarborough. 
Woodd, C. H. L., Esq., F.G.S., S.A., &c., Hillfield, Hampstead, N.W. 
Woodward, Charles, Esq., F.R.S., 10, Compton Terrace, Islington, N. 
Woodward, Henry, Esq., F.G.S., Z.S., British Museum, W.C. 

Worcestershire Natural History Society, Foregate, Worcester. 

Wright, F. Beresford, Esq., Aldercar Hill, Langley Mill, Derbyshire. 

Wright, Joseph, Esq., F.G.S., 7, Donegall Street, Belfast. 

Wright, Thomas, M.D., F.R.S.E., G.S., Local Secretary, St. Margaret’s Terrace, Cheltenham. 
Wurzburg, the Royal University Library of. 


Yates, James, Esq., M.A., F.R.S., L.S., G.S., Lauderdale House, Highgate, N. 
Yorkshire Naturalists’ Club, York (by Ed. Smallwood, Esq.). 

Yorkshire Philosophical Society, York. 

Young, James, Esq., Burr Street, Tower Hill, E. 


Zoological Society of London, 11, Hanover Square, W. 


18 


CATALOGUE OF WORKS 


ALREADY PUBLISHED BY 


THE PALZONTOGRAPHICAL SOCIETY: 


Showing the Ornur of publication; the Yuars during which the Society has been in 


operation ; and the Contents of each yearly Volume. 


Vol. I. Issued for the Year 1847 The Crag Mollusca, Part I, Univalves, by Mr. S. V. Wood, 21 plates. 


Sy UE 


> Want 


7 AY 


plates. 


Tae ; The Reptilia of the London Clay, Part I, Chelonia, &c., by Profs. Owen and Bell, 38 
8 
The EHocene Mollusca, Part I, Cephalopoda, by Mr. F. EH. Edwards, 9 plates. 


The Entomostraca of the Cretaceous Formations, by Mr. T. R. Jones, 7 plates. 
The Permian Fossils, by Prof. Wm. King, 29 plates. ae 
1849 é The Reptilia of the London Clay, Part Il, Crocodilia and Ophidia, &c., by Prof. Owen, 
18 plates. ; 
| The Fossil Corals, Part I, Crag, London Clay, Cretaceous, by Messrs. Milne-Hdwards 
L and Jules Haime, 11 plates. 


¢ The Crag Mollusca, Part II, No. 1, by Mr. S. V. Wood, 12 plates. 
J The Mollusca of the Great Oolite, Part I, Univalves, by Messrs. Morris and Lycett, 15 
1850 plates. 
[pike Fossil Brachiopoda, Part ITI, No. 1, Oolitic and Liassic, by Mr. Davidson, 15 
plates. 


The Fossil Corals, Part II, Oolitic, by Messrs. Milne-Edwards and Jules Haime, 19 
plates. 


The Reptilia of the Cretaceous Formations, by Prof. Owen, 39 plates. 
1851 
The Fossil Lepadidx, by Mr. Charles Darwin, 5 plates. 


19 


CATALOGUE OF WORKS—Continued. 


{ The Fossil Corals, Part III, Permian and Mountain-limestone, by Messrs. Milne- 
Edwards and Jules Haime, 16 plates. 


Vol. VJ. Issued for the Year 3 The Fossil Brachiopoda, Part I, Tertiary, by Mr. Davidson, 2 plates. 
52 


ee WALL 


5) MAILE = 


xb. 


9 | The Fossil Brachiopoda, Part II, No. 1, Cretaceous, by Mr. Davidson, 5 plates. 
The Fossil Brachiopoda, Part III, No. 2, Oolitic and Liassic, by Mr. Davidson, 5 plates. 
L The Hocene Mollusca, Part IT, Pulmonata, by Mr. F. E. Edwards, 6 plates. 
The Radiaria of the Crag, London Clay, &c., by Prof. E. Forbes, 4 plates. 


( The Fossil Corals, Part IV, Devonian, by Messrs. Milne-Edwards and Jules Haime, 10 
| 
plates. 
| The Fossil Brachiopoda, Introduction to Vol. I, by Mr. Davidson, 9 plates. 
1853 4 The Mollusca of the Chalk, Part I, Cephalopoda, by Mr. D. Sharpe, 10 plates. 
The Mollusca of the Great Oolite, Part II, Bivalves, by Messrs. Morris and Lycett, 8 
plates. 
| The Mollusca of the Crag, Part II, No. 2, Bivalves, by Mr. 8. V. Wood, 8 plates. 
L The Reptilia of the Wealden Formations, Part I, Chelonia, by Prof. Owen, 9 plates. 


[ The Fossil Brachiopoda, Part II, No. 2, Cretaceous, by Mr. Davidson, 8 plates. 
The Reptilia of the Wealden Formations, Part II, Dinosauria, by Prof. Owen, 20 plates. 
| The Mollusca of the Great Oolite, Part III, Bivalves, by Messrs. Morris and Lycett, 7 
plates. 
*1854 2 The Fossil Corals, Part V, Silurian, by Messrs. Milne-Edwards and Jules Haime, 16 
lates. 
The Fossil Balanide and Verrucide, by Mr. Charles Darwin, 2 plates. 
The Mollusca of the Chalk, Part II, Cephalopoda, by Mr. D. Sharpe, 6 plates. 
The Eocene Mollusca, Part III, No. 1, Prosobranchiata, by Mr. F. E. Edwards, 8 
L plates. 


( The Mollusca of the Crag, Part II, No. 3, Bivalves, by Mr. 8. V. Wood, 11 plates. 
| The Reptilia of the Wealden Formations, Part III, by Prof. Owen, 12 plates. 
7 The Eocene Mollusca, Part III, No. 2, Prosobranchiata, continued, by Mr. F. E. 
T1855 4 Edwards, 4 plates. 
| The Mollusca of the Chalk, Part III, Cephalopoda, by Mx. D. Sharpe, 11 plates. 
The Tertiary Entomostraca, by Mr. T. R. Jones, 6 plates. 
The Fossil Echinodermata, Part I, Oolitic, by Dr. Wright, 10 plates. 


The Fossil Echinodermata, Part II, Oolitic, by Dr. Wright, 12 plates. 

The Fossil Crustacea, Part I, London Clay, by Prof. Bell, 11 plates. 

The Fossil Brachiopoda, Part IV, Permian, by Mr. Davidson, 4 plates. 

The Fossil Brachiopoda, Part V, No. 1, Carboniferous, by Mr. Davidson, 8 plates. 
| The Reptilia of the Wealden Formations, Part IV, by Prof. Owen, 11 plates. 
L The Reptilia of the London Clay (Supplement), by Prof. Owen, 2 plates. 


1856 


* This Vol. is marked on the outside 1855. f This Vol. is marked on the outside 1856. 


20 


CATALOGUE OF WORKS—Continued. 


( The Fossil Echinodermata, Part ITI, Oolitic, by Dr. Wright, 14 plates. 
FOREX. Taened forithe aes J The Fossil Brachiopoda, Part V, No. 2, Carboniferous, by Mr. Davidson, 8 plates. 
Vo]. XI. Issued for the Year J me Reptilia of the Cretaceous Formations (Supplement 1), by Prof. Owen, 4 plates. 
[ The Reptilia of the Wealden Formations (Supplement No. 2), by Prof. Owen, 8 plates. 
The Polyzoa of the Crag, by Prof. Busk, 22 plates. 


( The Fossil Echinodermata, Part IV, Oolitic, by Dr. Wright, 7 plates. 
| The Eocene Mollusca, Part III, No. 3, Prosobranchiata continued, by Mr. F. E. 
] Edwards, 6 plates. 

ip OSE * 1858 < The Reptilia of the Cretaceous Formations (Supplements No. 2, No. 3), by Prof. Owen, 
| 7 plates. 


The Reptilia of the Purbeck Limestones, by Prof. Owen, 1 plate. 
The Fossil Brachiopoda, Part V, No. 3, Carboniferous, by Mr. Davidson, 10 plates. 


The Reptilia of the Oolitic Formations, No. 1, Lower Lias, by Prof. Owen, 6 plates. 
The Reptilia of the Kimmeridge Clay, No. 1, by Prof. Owen, 1 plate. 
The Hocene Mollusca, Part IV, No. 1; Bivalves, by Mr. 8. V. Wood, 18 plates. 


{ The Fossil Brachiopoda, Part V, No. 4, Carboniferous, by Mr. Davidson, 20 plates. 
, 2ST. 1859 


( The Fossil Brachiopoda, Part V, No. 5, Carboniferous, by Mr. Davidson, 8 plates. 
| The Reptilia of the Oolitic Formation, No. 2, Lower Lias, by Prof. Owen, 11 plates. 
exlive s3 1860 1 The Reptilia of the Kimmeridge Clay, No. 2, by Prof. Owen, 1 plate. 
The Fossil Estheriz, by Prof. Rupert Jones, 5 plates. 
L The Fossil Crustacea, Part II, Gault and Greensand, by Prof. Bell, 11 plates. 


The Fossil Echinodermata, Vol. II, Part I (Oolitic Asteroidea), by Dr. Wright, 13 
NOE 5 1861 plates. 


Supplement to the Great Oolite Mollusca, by Dr. Lycett, 15 plates. 


The Fossil Echinodermata, Cretaceous, Vol. I, Part I, by Dr. Wright, 11 plates. 
| The Trilobites of the Silurian, Devonian, &c., Formations, Part I, by Mr. J. W. Salter, 
6 plates. 
peovel , 1862 4 The Fossil Brachiopoda, Part VI, No. 1. Devonian, by Mr. Davidson, 9 plates. 
| The Hocene Mollusca, Part IV, No. 2, Bivalves, by Mr. 8. V. Wood, 7 plates. 
L The Reptilia of the Cretaceous and Wealden Formations (Supplements), by Prof. Owen, 
10 plates. 


The Trilobites of the Silurian, Devonian, &c., Formations, Part II, by Mr. J. W. 
Salter, 8 plates. 
4 DL OVATLE, +, 1863 4 The Fossil Brachiopoda, Part VI, No. 2, Devonian, by Mr. Davidson, 11 plates. . 
The Belemnitide, Part I, Introduction, by Prof. Phillips. 
\ The Reptilia of the Liassic Formations, Part 1, by Prof. Owen, 16 plates. 


Vol. XVIII. Issued for the Year 


7) PAD 


5 OO: 


20-day 


Sr, DO. 


el T.* 


21 


CATALOGUE OF WORKS—Continued. 


¢ The een Echinodermata, Vol. II, Part II (Liassic Ophiuroidea), by Dr. Wright, 
plates. 
| The Trilobites of the Silurian, Devonian, &c., Formations, Part ITI, by Mr. J. W 


J Salter, 11 plates. 
The Belemnitidx, Part II, Liassic Belemnites, by Prof. Phillips, 7 plates. 
1864 | The Pleistocene Mammalia, Part I, Introduction, Felis speleea, by Messrs. W. Boyd 
Dawkins and W. A. Sanford, 5 plates. : 
Title-pages, &c., to the Monographs on the Reptilia of the London Clay, Cretaceous 
and Wealden Formations. 


The Crag Foraminifera, Part I, No. 1, by Messrs. T. Rupert Jones, W. K. Parker, and 
H. B. Brady, 4 plates. 
1865 Supplement to the Fossil Corals, Part I, Tertiary, by Dr. Duncan, 10 plates. 
The Fossil Merostomata, Part I, Pterygotus, by Mr. H. Woodward, 9 plates. 
The Fossil Brachiopoda, Part VII, No. 1, Silurian, by My. Davidson, 12 plates. 


( Supplement to the Fossil Corals, Part IV, No. 1, Liassic, by Dr. Duncan, 11 plates. 
| The Trilobites of the Silurian, Devonian, &c., Formations, Part IV (Silurian), by Mv. 
1866 J. W. Salter, 6 plates. 
The Fossil Brachiopoda, Part VII, No. 2, Silurian, by Mr. Davidson, 10 plates. 
The Belemnitide, Part III, Liassic Belemnites, by Prof. Phillips, 13 plates. 


( Flora of Carboniferous Strata, Part I, by Mr. E. W. Binney, 6 plates. 
| Supplement to the Fossil Corals, Part IV, No. 2, Liassic, by Dr. Duncan, 6 plates. 
| The Fossil Echinodermata, Cretaceous, Vol. I, Part II, by Dr. Wright, 14 plates. 
1867 } The Fishes of the Old Red Sandstone, Part I, by Messrs. J. Powrie and E. Ray 
Lankester, 5 plates. 
| The Pleistocene Mammalia, Part II, Felis spelea, continued, by Messrs. W. Boyd 
Dawkins and W. A. Sanford, 14 plates. 


Supplement to the Fossil Corals, Part II, No. 1, Cretaceous, by Dr. Duncan, 9 plates. 
The Fossil Merostomata, Part II, Pterygotus, by Mr. H. Woodward, 6 plates. 
The Fossil Brachiopoda, Part VII, No. 3, Silurian, by Mr. Davidson, 15 plates. 
1868 < The Belemnitide, Part IV, Liassic and Oolitic Belemnites, by Prof. Phillips, 7 plates. 
The Reptilia of the Kimmeridge Clay, No. 3, by Prof. Owen, 4 plates. 
The Pleistocene Mammalia, Part III, Felis spelwa, continued, with F. Lynx, by 
L Messrs. W. Boyd Dawkins and W. A. Sanford, 6 plates. 


Supplement to the Fossil Corals, Part II, No. 2, by Dr. Duncan, 6 plates. 
The Fossil Echinodermata, Cretaceous, Vol. I, Part III, by Dr. Wright, 10 plates. 
The Belemnitidw, Part V, Oxford Clay, &c., Belemnites, by Prof. Phillips, 9 plates. 
1869 4 The Fishes of the Old Red Sandstone, Part I (concluded), by Messrs. J. Powrie and 
E. Ray Lankester, 9 plates. 
The Reptilia of the Liassic Formations, Part II, by Prof. Owen, 4 plates. 
The Crag Cetacea, No. 1, by Prof. Owen, 5 plates. 


* These Volumes are issued in two forms of binding, first, with all the Monographs stitched together and enclosed in 
one cover; secondly, with each of the Monographs separate, and the whole of the separate parts placed im an envelope 
The previous volumes are not in separate parts. 


co) 
~ 


LIST OF MONOGRAPHS 


Completed, in course of Publication, and in Preparation. 


MONOGRAPHS which have been CompLetEp :— 


The ‘Tertiary, Cretaceous, Oolitic, Devonian, and Silurian Corals, by MM. Milne-Edwards 
and J, Haime. 

The Polyzoa of the Crag, by Mr. G. Busk. 

The Tertiary Echinodermata, by Professor Forbes. 

The Fossil Cirripedes, by Mr. C. Darwin. 

The Tertiary Entomostraca, by Prof. T. Rupert Jones. 

The Cretaceous Entomostraca, by Prof. T. Rupert Jones. 

The Fossil Estheriz, by Prof. T. Rupert Jones. 

The Tertiary, Cretaceous, Oolitic, Liassic, Permian, Carboniferous, and Devonian Brachiopoda, 
by Mr. T. Davidson. 

The Mollusca of the Crag, by Mr. 8. V. Wood. 

The Great Oolite Mollusca, by Professor Morris and Mr. J. Lycett. 

The Cretaceous (Upper) Cephalopoda, by Mr. D. Sharpe. 

The Fossils of the Permian Formation, by Professor King. 

The Reptilia of the London Clay (and of the Bracklesham and other Tertiary Beds), by 
Professors Owen and Bell. 

The Reptilia of the Cretaceous, Wealden, and Purbeck Formations, by Professor Owen. 


MONOGRAPHS in course of PuBLICATION :*— 


The Flora of the Carboniferous Formation, by Mr. E. W. Binney. 

The Crag Foraminifera, by Messrs. T. Rupert Jones, W. K. Parker, and H. B. Brady. 
Supplement to the Fossil Corals, by Dr. Duncan. 

The Echinodermata of the Oolitic and Cretaceous Formations, by Dr. Wright. 

The Fossil Merostomata, by Mr. H. Woodward. 


* Members having specimens which might assist the authors in preparing their respective 
Monographs are requested te communicate in the first instance with the Honorary Secretary. 


ro) 
Go 


MONOGRAPHS in course of Pusiicatton—Continued. 


The Trilobites of the Mountain-Limestone, Devonian, and Silurian Formations, by Mr. J. W 
Salter.* 

The Malacostracous Crustacea, by Professor Bell. 

The Eocene Mollusca, by Messrs. F. E. Edwards and 8. V. Wood. 

The Silurian Brachiopoda, by Mr. Davidson. 

The Belemnites, by Professor Phillips. 

The Fishes of the Old Red Sandstone, by Messrs. J. Powrie and E. Ray Lankester. 

The Reptilia of the Kimmeridge Clay, by Professor Owen. 

The Reptilia of the Liassic Formations, by Professor Owen. 

The Pleistocene Mammalia, by Messrs. Boyd Dawkins and W. A. Sanford. 

The Cetacea of the Crag, by Professor Owen. 


* Unfinished through the death of the Author. 


MONOGRAPHS which are in course of PREPARATION :+}— 


The Flora of the Tertiary Formation, by Mr. W. S. Mitchell. 

The Cretaceous Foraminifera, by Messrs. T, Rupert Jones, W. K. Parker, and H. B. Brady 

The Foraminifera of the Lias, by Mr. H. B. Brady. 

The Graptolites, by Professor Wyville Thomson. 

The Polyzoa of the Chalk Formation, by Mr. G. Busk. 

The Paleozoic Polyzoa, by Dr. Duncan. 

The Crinoidea, by Professor Wyville Thomson. 

The Post-Tertiary Entomostraca, by the Rev. H. W. Crosskey and Messrs. G. S. Brady and 
D. Robertson. 

The Wealden, Purbeck, and Jurassic Entomostraca, by Messrs. T. Rupert Jones and G. S. 
Brady. 

The Bivalve Entomostraca, of the Carboniferous Formations, by Messrs. T. Rupert Jones and 
J. W. Kirkby. 

The Trigoniz, by Dr. Lycett. 

The Post-Tertiary Mollusca, by Mr. J. Gwyn Jeffreys. 

The Cretaceous Mollusca (exclusive of the Brachiopoda), by the Rev. T. Wiltshire. 

The Purbeck Mollusca, by Mr. R. Etheridge. 

The Inferior Oolite Mollusca, by Mr. R. Etheridge. 

The Rheetic Mollusca, by Mr. R. Etheridge. 

The Liassic Gasteropoda, by Mr. Ralph Tate. 

The Ammonites of the Lias, by Dr. Wright. 


+ Members having specimens which might assist the authors in preparing their respective 
Monographs are requested to communicate in the first instance with the Honorary Secretary 


Dates of the Issue of the Yearly Volumes of the 
Paleontographical Society. 


The Volume for 1847 was issued to the Members, March, 1848. 


5 1848 5 5 sa July, 1849. 

5 1849 , $5 Bs August, 1850. 

ss 1850 s - ay June, 1851. 

e 1851 5 ~ Ss June, 1851. 

P 1852 53 a A August, 1852. 

& 1853 5 “f ss December, 1853. 
% 1854. ‘ . - May, 1855. 
1855 ms AS ee February, 1857. 
. 1856 hs . April, 1858. 

as 1857 5; i =p November, 1859. 
a 1858 a - - March, 1861. 

_ 1859 - e 33 December, 1861. 
ee 1860 A . May, 1863. 

s 1861 55 5 ¥ May, 1863. 

> 1862 BB , August, 1864. 

is 18638 a 3 . June, 1865. 

Be 1864 PA FP pe April, 1866. 

Bs 1865 “ ie a December, 1866. 
' 1866 oe es = June, 1867. 

a 1867 e " R June, 1868. 

A 1868 4 As . February, 1869. 
” 1869 5 3 January, 1870. 


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— 


PALHONTOGRAPHICAL SOCIETY. 


INSTITUTED MDCCCXLVII. 


VOLUME FOR 1869. 


LONDON: 


MDCCCL 


A MONOGRAPH 


OF THE 


BRITISH FOSSIL CORALS. 


SECOND SERIKS. 


BY 


P. MARTIN DUNCAN, M.B. LOND., F.R.S., 


FELLOW OF, AND SECRETARY TO, THE GEOLOGICAL SOCIETY. 


Being a Supplement to the 
‘Monograph of the British Fossil Corals, by MM. Mitne-Epwarps aad Jutes aime. 


PART Uh, No. 2: 


CorAts FROM THE Upper GREENSAND OF HALDON, FROM THE GAULT, AND THE LOWER 
GREENSAND. 


Pages 27—46 ; Plates X—XYV. 


LONDON: 
PRINTED FOR THE PALAXONTOGRAPHICAL SOCIETY. 


1870. 


PRINTED BY 
J. E. ADLARD, BARTHOLOMEW CLOSE. 


CONTENTS. 


PAGE 
Corats FROM THE Upper Greensand or HaLpon . : , = Py; 


Corats FROM THE GauLT; Description oF New, anp Notes on Op Species. 31 
List oF SPECIES FROM THE GAULT 


: : ; : : . 38 
CoraLs From THE LOWER GREENSAND; Descriprion or New, anp Novices or 
Op Species . : : 3 : : , , . 39 
List or New Species “Shoes : . F ; . 43 
List or Sprectes FROM THE LOWER GREENSAND . : : P . 43 
List oF Seecies rroM THE Cretaceous Formations. : : . 44 


SAD Ae eae) te 


A MONOGRAPH 


OF THE 


BRITISH FOSSIL CORALS. 


(SECOND SERIES.) 


Part II.—wNo. 2. 


J.—CoraALs FROM THE Upprr GREENSAND oF HALDON. 


Some time after the ‘ Supplement to the Monograph of the Fossil Corals of the Upper 
Greensand’ was published several very interesting specimens of fossil Corals were sub- 
mitted to examination from the deposit at Haldon, in Devonshire.’ It was necessary to 
describe them, for they had not been previously noticed, and this could not be done before 
the Corals from the Red Chalk were published. The Corals from Haldon should have 

been described amongst those of the Upper Greensand. It is, of course, evident that the 
list of Upper Greensand species (p. 23) is incomplete. 


MADREPORARIA APOROSA. 


Famity—ASTRAIDA. 
Sub-family—KvsMI11n &. 
Genus—PLACOSMILIA. 


1. Pracosmitia cunzirormis, Ld. and H. PI. X, figs. 1—5. 


The corallum is much compressed, and deltoid in shape. 
The costz are delicate, close, slightly prominent, and subequal. 


‘ Mr. Vicary, of Exeter, had collected the fossils himself, and pointed out to me their siliceous condition 
of fossilization. 


3) 


28 BRITISH FOSSIL CORALS 


The calicinal fossa is very narrow, long, and shallow. 

The septa are close, alternately thick and thin. They number (in full-sized calices) 176. 

The columella is lamelliform and indistinct. 

Locality. Waldon. In the Collection of William Vicary, Esq., F.G.S., Exeter. 

The specimen figured in Pl. X is a young corallum, and has only five cycles of septa. 
Its granular costee and the peculiar striation of its septa are very characteristic. 

The height of the specimen is $ inch, and the length of the calice is rather more. The 
breadth is 53;ths inch. 

The Placosmilie hitherto described are from the Craie tuffeau and the Hippurite 
Chalk of Soulage and Bains de Rennes (Corbieres), Les Martigues, Uchaux, Obourg near 
Mons, and Gosau. 


2. Puacosmitia Parxinsoni, Hd. and H. PI. X, figs. 6 and 7. 
PLACOSMILIA CONSOBRINA, Reuss. 


The corallum is tall, compressed, conical, and slightly curved. 

The coste are fine and separated by decided intercostal spaces. 

The calice is subelliptical in shape. 

The fossa is narrow and shallow. 

The columella is feebly developed. 

There are five cycles of septa, and the laminz are very unequal. 

Locality. Haldon. In the Collection of William Vicary, Esq., F.G.S., Exeter. 

The specimen from Haldon is somewhat rolled and worn. ‘The height is =5ths inch. 
The breadth of the calice is ;3,ths inch, and its length is 7%oths inch. 

Placosmilia Parkinsoni has been found at Gosau, in the Corbiéres, and at Uchaux. 


3. Puacosminia MAGNIFICA, Duncan. Pl. X, figs. 11—13. 


The corallum is compressed, short, very elongate, and the calicular margin is curved 
and rounded. 

The calice is very long, curved, rounded at each end, compressed, very open, and 
shallow. 

The septa are unequal, distant, large, and curved ; they correspond to costa of the 
same size. ‘There are five cycles of septa. 

The columella is lamellar, very much developed, thick, continuous, long, and slightly 
prominent in the calicular fossa. 

The coste are unequal and distant. 


FROM HALDON. 29 


The exotheca is inclined and very strongly developed. 

Height of the corallum, 1} to 14 inch. Length of the calice, 23 to 3,/5th inches. 
Breadth of the calice, ;%ths to 15th inch. 

Locality. Waldon. In the Collection of William Vicary, Esq., F.G.S., Exeter. 

This fine species is strongly Placosmilian, and might be taken as the type of the 
genus. 


Genus—PrEPLOSMILIA. 
PrrLosMILIa DEpRESSA, 7. de Fromentel. PI. X, figs. 8—10. 


The corallum is not very tall, and shows traces of epitheca. 

The calice is shallow and round. 

The septa are well developed and thin. ‘There are more than four cycles, and 
probably a fifth exists in full-grown individuals. 

The columella is very thin and narrow. 

Height, $ inch. Breadth of calice, {ths inch. 

Locality. Ualdon. In the Collection of William Vicary, Esq., F'.G.S., Exeter. 

M. de Fromentel, ‘ Pal. Franc., Terr. Crét.,’ pl. 46, fig. 1, 1863, and page 241, 
states that his specimens came from the Upper Greensand of Mans. 

The specimen from Haldon is fragmentary, and its columella is defective, but it is 
so like M. de Fromentel’s delineation of Peplosmilia depressa that there is no doubt about 
its being of that species. 


Division—ASTRRACER. 
Genus—ASTROCG@NIA. 


ASTROC@NIA DECAPHYLLA, Hd. and H. PI. XI, figs. 1—6. 


This species, described by MM. Milne-Edwards and Jules Haime (‘Ann. des 
Sci. nat.,’ 3me série, t. x, p. 298, 1849) was subsequently named Astrea reticulata by 
D’Orbigny (1850), and was noticed as Astrocenia magnifica by Reuss in his great work 
on the Corals of Gosau (‘ Denkschr. der Wien Akad. der Wissensch.,’ t. vu, p. 94, pl. 8, 
figs. 4—6, 1854). 

Reuss’s admirable delineation of the species enables the British form to be recognised 


30 BRITISH FOSSIL CORALS 


at once, and it even possesses the curious transverse arrangement of the walls of some 
calices which renders the comprehension of Reuss’s sixth figure rather difficult. 

The Astrocenia have been fully considered in the ‘ Monograph of the Liassic Corals’ 
(Pal. Soc., 1867). 

Astrocenia decaphylla is a rather variable species, on account of the preponderance or 
deficiency, as the case may be, of ccenenchyma. ‘The size of the costz is limited by the 
eoenenchyma, and when this is very deficient they are almost rudimentary. 

There are ten principal and ten secondary septa; the secondary are the smallest, and 
do not reach the styliform columella like the primary. They are slightly spined towards 
their mner margin. ‘The coste are small. The columella is well developed, and is 
essential and styliform. The shape of the calices varies; im some places they are 
circular, and in others polygonal. | 

Locality. Waldon. In the Collection of William Vicary, Esq., F.G.S., Exeter. 

The British specimens are not to be distinguished from those of the Hippurite Chalk 
of Gosau, or of the Craie tuffeau of Corbiéres. 

Astrocenia decaphylla was a very persistent form. It resembles in some of its peculiar 
structures the Astroccenias of the Lias, and a specimen from the Miocene coralliferous 
strata of Jamaica! cannot be distinguished from the form from Gosau. 


Genus—IsastREA. 
IsastR@A Haxponrnsis, Duncan. Pl. XI, figs. 7 and 8. 


The corallum is hemispherical. 

The calices are large, irregular in size, very deep, and rather quadrangular. 

The wall is thin. 

The septa are crowded, small, long, and there are five cycles of them in the largest 
calices. 

There is a disposition to serial growth in some calices. » 

Diameter of the largest calices, nearly 4 inch. 

Locality. aldon. In the Collection of William Vicary, Esq., F.G.S., Exeter. 

The depth and size of the calices, their thin walls, and the numerous septa, distinguish 
this species, whose closest allies are /sastr@a lamellosissima, Michelin, sp., from the Craie 
tuffeau of Uchaux, Jsastrea Haidingeri, Ed. and H., from the same formation at 
m the Eastern Alps, and Jsastrea tenuistriata, M‘Coy, sp., of the Inferior 


oR 
Piesting, 


Oolite. 


1 Duncan, ‘ West Indian Corals,’ ‘ Quart. Journ. Geol. Soc.,’ Nov., 1863, vol. xix, page 440. 


FROM THE GAULT. 31 


List or Uprrr Greensanp Corats rrom Hanpon. 


1. Placosmilia cuneiformis, Ed. and H. 
2. fi Parkinsoni, 

3. PF magnifica, Duncan. 

4. Peplosmilia depressa, K. de ¥romentel. 
5. Astrocenia decaphylla, Kid. and HH. 

6. Tsastrea Haldonensis, Duncan. 


Peplosmilia Austeni, Kd. and H., and Favia stricta, Hd. and H., are also found at 
Haldon. They have been already noticed as Upper Greensand forms. 


I].—Corats From THE GAULT. 


Only six well-marked species of Corals were known to MM. Milne-Edwards and 
Jules Haime as having been found in the Gault. They were all simple or solitary forms, 
and such as one would expect to find in moderately deep water. It is evident that the 
area occupied by the English Gault was not the Coral tract of the period. The resem- 
blance of the Coral-faunas of the Gault and the London Clay is somewhat remarkable, 
and probably the physical conditions of the area during the deposition of the strata were 
not very dissimilar. 

The following pages contain the descriptions of some species which were not known to 
MM. Milne-Edwards and Jules Haime, and some notices of the most important forms 
they described. 


MADREPORARIA APOROSA. 
Famtty—TURBINOLIDA. 
Sub-family—CarYOPHYLLIN&. 
Division—CaRyYOPHYLLIACES. 


Genus—CARYOPHYLLIA. 


MM. Milne-Edwards and Jules Haime have changed the generic term Cyathina into 
that of its predecessor Caryophyllia ; consequently Cyathina Bowerbanki, Wd. and H., is 
now called Caryophyllia Bowerbanki, Ud. and H. (‘ Hist. Nat. des Corall.,’ vol. ii, p. 1%). 


32 BRITISH FOSSIL CORALS 


A very interesting variety of this species is in the Rev. T. Wiltshire’s Collection, and 
has its costae running obliquely to the long axis of the corallum. They are pol 
granulated (Pl. XII, figs. 8, 9). 


Division—TrocHocyaTHACE. 


Genus—TROCHOCYATHUS. 
1. Trocnocyatuus Harveyanus, Wd. and H. 


This species was described by MM. Milne-Edwards and Jules Haime in their‘ Mono- 
graph of the British Fossil Corals,’ Part I, p. 65. They associated it with two species, 
which are, as they suggest, indistinguishable, viz. Trochocyathus Koenigi and Trochocya- 
thus Warburtom. ‘The first of these species is the Zurbcnolia Koenigi of Mantell. 

An examination of a series of specimens attributed to Trochocyathus Harveyanus, Kd. 
and H., and the consideration of the value of the Trochocyathi just mentioned, have led me 
to recognise five forms of Zrochocyathi breves, all closely allied and well represented by 
the original type of Zrochocyathus Harveyanus, Bd. and H. When placed ina series with 
this Zrochocyathus at the head, there is a gradation of structure which prevents a 
strictly specific distinction being made between the consecutive forms; but when the 
first and the last forms are compared alone, no one would hesitate to assert that there is 
a specific distinction between them. All the forms are simple, short, and almost hemi- 
spherical ; all have four cycles of septa, and the same proportion of pali. These are the 
primary and most essential peculiarities of the genus. 

‘The cost differ in their size, prominence, ornamentation, and relation to the septa in 
some of the forms ; and the exsert nature of the septa, their granulation, and the size of the 
corallum, also differ. The structural differences are seen in many examples, and are 
therefore more or less persistent ; nevertheless it is found that, whilst several specimens 
have the septa springing from intercostal spaces instead of from the ends of the costz, one 
or more, having all the other common structural peculiarities, present septa arising from 
the costal ends. This method of origin can hardly constitute a specific distinction. I 
propose to retain Zrochocyathus Harveyanus as the type of a series of forms the sum of 
whose variations in structure constitutes the species. 


Variety \ (Pl. XIII, figs. 1, 2).—The corallum is nearly double the size of the type; its 
septa are rather exsert, and are very granular. 

The coste are very prominent, ridged, marked with numerous small pits, and are 
continuous with the septa. 


FROM THE GAULT. 33 


The epitheca is waved and well developed. The spaces between the larger cost are 
more or less angular. 

The peduncle is large. 

Locality. Gault, Folkestone. In the British Museum. 


Variety 2 (Pl. XII, figs. 3, 4).—The corallum is as large as that of variety 1, but it 
is more conical. 

The costee are less pronounced, and the septa, which are more granular than those of 
variety 1, arise from the intercostal spaces. The costal ends are very elegant in shape, 
and form a margin of rather sharp curves, side by side. 

Locality. Gault, Folkestone. In the British Museum. 


Variety 3 (Pl. XII, figs. 1, 3, 4; and Pl. XIII, fig. 13).—The corallum is rather flat, 
but hemispherical. 

The septa are not exsert, and they arise from the costal ends. 

The coste are equal; none are more prominent than others. They are all rather 
broad, flat, and beautifully ornamented with diverging curved lines. ‘Their free ends are 


equal and curved. 
Locality. Gault, Folkestone. In the Collection of the Rev. T. Wiltshire, F'.G.S. 


Variety 4.—TYhe corallum and coste are like Variety 3, but the septa arise from the 


intercostal spaces. 
Locality. Gault, Folkestone. In the Collection of the Rev. T. Wiltshire, F.G.S. 


Variety 5 (Pl. XII, fig. 2)—The corallum is rather more conical inferiorly than in 
Varieties 3 and 4. 

The septa are exsert, and project slightly beyond the costal margin. 

The costz are all rudimentary. 

The epitheca is well developed, and reaches up to the septa. 

Locality. Gault, Folkestone. 


The forms may be distinguished as follows : 

The type. 

Variety 1. 
2. 


c 
(Variety 3. 
he 
ee 


With more or less ridged cost 


With nearly equal flat costze 


Coste rudimentary Variety 5. 


The type. 
Septa arising from the costal ends Variety . 


ome : iety 2. 
Septa arising from the intercostal spaces ai r 


34, BRITISH FOSSIL CORALS 


All the forms have four cycles of septa and pali before the first, second, and third 
orders. 
An ill-developed and monstrous form is shown in Pl. XIV, figs. 1—5. 


2. TrocnocyatHus WiutsHire!, Duncan. Pl. XIV, figs. 10O—12. 


The corallum is straight, conical, and either cylindrical above or compressed. Its 
base presents the trace of a peduncle for attachment. 

The epitheca is scanty and in transverse masses. 

The coste are distinct and subequal. 

The calice is very open and rather deep. 

The septa are unequal, hardly exsert, and broad at the margin of the calice. There 
are four cycles of septa, and six systems. 

The pali are large, and are placed before all the cycles except the last. 

The columella is rudimentary. 

Height, ;3,ths inch. Breadth of calice, ;%ths inch. 

Locality. Gault, Folkestone. In the Museum of the Royal School of Mines, and 
in the Collection of the Rev. T. Wiltshire, F.G.S. 

This species is closely allied to Zrochocyathus conulus, Phillips, sp. The compressed 
calice, the rudimentary columella, and the shape of the corallum, distinguish the new 
species from Zrochocyathus conulus. 


Genus—LEPTOCYATHUS. 
1. Leprooyataus Graciiis, Duncan. P1*XIFL, figs. 5—8. 


The corallum is small, flat, and circular in outlgas 

The cost are very prominent, and join exsert septa. The primary and secondary 
coste are very distinct, and the ‘offfs’less so. All the coste unite centrally at the base. 
Many are slightly curved. 

The septa are thick externally, ver Yoepiausal, an internally, and the largest are more 
exsert than the others. There are six systems and four cycles of septa. 

The pali are small and exist before all the septa. 

The columella is very rudimentary. 

The calicular fossa is rather wide and shallow. 

Height, hardly 7'5th inch. Breadth, 5% ths inch. 

Locality. Gault, Folkestone. In the British Museum. 

This species is very closely allied to Leptocyathus elegans, Kd. and H., of the London 


FROM THE GAULT. 35 


Clay. Leptocyathus elegans has not a flat base, and it has very granular septa. Moreover, 
its coste are large and small in sets. Nevertheless the alliance is of the closest kind. 


Genus—BaTHYCYATHUS. 


MM. Milne-Edwards and Jules Haime described a species of this genus in their 
‘Monograph of the British Fossil Corals,’ Part I, pp. 67, 68. Two specimens in the 
Collection of Rev. T. Wiltshire present all the appearances recognised by those distinguished 
authors. The cost are very granular, and not in a simple row. In one specimen the 
breadth of the base is very great (Pl. XII, figs. 5—7). 


Famity—TURBINOLID A. 
Sub-Family—TorBinouin &. 
Division-—TvBInoLiace’. 
Genus—SMILOTROCHUS. 


Some species of this genus were described amongst the Corals from the Upper Green- 
sand,! and one was noticed as belonging to this geological horizon which should have 
been included with the Lower Greensand forms. 

The Upper Greensand Smlotrochi are— 


Smilotrochus tuberosus, Kd. and H. 
if elongatus, Duncan. 
* angulatus, _,, 


There are four species of the genus found in the Gault, which are all closely allied. 
One of them cannot be distinguished: from Smilotrochus elongatus of the Upper 
Greensand. 

The specimens of this species found in the Upper Greensand are invariably worn and 
rolled, and are generally in the form of casts; but in the Gault the structural details are 
well preserved, and even the lateral spines on the septa are distinct. 

The Gault forms are shorter and more cylindro-conical and curved than those from the 


Upper Greensand. 
1 See ante, p. 19. 
6 


36 BRITISH FOSSIL CORALS 
The species of the genus Siilotrochus from the Gault are as follows : 


1. Smilotrochus elongatus, Duncan. 


* cylindricus, ,, 
- granuatus, ,, 
cnsignis, ¥ 


1. SmiLorRocuus ELoNGaTUS, Duncan. Pl. XII, figs. 10—16; Pl. XIII, figs. 10—12; 
and Pl. XIV, figs. 13—15. 


This species is described at page 19 of the first number of this Part, and is figured in 
Pl. VI, figs. 1—6. 

Locality. Folkestone. In the Collection of the Royal School of Mines. 

The lateral spines of the septa are very well marked, and the costz are equal in size in 
this species. Its septal number varies, on account of the very late perfection of the fourth 
cycle of septa. 


2. SminoTRocHus cyLinDRicus, Duncan. PI. XIV, fig. 16. 


The corallum is small, cylindrical, nearly straight, and has a truncated base. 

The cost are equal, very distinct above, and rudimentary below and in the middle. 
‘They are marked with a few large granules in one series. 

The septa are subequal, very exsert, thin, close, and marked with large granules, 
few in number. ‘The septa are in six systems, and there are three cycles. 

Height, ;8ths inch. Greatest breadth, rather less than ;’pths inch. 

Locality. Gault, Folkestone. In the Collection of the Rev. T. Wiltshire, F.G.S. 


3. SMILOTROCHUS GRANULATUS, Duncan. Pl. XIV, fig. 17. 


‘The corallum is conico-cylindrical in shape, and has a more or less truncated base. 

The cost are subequal, prominent, very granular, and distinct superiorly. 

The septa are subequal, thick, and very granular. The septa are in six systems, and 
there are three cycles. 

Height, 5ths inch. Breadth, 535ths inch. 

Locality. Gault, Folkestone. In the Collection of the Rev. T. Wiltshire, F.G.S. 


FROM THE GAULT. 37 


4. SmiLorrocnus insienis, Duncan. PI. XII, fig. 17 ; and Pl. XIV, fig. 18. 


The corallum is trochoid, short, and has a wide calice, and a conical and rounded 
base. 

‘The calice is circular in outline; the fossa is deep and small, and the septa are wide, 
exsert, curved above, and so marked with one row of granules that their free margin 
appears to be spined. ‘There are three cycles of septa, and the orders are nearly equal as 
regards size. 

The costz are large, prominent, broad at their base, and are marked with one row of 
cranules on the free surface. 

Height, oths inch. Breadth of calice, 7’pths inch. 

Locality. Gault, Folkestone. In the Collection of the Rev. T. Wiltshire, F.G.S. 

An analysis of the genus will be found after the description of the species from the 
Lower Greensand. 


There is a compound or aggregate Madreporarian found in the Gault of Folkestone. 
It has much endotheca, and resembles worn specimens of the well-known /folocystis 
elegans of the Lower Greensand. ‘The specimens are not sufficiently well preserved for 
indentification with any genus. 


Family—FUNGID A. 

Sub-family—FuNGIN&. 

Genus—MIcRABACTIA. 
1. Micrapacta Frrroni, Duncan. PI. XIV, figs. 6—9. 


The corallum is nearly hemispherical in shape. Its base is flat, and extends beyond 
the origin of the septa in a sharp and uninverted margin. The breadth of the base exceeds 
the height of the corallum. 

The coste are flat, straight, convex externally at the calicular margin, and equal. 

The septa are unequal, much smaller than the costa. ‘There are four cycles of septa, 
im six systems. 

‘The synapticule between the septa are large. 

Height, ths. Breadth, nearly $ inch. 

Locality. Gault, Folkestone. In the Collection of the Rev. ‘I’ Wiltshire, F.G.S. 


38 


BRITISH FOSSIL CORALS 


The flat base, the flat costa, and the limitation of the septal number to four cycles, 
distinguish this species from MJicrabacia coronula’ of the Upper Greensand, and from 
Micrabacia Beaumontii*, Ed. and H., of the Neocomian. 


List or New SprEcIES FROM THE GAULT. 


Variety of Caryophylha Bowerbanki, Ed. and H. 

Five varieties of Zrochocyathus Harveyanus, Ed. and H. 
Trochocyathus Wilishiret, Duncan. 

Leptocyathus gracilis, a 


Smilotrochus elongatus, 


29 


= granulatus, ,, 
Pe MSIGNIS, ¢ 
be. cylindricus, ,, 


Micrabacia Fittoni, 


Il].—List oF SpEcIES FROM THE GAULT. 


. Caryophyllia Bowerbanki, Kd. and H., and one variety. 
. Trochocyathus conulus, Phillips, sp. 


e Wiltshiret, Duncan. 
- Harveyanus, Kid. and H., and five varieties. 


. Bathycyathus Sowerby, Ed. and H. 
. Leptocyathus gracilis, Duncan. 

. Cyclocyathus Filton, Kd. and H. 

. Smilotrochus elongatus, Duncan.? 


a) granulatus, of 
es cylindricus, ,, 
~ MSignis, ns 


. Trochosmilia sulcata, Ed. and H. 
. Micrabacia Fittoni, Duncan. 


' «Hist. Nat. des Coral.,’ vol. iii, p. 30. 
2 Ibid., p. 30. 
$ Common to the Gault and Upper Greensand. 


FROM THE LOWER GREENSAND. 39 


IV.—Corats From THE Lower GREENSAND. 


One species of Coral was described by MM. Milne-Edwards and Jules Haime from the 
Lower Greensand, in their ‘ Monograph of the British Fossil Corals.’ 

Fitton had noticed a compound Coral in the Lower Greensand, and named it Astrea 
in his “ Essay on the Strata below the Chalk,” ‘Geol. 'Trans.,’ 2nd series, vol. iv, p. 352, 
1843. In 1847 he called the species Astrea elegans, and Lonsdale separated it from the 
Astreide wnder the name Cyathophora ? elegans in 1849 (* Quart. Journ. Geol. Soc.,’ 
vol. v, pt. 1, p. 83, pl. iv, figs. 12, 15, 1849). 

MM. Milne-Edwards and Jules Haime recognised the quadrate arrangement of the 
septa of this species, and classified it amongst the Rugosa, in the family Stauwride. The 
Holocystis elegans, Fitton, sp., is a very good species, and specimens are found varying in 
the size of the corallum and of the calices. 

Since the publication of their ‘ Monograph on the British Fossil Corals,’ MM. Milne- 
Edwards and Jules Haime have named a species from Farringdon Smilotrochus Austeni 
(‘ Hist. Nat. des Corall.,’ vol. ii, p.71). Ihave noticed it inadvertently in my description 
of the Upper Greensand Corals, p. 19, and Pl. VII, fig. 12. In order to complete this 
part it is introduced here again. 


Famity—TURBINOLIDA. 
Division—TuURBINOLIACES. 
Genus—SMILOTROCHUS. 


1. SmiLorrocuus Avsteni, Hd. and 7. Pl. VII, fig. 12. 


The corallum is regularly cuneiform, very much compressed below, and slightly 
elongate. 


1 The following authors have written upon the Fossil Corals of the Gault : 
MM. Milne-Edwards and Jules Haime, ‘ Monograph of the British Fossil Corals ; Pal. Soc. 
is ‘Hist. Nat des Coralliaires.’ 
Phillips, ‘ Tlust. of Geol. of Yorkshire.’ 
Mantell’s ‘ Geol. of Sussex,’ Lonsdale in. 
Fleming, ‘ British Animals.’ 
The authors who have written upon the Corals of the Lower Greensand are— 
MM. Milne-Edwards and Jules Haime, opp. citt. 
Fitton, ‘ Quart. Journ. Geol. Soc.,’ vol. iii, p. 296, 1847. 
Lonsdale, ‘ Quart. Journ. Geol. Soc.,’ vol. v, p. 83. 
M. de Fromentel has paid especial attention to the French Neocomian Corals; and C. J. Meyer, 
Esq., F.G.S., has enabled me to study the most interesting species in his collection. 


4.0 BRITISH FOSSIL CORALS 


‘The calice is elliptical ; the summit of the larger axis is rounded. 

Forty-eight costa, subequal, straight, fine, and granular. 

Height of the corallum, about 4rd inch. 

Locality. Farringdon. 

MM. Milne-Edwards and Jules Haime do not mention where their specimen is 
deposited. Mr. Vicary, of Exeter, has a fine specimen of this Coral. 

The genus Simi/otrochus has become of some importance in the paleontology of the 
Cretaceous rocks. ‘The species are distributed as follows in Great Britain : 


Smilotrochus tuberosus, Eid. and H. 
e elongatus, Duncan Upper Greensand. 
if angulatus, ,, 


elongatus, 5, \ 
granulatus, ,, | 

oJ Cae Gault. 
am INSLINGS, - 


cylindricus, ,, 


Austeni, Ed. and H. Lower Greensand. 


Smilotrochus elongatus, Duncan, is found in the Gault and Upper Greensand. 
Smilotrochus Hagenowi, Hd. and H., is a fossil from the Maestricht Chalk (Ed. and H., 
‘Hist. Nat. des Corall.,’ vol. ii, p. 71). Smzlotrochus irregularis, K. de Fromentel, is a 
small cornute form, with rounded primary coste and rather an open calice ; it is from the 
Chalk (‘ Pal. Frang.,’ tome vii, livraison 4, Zooph., pl. 1x). 


Sub-family—CaRYoPHYLLINE. 
Division—CaRYoPHYLLIACES. 


Genus—BRACHYCYATHUS. 
1. Bracuycyatuus Orsienyanus, Ed. and H. PI. XV, figs. 8, 9. 


‘The corallum is very short. 

The coste are indistinct. 

The septa are long, very slightly exsert, granulated from below upwards, and there 
are four cycles in six systems. The primary and secondary septa are equal. The 
tertiary are a little longer than those of the fourth cycle. All are thin and straight. 

The pali are like continuations of the tertiary septa before which they are placed. 
They are granular. 

Height, =45th mch. Breadth, =®ths inch. 


FROM THE LOWER GREENSAND. Al 


Locality. Yast Shalford, Surrey. Base of the Lower Greensand; found with 
Cerithium Neocomiense, D’Orb.; Exogyra subplicata, 'qm.; Arca Raulini, Leym.; Tere- 
bratula sella, Sow. In the Collection of C. J. A. Meyer, Esq., F.G.S. 

The specimen upon which the genus was founded was found in the Neocomian forma- 
tion of the Hautes Alpes, at St. Julien, Beauchéne. I have added to the original descrip- 
tion, as some portions of the English specimen are better preserved than the type. 


Famiry—ASTRAIDZ. 
Sub-family— “USMILIN &. 
Division—VRoCHOSMILIACE. 


Genus—TROCHOSMILIA. 
TrocHosMitia Mryert, Duncan. Pl. XV, figs. 1—7. 


The corallum is small, cylindrical or cylindro-conical. Its base may be wide or very 
small, and was adherent. 

The epitheca is complete. 

The coste are very small, and are occasionally seen where the epitheca is worn. 

The calice is rather deep. 

The septa are crowded, unequal, spined near the axis, and form six systems. ‘I’here 
are four cycles of septa. | 

The calice is usually circular in outline, but it is occasionally compressed. 

The axial space is small. 

The endotheca is very scanty. 

Height, ;‘5ths inch. Greatest breadth, ths inch. 

Variety——The corallum is short, broad, cylindrical, slightly constricted centrally, and 
has a broad base. 

Height, 35thsinch. Breadth, soths inch. 

Locality. Bargate Stone; upper division of the Lower Greensand. Guildford, 
Surrey. Found with Avicula pectinata, Sow. In the Collection of C.J. A. Meyer, 
Esq., F.G.S. 

These small Zrochosmilie are common in the Bargate Stone, where they were 
discovered by Mr. Meyer, from whom I have obtained the names of the associated fossils. 
The presence of epitheca would apparently necessitate these fossils being placed in a new 
genus, but, after a careful examination of the bearings of the absence or presence of 


42 BRITISH FOSSIL CORALS 


epithecal structures upon the natural classification of simple Corals, I do not think the 
point sufficiently important to bring about the separation of Mr. Meyer’s little Corals 
from the Zrochosmilie. They form (i.e. the type and the variety) a sub-genus of the 
Trochosmilie. 


Sub-family—AstREINZ. 


Division—AStTRACER. 


Genus—ISASTREA. 


Isastr@#A Morris, Duncan. Pl. XV, figs. 10O—12. 


The corallum is flat and very short. The corallites are unequal, and usually five- 
sided. 

There is no columella. 

The wall is thin. 

The septa are slender, unequal, and most of them reach far inwards. There are in 
the perfect calices three cycles of septa in six systems. Usually some of the septa of the 
third cycle are wanting. 

Breadth of a calice, rather more than 3/5th inch. 

Locality. Bargate Stone, Guildford, Surrey ; with Zerebratella Fittoni, Meyer. In 
the Collection of C. J. A. Meyer, Esq., F.G.S. 

This small /sastr@a is usually found as a cast, and the restored drawing is taken from 
an impression. ‘The central circular structure is due to fossilization. 

The species is closely allied to Lsastre@a Guettardana, Ed. and H., of the Lower Chalk 
of Uchaux. 


Family—FUNGIDA.. 
Sub-family—LoPHosERINE. 
Genus—TURBINOSERIS. 
Genus nov.—Tursinoseris. The corallum is simple, more or less turbinate, or 


constricted midway between the base and calice. The base is either broad and adherent, 


or small and free. 


FROM THE LOWER GREENSAND. 43 


There is no epitheca, and the costa are distinct. 
There is no columella, and the septa unite literally, and are very numerous. 


Tursinoseris Du-FRomrnrEni, Duncan. Pl. XV, figs. 13—18. 


The corallum is tall, and more or less cylindro-turbinate. 

The calice is shallow, and circular in outline. 

The septa are very numerous, long, thin, straight, and many unite laterally with longer 
ones. ‘I'here are 120 septa, and the cyclical arrangement is confused. 

The synapticulz are well developed. 

There is no columella, and the longest septa reach across the axial space. 

The costz are well developed, and often are not continuous with the septal ends. 

Height, 14%>ths inch. Breadth of calice, 142;ths inch. 

Variety —With a constricted wall and large base. 

Locality. Atherfield, in the Lower Greensand. In the Collection of the Royal School 
of Mines. 

The necessity for forming a new genus for this species is obvious. It is the neighbour 
of Zrochoseris in the sub-family of the Lophoserine. ‘This last genus has a columella, 
and the new one has none. 

The species has not been hitherto described, but it has been familiarly known as a 
Montlivaitia ; but the synapticule between the septa and costa determine the form to 
belong to the Fungide. 


V.— List or New Specius FROM 'rHE Lower GREENSAND. 


1. Brachycyathus Orbignyanus, Ed. and H. 
2. Trochosmilia Meyer, Duncan. 

3. Lsastrea Morrisi, os 

4. Turbinoseris De-Fromenteli, ,, 


VI.—List or THE Sprecius From tHE Lower GREENSAND. 


. Brachycyathus Orlignyanus, Kid. and H. 
. Smilotrochus Austeni, Ed. and H. 
. Trochosmilia Meyeri, Duncan. 


me Oo wm 


. Isastrea Morrisii, 


be) 


or 


. Turbinoseris De-Fromonteli, ,, 


for) 


. Holocystis elegans, Lonsdale, sp. 
7 


BRITISH FOSSIL CORALS 


VIT.—List oF THE SPECIES FROM THE CrEeTacrEous FORMATIONS. 
A. Upper and Lower White Chath. 


. Caryophyllia cylindracea, Reuss, sp. 
: < Lonsdalei, Duncan. 

2 Tennanti, = 
. Onchotrochus serpentinus,  ,, 


1 
2 
3 
4, 
5. Trochosmilia lava, Ed. and H., sp., and three varieties. 
6 
i 
8 
9 


ms cornucopie, Duncan. 
a Wiltshiret, - 
fe Woodwardi, ,, 
i ra granulata, *s 
10. fe cylindrica, zy 


11. Parasmilia centralis, Mantell, sp., and two varieties. 


12. - cylindrica, Ed. and H. 
13: Fittoni, A 

14. . serpentina, d 

15. - monilis, Duncan. 

16. =< granulata, ,, 


17. Diblasus Gravensis, Lonsdale. 
18. Synhelia Sharpeana, Ed. and H. 
19. Stephanophyllia Bowerbanki, Ed. and H. 


B. Upper Greensand. 


20. Onchotrochus Carteri, Duncan. 
21. Smilotrochus tuberosus, Ed. and H. | 
22. elongatus, Duncan. 

23. - angulatus, ,, 

24. Peplosmilia Austeni, Wd. and H. 

25. Cyathophora monticularia, D’Orbigny. 
26. Favia stricta, Ed. and H. 

27. ,, minutissima, Duncan. 

28. Thamnastrea superposita, Michelin. 
29. Micrabacia coronula, Goldfuss, sp. 

30. Placosmilia cuneiformis, Bd, and H. 
ol. ns Parkinsont, 59 

32. x magnifica, Duncan. 

33. Peplosmilia depressa, E. de Fromentel. 
34. Astrocenia decaphylla, Ed. and H. 

35. Isastrea Haldonensis, Duncan. 


FROM THE CRETACEOUS FORMATIONS. A5 


c. Red Chalk of Hunstanton. 


36. Cyclolites polymorpha, Goldfuss, sp. 

37. Podoseris mammiliformis, Duncan. 

38. eg elongata, < 

39. Micrabacia coronula, Goldfuss, sp., and variety. 


pv. Gault. 


40. Carophyllia Bowerbanki, Hd. and H., and a variety. 

Al. Trochocyathus conulus, Phillips, sp. 

A2, a Wiltshirei, Duncan. 

43. . Farveyanus, Ed. and H., and five varieties. 
44, Bathycyathus Sowerbyi, Ed. and H. 

45. Leptocyathus gracilis, Duncan. 

A6. Cyclocyathus Fittoni, Kd. and H. 

A7. Smilotrochus elongatus, Duncan. 


48, ¥; granulatus, ,, 
49, - IMSLgNIS, es 
50. * cylindricus  ,, 


51. Trochosmilia sulcata, Ed. and H. 
52. Micrabacia Fittoni, Duncan. 


gk. Lower Greensand. 


53. Brachycyathus Orbignyanus, Ed. and H. 
54. Smzlotrochus Austeni, 

55. Lrochosmilia Meyeri, Duncan. 
56. Lsastrea Morrisii, 
57. Zurbinoseris De-Fromenteli, Duncan. 
58. Holocystis elegans, Lonsdale, sp. 


> 


3? 


Micrabacia coronula is common to the Upper Greensand and the Red Chalk. 
Smilotrochus elongatus is found in the Gault and in the Upper Greensand. 

The number of species of Madreporaria in the British Cretaceous formations 1s 
therefore fifty-six. 

MM. Milne-Edwards and Jules Haime had described twenty-three species before this 


46 BRITISH FOSSIL CORALS. 


series was commenced. Of these I have ventured to suppress Parasmilia Mantelli, 
Trochocyathus Koenigi, and Trochacyathus Warburtoni. 

The Coral-fauna of the British area was by no means well developed or rich in genera 
during the long period during which the Cretaceous sediments were being deposited. The 
Coral tracts of the early part of the period were on the areas now occupied by the Alpine 
Neocomian strata, and those of the middle portion of the period were where the Lower Chalk 
is developed at Gosau, Uchaux, and Martigues. 

There are no traces of any Coral reefs or atolls in the British Cretaceous area, and its 
Corals were of a kind whose representatives for the most part live at a depth of from 5 to 
600 fathoms. 


PLATE X. 


CORALS FROM THE UPPER GREENSAND OF HALDON. 


. The corallum of Placosmilia cuneiformis, Ed. and H. (P. 27.) 
. Part of a septum, magnified. 

. The coste, magnified. 

. Oblique view of the costz, magnified. 

. The calice, magnified. 

). The corallum of Placosmilia Parkinson, Ed. and H. (P. 28.) 


i. The calice, magnified. 


8. The corallum of Peplosmilia depressa, EK. de From. (P. 29.) 


10. 


Ale 
12. 


. The costa, magnified. 


The calice, magnified. 


ou corallum of Placosmilia magnifica, Duncan. (P. 28.) 


die BE hohe GERM & 


ee . 
AE. 


fDi 3) 


o 


pba. falta 


ies i ek: 
i Sp sme \ 


PLATE XI. 


. The corallum of Astrocaenia decaphylla, Ed. and H. (P. 29.) 
. The same, magnified. 

. The upper part of a calice, magnified. 

. The corallum of a variety. 


. The upper part of a calice, magnified. 


. The corallum, magnified. 


‘|The corallum and calices of /sastrea Haldonensis, Duncan. (P. 30.) 


DD 


Ge, LY / 
iy 4, 


- 
VAlLe@ 
Ao, 


(A 


a 


a 


Ziff 


{ 


iy 


43 


\ 


Ni 


i! 


4 


F) 


x] 


PLATE XII. 


CORALS FROM THE GAULT. 


Fic. 
aa Varieties of Zrochocyathus Harveyanus, Kd. and H. (P. 33.) 


. Magnified view of the ends of the cost of one of the varieties. 
. A longitudinal section of a variety, slightly magnified. 

. A variety of Bathycyathus Sowerbyi, Ed. and H. (P. 35.) 

. Its costae, magnified. 

. The corallum of Bathycyathus Sowerbyi, Ed. and H. (P. 35.) 

. A variety of Caryophyllia Bowerbanki, Kd. and H. (P. 32.) 

9. Its coste, magnified. 


10 to 16. Views of Smilotrochus elongatus, Duncan. (P. 36.) 


anrnn & W 


12. Cost, magnified. 
14. The calice of a young specimen, magnified. 
16. The coste, magnified. 


17. Corallum of Smilotrochus insignis, Duncan. (P. 37.) 


Sot | 

| 

| 

| 

| 

| | 

| 

| | 

| a 

| as © 

E 
= 
K 
C 

Hh | fa 

fi 

| C 

| F 

< 
6 
C 
io 

| 

| 


é& 


PLATE XIII. 


CORALS FROM THE GAULT. 


1. A variety of Zrochocyathus Harveyanus, Hd. and H. The base. (P. 32 and 33.) 
2. Coste and septa, magnified. 
3. A variety of the same species. 
4. Coste and septa, magnified. 
13. A transverse section, magnified. 
5. Leptocyathus gracilis, Duncan. Under surface. (P. 34.) 
6. The under surface or base, magnified. 
7. A transverse section, magnified. 
8. A side view, magnified. 
9. Smilotrochus insignis, Duncan. (P. 37.) | 


10. 
11. ‘Young of Smilotrochus elongatus, Duncan. (P. 36.) 


12. 


re 
oOanNRr DaAPhwW NH — 5 


— et 
iS) 


PLATE XIV. 


CORALS FROM THE GAULT. 


|| Abnormal form of Trochocyathus Harveyanus, Ed. and H. (P. 34.) 


| Mansi views. 


. Base of Micrabacia Fittoni, Duncan. (P. 37.) 

. The same, magnified. 

. Side view of the corallum, magnified. 

. Junction of septa and cost, magnified. 

. Corallum of Trochocyathus Wiltshire’, Duncan. (P. 34.) 

. Magnified view. 

. The calice, magnified. 

. Smilotrochus elongatus, Duncan. Adult form. (P. 36.) 

. The same, magnified. 

. The calice, magnified. 

. Smulotrochus cylindricus, Duncan. Corallum, magnified. (P. 36.) 
. Smilotrochus granulatus, Dancan. Corallum, magnified. (P. 36.) 


. Smilotrochus insignis, Duncan. Corallum, magnified. (P. 37.) 


a 


De Wilde lith, 


x 
¥ 
~ 


Say 
eS 


Hints 


aA, 


GAULT 


: 
: 


PLATE XV. 


CORALS FROM THE LOWER GREENSAND. 


Fie 


| 
| 


ile 
Cis of Trochosmia Meyeri, Duncan. (P. 41.) 


- | 


Calices, magnified. 


oe ww 


6. Variety with broad base. 
7. Its calice, magnified. 
8. Part of the corallum of Brachycyathus Orlignyanus, Ed. and H. (P. 40.) 
9. Longitudinal view of the septa and pali, magnified. The notch indicates the 
commencement of pali attached to tertiary septa. 
10. Corallum (cast) of /sastrea Morrisii, Duncan. (P. 42.) 
11. The cast, magnified. 
12. Impression, magnified. 
13. The corallum of Turhinoseris De-Fromenteh, Duncan. (P. 43.) 
14. A variety. 
15. Synapticule and septa, magnified. 
16. Calice, size of life. 
17. Coste, magnified. 


18. The unusual appearance of septa ending in intercostal spaces, magnified. 


id aie 


PALAONTOGRAPHICAL SOCIETY. 


INSTITUTED MDCCCXLVII. 


VOLUME FOR 1869. 


LONDON: 
MDCCCLXX. 


PAU 


Pi 


Wty 


i) 


7 
, he 
7 8 


A MONOGRAPH 


ON THE 


BRITISH FOSSIL 


i 
i 
’ 


~ECHINODERMATA 


FROM 


THE CRETACEOUS FORMATIONS. 


BY 


THOMAS WRIGHT, M.D., F.R.S. EDIN., F.G.S., 


CORRESPONDING MEMBER OF THE ROYAL SOCIETY OF SCIENCES OF LIEGE, THE SOCIETY OF 
NATURAL SCIENCES OF NEUFCHATEL, AND SENIOR SURGEON TO 
THE CHELTENHAM HOSPITAL. 


VOLUME FIRST. 


PART THIRD. 


ON THE DIADEMAD. 


Paces 113—136; Puares XXII—XXIX, XXIXa, XX1Xs. 


LONDON: 
PRINTED FOR THE PALHONTOGRAPHICAL SOCIETY. 
1870. 


PRINTED BY 
J. EB. ADLARD, BARTHOLOMEW CLOSE. 


FROM THE GREY CHALK. 113 


The pores are scarcely bigeminal near the summit. The primary tubercles are 
numerous, nearly uniform in size, and closely set together; the mouth-opening is very 
small, and lies in a deep depression. 

Pseudodiadema variolare has the test in general more depressed, the base wider, and 
the pores more distinctly bigeminal in the upper fourth of the zones; the base is wider, 
more convex, and less contracted than in Ps. Brongniarti. 

Locality and Stratigraphical Position.—Pseudodiadema Brongniarti has been col- 
lected from the Grey Chalk near Folkestone, from which stratum all the large fine speci- 
mens in the British Museum, and those in the Rev. 'T. Wiltshire’s cabinet figured in 
this Monograph, have been obtained. The Red Chalk of Hunstanton Cliff has yielded 
a few examples, two of which, from Mr. Rose’s and the Rev. 'T. Wiltshire’s collections, 
are figured in Pl. XXI B. Forms referred to this species have been collected from 
the Chalk-marl of Maiden Bradley, Dorset, and the Chloritic Marl, Somerset. 

Foreign Localities—Professor Pictet, of Geneva, kindly gave me several type speci- 
mens of this Urchin collected from the Gault of the Perte du Rhdéne (Ain), which so much 
resemble Professor Agassiz’s figures of this species from the same locality that my speci- 
mens might have been the originals of the drawings in his ‘ Hchinodermes foss. de la 
Suisse.’ It is found likewise at Escragnolle (Var), Montagne des Fis (Savoie), where it is 
an abundant fossil in the Etage Albien or the Gault. M. Desor, in addition to these 
localities, gives Clar and ‘La Presta, as places where this species is abundant. 

History.—¥ rst figured by M. Al. Brongniart’ in his ‘ Description de la Perte du Rhone,’ 
under the name Crdariles variolaris (7), as a characteristic fossil of the Craie Marneuse ; 
afterwards (1840) it was described and figured by Professor Agassiz in the ‘ Echinodermes 
foss. de la Suisse’ as Zetragramma Brongniarti ; afterwards (1856) it was removed by 
M. Desor into his genus Pseudodiadema, where it now remains. 


Genus—Pepinopsis, Cotfeav, 1863. 


Test large, round, inflated, sometimes subconical. Poriferous zones wide and straight ; 
the pores bigeminal throughout, forming at the upper surface and ambitus two distinct 
_ rows, which become more blended together at the infra-marginal region, and are distinct 
at the base. ‘Tubercles of both areas small, and nearly the same size; summit of the 
_ boss finely crenulated, and the mammillon perforated ; the tubercles disposed in regular 
rows, the number varying in the different species, and always diminishing as they 
approach the summit; coronal plates long, narrow, and granular; mouth-opening large, 
peristome moderately developed, circumference slightly incised, apical disc small, sub- 
circular, elements feebly united, absent in the specimens known. 


' “Description géologique des Environs de Paris,’ troisiéme édition, 1835, p. 174, pl. m, fig. 9. 


1) 


114. PEDINOPSIS 


This genus was established by M. Cotteau in his memoir! on ‘Les Hchinides 
des Pyrénées,’ and placed near Pseudodiadema, with which it has many affinities; the 
tubercles are crenulated and perforated, and the pores bigeminal throughout, a character 
which is in part possessed by Ps. Brongniarti, variolare, &c. It has affinities with 
Salmacis in the number of its pores ; in this genus, however, they are arranged in triple 
oblique pairs, whilst in Pedinopsis they are regularly bigeminal throughout. 

The original specimen was collected from the Neocomian strata of Aude, where it is 
rare ; another of the same species has been found at Caussols (Var); the one which I 
now figure was obtained from the Chloritic Marl near Chardstock, Somerset. 


Prepinopsis Wresti, Wright, nov. sp. Pl. XIV, fig. 1; Pl. XXIX a. 


Test moderate in size, circular, inflated at the sides, convex on the upper surface, and 
flat at the base ; ambulacra with two complete marginal rows and two inner incomplete 
rows of tubercles; inter-ambulacra with six rows of turbercles at the ambitus, the two 
central of which are complete, and the four lateral incomplete ; tubercles nearly all of the 
same size; pores uniformly bigeminal throughout. 

Dimensions.—Height one inch; transverse diameter one inch and seven tenths ; 
mouth-opening six tenths of an inch. 

Description.—This rare Urchin forms the type of a new genus, established by my friend 
M. Cotteau for a similar rare species from the Neocomian of Caussols (Var). ‘The test is 
of moderate size, circular at the ambitus, inflated at the sides, convex on the upper 
surface, and flattened at the base. The ambulacral areas are moderately wide, and retain 
their proportional diameter throughout the area; they have two rows of marginal 
tubercles of small size, which extend from the peristome to the disc (PI. XXIX a, fig. 1), 
and are very regular both in size and arrangement ; between the base and upper surface two 
other rows occupy the centre, so that this area at the ambitus is furnished with four rows 
of tubercles (Pl. XXIX a, fig. 1g), the two inner rows of which disappear at the lower 
seventh (fig. 1 /) and upper third of the areas (fig. 1 d) ; the tubercles are nearly all of the 
same size; around the base of the bosses some fine granules are sparsely distributed, 
which form imperfect circlets around them (Pl. XXIX a, figs. 1 d,e,f,g, 4). None of 
the marginal rows in the specimen is complete, so that the exact number in each series 
cannot be ascertained ; but as thirty can be counted in one incomplete column, six more 
may be fairly estimated as wanting (fig. 1 a, 4, ¢). 

The poriferous zones are moderately wide, and very uniform in diameter throughout, 


1 « Fchinides foss. des Pyrénées,’ p. 16. Extrait du ‘Congrés scientifique de France,’ 28° session, 
tenue a Bordeaux, t. iii, pl. ix, 1863. 


FROM THE CHLORITIC MARL. 115 


expanding, however, near the mouth, where an increased number of holes seem almost 
completely to encircle the peristome (fig. 1 4); throughout the zones the pores are very 
regularly bigeminal, and are grouped into distinct ranges, the small plates forming these 
zones being beautifully dovetailed into each other (figs. 1 d, gy, 4); by this arrangement 
there are from six to seven pairs of holes opposite each ambulacral plate, which gives 
36 < 6 = 216 pairs of holes in each zone. 

The inter-ambulacral areas are rather more than double the width of the ambulacral ; 
the individual plates are long and narrow, being only a little deeper than those of the 
latter ; two rows of tubercles occupy the centre of the plates, and extend very regularly in 
size and disposition from the peristome to the disc; on the zonal side of this central row 
a second row extends from the peristome over three fourths of the area, and between the 
central row and the median suture there is another row of the same length ; in addition 
to these six rows a few additional tubercles are introduced at the zonal and median sides 
of the widest ambital region of the area. I have shown this character in figs. 1 g and 4, 
and the gradual disappearance of the tubercles in the upper part of the area is seen in figs. 
1 a and ¢, and in its lower part in figs. 1 d and 4; between the tubercles a number of 
small granules are sparsely distributed over the surface of the plates. 

The tubercles in this genus are very uniform in size and structure throughout both 
areas ; the boss rises suddenly from the surface of the plate, without any areolar depres- 
sion ; its summit is very finely crenulated only where it closely embraces the mammillon, 
which is small, prominent, and perforated at the summit; in the widest part of the 
areas the granules form imperfect circlets around the tubercles, and an increased orna- 
mentation at the ambitus and base ; at the upper surface they become more sparse in the 
ambulacra (fig. 1 7), and are almost entirely absent in the inter-ambulacra (figs. 1 a, c). 

The mouth-opening is nearly circular, about one third the diameter of the test (fig. 4) ; 
the peristome is slightly incised and unequally divided, the ambulacral being much longer 
than the inter-ambulacral lobes, allowing a wider space for the development of pedal pores 
around the peristome. 

The apical disc is absent, and the upper part of the test is unfortunately broken off, 
so that no indication of the size or form of the disc remains imprinted on the mould. 

Affinities and Differences.—Vhe only two species of this genus at present known 
are Pedinopsis Meridanensis, Cotteau, from the Neocomian of Aude; and P. Wiesti, 
Wright, from the Chloritic Marl near Chardstock. These Urchins resemble each other 
very much; P. Meridanensis (Pl. XXIX a, fig. 2') has a greater number of tubercles ; 
and they are likewise larger and more regularly disposed on the plates; the poriferous 
zones are wider, and the holes larger than in P. Wiesti. The bigeminal character of the 
zones, the smallness of the tubercles, and the thinness of the shell, are special characters 
by which this Urchin can be readily distinguished from all other congeners. 


1 Copied from M. Cotteau’s ‘ Paléontologie Frangaise,’ pl. 1125, for comparison with the English 
species. 


116 ECHINOCYPHUS 


Locality and Stratigraphical Position —This specimen was collected by Mr. Wiest, from 
the Chloritic Marl near Chardstock, Somerset, where it is associated with several species 
of Hehinide that are characteristic of the Upper Greensand formation. 


Ecuinocyputs, Cotteau, 1860. 


GLYPHOCYPHUS (pars), Desor, 1856. 
CypHosoma (pars), Woodward, 1857. 
Ecuinocyrputs, Cotteau, 1860. 


Test small, circular, moderately high, more or less inflated on the upper surface, very 
concave at the base. Poriferous zones straight, and composed of simple pores throughout. 
Ambulacral and inter-ambulacral tubercles nearly the same size in both areas, bosses 
crenulated, mammillon not perforated. In many specimens the ambulacra have only one 
row of tubercles instead of two, their normal number. 

The inter-ambulacral plates are marked at the base with more or less well-marked 
sutural and horizontal impressions. Peristome moderately large, subcircular, sunk in a 
depression, and provided with slight lobes. Apical disc unknown, opening elongated 
and subpentagonal, as indicated by the impression. 

‘This genus was established to receive certain species referred by some authors to the 
genus Glyphocyphus, by others to Cyphosoma. These species, according to M. Cotteau, 
are distinguished from @lyphocyphus by their imperforate tubercles, their horizontal sutural 
impressions, and less solid apical disc. ‘Their crenulated and non-perforated tubercles 
bring them into near relation with certain species of Cyphosoma, as C. Delamarret and 
C. magnificum, var. sulcatum, which show at the base of their mter-ambulacral plates some 
traces of sutural impressions ; but these feeble depressions, however, cannot be compared 
with the deep horizontal grooves that characterize Lehinocyphus, and impart to the few 
species composing this group the physiognomy of Glyphocyphus, with which M. Desor 
placed them. 


KcHINOCYPHUS DIFFICILIS, 4gass. Pl. XXII, figs. 1 a, 6, 2 a, 6, ¢, d, 4. 


CYPHOSOMA DIFFICILE, Agass. Catal. Syst. Ectyp., Mus. Neocom., p. 12, 1840. 
= — Agassiz et Desor. Catal. rais. des Echinid., Ann. Sc. Nat., 
3e série, tom. vi, p. 352, 1846. 
— a Bronn. Index Paleontologicus, p. 381, 1848. 


DIADEMA ROTULARE, McCoy. Mesozoic Radiata, Ann. of Nat. History for 
December, 1848. 
a Maccoy1, Forbes. Catalogue of British Fossils, Morris, 2nd ed., 
p. 76, 1854. 
— ROTATUM, Forbes. Memoirs of Geological Survey, Decade V, p. 3, 


1856. 


FROM 'TTHE GREY CHALK. C7 


GLYPHOCYPHUS DIFFICILIS, Desor. Synops. des Echinides foss., p. 104, 1857. 

CYPHOSOMA DIFFICILE, Woodward (pars). Mem. of Geol. Survey, Appendix 
to Decade V, p.3, 1858. 

GLYPHOCYPHUS DIFFICILIs, Dujardin et Hupé. Uist. Nat. des Zooph., Eechinid., 
p. 513, 1862. 

ECHINOCYPHUS DIFFICILIs, Cotteau. Pal. Frangaise, Ter. Crétacé, t. vii, p. 708, 
pl. 1174, 1866. 

— ROTATUS, Cotteau (pars). Ibid., t. vii, p. 711, pl. 1174-75. 


‘Test small, subcircular, moderately inflated on the upper surface, flattened at the 
base, concave around the mouth, and rounded at the sides; poriferous zones narrow, 
straight, and subfiexuous at the ambitus; pairs of pores in a single series ; ambulacral 
areas narrow above, enlarged at the ambitus, with two rows of tubercles, one of which is 
often abortive; inter-ambulacral areas wide, having two rows of tubercles; plates pos- 
sessing small, strongly radiated areola, and divided by deep sutural impressions ; apical 
disc equal to the oral opening, flat, and finely granulated. 

Dimensions.—Transverse diameter eight tenths of an inch ; height four tenths of an inch. 

Description —This Urchin has long been mistaken for a Diadema, and is catalogued in 
some lists of Upper Greensand fossils as D. rotatum. Small specimens, measuring from four 
to five lines in diameter, are abundant in the Upper Greensand of Warminster, and may 
readily be distinguished from Diademas by the sutural impressions on their plates, the 
small radiating ribs on many of the large areole, and the irregularity of the two rows of 
ambulacral tubercles, one of which is often abortive. The larger and taller specimens 
agree with the published mould of Professor Agassiz’s type specimen of Cyphosoma difficile, 
to which this Urchin is now referred. 

Almost all the examples I have collected from the Upper Greensand are small and 
well preserved; the few I have from the Lower Grey Chalk are larger, and show the 
sculpturing of the plates better. The test is subcircular, or slightly pentagonal, mode- 
rately convex on the upper surface, flattened at the base, and rounded at the border. 
The poriferous zones are narrow, straight above and below, and subundulated at the 
ambitus. ‘I'he pores are small, and form a single series of pairs throughout from the 
mouth to the disc (fig. 2 7); the pairs of pores are separated horizontally by small promi- 
nent ribs, more or less granular, which correspond to the poriferous plates, and are pro- 
longed to the base of the tubercles. The ambulacral areas are narrow, and provided with 
two rows of tubercles; in some specimens from the Grey Chalk these rows are regular 
and the tubercles of equal size, but in many specimens from the Upper Greensand, 
as in figs. 2 a, 6, c, d, one row becomes abortive, and the other only is developed ; 
more than half the specimens that have passed through my hands have been thus 
formed. The inter-ambulacral areas are wide, with two rows of tubercles a little 
larger than those occupying the ambulacra; in large specimens there are ten or twelve 
tubercles in each row, the number depending on the age of the individual. Between 
the ambitus and peristome the areolas are well developed, and surrounded by 


118 ECHINOCYPHUS 


circles of large sub-elliptical granules, that are prolonged in a radiated manner towards 
the base of the mammillon (fig. 2d). The coronal plates at their lower border are 
marked by a horizontal depression, as if the lower half of the plate had been scooped out 
(fig. 2 a) at the expense of a portion of the granular circle. The boss is prominent, 
and its summit is marked with feeble crenulations. The mammillon is always large and 
unperforated. The miliary zone is wide in its upper part, and the surface of the plates is 
covered with a very fine granulation; at the ambitus it is filled with two or three rows of 
large granules, which diminish below into a single series, and take a zigzag direction 
towards the peristome. 

The small mouth-opening, about one third the diameter of the test, is lodged in a 
concave depression (fig. 2 4). The peristome is circular, and nearly equally lobed, and 
the incisions are shght. 

‘The apical disc, which is very rarely preserved (Pl. XXII, fig. 4), equals the oral 
opening in size, and is flat and finely granulated; the cordate ocular plates are rather 
large, and perforated close to the ambulacral margin; the large ovarial plates are per- 
forated, and the madreporiform body occupies the right anterior plate ; the single posterior 
plate is wanting. 

Affinities and Differences.—Yhis species very much resembles Cyphosoma rotatum, Forb., 
of which it proves to be a variety ; the irregularity of the two rows of ambulacral tubercles, 
one of which is often abortive, is not, according to our observations, a persistent character, 
and for this reason we regard £7. diffcilis and LF. rotatus as varieties of one form. M. 
Cotteau, however, considers them distinct, and takes as a diagnostic character the single 
row of tubercles in the ambulacral areas, as this difference does not appertain to age, 
seeing that it is found in small as well as in large specimens. chinocyphus dificilis, 
Ag., may be confounded with G/yphocyphus intermedius, Cott., which has a similar dis- 
position of the ambulacral tubercles, only that they are perforated in Glyphocyphus 
and unperforated in Lehinocyphus. 

Locality and Stratigraphical Position.—I have several specimens which I collected 
from the hard Grey Lower Chalk near Folkestone, and numerous smaller examples from 
the Upper Greensand near Warminster ; from the latter formation a very large specimen 
was obtained, now in the cabinet of G. E. Sloper, Esq., Devizes. ‘The late Dr. S. P. 
Woodward states that it measures nine lines in diameter, and four and a half in height ; 
both rows of ambulacral areas are well developed, and the miliary granules form a 
prominent framework to the tubercles. 

In France, M. Cotteau records this species from the Htage Cénomanien of Vellclaire 
(Haute-Sadne), and Présagny (Eure), where it is rare. 

History.—First catalogued as Diadema rotulare, Ag., from Durdle Door, Dorset, by 
Prof. McCoy, in his ‘ Mesozoic Radiata,’ in 1848, and afterwards by Prof. E. Forbes, as 
Diadema rotatum, from the Upper Greensand of Warminster, in the second edition 
of the ‘Catalogue of British Fossils,’ 


FROM THE GREY CHALK. 119 


Ecurnocyruus MeEsPILiA, Woodward. Pl. XXII, figs. 3 a, 4, ¢, d. 


CyPHOSOMA MESPILIA, Woodward. App. Decade V, Mem. of the Geol. Survey, 
p. 3, 1856. 


Test small, circular, inflated or depressed, convex above and below, with subequal 
apertures ; poriferous zones straight, simple, pairs of pores very oblique; tubercles nearly 
equal, prominent, imperforate and crenulated; areola small, radiated, with elongated 
miliary granules, sutural impressions slight; inter-ambulacra wide, with two rows of 
tubercles, seven to eight in each; ambulacra narrow, with seven to eight tubercles, alter- 
nate, and irregular. 

Dimensions.—Height three lines ; transverse diameter five lines. 

Description.—This species was discovered by my late friend, Dr. S. P. Woodward, 
who found it in the Lower Hard White Chalk along with Cyphosoma simplex, Korb., and 
Salenia granulosa, Forb. It is a pretty little Urchin, with inflated sides ; the test is nearly 
convex above and below, the poriferous zones are straight and simple, the unigeminal 
pores are very oblique, the ambulacral areas are narrow, and the tubercles, seven or eight 
in number, are alternate and irregular (fig. 3), one of the rows being partially abortive. 
The inter-ambulacral areas are wide, and have two rows of prominent tubercles, seven to 
eight in each row (fig. 3), which occupy the centre of the plates, and are surrounded by 
small radiated areole, formed by an oblique arrangement of the elongated granules 
thereon, and imparting an ornamented character to the test. The mouth-opening is more 
than one third of the diameter of the test, the peristome is nearly equally lobed, and the 
incisions are wide and deep. ‘The discal opening is larger than the oral in diameter. 

Affinities and Differences—TYhis Urchin very much resembes £. dificilis; it has, 
however, a more inflated test, with stronger radii on the areola, and larger oral and discal 
apertures. Its author considered it “a very distinct species,’ and called my especial 
attention to it. A careful comparison of the accurate figures of both species drawn on 
Pl. XXII will enable the student to appreciate the affinities and differences between them 
better than any description, however elaborate. 

Locality and Stratigraphical Position —E. mespilia has been collected only from the 
Lower Chalk, in the hard beds of which it has been found, associated with Cyphosoma 
simplex and Salenia granulosa. The only specimens I have seen are in the British 
Museum. 


120 GLYPHOCYPHUS 


Genus—GuyPHocyenus, Haime, 1853. 


ARBACIA (pars), Agassiz, 1836. 
Ecutnopsis (pars), dgassiz, 1646. 
HEMIDIADEMA, Agassiz, 1846. 
TEMNOPLEURA (pars), Sorzgnet, 1850. 
GuiyrHocyenus, Jules Haime, 1853. 
— Desor, 1856. 
a Cotteau, 1859. 


The Urchins forming this group have a small circular test, more or less inflated above 
and concave below. .'The poriferous zones are straight, depressed, and composed of uni- 
geminal pores throughout. The primary tubercles of both areas are nearly of the same 
size. ‘The boss, with its crenulated summit, is surrounded by a distinct areola, and sur- 
mounted by a small mammillon minutely perforated at the vertex. ‘The ambulacral and 
inter-ambulacral plates are marked with sutural depressions, which are wider and deeper 
on the plates of the latter areas; the surface is covered with fine, close-set homogeneous 
granules, nearly uniform in size over the whole test (Pl. XXIX B, figs. 1 and 2). The 
apical disc is a small annular structure, solidly united to the test, and forming a narrow, 
slightly elongated, oblong ring, composed of five ovarial and five ocular plates, articulated 
alternately together on the same line around the vent aperture or periprocte (fig. 2 0). 
The ovarial plates are the largest; they have a triangular shape, with tubercles external 
to the annulus, and a large hole at the apex; the right antero-lateral plate is the largest, 
and supports the madreporiform body. The rhomboidal ocular plates are wedged in 
alternately on the same line with the ovarials; and the portion external to the annulus is 
covered with numerous granules. Mouth-opening is small, subcircular, and sunk in a 
slight depression ; peristome decagonal, divided into lobes by feeble incisions. 

The genus Glyphocyphus forms among the Diapnmap& with crenulated and perforated 
tubercles a very small group, in which the ambulacral and inter-ambulacral plates are 
marked with angular impressions, and their surface covered with prominent close-set 
granulations. The solidity of the apical disc, and its firm articulation to the coronal 
plates, with the regularity as regards size and arrangement of the tubercles in both 
areas, form an assemblage of characters which readily distinguish the small Urchins of this 
group from their congeners. A considerable difference of opinion has prevailed amongst 
Echinologists regarding the true characters of the forms now comprised in this genus, 
arising, doubtless, from the smallness of the individuals themselves, the imperfect pre- 
servation of most of the specimens, and the destruction of many of the essential structures 
that distinguish them. M. Cotteau, however, had lately at his disposal a fine series of 


FROM THE GREY CHALK. 12] 


beautiful and perfect specimens of Glyphocyphus radiatus, collected by the late M. Triger 
from the Terrain Crétacé of the department of the Sarthe, and from the study of these 
M. Cotteau has been enabled to clear up doubts that had previously existed regarding the 
structure of many of the species. This genus is nearly allied to Achinocyphus, which 
presents a similar facies of lateral impressions on the ambulacral and inter-ambulacral 
plates, but is distinguished from it by having the mammillons of the tubercles perforated, 
whereas in Lchinocyphus they are imperforate. The apical disc is, likewise, solidly united 
to the coronal plates in Glyphocyphus, but slenderly so in Lehinocyphus. M. Agassiz 
proposed the genus Hemidiadema for Urchins differing from the Diademas in one 
character, that the ambulacral areas possessed only a single row of tubercles, and cited 
as the type of this group Hemidiadema rugosum, Agass., from the Upper Greensand (Htage 
Cénomanien) of Grandpré, Ardennes, a small species having the ambulacral tubercles 
as large and even larger than the inter-ambulacrals ; this Urchin, it now appears, is a true 
Glyphocyphus, with a single row of tubercles in its ambulacral areas—a character which 
is only specific, and not generic, as far as we at present know. 

The genus Lchinopsis, Agass., in which certain species of Glyphocyphus have been 
placed, consists, according to its author, of ‘ small, subconical, inflated Urchins, with the 
ambulacra nearly as large as the inter-ambulacra, and both ornamented with tubercles 
perforated but not crenulated. Mouth small, with feeble incisions, differing from the 
Diademas by the absence of crenulations on the tubercles.” To this genus was referred 
Echinopsis contexta, Ag., L. latipora, Ag., EB. depressa, Ag., all of which are forms of 
Glyphocyphus radiatus and Glypticus Koninckii, Forb. Lchinopsis pusilla, Roem., and 
Temnopleurus pulchellus, Coquand, must now likewise be added to the list. 


GiypyocypHus rapiatus, Haninghaus. Pl. XXIX B, figs. 1, 2, a, 4, ¢, d, e. 


ECHINUS RADIATUS, Heninghaus in Goldfuss. Petref. Germaniz, p. 124, tab. x], 
fig. 13, 1826. 
— — Agassiz. Prod., Mém. Soc. Nat. des Se. de Neufchatel, 
t. i, p. 196, 1836. 
— —— Desmoulins. Etudes sur les Echinides, p. 292, No. 53, 
1837. 
EcHINOPSIS LATIPORA, Agassiz. Catal. Syst. Ectyp. foss., Mus. Neoc., p 9, 1840. 
— CONTEXTA, Agassiz. Ibid. 
~- DEPRESSA, Agassiz. Ibid. 
ECHINUS RADIATUS, Dujardin. In Lamarck’s Anim. sans Vert., 2e éd., t. iii, 
p. 371, 1840. 
ARBACIA RADIATA, Roemer. Norddeutschen Kreidegebirges, p. 30, 1840. 
ECHINOPSIS PUSILLA, Roemer. Ibid., pl. vi, fig. 10, 1840. 
CYPHOSOMA RADIATUM, Agassiz et Desor. Ann. Sc. Nat., 3e série, t. vi, p. 352. 


Cat. rais. des Echinides, 1846. 
16 


122 GLYPHOCYPHUS 


EcuINOPSIS LATIPORA, Agassiz et Desor. Ibid., p. 351, 1846. 
a CONTEXTA, Agassiz et Desor. Ibid. 
— DEPRESSA, Agassiz et Desor. Ibid. 
— CONTEXTA, Bronn. Index Paleontologicus, p. 447, 1848. 
— DEPRESSA, Bronn. Ibid. 
— LATIPORA, Bronn. Ibid. 
— PUSILLA, Bronn. Ibid. 
ARBACIA RADIATA, Bronn. Ibid., p. 91, 1848. 
EMNOPLEURUS PULCHELLUS, Sorignet. Ours. fossiles du Dép. de l’Eure, p. 31, 1850. 
Guypricus KonINnckKII, Forbes. In Dixon's Geology of Sussex, p. 340, tab. xxv, 
fig. 30, 1850. 
EcHINOPSIS PUSILLA, Forbes. In Dixon, ibid., tab. xxv, fig. 31, 1850. 


— Giebel. Deutschlands Petrefacten, p. 320, 1852. 
GLYPHOCYPHUS PULCHELLUS, d’Archiac et Jules Haime. Descript. Nummilites del’ Inde, 
p. 202, 1854. 


ECHINOPSIS PUSILLA, Morris. Catal. of Brit. Fossils, 2nd. ed., p. 78, 1854. 
— LATIPORA, Pictet. Traite de Paleontol., 2e éd., p. 242, 1846. 
— CONTEXTA, Pictet. Ibid. 
— DEPRESSA, Piectet. Ibid. 

— PUSILLA, Pictet. Ibid. 

CYPHOSOMA RADIATUM, Pictet. Ibid. 


GLYPHOCYPHUS RADIATUS, Desor. Synopsis des Echinides fossiles, p. 103, tab. xvii, 
figs. 1—3, 1856. 
“= — Cotteau et Triger. Fichinides du dép. de la Sarthe, p. 185, 
pl. xxviii, figs. 7—12, 1859. 
— —_ Dujardin et Hupé. Hist. nat. des Kehinod., p. 513, 1862. 
TEMNOPLEURUS PULCHELLUS, Coquand. Geol. et Pal. dela prov. de Constantine, p. 294, 
1863. 
GLYPHOCYPHUS RADIATUS, Cotteau. Paléont. Franc., Ter. Crétacé, t. vii, p. 535, 
pl. 1127, 1128, 1863. 


Test small, circular, inflated, subglobular, rounded at the border and concave at the 
base; ambulacra very narrow, with two rows of tubercles, alternate, irregular. Poriferous 
zones narrow, straight ; simple pores in single oblique pairs. Inter-ambulacra with two 
rows of small tubercles, areola radiated, plates grooved at the lower border; apical disc 
solid, forming a narrow, elongated, subpentagonal ring, having the genital and ocular 
plates almost of equal length ; oculars finely granulated ; vent large, oblong. 

Dinensions.—Height five twentieths of an inch; transverse diameter eight twentieths 
of an inch. 

Description —This small Urchin has been a great puzzle to Palaontologists, as proved 
by the long list of synonyms given above. The test is circular, inflated, and subglobular; 
rounded at the sides, and concave towards the base (figs. 1 a, 4). The ambulacral areas 
(fig. 2 e), about half the width of the inter-ambulacral, have two rows of small tubercles, which 
form an alternate series on each side of the area; they are largest at the ambitus, smaller 


FROM THE GREY CHALK. 123 


at the base and upper surface ; they are feebly developed throughout, and finely perforated 
at the summit ; the areola are regularly spaced out and placed on the border of the poriferous 
zones, which are straight, slightly depressed, and formed of small round pores, disposed 
in single oblique pairs throughout, from the peristome to the disc; the pairs of pores 
are separated from each other by a horizontal granular ridge of the test, which is more or 
less apparent in different specimens; in some examples one of the rows of tubercles 
becomes abortive ; the surface of the area is filled in with numerous small granules, and the 
ambulacral plates are slightly marked with impressions on the line of the median suture. 
The inter-ambulacral areas, fig. 2d, are double the width of the ambulacral, and provided with 
two rows of tubercles, similar in structure to, but larger in size than, those of the ambulacra; 
the areolz occupy the centre of the plates, and are surrounded with a close-set granulation ; 
two of the granules, elongated in a vertical direction, unite the contiguous areola, which 
imparts a moniliform character to well-preserved tests of this pretty little Urchin ; the plates 
are marked with impressions more or less deep at the inner and outer angles, and on each 
side of the vertical! filament at the lower part of each plate (fig. 1 ¢, fig. 2c). 

The mouth-opening is small, situated in a slight depression, and the peristome is 
delicately incised into ten unequal lobes (fig. 2 a). 

The vent is large, sub-elliptical in shape, and contracted behind; the apical disc 
forms a narrow, elongated, subpentagonal rg, somewhat peculiar in its structure, for 
instead of the small ocular plates being wedged in between the larger genitals, they 
are arranged alternately with them on the same line, and form a strong ring around the 
elliptical vent ; the genital plates are a little larger than the oculars, and have two small 
tubercles on their surface; the oculars are covered with a fine granulation, and the 
madreporiform tubercle is conspicuous by its spongy surface (fig. 2 4). 

This Urchin varies much in size; the specimens figured by Goldfuss, Desor, and 
Dixon, with those I possess, are all small. M. Cotteau has given elaborate details of a 
larger specimen found in the Department of the Sarthe, in which the characters of the test 
are admirably exhibited ; they are beautifully figured, both in his fine plates on the Hchinides 
of the Sarthe, and likewise in those in the ‘ Paléontologie Frangaise,’ some of which I have 
copied in Pl. XXIX 8, as none of my specimens have the characters so well preserved as 
in the perfect fossil test figured by my friend. 

Affinities and Differences.—Glyphocyphus radiatus is so rare an Urchin in the English 
Chalk that it is not likely to be mistaken for any other. It is smaller and more globular 
than Lchinocyphus dificilis, which it most resembles, and has the tubercles perforated, the 
miliary zone wider and more granular, the disc smaller and more solidly united to the 
coronal places than im any Lehinocyphi. 

From a careful examination of the ample materials at his disposition, M. Cotteau con- 
cludes that the large examples forming the type of the species represent Lchinopsis 
contevta, Ag., and exhibit natural impressions either deep and angular or linear and 
attenuated. Some specimens, less inflated, and possessing a stronger development of the 


124 ECHINOTHURIA 


vertical filament form /. latipora, Ag., and depressed examples with the median impres- 
sion almost absent are #. depressa, Ag. A careful study of the original types has, how- 
ever, convinced my learned friend that they are all varieties of one form, and not distinct 
species. 

Locality and Stratigraphical Position—This species is found very rarely in the hard 
beds of the Lower Chalk near Lewes, in Sussex, and in the Grey Chalk near Folkestone. 
From this stratum my best specimen was collected by Captain Cockburn, R.A., to whose 
kindness and liberality I am indebted for the example. 

The Foreign Localities given by M. Cotteau are Villers-sur-Mer, Bruneval, Saint- 
Jouin, Vaches-Noires, Dives (Calvados); Fécamp, le Havre, Rouen (Seine-Inférieure) : 
Gacé, La Perriére (Orne) ; Nogent-le-Bernard (Sarthe); Saint-Fargeau (Yonne); la 
Bedoule, Cassis (Bouches-du-Rhéne). In all these localities it is very rare, and occurs 
in the Etage Cenomanien, in the zone of Scaphites equalis, which is the equivalent of the 
English Lower Grey Chalk. 

Listory.—This species was found in the Lower Chalk of Essen and Gehrden, West- 
phalia, and was first figured and described by Goldfuss in his ‘ Petrefacta Germaniz.’ 
Since that time it has passed through a series of changes which are most correctly read 
in the long list of synonyms introductory to this article, and to which I commend the 
reader’s especial attention. 


Genus—Ecutnornuria,’ Woodward, 1863. 


“ Test globular ?, diameter of compressed specimen four inches, thickness half an inch, 
lantern projecting half an inch; composed of ten segments or double series of imbricating 
plates, ornamented with obscure miliary granules and small spine-bearing tubercles, a few 
larger than the rest ; ¢afer-ambulacral plates narrow, slightly curved, with the convex edge 
upwards and overlapping; the alternate plates bearing one large extero-lateral tubercle, 
perforated, and surrounded by a raiséd ring and smooth areola; largest plates measuring 
six lines in length, the smallest three lines or less (the longest in second specimen equal- 
ling seven lines) ; ambu/acral plates seven lines long, equalling the breadth of the exposed 
portion of eight plates, similar to the former, but curving and imbricating downwards 
towards the dental orifice, and having two small plates, each perforated by a pair of pores, 
intercalated in a notch of the middle of the lower margin; a third pair of pores perfo- 
rating the plate itself a little external to the centre; primary tubercles few, irregularly 
distributed. 


1 “ Etymologists need not trouble themselves about the derivation of this name; it is intended merely to 
express the dilemma in the writer’s mind, arising from imperfect knowledge, but which he believes to have 
no foundation in nature.” —‘ Geologist,’ vol. vi, p. 330. 


FROM THE WHITE CHALK. 125 


“« Spines of three kinds ; those adhering to the plates minute and striated ; fragments of 
larger spines (not certainly belonging to the species) striated, annulated, and furnished 
with a prominent collar to the articular end (fig. 4) ; the third kind minute, clavate, and 
truncate, articulated to a slender stalk ” (fig. 5). 


Ecutnotuvuria rioris, Woodward.\—Pl. XXIX B, figs. 3—5. 
EcuINotTuuria FLORIS, Woodward. ‘ Geologist, vol. vi, pp. 327—330, 1863. 


“The fossils represented” in Pl. XXIX B “are probably only fragments of the original 
structure, and possibly only the smaller and less essential portions of the whole. Never- 
theless, I have determined to publish some account of them, although at the risk of 
committing an extravagant error, as a last resort towards obtaining more complete ex- 
amples or suggestions for their more correct interpretation. 

“Both specimens have been presented to the British Museum; one by J. Wickham 
Flower, Esq., of Park Hill, Croydon, the other by the Rev. Norman Glass, of London. 

«The first example was obtained, at least sixteen years ago, from the Upper Chalk of 
Higham, near Rochester, and was submitted to Professor E. Forbes, in whose custody it 
remained for several years. It was originally shown to me in connection with the 
anomalous Cirripede Zoricu/a, then newly discovered by Mr. Wetherell. The resemblance 
between them is certainly curious ; but there is no real relationship. Mr. Flower’s fossil 
exhibits distinct traces of the crystalline structure peculiar to petrified Echinodermata, 
and the pairs of pores in the ambulacral plates are equally characteristic of the Echinide. 
Mr. Darwin also has examined this fossil and rejected it from his province of inquiry. 

** Professor Forbes could not make up his mind to describe the specimen, and ulti- 
mately it was returned to Mr. Flower, with whom it remained until the publication of a 
note on the genus Profo-echinus, by Major ‘Thomas Austin, in the ‘Geologist’ for 1860 
(vol. ii, p. 446), when it was entrusted to me for the purpose of considering whether it 
had any special affinity with this new type, and for description in the same journal. 

“The Proto-echinus was obtained from the Carboniferous Limestone of Hook Head, 
Wexford, and is but a fragment of a single ambulacrum, consisting of three series of 
plates at the wider end and two at the other extremity, with apparently a single terminal 


1 ‘On Eechinothuria floris, a new and anomalous Echinoderm from the Chalk of Kent.’ By S. P. 
Woodward, F.G.S. I have printed this paper from the ‘Geologist’ entire (altering the references to 
figures), as a contribution to British Echinology by my late esteemed friend. Dr. Woodward took so 
warm an interest in my work, and afforded me such valuable assistance, by the loan of specimens for 
figuring, that it affords me very great pleasure to acknowledge here his uniform kindness, and connect 
his name with a Monograph to which he contributed important aid. 


126 ECHINOTHURIA 


plate. Each plate is perforated by a pair of pores. It differs from Hehinothuria in 
every particular. 

“The question presented to me by Mr. Flower’s fossil was, whether to consider it part 
of the envelope of a new kind of Holothuria, or whether it might be no more than a fragment 
of the oral disc of some great unknown Zehinus. Portions of the imbricating scaly 
armour of a Psol/ws had been met with when examining the fossils of the Boulder Clay 
collected by Mr. J. Richmond, of Rothsay; but in Pso/ws, while the greater part of the 
body is clothed with fish-like scales, the ambulacra are only developed on one side, form- 
ing a creeping disc, the scales of which are small and not imbricated. On the other hand, 
the peristome of the largest known Echinite from the Chalk is less than an inch in 
diameter; and the largest recent Sea-urchin in the Museum has an oral disc not more 
than two inches wide, whereas the fossil is a segment of a disc which must have been at 
least four inches across. This objection, on the score of size, was, however, less felt, 
because the Cyphosomas and Diademas of the Chalk have larger oral and apical orifices 
than any other Urchins, and the character of their apical disc was unknown, being only 
preserved in a few minute specimens of C. dificile, from Chute Farm. Moreover, there 
were indications in the Upper Chalk of a great Dzadema, of which nothing more had been 
obtaimed than scattered plates and fragments of spines. ‘This species is referred to in 
Decade V of the ‘ Geological Survey’ (Article “ Diadema,” Section C, spines tubular, annu- 
lated). Mr. Wetherell obtained a mass of Chalk containing above one hundred fragments 
of spines, which are hollow, striated and annulated, as in the recent D. calamaria. From 
the plates mingled with the spines we ascertained that the ambulacral pores presented 
the usual characters, being arranged in single file, and a little crowded near the peristome ; 
but many of the plates presented only their smooth mner surfaces. A smaller mass of 
Chalk, in Mr. Wiltshire’s cabinet, contains similar plates and spines, mingled with a few 
true scales and minute truncated spines like those of Hehinothurta. The Diadema spines 
were erroneously referred by Professor E. Forbes to the genus Micraster (Decade III, pl. 10, 
fig. 15 ; bad, for they are not spiral). ‘They are also figured by Dixon, in his ‘ Geology 
of Sussex,’ and described by Forbes as “spines of a Cidaris.” Diademas possessing spines 
of this character are known to occur in the Upper Cretaceous strata of France; and 
Dr. Wright has lately obtained a small specimen from the Chloritic Marls of Dorsetshire. 
In these the apical disc is quite small. 

“ A more serious difficulty, m comparing Mr. Flower’s fossil with the oral dise of any 
Hchinite, was presented by the arrangement of the plates ; in the recent Echinidee pees 
they are all directed towards the dental orifice, but here the alternate series take opposite 
‘dips,’ the ambulacral plates overlapping one way and the others in a contrary direction. 

“ Last year, while I was still hesitating about the publication of Mr. Flower’s fossil, a 
second specimen was obtained from Charlton, in Kent, by the Rev. N. Glass, who has 
cleared it from the matrix with great skilland patience. . . . . At first sight this specimen 
would scem to solve the problem, by supplying the peristome and lantern of the same 


FROM THE WHITE CHALK. 127 


great Cyphosoma or Diadema, of which Mr. Flower’s specimen might be the apex or 
periproct. But a closer examination confirms the objections already stated, and gives 
increasing probability to the other conjecture (if, indeed, it does not compel us to adopt 
it), however difficult it may be to realise the notion of an Hchinite having no proper 
‘test,’ and clothed entirely with imbricating scales like those of the peristome of Cidaris. 

“Tn Mr. Flower’s specimen the imperforate plates imbricate towards the centre (or apex), 
where the smaller ends of the several series converge. In Mr. Glass’s specimen they slope 
away from the centre (or mouth), that is, also towards the apex. The perforated or ambulacral 
plates, which overlap one another outwardly (¢. e. downwards) in specimen, fig. 3 a, are seen 
sloping towards the dental cone and reclining upon it. ‘The portion of an ambulacrum, fig. 
5 a, consists of four plates, diminishing in size from ¢ to a, ina line not accurately directed 
towards the centre. This portion exhibits the zzferior surface of the plates, known by their 
curved surfaces, destitute of ornamental granules ; it is not, however, the oral end of one of 
the segments turned over, a thing scarcely possible to happen, for in that case the dip of 
the plates would be reversed ; but it must be the opposite (or apical) extremity of a series 
folded back upon its origin, and exposed to view by the damage which the surface of the 
specimen has sustamed. From this circumstance it seems probable that the whole fossil, 
when complete, was not elongated, nor even spherical, but somewhat depressed in a ver- 
tical direction, though doubtless admitting of a moderate amount of flexure. At the last 
hour, after making the drawing, I ventured to clear away the chalk from the side of Mr. 
Glass’s fossil, near where an ambulacral segment is seen to curve as if it might be con- 
tinued round to the other surface. This attempt was successful, for the ambulacrum 
and also the adjacent inter-ambulacral segment were found continuous, though crowded 
and displaced at the turning, falling again into regular order, and diminishing in size, 
though not so nearly complete as in Mr. Flower’s example. 

“ After this apparently conclusive demonstration, it appears desirable to give a name to 
the fossil, and to attempt a short description, although its rank and affinities are to us 
still matter of conjecture. At present it is one of those anomalous organizations which 
Milne-Edwards compares to solitary stars, belonging to no constellation in particular. 
The disciples of Von Baer may regard it as a ‘ generalised form’ of Echinoderm, coming, 
however, rather late in the geological day. ‘I'he publication of it should be acceptable 
to those who base their hopes on the ‘imperfection of the geological record,’ as it seems 
to indicate the former existence of a family or tribe of creatures whose full history must 
ever remain unknown.” 

Locality and Stratigraphical Position —Collected from the Upper Chalk of Higham, 
near Rochester. ‘The fine specimens in the Rev. ‘I’. Wiltshire’s cabinet were obtained 


from the Upper Chalk at Gravesend and Charlton. 


128 CYPHOSOMA 


Genus—CyPHosomA, Agassiz, 1840. 


Cypnosoma, Agassiz, 1840. 

Puymosoma, Haime, 1853. 
— Desor, 1858. 

Cypuosoma, Cotteau, 1863. 


Test moderate in size, circular or subpentagonal, slightly inflated at the sides. 
Poriferous zones well developed and undulated, composed of simple pores that in 
general are unigeminal throughout, and sometimes are bigeminal on the upper surface, 
and crowded a little together around the peristome. The poriferous plates are un- 
equal and irregular in their mode of arrangement. The primary tubercles are nearly 
equal in size in both areas, the areola are well developed, and sometimes marked with 
radiated striations; the bosses are prominent, and have their summits sharply crenu- 
lated ; the mammillon is large, prominent, and imperforate, and the general facies of the 
test shows a regular, and uniform development of all its several elements. The mouth- 
opening is large, the peristome slightly incised, and the oral lobes nearly equal. The 
discal opening is large and pentagonal, one angle of which extended far into the single 
inter-ambulacrum ; the elements of the disc were feebly united, as they are absent in all 
the specimens that have hitherto been collected. 

The spines are long, solid, subcylindrical, aciculate, or spatuliform ; sometimes they 
are straight and lanceolate, or bent, ramiform, or spoon-shaped ; all these varieties are 
figured in Pls. XXIV and XXVI. The stem is smooth and marked with fine longi- 
tudinal striae, the milled ring is prominent, the head distinct, and the rim of the aceta- 
bulum crenulated. 

The genus Cyphosoma is distmguished from all others by its prominent tubercles 
with crenulated bosses, and imperforate mammillons ; in the structure of these it resembles 
Echinocyphus and Temnopleurus, but is readily distinguished from these by the absence of 
the angular and sutural impressions which impart so marked a character to their tests. 

M. Desor has separated into the genus Coptosoma all those Cyphosomata from the 
Nummulitic formation (‘lertiary) with much undulated poriferous zones, tubercles with 
very large mammillons, and having the plate-sutures of the areas deeply incised; thus 
leaving the typical Cyphosomata as true Cretaceous fossils, which first appear in the 
Neocomian beds, and attain their maximum development in the upper stage of the 
White Chalk. 

The Cretaceous rocks of France are very rich in species of Cyphosoma, of which a 
very small proportion have hitherto been found in the British islands. 


FROM THE LOWER CHALK. 129 


A.—Species from the Lower Chalk. 
CyPHOSOMA GRANULOSUM, (o/dfuss, sp., 1826. Pl. XXIII, figs. 2 a, 0, ¢, d. 


CIDARITES GRANULOSUS, Goldfuss. Petrefact. Germanie, pl. xl, fig. 7, p. 122, 1826. 
DIADEMA GRANULOSUM, Agassiz. Prod., Mem. Soc. des Sc. Nat. de Neufchatel, t. i, 
p. 189, 1836. 


Ecuinus MILiert, Desmoulins (pars). Etudes sur les Behinides, p. 294, 
No. 68, 1837. 
EcHINUS GRANULOSUS, Dujardin. ‘non Lamarck’s Anim. sans Vertébres, 2e éd., 


t. iil, p. 372, 1840. 
CIDARITES GRANULOSUM, Geinitz. Charakt. der Schict. und Petref., p. 90, 1842. 


DIADEMA — Morris (pars). Catalogue of British Fossils, p. 51, 1843. 
CypHOsoMaA — Reuss. Verstein. der Bohm. Kreideform., p. 58, 1846. 
_ MILLERI, Agassiz et Desor (pars). Catalogue rais. des Echinid., 


p. 351, 1848. 

— — Bronn. Index Paleontologicus, p. 381, 1848. 

— — D’Orbigny. Prod. Pal. strat., t. ii, p. 273, Et. 22, 1850. 

-- — Broan. Lethea Geognost., Kreide-gebirges, pl. xxix, p. 186, 
1851. 

-- Kontet, Forbes. In Morris’s Catalogue of British Fossils, 2nd ed., 
p. 75, 1854. 

PHyMosoMA GRANULOSUM, Desor. Synops. des Echinid. foss., p. 87, 1856. 


CypHosoMa _ Woodward. Mem. Geol, Surv., Dec. V, p. 1, 1857. 
PHYMOsOMA — Hupé. Hist. Nat. Echinod., p. 508, 1862. 
CYPHOSOMA _ Cotteau. Pal. Francaise, Ter. Crétacé, t. vii, p. 685, 


pl. 1169, 1865. 


Test large, circular, depressed, convex on the upper surface, inflated at the sides, and 
flattened below ; poriferous zones wide and straight in the upper third, narrow and undu- 
lated at the ambitus and base; pores largely bigeminal in the wide upper third, and 
unigeminal in the rest of the zones; ambulacra narrow above, wide below, with two rows 
of large tubercles, twelve in each ; inter-ambulacra with two rows of primary tubercles in 
the middle of the area, secondary tubercles wanting in young specimens, and only slightly 
developed in the largest tests; miliary zone wide, depressed, and naked above, narrow 
and granular below ;. mouth-opening small and circular ; peristome with shallow entailles ; 
discal opening large, and widely pentagonal. 

Dimensions.—Height eight tenths of an inch; transverse diameter one inch and eight 
tenths. 

Description —This Urchin resembles C. Aénigi in so many points of structure, the 
absence or rudimentary condition of the secondary tubercles excepted, that it may pro- 
bably be only a variety of that species. It has, however, been treated by different sys- 

17 


130 CYPHOSOMA 


tematic authors as specifically distinct ; and im deference to their opmion, rather than in 
accordance with my own convictions, I have devoted this article to its description. 

The test large and circular, inflated at the ambitus, convex above, and flat below 
(Pl. XXIII, figs. 2 a, 4, ¢). 

The ambulacral areas are enlarged at the ambitus, moderately wide at the base, and 
very narrow in the upper part; by reason of the mcreased development of the poriferous 
zones in this region, from the ambitus to the peristome, the tubercles are as large as 
those in the inter-ambulacra, but in the upper third of the area they diminish rapidly 
in size, and on the four or five coronal plates they are quite rudimentary (Pl. XXIII, 
fig. 2a). The pores are arranged in oblique pairs, and are bigeminal from the ambitus 
to the summit (Pl. XXIII, fig. 2 ¢, 7), and unigeminal from the ambitus to the peristome, 
the zones forming a series of crescents around the areole of the ambulacral tubercles. 

The inter-ambulacral areas have two rows of primary tubercles, twelve to thirteen in 
each row, which vary gradually in size from the ambitus to both apertures; they are sur- 
rounded by well-developed areolze, confluent at the upper and lower borders, and surrounded 
by granules at the sides. The secondary tubercles are irregular, very small, and limited 
to the under surface (fig. 2 4). On one or two plates above the ambitus there are only 
one or two solitary tubercles, which are, however, inconstant in different specimens (fig. 2 d). 
The miliary zone is narrow and granular at the ambitus, and wide, depressed, and naked 
on the upper surface; the granules are unequal, of different sizes, and placed in semicir- 
cular groups around the lateral parts of the areole. Many of the granules are large and 
mammillated, and are nearly as large as the row of secondary tubercles, the small granules 
being compactly fitted in between them. ‘The coronal plates are marked by slight sutural 
impressions, which become more apparent in consequence of the nudity of the depressed 
upper surface of the areas. 

The discal opening is very large, pentagonal, and angular; and the single posterior 
ovarial plate is projected far into the area (fig. 2 a). ‘The mouth-opening is moderate in 
diameter (fig. 2 6); the peristome is circular, and divided into ten unequal lobes by shght 
incisions with reflexed borders. 

Affinities and Differences —This species differs from the typical forms of Cyphosoma 
Konigi in the absence of a regular series of secondary tubercles above the ambitus; but 
in the general character of the test and in most of its details it has close affinities with 
that species. Whether the spines of this Urchin exhibit any difference from those attached 
to the typical form of C. Kénigi (Pl. XXIV, fig. 1) remains to be determined by those who 
may be fortunate enough to discover a specimen with the spines adherent to the test. 

Locality and Stratigraphical Position —This Urchin is found in the Lower Chalk of 
Kent and Sussex. The specimen figured in P]. XXIII, belonging to the British Museum, 
was collected near Lewes. 

Foreign Localities. —Houguemarre (Eure), Orglande (Manche), from the Etage Sénonien, 
where it is rare (M. Cotteau) ; the type of Goldfuss’s figure was collected from the Chalk 


of Westphalia. 


FROM THE WHITE CHALK. 131 


B.—Species from the Upper Chalk. 


Cypnosoma Ko6nicr, Manftell. Pl. XXIII, figs. 1,2; Pl. XXIV, figs. 1—7; PI. 
MXV, fig! 3; Pl. XX VI, fig. 1. 


EcuINItTE. 
Crparis K6n1Gr, 
Ecuinus Miter, 


CIDARITES VARIOLARIS, 


Ecuinus Kontet, 

Crparites KO6nrer, 

Ecuinus K6ntet, 
— Mrriert, 


DrapEMA KONIGrI, 
CypHosoma MILLERTI, 
DIADEMA GRANULOSUM, 
CyrpHosoma MILLERI, 


—— ORNATISSIMUM, 


— MIttert, 

— MAGNIFICUM, 
DrapeMa KOontel, 
CypHosoma MILLERI, 


= ORNATISSIMUM, 


— MILLER, 


_ VARIOLARIS, 
Ecutnus Konier, 
Cyrnosoma KOniar, 


Parkinson. Organic Remains, vol. iii, pl. i, fig. 10, 1811. 

Mantell. Geology of Sussex, p. 180, 1822. 

Desmarest. Oursins fossiles, Dict. Se. Nat., t. xxxvii, 
p. 101, 1825. 

Goldfuss. Petref. Germaniz, p. 123, pl. xl, figs. 9 a, 4, 
1826. 

Fleming. British Animals, p. 479, 1828. 

Brongniart. Tableau des Terrains, p. 405, 1829. 

Woodward. Synopt. Tab. of Brit. Organ. Rem., p. 6,1830. 

De Blainville. Zoophytes, Dic. Sc. Nat., t. Ix, p. 210, 1830. 

De Blainville. Ibid. 

Agassiz. Prod., Mém. Soe. Se. Nat. de Neufchatel, t. i, 
p. 190, 1836. 

Desmoulins. Etudes sur les Bchinides foss., p. 294, 
No. 68, 1837. 

Desmoulins (pars). Ibid., p. 312, No. 10, 1837. 

Agassiz. Catal. Syst. Ectyp., p. 11, 1840. 

Morris. Catalogue of British Fossils, p. 51, 1843. 

Agassiz et Desor (pars). Catal. rais. des Echinides, 
p. 351, 1846. 

Agassiz et Desor. Ibid., p. 352, 1846. 

Graves. Essai Topogr. Géognost. Dep. l’Oise, p. 688, 1847. 

Graves. Ibid. 

Bronn. Index Palzeontologicus, p. 418, 1848. 

Bronn (pars). Ibid., p. 381, 1848. 

D' Orbigny. Prodrom. de Pal. strat., t. ii, p. 273, 1850. 

D Orhigny. Ibid., t. ii, p. 273, 1850. 
‘orbes. In Dixon’s Geol. of Sussex, pl. xxv, figs. 17, 
26, 27, 1850. 

Forbes. Ibid., pl. xxv, fig. 29, 1850. 

D'Archiac, Hist. des Progr. de la Géol., t. iv, p. 20, 1851. 

Forbes. Morris, British Fossils, 2nd ed., p. 75, 1851. 


— ORNATISSIMUM, Forbes. Ibid. 


Puymosoma KO6nrert, 
CypHosoma KOnrai, 


= ORNATISSIMUM, 


Desor. Synopsis des Bchinides fossiles, p. 86, 1856. 

Pictet. Traité de Paléontologie, 2e éd., t. iii, p. 243, 
1857. 

Pictet. Ibid. 


132 CYPHOSOMA. 


CypHosoma KOntet, Woodward. Mem. Geol. Surv., Decade V, Appendix, 
1857. 
PuymMosoMA. — Dujardin et Hupé. Hist. Nat. des Zooph., p. 508, 1862. 
CypHOSOMA = — Woodward. “On Kénig’s Sea-urchin,” Geologist, vol. v, 
p. 41, 1862. 
— — Cotteau. Paléontologie Francaise, t. vii, p. 678, pl. 1167, 
1168, 1863. 


Test large, subcircular, depressed, convex on the upper surface, inflated at the sides, 
almost flat on the under surface ; poriferous zones wide and straight in the upper third, 
narrow and undulated at the ambitus and base; pores bigeminal in the upper third, uni- 
geminal in the lower two thirds of the zone; ambulacra narrow above, wider below, with 
two rows of large tubercles, twelve in each; inter-ambulacra with two regular rows 
of primary tubercles in the middle, and two rows of secondary tubercles, irregular in size 
and distribution, on the zonal sides of the area; miliary zone wide, naked, and depressed 
above, narrow and granular below; mouth-opening small, peristome circular, lobes nearly 
equal; discal opening large, pentagonal; spines long, subcylindrical; upper third of 
the stem aciculate, spatulate, straight or bent; lower third sculptured with fine longitu- 
dinal lines. 

Dimensions.—Height seven tenths of an inch; transverse diameter two inches. 

Description.—This is one of the largest, most beautiful, and typical of our British 
Cyphosoma ; it was well figured by Parkinson, 1811, in bis ‘Organic Remains,’ as “an 
Echinite from Kent with its spine ;” he gave no description of the specimen, and it was 
reserved for Dr. Mantell, 1822, to give it a specific place among our Cretaceous 
Urchins. 

The long list of synonyms prefixed to this article exhibits the historical phases through 
which it has passed, and the numerous admirable and accurate drawings with which our 
excellent friend Mr. C. R. Bone has enriched our Monograph will make the determination 
of Cyphosoma Konigi a matter of ease and certainty to all future observers. 

‘The specimen figured Pl. XXIII, figs. 1 a—g, belongs to the British Museum. The 
large test is subcircular, slightly convex above, inflated at the sides, and flattened below 
(fig. 1 ¢); the surface is highly ornamented, the tubercles are nearly all of the same size 
and regular in their disposition, and the granules are large and conspicuous at the 
base. 

The ambulacral areas are narrow above, wide at the ambitus, and contracted at the 
base ; they possess two rows of tubercles, 12 or 13 in each row; those at the ambitus 
are very large, and nearly equal in size the inter-ambulacral tubercles ; on the upper third 
of the area they diminish rapidly in magnitude, and from the ambitus to the peristome 
are much larger. 

At the ambitus the areola are wide and confluent, but on the upper surface they are 
narrow and separated only by a line of granules; the zone which divides the two series is 


FROM THE WHITE CHALK. 133 


contracted, and provided with two rows of granules closely set together, unequal in size. 
some of them being mammillated at the angles of the plates. 

The poriferous zones are narrow and undulated at the base and ambitus, and wide and 
straight at the upper surface ; the pores are small, and unigeminal from the peristome to 
the ambitus (fig. 1 e), and distinctly bigeminal on more than the upper third of the 
zones (fig. 1 d); near the peristome the pairs are doubled (fig. 1 g). 

The inter-ambulacral areas are about one third wider than the ambulacral, and furnished 
with primary and secondary tubercles ; the primary series consists of two rows, twelve in 
each, occupying the centre of the plates; the tubercles are smaller on the upper than on 
the lower third of the area; the secondary tubercles are large and unequal in size; they 
form a series between the poriferous zones and primary tubercles, some of which they 
resemble in magnitude, especially those extending from the ambitus to the coronal plates ; 
between the ambitus and peristome they are much smaller, and in many specimens are 
not more developed than large granules on mammillated eminences. 

The miliary zone is very wide, naked, and depressed at the upper surface, 
becoming gradually narrower at the ambitus, and much contracted at the base; the granules, 
unequal in size, are fine, abundant, set closely together, and arrarged in semicircles 
around the primary and secondary tubercles ; some of the granules, much larger than the 
others, are set on mammillated eminences, and may easily be mistaken for the small 
secondary tubercles placed near them ; this mingling together of small tubercles and large 
mammillated granules imparts a highly ornamented character to the infra-marginal region 
of this species. 

Discal opening large, pentagonal, and subangular, the elements wanting im all the 
specimens hitherto found. Mouth-opening small, circular, the peristome feebly incised, 
the border reflected, the entailles having the border elevated and opening upwards. 

The spines exhibit some remarkable variations from the typical form of structure. 
Some of the most curious of these I have figured in Pls. XXIV and XXVI. The 
typical spines, as seen in those 7 sift in the specimen belonging the British Museum 
(Pl. XXIV, fig. a), are elongated, subcylindrical, and sometimes aciculate at the summit 
(fig. 1 d, and fig. 3 a), or spatuliform with carinz, as Pl. XXIV, fig. 1 ¢, or spoon- 
shaped, as Pl. XXVI, figs. 1 a, 4, the lower part or collerette being long and distinct, 
and covered with fine longitudinal lines (Pl. XXIV, fig. 3 4), much stronger than those 
observed on the stem; the head is well developed, the milled ring very prominent, flat, 
and deeply striated, and the rim of the acetabulum finely crenulated (PI. XXVI, figs. 1 
a, ¢, figs. 2, 4, 6), with other varieties of spines figured in this plate. 

The test of this Urchin presents many variations of form from the typical shape seen 
in the specimen figured in Pl. XXIII, fig. 1 @, 4, and considered to be its normal form. 
In Pl. XXIV, fig. 7, I have figured a remarkable monstrosity of this species from 
the British Museum Collection; it is inversely conical, like Psewdodiadema tumidum ; 
measures three quarters of an inch in height, and is one inch and one third in diameter 


134. CYPHOSOMA 


above, contracting below down to the oral opening, which, at first sight, appears to be the 
In Pl. XXII, figs. 5 a, 4, c, I have given drawings of a young test 
from the National Collection. 

Affinities and Differences —This species forms one of the most typical forms of the 
genus Cyphosoma, well characterised by its large size, round and inflated at the 


summit of the test. 


ambitus, highly ornamented at the base, and having the poriferous pores distinctly 
bigeminal in all the upper part of the zones; the secondary tubercles are large above the 
ambitus and small at the base; the spines are strong, elongated, and furnished with a 
striated collar, having the summit sometimes flattened and carimated, or expanded and 
spatulate, as seen in the different figures. 

The nearest affinities of C. Adnzgi are with C. granulosum, from which it differs in 
having large secondary tubercles above the ambitus and a more ornamented test at the 
base. Jn size and height, and in the disposition of its tubercles on the upper surface, as 
well as in the bigeminal arrangement of the pores in the upper part of the zones, it much 
resembles C. magnificum, Agass., from the Htage Sénonien inférieure of the south-west 
of France. 

Locality and Stratigraphical Position —C. Kénigi is found in fine preservation in the 
Upper Chalk of Kent, Sussex, Norfolk, Wiltshire, and Yorkshire. 
lected in the Ktage Sénonian at Thuison (Somme); Tartigny (Oise); Vernonnet, La 
Villette, Hougouemarre (Eure); where it is rare. It is likewise found at Rigen, and 


In France it is col- 


Cesfeld, near Dusseldorf. 
CypHOSOMA CoROLLARE, Klein. Pl. XXVI, figs. 7, 8, 9, 10. 


EcHINITES ORBICULATUS, Lister. Hist. Animal. Angliz, p. 220, pl. vil, fig. 19, 1678. 


CIDARIS COROLLARIS, Klein. Natur. Dispos. Echinoderm,, pl. viii, fig. c, p. 20, 
1734. 
— — Baier. Oryctographia Norica, p. 70, pl. ii, fig. 36, 1759. 


Leske. » Klein, Nat. Dispos. Echinoderm., p. 138, pl. viii, 
fig. c, 1778. 


— _CORONALIS, Gmelin. Linné’s System. Natur., p. 3177, 1788. 
ECHINUS SAXATILIS, Parkinson. Organic Remains, pl. iii, fig. 1, 1811. 
CIDARIS COROLLARIS, Parkinson. Ibid., pl. i, fig. 7, 1811. 

—  SAXATILIS, Mantell. Geology of Sussex, p. 180, 1822. 
—  COROLLARTS, Mantell. Ibid., p. 181, 1822. 


ECHINUS SAXATILIS, 


CIDARIS COROLLARIS, 
EcHINUS COROLLARIS, 


Fleming. History of British Animals, p. 479, 1828. 
Brongniart. Tableau des Terrains, p. 405, 1829. 

‘oodward. Synop. Table of Brit. Org. Remains, p. 6, 1830. 
De Blainville. Zoophytes, Dict. des Sc. Nat., t. lx, p. 210, 


1830. 
Agassiz. Prodrome des Radiaires, p. 188, 1836. 
Desmoulins. Etudes sur les Eichinides, p. 298, 1837. 


FROM THE UPPER CHALK. 135 


CIDARIS COROLLARIS, Morris. Catalogue of British Fossils, p. 49, 1843. 

— SAXATILIS, Morris. Ibid., p. 50, 1843. 
CYPHOSOMA COROLLARE, Agassiz et Desor. Catal. Rais. des Echinides, p. 351, 1846. 
CIDARIS COROLLARIS, Bronn. Index Paleeontologicus, p. 298, 1848. 

—  SAXATILIS, Bronn. Ibid., p. 300, 1848. 


CypHosoma coroLLare, D’Orbigny. Prodrome, t. ii, p. 273, Bt. 22, 1850. 

_ —- Forbes. In Dixon’s Geol. of Sussex, p. 340, 1850. 

—_ — Forbes. In Morris’s Catalogue of Brit. Foss., p. 75, 1850. 
PHYMOSOMA — Desor. Synop. des Echinides fossiles, p. 88, 1856. 
CYPHOSOMA SAXATILE, Desor, Ibid., p. 87, 1856. 

—— COROLLARE, Pictet. Paleontologie, 2e éd., t. iii, p. 243, 1857. 

= — Woodward. Mem. Geol. Surv., Decade V, p. 2, 1858. 

— 7 Leymerie et Raulin. Géol. du Départ. de ?Yonne, p. 621, 

1858. , 
PHYMOSOMA SAXATILE, Dujardin et Hupé. Hist. des Echinoderm., p. 508, 1862. 
—  .COROLLARE, Dwardin et Hupé. Ibid. 
CypHosoMA PERFECTUM, Cotteau et Triger. Wchinides de la Sarthe, p. 261, pl. xlii, 
figs. 13—16, 1860. 
— — Cotteau. Echinides foss. des Pyrenées, p. 24, 1863. 
i COROLLARE, Cotteau. Pal. Frangaise, t. vil, p. 669, pl. 1165, 1864. 


Test small, circular, depressed ; ambulacra wide, two rows of tubercles, nine to ten in 
each, gradually diminishing in size towards the poles ; inter-ambulacra moderate, with two 
rows of tubercles, nine in each, four ambital, large; areolee defined by rows of granules ; 
miliary zone moderate, granular below, becoming smooth above ; poriferous zones narrow, 
undulated, pores unigeminal, and crowded near the summit ; base concave; oral opening 
small, one third of an inch in diameter; discal opening large, pentagonal, half an inch 
im diameter; spines long, slender, spatulate, one fourth longer than the diameter of the 
test. 

Dimensions.—Transverse diameter one inch; height one third of an inch; in general 
the specimens are not so large. 

Description —This is one of our most common Cyphosomata. In certain localities it is 
a small, circular Urchin, rarely exceeding an inch in diameter and about three or four 
lines in height ; it is convex and depressed above, and flat or subconcave below ; the areas 
are nearly equal in width, the tubercles very much alike in form and size, the areole are 
encircled with granules, and the structure of the different divisions of the test is very 
uniform throughout. 

The ambulacral areas (Pl. XX VI, fig. 10), contracted above by the poriferous zones, 
have two rows of primary tubercles supported on large bosses, and arranged in alternate 
series on each side of the area;,in adult specimens there are nine or ten tubercles in a 
row; the areolz are wide, and a single row of granules (rarely double) separates them 
from each other. 


The poriferous zones are narrow, and much undulated at the ambitus and infra-mar- 


136 CYPHOSOMA 


ginal region ; the small pores are arranged in oblique pairs at the sides, and at the upper 
surface become bigeminal; fig. 10 shows an ambulacral area with its poriferous 
zones magnified six diameters ; fig. 7 6 exhibits the upper part of the area magnified six 
times, with the bigeminal arrangement of the pores in the upper part of the zones. 

The inter-ambulacral areas are furnished with two rows of tubercles nearly identical 
with those of the ambulacral ; they are, however, a little larger than the latter in the upper 
surface, and are surrounded by areole that are placed wider apart. The secondary 
tubercles are very small and unequal in size; they form a series near the zones, one mam- 
millated tubercle rismg in each plate from the midst of numerous granules. They are 
most apparent at the lower surface and the ambitus, and are rare on the upper surface. 

The miliary zone is wide, naked, and depressed in the upper surface, forming at the 
ambitus and lower surface a zigzag line, which defines the contour of the plates. The 
granules are large, and those surrounding the primary tubercles at the ambitus send pro- 
Jongations into the areole, which impart a radiated character to the structure of these 
parts. ‘The intermediate granules are more or less abundant, and form circles or semi- 
circles around the areola. The small mouth-opening is lodged in a concave depression 
in the base. The peristome is circular, and divided by feeble incisions, the lobes of 
which are nearly equal. The apical disc is absent. The opening is large, pentagonal, 
and angular, and in our large specimen is about five lines in diameter. 

The spines are long, slender, and spatulate, one fourth longer than the diameter of the 
test, to which they are sometimes found adherent and 7m siti. 

Affinities and Differences.—C. corollare is readily recognised by its moderate size, 
depressed circular test, sometimes subpentagonal; the pores are unigeminal on the sides 
and bigeminal on the upper surface, with small secondary tubercles at the base near to the 
zones ; the miliary zone is nude and depressed near the summit ; the mouth-opening is small, 
and lodged in a concave depression. By these characters it may be distinguished from_ 
C. fara and young examples of C. granulosum, both of which it resembles much. 

Locality and Stratigraphical Position —Vhis Urchin is very common in the upper 
flinty Chalk of Brighton, Gravesend, and Woolwich. In France M. Cotteau enumerates 
the following localities in which it is rarely found in the Htage S¢nonien :—Senneville, 
Saint-Pierre en Port (Craie supérieure, M. Hebert) ; Seine Inférieure, La Herelle (Oise) ; 
Pinterville (Eure); Villeneuve-sur-Yonne (Yonne); Meudon (Seine-et-Oise) ; Sarlat 
(Dordogne) ; Tercis (Landes). 

History.—TVhe history of this species, one of the oldest Cyphosomas, is very difficult 
to trace, as our table of synonyms has already exposed. Lister and Klein have both 
figured it, and Parkinson has given a very good figure of it under the name of Hehinites 
saxatilis in his ‘Organic Remains.’ Klein’s name, however, has the priority, and 


therefore is retained. 


Fig. 


[9 fe} 


Fig. 


co) 


PLATE XXII. 


Echinocyphus from the Grey Chalk. 


1 a. Ecuinocyrnus pirriciuis, Agassiz. ‘Test, natural size. British Museum. 


(P. 116.) 


1 6. Ambulacral and inter-ambulacral plates, magnified four diameters, do. —_Do. 


2 a. ECHINOCYPHUS DIFFICILIS. 


wo w ww 
a 


Q 


29 


Upper surface, magnified three diameters. The Rev. 
T. Wiltshire, F.G.S. 

Under surface, do. do. Do. 

Lateral view, do. do. Do. 

Ambulacral and inter-ambulacral plates, magnified 
six times. 


3a. Ecarnocypnus muspitia, Woodward, sp. ‘est, natural size. (P. 119.) 


3) 0. my 
3 e. 3 
B) OF ty 
3 eC. 5 


2? 


Or 
g & 


o> 


or Or 


SS 


29 


9) 


.4.  HKcHINocYPHUS DIFFICILIS. 


British Museum. 
Upper surface, magnified three diameters. 
Under surface, do. do. 
Lateral view, do. do. 
Ambulacral and inter-ambulacral plates, 
magnified eight diameters. 


Apical disc, magnified four times. 


. 5 a. Cypuosoma Konier, Mantell. Young test, natural size. British Museum. 


(Peigis) 
Under surface, do. Do. 
Lateral view, do. Do. 
Upper third of ambulacrum, magnified four 
times. 


eevee’ 
2 \ aie 
we a Se, 


C RBone. deletlith. 


PLATE XXIII. 
From the Upper White Chalk. 


Fig. 1 a. Cypnosoma Kéniai, Mantell. Upper surface, natural size. (P. 181.) 


L 6. - 2. Under surface, natural size. British Museum. 

xe? - - Lateral view, do. 

ah a - Upper part of ambulacral area, magnified three 
times. 

be: _ - Portion of the areas and pores, magnified three 
times. 

ay A i : Boss and mammillon, highly magnified. 

leas re a dnferior portion of an ambulacrum, magnified 


three times. 


Nig. 2 @. Cypnosoma GranuLosuM, Goldfuss. Upper surface, natural size. (P. 129.) 


276. £ a Under surface, natural size. British 
Museum. 
2G. ss a Lateral view, showing the depression 


of the upper surface. 
2d. 2 » Portion of the areas and pores, magnified 
four times. 


Fig. 3. Cypnosoma Konic1, Mantell. Hlevated variety. British Museum. 


CRBone del et lith 


PLATE XXIV. 
From the Upper White Chath. 


Fig. 1 a. Cypnosoma Konter, Mantell. A fine specimen, with the spies 7 siti, 
natural size. British Museum. (P. 131.) | 


L 4. Fe Ramiform spine, natural size. 
ve: . f . do., do., magnified three times. 
Ld. . - Bent spine, magnified twice. 

Fig. 2. " Test, with spines. Rev. 'T. Wiltshire, F.G.S. 

Natural size. 

3a ‘3 - Bent spine 
3b. ~ do., magnified six times > British Museum. 
4. fe ie Spatulate, do. 
6. © : Inter-ambulacral plate, magnified six times. 
ie - 3 Elevated variety. British Museum. 


Bone, de] et lith. 


Fie. 


Fig 


L 3b. 
LG: 


ld. 


pone 
Dua: 


2) 


> 


° 


PLATE XXV. 


Cyphosomas from the Upper White Chalk. 


3) 


. CypHosoma Konier, 


Mantel. 


1 a. CYPHOSOMA MAGNIFICUM, Agassiz. Natural size. Rev.'l. Wiltshire. (P. 137.) 


Upper surface, magnified twice. Do. 
Under surface, do. Do. 


Lateral view, do. Do. 


Upper part of areas, magnified four diameters. 
Base of ambulacrum, do. do. 


Tubercle, magnified six diameters. 


Mould in flint. Upper surface. (P. 131.) 
Do. Under do. 


, 
4ou90 


“- 
ee) ry] a 


PYRO? 


C.R Bone. del et ith. 


PLATE XXVI. 
Cyphosomas from the Upper White Chalk. 


Fig. 1 a, 6, c. CypHosoma Konter. Spoon-shaped spines, magnified. The Rev. ‘I’. 
Wiltshire, F.G.S. 


2 i % Club-shaped spine, do, Do. 
SB Pr : Awl-shaped spine, magnified six times. Do. 
AG bb. . Conical and spoon-shaped, do. Do. 
Fig. 7 a. CypHosoMa CorROLLARE, Klein. Test, natural size. British Museum. (P. 134.) 
7 6. 3 i Upper third of an ambulacrum, magnified six 
times. 
(ee © ,; Inter-ambulacral plates, do. do. 
8 a. % cf Upper surface of another test, magnified twice, do. 
8 6. s % Lateral view of do., do., do. 
9. mn Under surface of _ do., do., do. 
10. + ‘ Ambulacral area, magnified six times. The 


Rev. ‘I. Wiltshire, F.G.S. 


C.RBone, del et lith. 


PLATE XXVII. 
From the Upper White Chalk. 


Hig. 1 a, Cypnosoma Wetnerenit, Forbes. ‘Test, natural size. Museum of the Royal 
School of Mines. (P. 1839.) 


1: - ‘The upper surface of do., magnified twice. 

le P, e The under surface of do., do. 

evs 4 a The lateral view of do., do. 

le. - " Areal plates and zones, magnified six times. 

Baya e i Lateral view of a single tubercle, mag- 
nified. 

lg. * E; Upper portion of an ambulacrum, magnified 
six times. 

ee 


e - Inferior portion of do., do. 


ea re 
OAR ¢ GS 
Ly og ' y ~ 
% s 
a 


, PHO 


C R.Bone, del et lith. 


PLATE XXVIII. 
From the Upper White Chalk. 


Fig. 1 a. CypHosoma sparutireruM, Morbes. Test, natural size. British Museum. 


(P. 141.) 
1 2. 4 - Upper surface of do., magnified twice. 
Le: ss Under surface of do., do. do. 
1 d. .: es Lateral view of do., do. do. 
ae: o a Inter-ambulacral plates, do. six times. 
lf - aa Ambulacrum entire, do. do. 
1 g. Fs a Inter-ambulacral plates, do. do. 
1h: B rr Lower portion of an ambulacrum, mag- 


nified six times. 


N 


Ms 


= 
) 
7 


GeP et ec O/ 


Fr 
= 
7 
2 
4 
» 
ao 
= 
= 
Ss 
mo 
pa 
oOo 


PLATE XXIX. | 


From the Upper and Lower Chalk. 


Fig. 1 a. CypHosoma spatunireruM, var. The Rev. ‘I’. Wiltshire, F.G.S. (P. 141.) 


1 6. ss m Upper surface of do., magnified twice. 
ae: 5 4 Lateral view of do., do. do. 
1d. ri he Areal plates and zones, do. six times. 


2 a. CypHosoma siMPLEX, Forbes. Royal School of Mines. (P. 148.) 


2 6. i 5 Upper surface of test, magnified two and a 
half times. 

2G: 5: 7 Upper portion of ambulacra, do. six times. 

Bis : - Another test, magnified three times. 

3 4. -) s Radiated areola, magnified eight times, Museum 


of the Royal School of Mines. 


€ RBone.deletlith. 


PAE XOXO: 
From the Chloritic Mart. 


Fig. 1 a. Pepinopsis Wisi, Wright. Test, natural size, upper. From the Collection of 
W. Wiest. (P. 114.) 


Ls, Be; a, Under surface of do., natural size. 

l ¢. PY a Lateral view of do., do. 

Lia: : " Upper third of an ambulacrum, magnified six 
times. 

le. ee a Inter-ambulacrum plates, do., do. 

hve be m Do. dom |) doy do. 

We ” , Areal plates and zones, _—do., do. 


2.  Prprnopsis Meripanensis, Cotfeau. Copied trom Pl. 1125, fig, 2, of the 
‘ Paléontologie Frangaise,’ for compari- 
son with P. Wiesti. 


foe 


Go Oy 


22 


sai 


nf? Dig oe 


a) Fe 
3299/9909 


»o% 9 59 0,938 


ay Psa o¥ 
2779 2.008 


a  —- 


VAN 


CR.Bone del.et lith 


PLATE XXIX s. 


From the Upper and Lower Chalk. 


Vig. | a. GLypHocYPHUS RADIATUS, Heninghaus, sp. ‘Test, upper surface, magnified 
twice. (P. 121.) 


1p: a 2 Lateral view, magnified twice. 
Le. > i$ Areal plates and zones, do. ten times. 
Fig. 2a ts " Under surface, do. twice. 
2 6. » “ Apical disc (from M. Cottean’s figure), magnified. 
2¢ $5 3! Areal plates and zones (do. do.) do. 
2 d. o " Inter-ambulacrum (do. do.) do. 
2 e. - Pe Ambulacrum (do. do.) do. 


Pig. 3 a. Ecutnornuria ELoris, Woodward. Portion of test, natural size, in the British 
Museum. (P. 125). 


3 6, »9 y Poriferous zones, magnified four times, do. 
Fig. 4 a. r a Do. do., do. do. 

4 6. ba _ Portion of spine, do. six times. 

Ae, 5 0 Do,  “dox do. do. 

Ad. Sy x Areal plates, do. four times. 

Dd. i * Ovarial plates, natural size. 

5 6 : ” Jaws, forming ‘“ the Lantern of Aristotle,” 


natural size. 


CRBone delet jith 


— LN Ni 


— 


PALAONTOGRAPHICAL SOCIETY. 


INSTITUTED MDCCCXLVIL. 


VOLUME FOR 1869. 


CCCLXX. 


ti 


A MONOGRAPH 


OF 


BRITISH BELEMNITID © 


BY 


OHeN PML LELIPS, 


M.A. OXON., LL.D. DUBLIN, LL.D. CAMBRIDGE, D.C.L. OXON., F.R.S., F.G.S., ETC., 
PROFESSOR OF GEOLOGY IN THE UNIVERSITY OF OXFORD. 


BAR. Vi, 
CONTAINING 


Paces 109—128; Pratres XX VIJTI—XXXVI. 


LONDON: 
PRINTED FOR THE PALHZONTOGRAPHICAL SOCIETY. 
1870. 


PRINTED BY 
J. E, ADLARD, BARTHOLOMEW CLOSE. 


BELEMNITES OF THE OXFORD CLAY. 109 


BELEMNITES OF THE OXFORD CLAY. 


In passing upward from the thin-bedded rocks of Stonesfield, we find few or no 
Belemnites for a considerable thickness of the Oolites. Through the whole series of the 
Great Oolite, Bradford Clay, Forest Marble, and Cornbrash, Belemnites, if ever found, 
are very rarely seen in the South of England. In the North of England, the doubtful 
“Grey Limestone,” as I termed it, of the Yorkshire Coast, contains Belemnites, but they 
are of the type of Inferior Oolite, and with Ammonites Blagdeni, Am. Humphreysianus, 
and dm. Parkinsoni, wust be held to carry that rock to the “ Lower Badonian” stage. Is 
the Great Oolite of the South of England wholly devoid of Belemnites, except in its 
lowest member, the Stonesfield slate? I can only reply that no specimen has occurred 
to my personal observation. Does the Bradford Clay contain any Belemnites? Only 
one notice is on record, and that is in the now rarely seen volume, published by W. 
Smith, under the title of ‘Stratigraphical System of Organized Fossils,’ 1817. In that 
work, page 79, occur these words :—“ Multilocular bivalves. Belemnites small, slender ; 
Stoford.” As my boyish hand wrote the words—the place being familiar to me, I have 
no reason to doubt the accuracy of the record. ‘The specimen was transferred to the 
British Museum. No Belemnite is mentioned in the Forest Marble beds, nor, so far as 
I now remember, has any one been quoted from the Cornbrash, except by error. In the 
first edition of my work on the ‘Geology of the Yorkshire Coast,’ 1829, I remarked 
(p. 145) “No Belemnites have been found in the Cornbrash of Yorkshire ;” and again 
(p. 146), “The Cornbrash is the only conchiferous stratum in the eastern parts of 
Yorkshire from which Belemnites are excluded.” In consequence of some notice reaching 
me of a specimen found in the Cornbrash of Yorkshire between 1829 and 1835, I 
modified the expression in the Second Edition, so as to call attention to the extreme rarity 
of the occurrence. If any Paleontologist whom these remarks may reach should find 
himself able to furnish me with specimens of Belemnites from beds between the Corn- 
brash and Stonesfield slate, of any part of England, he would oblige me much by a sight 
of them. 

There being then, as appears, this great blank in regard to Belemnites (the remark is 
almost equally good for Ammonites, but in this case we must exclude the Cornbrash), 
through a considerable range of conchiferous strata, it becomes a matter of great interest 
to compare the several Oxonian forms which now appear, with the numerous Badonian 
species which have disappeared. Are these the same species matured in some other 
part of the sea, modified there through a long succession of transmitted forms, and again 
brought into the Oolitiferous ocean? We may consider this question after the facts 
have been collected and studied. 

Among the Belemnites of the Oxford Clay and the Kelloway Rock (a sandy member 

17 


110 BRITISH BELEMNITES. 


which is not seldom absent from the section), four principal forms appear to have reached 
maturity in the area of England, which may, for convenience, be termed ‘ Hasfati,’ 
including B. hastatus of Blainville; ‘ Canaliculati,’ including B. sulcatus of Miller; 
‘Tornatiles” including £B. Puzosianus of D’Orbigny; and ‘ Lvcentrici,’ including 
B. abbreviatus of Miller. When we endeavour to trace the history of these several forms, 
from the youngest examples, we experience in more than the usual degree the difficulty 
of obtaining a series of all ages for any one of the species. 

However carefully we may collect, in many favourable localities, it is nearly impractic- 
able to fill up all the terms of the series ; and though scores of young ones have been col- 
lected by my own hands from different localities, it is only in a few instances, and by the 
aid of my pupils, that I have succeeded in proving to my own satisfaction the real 
progress of these several forms towards maturity. Nor does the method of examination 
by sections of the older individual succeed im this case so well as in some others, because 
of the frequently very close texture of the sparry substance, and its more complete con- 
densation into an amber-coloured mass, than is usual in the earlier deposits. For this 
reason the form of the very young shell cannot always be even approximately known by 
examining polished or natural sections. As far as can be judged from these sections, 
however, there is reason to think that most of the Belemnites of the Oxford Clay began 
life in a more or less hastate, or else lanceolate, shape; and this seems to be confirmed 
by the fact, as I believe it to be, that no very small specimen has ever been observed in 
the Oxford Clay or Kelloway Rock in England, except in one of these shapes. Extending 
our view to Scotland, we find a somewhat different result. The Belemnites of the 
Cromartie coast have been collected very successfully at Hathie and Shandwick by Lieut. 
Patterson, of Ripon, to whom I am obliged for the sight of his fine series, and for 
photographs of many specimens. 

T'wo species at least have there arrived at maturity ; one, a peculiar elongated spicular 
Belemnite, whose guard sometimes reaches the length of ten inches, is found in Mr. 
Patterson’s series of all sizes down to one inch: it is only in this very small and very 
slender specimen that any approach to a fusiform shape of the guard can be recognised, 
and then only in a very slight degree. Another of these species makes an approach to 
B. sulcatus of Miller, and is longitudinally grooved up to the point, at least in all the 
smaller specimens (Shandwick).’ There is nothing in the smallest of these at all com- 
parable to the clavate forms common in the Oxford Clay of England, though a slightly 
hastate shape can be recognised among them. The strata from which these Belemnites 
come have been called Lias, but what Ammonites and Conchifera I have seen from them are 
of the Oxonian type of life. 

Two of the four Oxonian groups have been already mentioned in the Badonian 


1 In Lieut. Patterson’s Collection is one specimen of a decidedly Liassic Belemnite of the group of 
B. elongatus (Miller), which is placed among the Shandwick fossils. 


BELEMNITES OF THE OXFORD CLAY. PM 


Oolites (Hastati and Canaliculati) ; the others (Zornatiles and Excentric?) now first make 
their appearance, to replace (perhaps we may say) the Gigantei, which they, however, 
resemble in no particular except size. 


BELEMNITES HASTATUS, Blainville. Pl. XXVIII, figs. 67—70. 


fteference. Var.a. Belemnites hastatus, Blainville, ‘Mém. sur les Bélemn.,’ p. 71, 

pl. 2, fig. 4, 4a. 1827. 

Belemnites semihastatus, Blainv.,‘ Mém. sur les Bélemn.,’ p. 72, 
pl. ui, fig. 5, 5a—5y. 1827. 

Belemnites gracilis, Phillips, ‘ Geol. of Yorkshire,’ vol. i, p. 138, 
pl. v, fig. 15. 1829. 

Belemnites hastatus, D’Orb., ‘Terr. Jurass.,’ p. 121, pl. xviii, 
fig. 1,9. 1842. (Exclude some of the synonyms.) 

Belemnites hastatus, Quenstedt, ‘ Cephalopoda,’ p. 442, pl. xxix, 
fig. 27—39. 1849. 

Belemnites semihastatus rotundus, Quenstedt, ‘ Cephal.,’ p. 440, 
pl. xxix, fig. 8—1l1l. 1849. 

Belemnites semihastatus depressus, Quenstedt, ‘ Cephal.,’ p. 440, 
pl. xxix, fig, 12—18. 1849. 


Guarp. Very elongate, smooth, hastate, with an acute apex (by decay of lamine 
about the alveolar apex it becomes fusiform) ; in all stages of life depressed and expanded 
laterally in the post-alveolar region ; cylindrical or somewhat compressed about the alveolus ; 
ventral surface marked by a distinct groove, which is extended forward over the alveolar 
cavity, and backwards toward the apex, about half the length of the axis of the guard, so 
as to leave much of the expanded part free from groove, or with merely a faint indication 
or trace of groove. In young specimens this part appears swollen on the ventral aspect 
(see fig. 69, 2?, 2+). 

Transverse sections show the lamin of the guard thickening on the dorsal aspect of the 
alveolar cavity, so as to make the section circular, or a little oblong there ; the sectional out- 
line is depressed and reniform in the post-alveolar region, till the bulbous part is reached, 
which has an oval section gradually growing circular toward the conical or slightly sub- 
mucronate point. The axisis nearly central. In some excellent specimens are small faint 
longitudinal bent or sigmoidal furrows one on each side of the guard. They begin nearer 
to the ventral than the dorsal surface, and bend upward before losing their distinctness 
(figs. 6827’, 672). 

Dimensions. In the Oxford district within a few miles of the city, the clay-pits have 
been well searched for these Belemnites, and with great success as far as the young 


112 BRITISH BELEMNI'TES. 


forms are concerned. Only one specimen has yet been met with corresponding in size to 
the figures of D’Orbigny and Quenstedt already referred to. This is in the collection of 
Mr. James Parker, from the Cowley clay pit at its deepest part, about 80 feet below the 
Calcareous Grit. This specimen has a total length of 6°75 inches ; greatest breadth 0°65 ; 
least 0-40 ; greatest depth 0°60; least 0°40; axis = 5°5 inches. Between this fine and 
solitary specimen and very many examples 2! inches long, no intermediate magnitudes 
have yet, near Oxford, been found; the smallest specimens, like oat grains, are about half 
an inch long, and then by decay of the lamine: about the alveolar apex acquire the aspect 
of the so-called ‘ Actinocamax’ of Miller. 

Proportions. In young specimens the axis is about seven times as long as the greatest 
post-alveolar breadth, and about ten times as long as the breadth at the alveolar apex : in 
an old specimen, the axis is fourteen times as long as the alveolar breadth, and nine times 
as long as the greatest post-alveolar breadth. 

It was with great pleasure that I received from the Oxford Clay of Eyebury, near 
Northamptonshire, a specimen found by Mr. Leeds (Pl. XXVIII, fig. 68), which happily 
fills up the blank in the history of the species, by a form of intermediate magnitude, not 
elsewhere recognised. We are thus assured of the persistence of the hastate form in this 
species through all stages of growth yet observed, from the very young to the apparently 
full-grown individuals. 


Puracmocons. Very few indications of this part of the fossil have been as yet seen 
by me. D’Orbigny, who had fine specimens at his disposal, figures the phragmocone of 
one (‘ Terr. Jurassiques,’ pl. viii, fig. 1) m the sheath, and represents it as having an angle 
of about 15° (in the description it 1s said to be 11° to 18°), with chambers whose 
diameter is only four times their depth. Quenstedt (‘Cephal.,’ pl. 29, fig. 8 a, 9) presents 
the phragmocone of 2B. semihastatus rotundus (regarded as a variety of B. hastatus by 
D’Orbigny), with septal intervals equal to one fifth of the diameter, and an angle of 13°. 
These may be regarded as good characters for discriminating between this specific group 
and that of B. ari-pistillum of Stonesfield. The septa are more nearly round than in the 
figure of D’Orbigny. 


Localities. In Oxford Clay, Weymouth ; in the middle part of the clay north of the 
town; on the shore in the upper part of the clay; and south of the town. Oxford, in 
the lower or middle part of the clay, with Ammonites Duncani, at Summertown, one mile 
to the north; in the upper part of the clay at Cowley Field, half a mile to the south-east ; 
and at Long Marston, in the upper part of the clay, one mile to the north-east (PAilips). 
Eyebury, near Peterborough, in the lower part of the clay (Leeds). St. Ives (Walker). 
Scarborough Castle Hill (PAzdips). In Calcareous Grit, Scarborough (Bean). 

Observations. D’Orbigny collects under one title the two fossils to which Blainville 
assigned the names of Belemnites hastatus and B. semihastatus. he differences between 
them were far from clear in the earlier author’s descriptions or figures. Quenstedt 


BELEMNITES OF THE OXFORD CLAY. 113 


retains the distinct names, and is, in this respect, in agreement with other German 
writers. Collections in Germany follow this model in their arrangements. D’Orbigny 
joins to these Belemnites fusiformis of several writers, B. ferrugineus of Voltz, and 
B. gracilis of Raspail. 

In considering the varieties to which the species seems liable, we find among English 
specimens, of lengths from half an inch to three inches, some differences in the general 
shape of the guard, which in some specimens is elegantly hastate (fig. 69, v°), in others 
more expanded and recurved at the apex (fig. 69, 7, wv’), in a few bulbiform 
(fig. 69, v°, ov’), m one deformed (fig. 70, u). ‘The ventral groove is generally 
absent from half of the length of the axis of the guard; in rare cases (fig. 69, v’) it is 
interrupted, sometimes it leaves more than half of the length of the axis of the guard 
smooth. I do not observe lateral furrows on any of the small specimens, and it seems 
rarely absent from any of the larger ones. In the older form, which exceeds six inches 
in length, the undulated, or somewhat oblique faint double furrow, is traceable nearly as 
D’Orbigny has represented it (‘'Terr. Jur.,’ pl. xviii, fig. 4). 


BELEMNITES HASTATUS, var. BULBosUS. Pl. XXVIII, fig. 69, v', v7. 


The variety to which attention is now called, is more than any other remarkable for 
the retral expansion of the guard and the swollen ventral outline of the expanded part. 
In eleven specimens before me, including individuals from 3 inch to 23, the characters were 
nearly uniform. ‘The broad part of the guard is about jrd of the axis from the apex ; thie 
groove usually terminates at half the length of the axis from the apex; but in one 
specimen (fig. 69, v’) it is interrupted and farther extended. 

The sections are nearly round in and for a small space behind the alveolar portion, 
but everywhere further back they are,clliptical. ‘The lamine over and a little behind the 
alveolar space are, as usual, pale and less calcareous than in the more solid part of the guard. 

Locality. Specimens of this very interesting form have been forwarded to me by 
Mr. J. F. Walker, of Sidney Sussex College, Cambridge, from the Oxford Clay of 
St. Ives, Cambridgeshire, where they are accompanied by B. Puzostanus of D’Orbigny, 
and 2B. sulcatus of Miller. 

I have examined in foreign collections a considerable number of specimens called ZB. 
hastatus and B. semihastatus, and considered the figures which are given as representing 
them. Iam unable to perceive differences among them or among the comparatively few 
English examples of full size, such as to require the employment of more than one 
specific name. At the same time there are differences; some have a nearly circular 
section across the expanded part of the guard, others a depressed contour there ; similar 
variations occur in the alveolar region. ‘lo the former the ‘ variety-name’ of rofwndus 
has been assigned, to the latter depressus. The somewhat flexuous lateral groove is 
absent in some and present in other examples not otherwise differing. 


114 BRITISH BELEMNITES. 


On a Group oF BELEMNITES, INCLUDING 2B. C4éNALICULATUS, SCHLOTHEIM (IN PART) ; 
B. suLcaTvs, MILLER (IN PART); B. ALTDORFENSIS, BLAINVILLE 2. ABSOLUTUS, 
Fiscurr ; B. Beavuonrranus, D’OrBIGNY. 


The canaliculated Belemnites above referred to are frequent in the Oxford Clay, and 
specially toward the lower part of it, as it occurs in England. They are found in the 
vicinity of Oxford, associated with Ammonites Duncani, in the parallel of the Kelloway 
~ Rock, or nearly so, for that rock is hardly traceable in this quarter. In the corresponding 
clay of Weymouth, Belemnites are found of the same general character, while at St. Neot’s 
specimens occur which cannot in the least particular be distinguished from Oxford 
specimens. 

Miller, while examining the Oxford Collection, certainly referred the channelled 
Belemnite of the neighbourhood to B. sulcatus ; but a frequent application of this name is 
to a species of the Lower Oolite, such as B. apiciconus. We find in Schlotheim 
B. canaliculatus corresponding to B. sulcatus of Miller, and, like it, cluding forms from the 
Inferior Oolite and the Oxford Clay. Blainville rightly separates them, and assigns to his 
B. Alidorfensis one of Miller’s figures (pl. viii, fig. 5, ‘ Geol. Trans.,’ 2nd series, vol. ii), and a 
part of B. canaliculatus of Schlotheim. Quenstedt employs the general title of B. canali- 
culatus for all these forms, and includes in it the Stonesfield fossils referred to B. Bessinus 
by Morris and Lycett. 

Belemnites having the same general character occur in the Oolitic series of Russia, with 
Ammonites of the Oxford Clay; and similar forms have come to my hand from the 
Himalaya. 

Among all these fossils there is so much of resemblance that in the sense of the term 
species, as it was employed by the earlier naturalists who thought with Linneus, they 
might be classified under one title, such as B. canaliculatus, the earliest on record, as Quen- 
stedt does. But this title is equally claimed for the grooved Belemnites of the Bath Oolite 
series, which contain several very distinguishable and characteristic forms. 

B, Altdorfensis of Blainville is supposed by this author to be identical with B. cana- 
liculatus of Schlotheim and B. su/catus of Miller, and is quoted from the ferruginous Oolite 
of Curey, near Caen. 

D’Orbigny disposes of the perplexity of this nomenclature by institutmg a new 
species, B. Beaumontianus, which he refers to the Lower Oxford Clay of Vaches-Noires. 
A fossil, corresponding to his figure, occurs at Loch Staffin im the Isle of Skye, according 
to Prof. E. Forbes. No other locality is given by Moris. 

Upon the whole I am disposed to preserve the name which Miller certainly imposed 
on the long-grooved fossils from the Oxford Clay; the more so as it will be seen that 
hardly any examples fit so exactly with the figure of B. Beaumontianus given by D’Orbigny 
as to render that a good general type of a variable species. 


BELEMNITES OF THE OXFORD CLAY. 115 


Few Belemnites appear to have had so large a distribution in time and space as the 
group allied to B. sulcatus of Miller and B. canaliculatus of Schlotheim, From the base 
of the Inferior Oolite to the middle of the Oxford Clay they are generally recognised in 
Europe ; specimens much like our examples from Oxford Clay are abundant in the country 
south of Moscow; others come to us from the Himalaya’, from Cutch’, South Africa’, 
New Zealand,‘ and Queensland.’ 


BeLemnites sutcatus, Adler. Pl. XXIX, XXX; figs. 71—75. 


Reference. Belemnites sulcatus, Miller, ‘ Geol. Trans.,’ 2nd series, vol. 11, p. 59, pl. vin, 
fig. 5 [excl. fig. 3 and 4]. 1823. 
Belemnites Beaumontianus, D’Orb., ‘ Pal. Frang., Terr. Jur.,’ p. 118, 
pl. xvi, fig. 7, 11 (on the plate it is called B. Altdorfensis). 1842. 


Guarp. Subcylindrical or conical im the alveolar region, more or less depressed in 
the post-alveolar region and deeply grooved ; the groove interrupted or expanded toward the 
apex, and gradually ceasing about the alveolar summit. Outline nearly straight on the 
ventral, more curved on the dorsal aspect; apicial region tapering, surface smooth or 
granulated. Sections show the axis to be nearest the ventral face, very excentric, and 
somewhat recurved. Near the apex the sections are almost circular or a little oblong. 

Greatest length observed 5:5 inches ; and of this the axis occupied 3 inches. Greatest 
diameter in this specimen 0°85, in a stouter specimen 1°05. 

Young. The very young form was more or less hastate (fig. 73 v'). Somewhat 
advanced in age is the very rarely seen form fig. 73 v, from near Oxford; next we have 
fig. 72 and 71 v, differing from full-grown specimens only in greater slenderness. 

Proportions. The normal diameter (not counting the groove) being taken at 100, 
the transverse diameter of the alveolar apex is, in full grown individuals, 108 ; the axis 
under 300; the ventral radius 40 ; the dorsal 60. In young specimens the axis is 500. 

Puracmocone. Slightly arched, very obliquely inserted ; septa nearly circular, un- 
usually approximate, their depth being about one eighth or even only one ninth of the 
diameter in the anterior part; sphericle distinct and rather large; angle 22°. The 
concave surface, within the septal edge, is a portion of a sphere, measuring 90° across. 

Observations. ‘There is some variety in the sections of the guard; some specimens 
showing more depression than others ; in some the groove is broader, in others it is deeper ; 
in a few the groove expands a little toward the apex (fig. 74 v), and also expands on the 
surface over the alveolus (fig. 75 v’) ; in some there are one or two lines parallel to the edges 

1 Specimen in my possession. 2 Sowerby, ‘ Geol. Trans.,’ 2nd Ser., vol. v, p.329. 

S$ Tate, in ‘Geol. Soc. Journal,’ 1867,p.151. * Hochstetter, Novara-Expedition. 

5 Specimens in the Collection of Mr. Charles Moore. 


116 BRITISH BELEMNITES. 


of the groove (fig. 75 v’ and v’). There is often a marked increase of depth and definition 
of the groove for half the length of the guard, measuring back from the alveolar region, as 
if in that part was a fissure (fig. 71, o’, v). A faint mtimation of the groove can almost 
always be traced to near the apex. 

Specimens occur with an external sheath of white fibrous matter, rough on the outer 
surface (Pl. XXIX, fig. 72m). One might fancy this to be a periostracon or capsule, 
but it is, I believe, really a concretionary deposit. The shell is sometimes granulated 
(Pl. XXIX, fig. 72 m). 

In figure 10, pl. xvi, of the ‘ Pal. Frane.’ the outline of the alveolar cavity, erroneously 
represented as somewhat transverse, should have been very nearly circular. The Oxford 
specimens are never so much depressed in the post-alveolar region as in fig. 9 of the same plate. 

The axis of the guard of this Belemnite, in some specimens obtained from the Oxford 
district, is hollow for a part of the length, as if the apices of the young laminz of the 
guard were, during life, removed, so that a sort of pipe, partially interrupted at intervals 
by the edges of these laminze, extended inwards from the perforated apex. Afterwards 
the sheaths successively formed covered them completely, and were not perforated. In some 
specimens (fig. 71s’, a’, a”) the axial canal is very narrow for a certain space above the 
alveolar cavity, then it enlarges in a fusiform shape, and again contracts to the mere line 
of junction of the opposite guard-fibres. This curious appearance will be further considered 
im connection with B. abbreviatus, 

Another very curious fact is observed in several of these fossils. On the ventral 
aspect, internally, are one or two cavities extended lengthways, through the substance of 
the guard, from a little in front of the alveolar apex to a greater distance behind. An 
explanation is found by the aid of cross sections: for these, taken a little behind the 
alveolar apex, show the cavities in question to be formed by the peculiar imflexion of the 
laminze of the guard on the ventral aspect. This inflexion becomes remarkable only after 
a certain age ; thenceforward grows continually deeper and deeper, always producing a 
groove, and sometimes by the contraction of this groove completely or partially enclosing 
longitudinal canals. 

Fig. 71s shows the arrangement of the lamine round the axis of the guard in 
conformity with this description. The axis is not tubular in this instance. ‘The lamine 
of the guard are crossed by the fibres nearly at right angles to the surface, and as this is 
a curve of contrary flexure about the ventral aspect, the fibres assume there remarkably 
arched directions. In these sections glistening dagger-shaped parts are present—they are 
merely the obliquely truncated prismatic cells of the so-called fibrous structure. It may 
be well to mention, that the specific gravity of most Belemnites (2°8) agrees with that 
carbonate of lime called arragonite, and not with ordinary calcite. 

The student of Homology will not fail to remark the analogy which this repetition of 
deep folds on the ventral aspect of Belemnites sulcatus offers to the more regular groove 
on the same aspect in Belemnitella. The groove of the latter group, however, is only on 


BELEMNITES OF THE OXFORD CLAY. Ly Wy 


the alveolar region, and reaches to its anterior edge, which is emarginate in consequence, 
while that of B. sulcatus belongs to the posterior part, dies out on the same region, 
and ceases nearly opposite the alveolar apex. ‘The canaliculated axis occurs in some 
examples of B. Bessinus of Stonesfield and in BZ. /ateralis of Speeton, but I have not 
yet seen it i any Liassic species. 

Belemnites perforatus, Voltz (Pl. VIII, fig. 2), from Cretaceous beds at Osterfeld, is 
canaliculated for the whole length of the axis of the guard; and specimens of B. guadratus 
and B. mucronatus from the Chalk frequently show this peculiarity, or else a condition of 
the central parts which suggests their easily acquiring it. 

Locality and distribution. Weymouth has yielded characteristic specimens of this 
species from the Oxford Clay, but they seem not to be plentiful there. 1 found only two 
or three fragments in the clay on the shore north of Weymouth, mixed with hundreds of 
the young forms of a hastate Belemnite. It is not mentioned among the fossils of the 
Oxford series known to Smith, who figures and describes the longer Belemnite known at 
Chippenham as B. Owenii. About Oxford we find it rather frequently, especially towards 
the middle and lower part of the clay deposit, with Ammonites Duncani, while in the 
upper part B. Owenit and B. excentricus occur more frequently, with Ammonites vertebralis. 
The young forms are very rare in these parts. Near St. Neot’s, again, they occur with 
Ammonites Duncani, but not plentifully, as I find by Mr. Walker’s communications, and 
again near Peterborough, as I learn from Mr. Leeds. I doubt the occurrence of the 
species in Yorkshire, and regard the mention of it in the first edition of my work on the 
geology of that county (1829) as requiring confirmation. 

The locality of D’Orbigny’s fossil is thus noticed: —‘ Elle a ¢té recueillée par 
M. Tesson dans les marnes Oxfordiennes des Vaches-Noires (Calvados); elle parait y 


étre rare.” 


On a Grove or BreLemnites ALLIED To BezeunirEs PuzosrtaANus OF D’OrBIGNY. 


In 1816 Wilham Smith figured, in his ‘Strata Identified,’ on the plate of Oxford 
[Clunch] Clay fossils, a long subcylindrical Belemnite from Dudgrove Farm, in Wilts. 
In his ‘ Stratigraphical System’ (1817) the fossil is described as “large, squarish, quickly 
tapering to the apex; diameter one inch at the largest end, length four or five inches.” 
The figure referred to represents the guard almost complete, with the alveolar cavity 
exposed. I remember the specimen, which is now in the British Museum. Some years 
later the species was recognised by my great relative in a fossil of the Kelloway Rock 
in Yorkshire, to which I gave the name of JB. éornatilis. Of this I had seen 
specimens when the first edition of my work on the ‘ Geology of Yorkshire’ was published 
(1829), and described the fossil as elongated. In the second edition it was named, 


with an equally brief description, but no figure (1835). In 1844 the rich deposit of 
18 


118 BRITISH BELEMNITES. 


Chippenham had yielded its treasures, and Belemnites of the same general aspect, with 
considerable portions of the phragmocone, and even extensions of the conotheca, had 
furnished to Prof. Owen the materials for a valuable essay on the structure of the 
shell and the relations of the animal (‘ Phil. Trans.,’ 1844). The fossil was named by 
Mr. Pratt B. Owenit. 

The same great record of science contains, in the volume for 1848, another “ Essay 
on the Belemnites of Chippenham,” by Dr. Mantell, in which the figures represent a 
variety of important facts previously unobserved. The Belemnite which he examined 
is here called £&. attenuatus—a name long before appropriated to a species found 
in the Gault, which had, however, been referred to a new genus, Belemunitella. 
D’Orbigny makes known to us a very similar form of Belemnite, also from the 
Oxford Clay, to which he gives the name of B. Puzosianus. Finally, Mr. Morris, in his 
excellent ‘Catalogue of British Fossils,’ 1854, employs the term B. Oweni, giving 
B. Puzosianus as a synonym, and under B. tornatilis proposes the question if it be not 
identical with B. Owenzt. 

The natural group thus noticed consists of Belemnitic guards of more than the usual 
length, with a generally cylindrical aspect, more or less compressed; always marked by 
a depression, often by a conspicuous groove, from the apex along the ventral surface 
for a third or half the length of the axis. 

To the forms best known must be added one or two more from the midland district 
of England, and as many from the Oolitic series of the coast of Cromarty. These are 
inordinately long, but in other respects correspond in general character with the more 
usual species. 

Looking back upon earlier groups of Belemnites, we find nothing so much like these 
as the long, somewhat compressed forms allied to B. tripartitus (see Pl. XI, fig. 28), 
in the Upper Lias. But all those Liassic forms have lateral grooves near the apex, often 
very conspicuous ; these of later ages, in the Oxonian strata, never. 

In regard to the synonymy, there can be little doubt about preserving Mr. Pratt’s 
name, B. Owenii, for the whole group; B. Puzostanus, D’Orbigny, having certainly to be 
associated with it, as a variety. 


BeLemnites Owrnul, Pratt. Pls. XXXI, XXXII, figs. 76—81. 


leeference. Belemnites (unnamed), Smith, ‘Strata Identified, 1816, and <‘ Strati- 
graphical System,’ p. 55, 1817. 
5. tornatilis, Phillips, ‘Geol. of Yorkshire, vol. 1, ed. 2, 1835 (no 
figure). 
B. Owenii, Pratt, ‘ Phil Trans.,’ 1844. 
B. attenuatus, Mantell, ‘ Phil. Trans.,’ 1848. 


BELEMNITES OF THE OXFORD CLAY. L19 


B. Oweni?, Quenstedt, ‘Cephalop.,’ pl. xxxvi, f. 9, 1849. 
B. Puzosianus, D’Orbigny, ‘ Pal. Frang., Terr. Jur.,’ p. 117, pl. xvi, 
f. 1—6, 1860. 


This frequent species, or group of species, varies much in several important 
characters. The degree of compression is by no means uniform, but I have seen no 
example of alveolar compression approaching to that represented in D’Orbigny, fig. 4, 
pl. xvi. All my specimens have in that part a slightly elliptical section. In some the sides 
of the guard are flattened or a little grooved (PI. XX XI, fig. 77); the apicial region is some- 
times unusually compressed, but generally follows the sweep of the sides ; the ventral groove 
near the apex varies from little more than a mere flattening (Pl. XXXII, fig, 79) to a 
broad furrow (fig. 76, v’), a sharp short rift (fig. 77, v’), a narrow groove (fig. 76, 0”, 
and fig. 78), and a deep lengthened canal (figs. 80 and 81). ‘The general figure, always 
long, varies in the proportion of length to diameter. 


Guarp. Very long, subcylindrical, more or less compressed, tapering evenly to a 
point, grooved on the ventral aspect from the apex through one third or more of the 
length of the axis; in perfect specimens this groove is often bistriated, or somewhat 
sharply bordered. 

Sections show the axis placed nearer to the ventral surface ; in young specimens the 
compression is considerable, growing less with age; there is sometimes a distinct lateral 
flattening on the middle part of the guard. 

Greatest length observed, in specimens from St. Neot’s, 10 inches, of which the axis 
is 6 inches; the diameter at the alveolar apex 1 inch. In another the diameter at the 
alveolar apex is 1} inch. 

In a very young state the pearly lamine about the alveolar apex are sometimes 
decomposed, and the guard assumes the delusive shape of ‘ Actinocamax. A drawing 
has been shown to me in which this fusiform guard, or ‘ossicle,’ is represented as 
separated from the ‘nucleus’ of the phragmocone, but I have seen no specimen of the 
kind. Prof. Owen figures (‘ Phil. Trans.,’ 1844, pl. 1, fig. 4) a very young individual, 
with the guard and alveolar chamber in their ordinary relations. The guard is in this 
state shorter in proportion than in after-life. 

Proportions. (na full-grown specimen from St. Neot’s the diameter from back to 
front, at the alveolar apex, being taken at 100, that from side to side is 90, the axis is 
600; ventral radius 45, dorsal 55. ‘The section is slightly oval, the ventral face rathe: 
broader than the dorsal. 

In a young specimen, 3 inches long, the proportions of the diameters are also 
100 to 90; the axis is more excentric than in the older specimen, the ventral radius 
being only 30, the dorsal 70, the axis 650. 


120 BRITISH BELEMNITES. 


Puracmocons. Known in a crushed state by specimens from near Chippenham. 
The uncrushed phragmocone has a slightly elliptical section. D’Orbigny gives a very 
elliptical section (55 to 45). The characteristic angle, as given by D’Orbigny, is 16° 30’. 
I have no good specimen of this part of the fossil. 

Locality. In the Oxford Clay of Wiltshire, Oxfordshire, Northamptonshire, Hunt- 
ingdonshire. In the Kelloway Rock of Yorkshire. In the Oxford Clay at Vaches-Noires 
(Calvados), and Marquise, near Boulogne. 

Observations. Specimens are often found invested with a sheath of white fibrous 
matter, externally rough and of granular aspect, within which the true shell is always 
smooth and shining. 


I remark the following varieties : 


1. Batemnites Owent (Puzostanus). Pl. XXXI, fig. 76; Pl. XXXII, figs. 78, 79. 


The guard is smooth and always compressed; the apicial furrow distinct or faint, 
never more than half the length of the axis; alveolar section elliptical. This is the ordi- 
nary form from the Oxford Clay of the Midland Counties. In one middle-aged specimen, 
corresponding to B. attenuatus of Mantell, lateral grooves extend all along the post-alveolar 
tract. 


2. BeLEMNITES OwEnrI (verRucosus). Pl. XXXI, fig. 77. 


The surface is ornamented with small, raised, smooth puncta, and undulations com- 
posed of these united. The distribution of these may be seen on the ventral face (v), the 
lateral (/’), and the dorsal (@’). On the first it wiil be noticed that the puncta disappear 
towards the apex, and diverge and disappear on the alveolar region. On the sides they 
show more of a tendency to gather in linear groups ; on the back this concurrence of the 
puncta makes short undulated ridges, which grow larger, but more dispersed, on the 
alveolar region. ‘The apex shows signs of very short plications. Only one specimen is 
known to me, found by Mr. J. E. Walker, at St. Neot’s, with Ammonites Duncani. The 
reader may compare the curious granulation in this specimen with that on B. infundibulum 
(Pl. I, fig. 3), with that on specimens of Belemnitella granulata, and with the diverging 
ornaments on a Sepiostarium. 

If further research should produce additional specimens, possibly there may be found 
reason to adopt a specific name for this fossil. But the surface-ornament being at present 
the only difference observed between this guard and ordinary specimens of B. Owenii of 
the same size, I prefer to mark it as a variety. 


BELEMNITES OF THE OXFORD CLAY. 12] 


3. Brremnires Owen (rorNnatiLis). Pl. XXXII, fig. 80. 


The guard is very smooth, less compressed than in the typical forms; more cylin- 
drical, with a longer, deeper, and narrower ventral furrow (fig. 80). ‘This furrow, indeed, 
occupies the greater part of the axial length of the guard; in middle-aged specimens an 
old specimen shows some trace of lateral flattenings. From the Kelloway Rock of 
Hackness and Scarborough. It is not unlikely that this may be found to deserve to be 


regarded as distinct specifically. 


BELEMNITES stTRicosus, n.s. Pl. XXXII, fig. 81. 


Guarp. Very long, slender, cylindro-conical, compressed, acuminated, smooth, with 
a distinct longitudinal furrow drawn from near the apex, on the ventral face, through 
two fifths of the length of the axis, and thence continued in a slighter depression 
towards the alveolar region. 

Transverse sections of the guard show an oval contour, the sides flattened; the 
ventral face broader than the dorsal; in the alveolar region the dorsal part of the shell is 
much thicker than the ventral part. 

Greatest length of the one specimen seen 7} inches, of which the axis is 63 inches ; 
greatest diameter #3 of an inch, 

Proportions. The diameter at the alveolar apex from back to front (3, of an inch) 
being taken at 100, that from side to side is about 80, ventral radius about 40, dorsal 


radius about 60, axis 1600. 


Puracmoconr. Unknown. ‘The alveolar section is nearly circular, the angle appears 
to be about 20°. 


Locality. In the upper part of the Oxford Clay, in Cowley Field, near Oxford; one 
specimen, presented to the University Museum by W. B. Dawkins, M.A., the first 
Burdett-Coutts Scholar. 

Observations. 'This remarkable fossil carries to extreme length the essential characters 
of the group of tornatile Belemnites, the cylindro-conical outlines, the slight compression, 
the apicial groove, and low angle of phragmocone. Having but one example to con- 
sider, I am unable to describe the variations due to age and accident, but it would be 
very agreeable to be furnished with evidence on these points. I have seen no foreign 
specimens corresponding with this species; but D’Orbigny’s fig. 3, pl. xvi, somewhat 
resembles it. A thin white external layer appears on the specimen, not the fibrous layer 
noticed in B. sulcatus and B. Owenii. 


122 BRITISH BELEMNITES. 


BELEMNITES sPicuLaRis, n.s. Pl. XX XVIII, fig. 82. 


Guarp. Cylindrical (hastate when young), tapering evenly to a point, much com- 
pressed to an oval section, with a faint ventral groove drawn from the apex through 
two fifths of the length of the axis; a few striz about the apex, especially on the dorsal 
aspect. 

Transverse section oval, the ventral face broader than the dorsal. Substance varied 
by bands of brown (sepia-tint) and honey-yellow spar. 

Greatest length observed 10 inches, greatest diameter 1 inch. Shortest specimen 1 inch 
long ; it is of the form Pl. XXXII, fig. 82 7. 

Proportions (old). Takmg the diameter at the aveolar apex at 100, the diameter from 
side to side is 90 +, the axis 1000; the excentricity of the axis variable, in some speci- 
mens small, in others the ventral radius = 40, the dorsal 60. 


Puracgmocone. Incompletely known. The section is elliptical, within a ring of the 
guard-fibres everywhere of nearly equal thickness ; the phragmocone section more elliptical, 
therefore, than the section of the guard. ‘The angle in one of Lieut. Patterson’s specimens 
appeared to be 18° at the apex, 15° in a more advanced part of the shell. The apex of 
this phragmocone was placed at about one third of the diameter from the ventral margin. 


Locality. Hathie Burn, and Shandwick, on the coast of Cromarty: collected in great 
abundance and in excellent condition by Lieut. Patterson, who gave me much informa- 
tion as to the circumstances under which he obtained the specimens and the accom- 
panying fossils. He further assisted my researches by presenting to me a set of 
photographic representations of much interest. 

The fossiliferous strata of the Mesozoic system on this coast have been usually described 
as Liassic, and on a first view of the shale and these Belemnitic fossils such an opinion 
might be readily adopted. The Belemnite now in question has analogy to some of the 
long species of the Upper Lias, such as 2. tripartitus, while the next to be mentioned 
seems to revive the memory of B. longissimus of Miller. Their affinity, however, is with 
the long species of the Oxonian stage in the Oolitic system. Among the accompanying 
fossils I observed in Lieut. Patterson’s collection Gryphea dilatata, large and small ; 
Perna ; Avicula Braamburiensis ; Pleurotomaria ; Ammonites resembling, if not identical 
with, 4. vertebralis, Sow., A. ewcavatus, Sow., A. flecicostatus, Phil., A. plicatilis, Sow., 
A. Gowerianus, Sow., A. biplexr, Sow; scales of Lepidosteus ; cervical vertebrae of 
Ichthyosaurus. The Belemnites form a bed in the shale. 

Observations. It is difticult to fix upon any definite characters by which to distin- 
cuish this Belemnite from B. Owenii, except the greater proportionate length of the axis 
and the faintness of the apici-ventral groove. ‘he slight strize about the apex are only 
seen on one or two specimens. 


BELEMNITES OF THE OXFORD CLAY. 123 


BELEMNITES OBELIscus, n.s. Pl. XXXIII, fig. 83. 


Gvuarp. Very long, almost uniformly tapering to a point, compressed, smooth, or with 
traces of longitudinal interrupted undulations. In some specimens a defined lateral 
flattening (Pl. XXXII, /’, 7°). No distinct apici-ventral groove. 

Greatest length observed 97; inch; greatest diameter in this specimen, just before 
the conical expansion, less than } an inch. In shorter specimens, 6! inches long, the 
corresponding diameter is nearly the same ; in still smaller examples, 3} inches long, the 
diameter is } of an inch. It seems as if two varieties exist, one much longer in propor- 
tion than the other. 

Proportions. The normal diameter at the alveolar apex being taken at 100, the trans- 
verse diameter is 84; the axis in the longer variety 2000 and more, in the shorter 1500 
The excentricity of the axis appears to be very small. 


Puraemocone. I have only been able to observe the cross section, which is less 
elliptical than the sectional outline of the guard, the guard-fibres being longer on the 
back and front than on the sides. In this the fossil is analogous to some Liassic forms. 


Locality. Hathie Burn and Shandwick, with the last-named species. 

Observations. Not only do the unequal proportions of different specimens suggest the 
idea of a sexual distinction, but the whole group, compared with B. spicularis, leads to 
reflections of the same order. The guard is colour-banded, as is that of B. spzcularis. 


On a Group oF BELEMNITES ALLIED TO BELEMNITES EXCENTRICUS OF BLAINVILLE 


Lister, in his ‘ Historia Anim. Angliz,’ pp. 226-228, has the following description oi 
a Belemnite of this group :—‘Titulus xxxii—Belemnites niger, maximus, basi forata.” 
Among the remarks on this species we find “ Perfricatum cornu combustum aut quoddam 
bitumen olet.” “In tota illa agri Eboracensis regione montosi, qui Blackmore appellatur, 
precipue abundant ; item in rivulo juxta Bugthorp, et alibi reperti sunt.” ‘The Black- 
more fossils belong to B. abéreviatus; a large fragment was above three inches in cireum- 
ference. Bugthorp is on the Lias. 

Lhwyd, in the ‘ Lithophylacium Britannicum,’ notices Belemnites of this group, from 
the vicinity of Oxford, No. 1667 :—Belemnites maximus oxyrrhynchus, four inches in 
girth where largest. Cowley, Bullington, Marsham, Stansford, Garford, the localities 
mentioned, indicate the species to have been what Miller called 2B. abbreviatus. 

Smith, in the ‘ Stratigraphical System,’ p. 50, describes a Belemnite as elongate, rathe1 
four-sided, from Wotton Basset and Shippon, in the Coral Rag, p. 43, and another, quite 
similar, from the Kimmeridge Clay of North Wilts. 

Miller described these forms as B. abbreviatus ; his followers have often assigned that 
name to a species from the Inferior Oolite. 


124 BRITISH BELEMNITES. 


Young and Bird, in their volume on the Yorkshire coast (ed. 1, 1822), notice a 
similar Belemnite, and give a figure (pl. xiv, fig. 4), and name it B. eacentralis, describing 
it as found in the “ Oolite, Upper (Speeton) Shale, and Chalk.” This is incorrect, but, as 
will be seen, the large Speeton Belemnite belongs to the same natural group. 

De Blainville, in 1827, describes and figures in his pl. iii, fig. 8, 8a, B. excentricus, 
froni Vaches-Noires, remarking that Miller’s B. abéreviatus much resembles it. 

D’Orbigny revives Young and Bird’s name for a species which he figures (pl. xvii) ; 
but im the text (p. 120) he makes no reference to those authors, and uses the name 
given by Blainville. 

In the second edition of the first volume of my work on the ‘Geology of Yorkshire’ 
I restored to the great Belemnite of the Malton Oolite the name assigned by Miller, and 
mentioned the large Speeton Belemnite as 2. /ateralis. An undescribed form in the 
Kimmeridge Clay of Oxfordshire, and another in the Tealby beds of Lincolnshire, will 
complete this series of excentral Belemnites, as far as I know them. 


BELEMNITES ABBREVIATUS, Miller. Pls. XXXIV, XXXV, figs. 84—93. 


Reference. B. niger maximus, Lister, ‘ Hist. Anim. Anghi,’ p. 226, 1678. 

Belemnites maximus oayrrhynchus, Lhwyd. (No. 1667.) 1699. 

B. excentralis (in part), Young and Bird, ‘ Geology of the Yorkshire 
Coast,’ pl. xiv, fig. 4, 1822. ; 

B. abbreviatus, Miller, ‘Geol. Trans., 2nd series, vol. ii, pl. vu, 
figs. 9, 10, 1828. 

B. excentricus, Blainville, ‘Mém. sur les Bélemn.,’ p.90, pl. in, f. 8, 1827. 

B. excentricus (also called excentralis), D’Orbigny, ‘ Pal. Frang., Terr. 
Jur.,’ p. 120, pl. xvu, 1842. 


Guarp. Cylindrical; sides flattened or somewhat hollowed longitudinally ; apex 
produced, compressed, sometimes incurved; ventral surface broader than the dorsal; a 
flattening near the apex, on the ventral surface. 

Very old specimens have the apicial region much compressed, produced, and incurved ; 
sides flattened by broad, shallow, longitudinal depressions, which continue over a part of 
the alveolar region, and are there gradually lost. 

Young specimens are slightly hastate, very young ones distinctly so, with little trace 
of the lateral hollow. 

Longitudinal sections show the axial line to be very excentric, especially so in the 
retral part of the guard, and in old specimens considerably curved. 

‘Transverse sections present a somewhat four-sided outline, the ventral surface being 
struck to a flatter curve than the dorsal, and the sides flat or a little concave. 

The length of a very large example is 8 inches; of another smaller, but extending 


BELEMNITES OF THE CORALLINE OOLITE, ETC. 12: 


wt 


farther along the phragmocone, 11 inches; the greatest diameter before the conical 
expansion of the sheath over the phragmocone 1} inch. The smallest which has occurred 
to me is little more than 1 inch long. 

Proportions in full-sized specimens. ‘Taking the dorso-ventral diameter at the alveolar 
apex at 100, the transverse diameter is about equal to it, the ventral radius is 32, the 
dorsal 68, the axis 250, justifying Miller’s title of B. absreviatus. In young specimens 
the axis = 300. Cross sections near the apex show a still greater excentricity, the axis 
curving towards the ventral surface. 


Puracmoconr. Conical, a little incurved towards the ventral line, with an almost 
perfectly circular section ; sides inclined at an angle of 18°, except near the apex, where 
it is greater (above 20°). The septa are numerous, lie at right angles to the axis, with 
plain unwaved edges, and are penetrated by a marginal siphuncle. 


Locality. Yn the Coralline Oolite and Calcareous Grit of Yorkshire at Malton, Scar- 
borough, &c. In the same strata at Heddington, Cowley, Bullington, &c., near Oxford ; 
near Calne, Weymouth, &c. In France, at Vaches-Noires and Marquise in Oxford Clay. 
D’Orbigny quotes it from near Moscow. In Oxford Clay, Cowley, near Oxford. In 
Kimmeridge Clay, Shotover. 

Varieties. In progress from youth to age, this Belemnite experiences considerable 
changes, as may be inferred from what has been said in respect of the guard. Besides 
these ordinary changes of form and proportion, it appears desirable to distinguish two 
types of general -shape, which occur in large specimens in some degree of relationship to 
the stages of the strata. 


a. BELEMNITES ABBREVIATUS (OXYRHYNCHUS). 


_ Large, cylindroidal, slightly bent, with incurved, produced, flattened apex. Viewed 
on the front or back, the sides are seen to contract rather suddenly from a cylindroid 
part to the apex in the architectural form known as ogee (fig. 84, ) ; viewed sideways, the 
dorsal outline is continued in a convex form to or nearly to the apex, while the ventral 
outline becomes concave under the apex (Pl. XXXIV, fig. 84, 7). 

Locality. The Coralline Oolite of Malton, Oxford, and Wilts. An abnormal specimerf, 
which places these characters in a strong light, is presented (Pl. XX XV, fig. 86) from 
near Oxford. 


B. BELEMNITES ABBREVIATUS (EXCENTRICUS). 


Large, conoidal, with sides almost straight, converging through the whole post-alveolar 
19 


126 BRITISH BELEMNITES. 


space of the guard, the dorso-ventral diameter being in that part much greater than the trans- 
verse diameter. The general figure is that of Belemnites explanatus (Pl. XXXVI, fig. 96). 

Locality. The Oxford Clay, upper part, at Cowley, near Oxford. The Calcareous 
Grit and Coralline Oolite near Oxford. 

This variety agrees well with the description and figure of Blainville (( Mém. sur les 
Bélemnites ’). D’Orbigny makes the phragmocone section to be more elliptical than it 
usually is, and the axis less excentric than usual ; there is also something about the outlines 
not as we commonly see them. 


Remarks ON SPECIMENS OF BELEMNITES ABBREVIATUS, VAR. EXCENTRICUS, IN THE 
CaBinet oF Mr. Werurre.u. 


The axial line of the guard is in many instances excavated into a canal which grows 
narrower towards the apex. ‘This is especially the case in specimens obtained from 
the Drift of Finchley, near London, from which a great variety of fossils of the Oxford 
Clay and other strata lying to the northward has been obtained by Mr. Wetherell. In 
the large collection of that gentleman are very many excellent examples of this struc- 
ture, and by careful study of them in comparison with other undisturbed specimens in 
the Oxford Clay, Calcareous Grit, and Oxford Oolite, we arrive at a clear view of a very 
curious subject, of which, at first sight, it might seem difficult to form a correct opinion. 

Fig. 88, Pl. XX XV, represents the surface produced by a splitting fracture through a 
Belemnite in Mr. Wetherell’s collection. Fractures of this kind are not infrequent in 
nature, and are easily produced by intention. The surface thus presented is usually flat 
and smooth in the ventral portion, as if a natural fissure existed there, but commonly 
uneven and more or less hackly m the dorsal portion. The hollow left by the spherule 
is sometimes traceable at the apex of the alveolar cavity, the phragmocone being generally 
absent in the specimens under consideration, but not seldom the alveolar cavity contracts 
gradually to the canal without any distinct enlargement at the alveolar apex. 

In the figure referred to the canal is seen to contract gradually until it finally dies 
out before reaching the apex. Examined with microscopic powers of 10 and upwards, 
the canal is found to be crossed by many ridges at nearly equal intervals, so as to 
suggest the appearance of an annulated or half-chambered canal, in continuation with the 
cavity or the spherule of the phragmocone (see fig. 91). The seeming septa of this canal 
are found by more careful research to be the truncated edges of the successive laminze of 
the guard (see figs. m’, m”), each conspicuous lamina giving origin to one septum. This 
appears quite certain under the lowest power of a good achromatic microscope, which 
discloses, moreover, that the laminz thus referred to appear to be often composed of two 
or three thinner layers, some dark, others paler, and probably more nacreous in substance. 


BELEMNITES OF THE KIMMERIDGE CLAY. 127 


Following the canal till it closes, the lamine are seen to lose their truncations, and to 
acquire the complete curvature. 

After careful study of many specimens, no doubt remains in my mind that the canal 
has been produced by the removal of the apices or terminal parts of the interior lamine 
of the guard. ‘This process began at the alveolar cavity; it happened during life, and 
was occasioned by decay and absorption of the apices in the earlier stages of life. 

That these special parts might be of somewhat different composition from the other 
parts of the lamin is suggested by some other cases in which terminal porosity and an 
axial canal have been noticed; and it is quite in agreement with two other circumstances 
to be observed in these fossils. First, it is to be remarked that the alveolar cavity in these 
Belemnites often appears marked by the undulated anterior edges of the lamine of the 
guard, which terminate in this cavity (see figs. 88, 91, 92), and show white, thin, sparry 
plates, in consequence of the removal of parts of the lamine. And again, some of 
the specimens show a curious appearance of a second canal gomg from the alveolar 
cavity (figs. 90 and 92) near its apex. This, being carefully studied, is found to be occa- 
sioned by the removal of some of the laminz of the sheath for a certain space inwards 
from the alveolar cavity, leaving a kind of slit where the removal has happened. 

In later life the deposited sheaths were, in general, not removed by decay or absorp- 
tion (see figs. 87, 88). 


ON THE BELEMNITES OF THE KIMMERIDGE CLAY. 


After diligent search in this clay near Oxford, where it is about 1 00feet thick, and is pretty 
well exposed in brickyards and in quarries of the Coralline Oolite, and after a careful search in 
the escarpment of Portland, I find, speaking generally, a remarkable accordance between 
its Belemnites and those of the Oxford Clay known as B. abbreviatus excentricus, B. Owenii, 
and B. hastatus. This analogy was, perhaps, to be expected, inasmuch as Ammonites of 
the groups of 4. vertebralis and A. biplex occur in both clays. 

Taking first the specimens allied to B.adéreviatus eacentricus,it would, I think, be difficult 
to assign characters of sufficient weight to claim a specific distinction, though in old speci- 
mens the ventral surface is more flattened towards the apex, and in young specimens the 
whole of the guard is depressed behind the alveolar region. In this respect the young 
forms closely resemble those of B. Souvichei (D’Orbigny, ‘Ter. Jur.,’ pl. xxii, figs. 4—8), 
which was found in beds referred to the Portland series at Hauvringhen (Pas de Calais), 
and at the Tour de Croi, near Boulogne. ‘These forms differ from those of the same age 
from the Oxford Clay and Coralline Oolite. 

Next, we may consider the longer forms, like B. Owenii, of the Oxford Clay. Of these 


128 BRITISH BELEMNITES. 


some appear tome quite indistinguishable from their analogues in the older deposit ; they 
occur of equal magnitude with them, but not in equal abundance, in the upper part of 
the Kimmeridge Clay of Shotover Hill, where it was cut through by the railway. One 
extremely lengthened variety of this Belemnite occurs at Shotover, reminding us of 
B. spicularis from the shore of Cromarty. 

Besides these is ayoung depressed Belemnite much like the young B. Aastatus of the 
Oxford Clay ; these occur near Oxford and in Portland Isle. 


BELEMNITES EXPLANATUS, n. s. Pl. XXXVI, figs. 94, 95, 96. 


Gvuarp (old). Conoidal, tapering gradually toa rather compressed apex ; sides more 
or less broadly channelled ; ventral aspect flattened and somewhat expanded, becoming 
concave towards the apex (a few dorsal strize about the apex are sometimes seen), 

Dimensions. Axis about 3 inches; diameter at the alveolar apex 0°85 inch. 

(Young.) Depressed, smooth, flattened on the ventral aspect, and hollowed, or 
marked by a narrow groove towards the apex, which is slightly curved ; sides more or less 
marked by a shallow continuous furrow (a very young form is almost fusiform). 

Dimensions. Axis in the smallest about 1 inch, with a diameter of 0°25 inch. 

Proportions. Axis in young specimens 400—450. The diameters at the alveolar 
apex 100 from front to back, 115 from side to side. In old specimens the axis is about 
350, the transverse diameter 107, the dorsal radius 64, the ventral 36. 


Locality. In the upper part of the Kimmeridge Clay of Waterstock, near Thame. 
Specimens of different ages— young (not middle-aged) and full-grown—have been 
presented to the Oxford Museum by Mrs. Ashworth. In the upper part of the same 
clay at Hartwell, near Aylesbury, with Cardium tnequistriatum, Astarte Hartwelliana, 
and Ammonites bipler. In the Kimmeridge Clay, upper part, where cut through in the 
railway-tunnel, at Wheatley, near Oxford. 


Observations. On many accounts this form of Belemnite is of interest in the study of 
the series to which it belongs. On the one hand its resemblance to the older type of 
B. abbreviatus (excentricus) of the Oxford Clay and Oolite, and on the other to that of 
Speeton, in Yorkshire (B. /ateralis), is such as to offer a most instructive example for 
study, in relation to the derivation of successive specific forms by hereditary trans- 
mission with modification. But this must be considered hereafter. 


Fig 


EXPLANATION OF PLATE XXVIII. 


67. BELEMNITES HASTATUS. 
Three views of a large specimen from the upper part of the Oxford Clay in 


Cowley Field, near Oxford ; in the Cabinet of Mr. Parker. Fig. 67, /, lateral 
view, showing the rather faint oblique groove ; v, the ventral aspect, with 
its deep characteristic furrow, suddenly followed by a shallower channel : 
d, dorsal aspect; 8’, the cross section of the alveolar cavity; s”, section 
across the expanded part of the guard. 


68. Two views of a specimen from the Oxford Clay of Northamptonshire, presented to 


the Oxford Museum by Mr. Leeds, B.A., of Exeter College. 

Lateral view, showing the oblique groove; v’, ventral aspect, showing the 
deep mesial groove expanding retrally; s’, the cross section of the 
alveolar cavity; s’, cross section of guard in the narrowest part: 


7 


s’”, cross section in the widest part; s””’, cross section near the apex ; 


$, a compressed phragmocone, partly covered by the guard. 


69. Young specimens from the Oxford Clay of St. Ives, presented to the Oxford 


hs 


Museum by Mr. J. F. Walker, of Sidney Sussex College, Cambridge. 
Lateral view, showing a recurved apex ; v’, ventral aspect of the same ; 
s', cross section of alveolar cavity; s’, cross section of enlarged guard : 
v*, ventral aspect of a younger individual; /* and »*, views of a still 
younger shell; v°, 7°, views of a shorter and more bulbous example ; 
v6, v', views of the bulbous variety, with interrupted ventral groove. 


70. uw. Deformity. The specimen is from the Oxford Clay of Cowley Field, near 


Oxford. Cabinet of Mr. Parker. 


, A 


)] 
J 


7 


See ee a AE emai ice * 


(oe 


ee lith, 


EXPLANATION OF PLATE XXIX. 


Fig. 
71. BELEMNITEsS SULCATUS. Specimens from the Oxford Clay at Summertown, 
near Oxford. 

v. Ventral aspect of a full-sized specimen ; the groove somewhat less dis- 
tinctly marked than in other cases, and slightly interrupted. 

v. A younger specimen, showing a more marked change of depth in the 
ventral furrow and its continuation over the alveolar region; s’, cross 
section of the alveolar region of the guard, depressed. 

s. Longitudinal section, from back to front, showing the phragmocone 
in situ, its apicial spherule, and a short lanceolate canal, formed by the 
decomposition of laminze, as shown in a’. 

s”. Another longitudinal section showing very similar facts; the canal 
somewhat more extended in a”. 

s. Cross section of the guard, showing the inflection of the lamine to form 
the ventral groove, and lacunee of a remarkable kind. 

72. Young specimens from Summertown, near Oxford; /, lateral; v, ventral aspect ; 
s, alveolar cross section, nearly round; s’, post alveolar section, nearly round ; 
m, magnified surface, the shell dotted with granules, and covered by a partially 
fibrous layer. Such a layer occurs on some Australian and on some Indian 
Belemnites. 

73. Youngest examples which occur near Oxford, at Cowley, and Long Marston ; 
v, ventral aspect, the groove distinct on the alveolar region; v’, the still younger 


shell; s, alveolar cross section. 


ee 


PL KAA. 


fl 


EXPLANATION OF PLATE XXX. 


Fig. 
74. BELEMNITES SULCATUS. 
Neot’s ; presented by Mr. J. F. Walker, of Sidney Sussex College, Cambridge ; 


/, lateral view, showing a very slight flattening; v, ventral aspect, showing the 


groove widening and growing shallow over the alveolar region, partially inter- 


Specimens of full size from the Oxford Clay of St. 


rupted toward the apex, with striation parallel to the groove; s, cross section 


of guard, slightly depressed. 


75. Specimens from Weymouth. v’, ventral surface, showing the groove growing 


wider and shallower both toward the apex and over the alveolus; 7’, dorsal 
aspect; s, cross section of guard nearly round Fig. 75, 7”, lateral view of 
another specimen, showing a slight flattening; v’, ventral aspect, the groove 
widening over the alveolar and apicial regions, with striz of decomposition 


parallel to the groove ; s”, alveolar cross section; s’”, cross section of guard. 


PL, ARS 
Imp B aon a Paris. 


1 


mam ane 


PB. Lackerbauer lith. 


EXPLANATION OF PLATE XXXI. 


Fig. 


76 


a | 
“I 


. BELEMNITES OWENHI, var. Puzostanus. Specimens from the Oxford Clay of 
St. Neot’s, presented by Mr. J. F. Walker. 

/, full-sized specimen seen laterally ; v’', middle-sized individual, showing the 

apicial groove distinct; v’, younger individual seen ventrally, with its 

apicial groove; /”, side view of the same; v’”, still younger example 


seen ventrally ; and /’”, seen laterally. 


. BELEMNITES OWENII, var. VERRUCOSUS. From St. Neot’s, presented by Mr. 


J. F. Walker. 
v’, seen ventrally ; 7’, laterally ; d’, dorsally ; s’, cross section in the alveolar 


region. 


i 


PL. XXX, 


Imp Beequeta Paris. 


' 


4 


P.Lackerbauer lith. 


Fig. 
. BELEMNITES OwENII, var. Puzostanus. From St. Neot’s, presented by Mr. 


79. 


80 


81 


EXPLANATION OF PLATE XXXII. 


Walker ; v, ventral face, with strongly marked apicial groove reaching to the 
point; /, lateral view. ‘The cross section is oval. 

BELEMNITES OWENII, var. Puzostanus. From St. Neot’s; v, ventral aspect, 
showing an almost evanescent apicial depression ; ¢, dorsal aspect; s, the cross 
section of the alveolar region. 

. BELEMNITES OWENII, var. TORNATILIS. From the Kelloway Rock of Hackness, 


near Scarborough; v, the ventral surface expanding anteriorly; cross section oval. 


. BELEMNITES PORRECTUS, n.s. From the Oxford Clay at Summertown, 
presented by Mr. Dawkins; v, ventral aspect, showing a strong, sharply cut 


apicial furrow, and its anterior extension ; cross section oval. 


Fig. ous 


TE SEEN REE 


EELS RS BCR ELBE a: tN AES ELEY AA AIRES A lt Ma i i A hi 8 CIAL 


‘ict nail 


, 


ackerbauer lith. 


— 


if 


EXPLANATION OF PLATE XXXIII. 


Fic. 
82. BELEMNITES SPICULARIS, 0. Ss. 
Specimens from Shandwick and Eathie, on the coast of Cromarty, collected 


by Lieut. Patterson ; v, ventral aspect of one of the larger specimens, showing 
the apicial groove; /, lateral view, showing the long side flattening, and a 
trace of short apicial grooves; d, the dorsal aspect, marked by some small 


strize ; /’, one of the youngest specimens in Lieut. Patterson’s series ; s, cross 


section of guard at the alveolar apex; s’, section of a smaller specimen 


toward the apex; s”, still nearer the point. 


83. BELEMNITES OBELISCUS, 0. s. 
Specimens collected by Lieut. Patterson at Shandwick and Kathie; /’, one of the 


longest examples, seen sideways; /”, one somewhat smaller, 7”, still smaller, 
and 7", one of the smallest observed ; v’”, ventral aspect of a small specimen, 
and v", of a still smaller; s’, alveolar cross section; s”, post-alveolar section 


across the guard; s’”, farther backward in the guard. 


PL. XXXIIT. 


D, 


1 


3ecquet a raris 


p 
Tap t 


a ae ow te hee A 


iN LE OE tpi PR ia NAS Ree: ais ar SMC vena Mieatnct ds 


a inate sc ER RTE SE 


% 


hinted aes 7 seein” 


in 


BS By Fy ge Se ee ee ee ee a a ae Rigor ante 


P.Lackerbauer ah. 


' 


EXPLANATION OF PLATE XXXIV. 


Fig. 


84. Large specimens of BELEMNITES ABBREVIATUS, from the Coralline Oolite of 
Yorkshire and Wiltshire. 
/. Side view, showing a broad lateral depression, and the incurved apex ; 
v’, ventral aspect, showing a flattening near the apex. The specimen 
is from Malton, in Yorkshire. 

@. Showing the phragmocone iz szfi, and the numerous septa, at right angles 
to the axis. The specimen is from the Calcareous Grit of Seend, in Wilt- 
shire. The drawing was made by Miss Anne Cunnington. 

‘. One of the septa seen axially; s”, cross section of the alveolar region ; 
s”, cross section behind the alveolus ; s”, section near the apex, to show 
the compression. 


$5. Younger individuals from Heddington, near Oxford, and Malton, in Yorkshire. 


seen laterally; ss”, alveolar section of the 


I". Middle-aged specimen 
same. 


example, a little hastate ; v‘, the youngest observed. 


/", A very young specimen seen sideways, a little hastate. 


Specimen seen ventrally, with distinct apicial depression; vo”, smaller 


& 
te 
3 
i) 

€ 

E 


Fig. 


86. 


87. 


88. 


90. 


oie 
92. 
93. 


EXPLANATION OF PLATE XXXV. 


A specimen of BELEMNITES ABBREVIATUS (seen laterally), compressed, and 
unusually bent at the apex; Heddington. 
Section of BELEMNITES ABBREVIATUS, with the phragmocone 7 sift. The 
bending of the axial line of the guard is not often so remarkable as in this case, 
even in old specimens ; it is characteristic of full growth. 


A specimen with less curvature at the point, and less flexure of the axial line. 
From the Drift of Finchley, in Mr. Wetherell’s collection. 


89. Natural section to show the decay of the axial lamine at their apex; Bullington, 


near Oxford. 


The following figures are taken from specimens in the Collection of Mr. 
Wetherell, from the Drift at Finchley. 


Section showing the formation of an axial canal and a vicinal fissure on the 
ventral side. 


Section of another specimen. 
Other sections, in which a small collateral ventral sht appears in the lamine. 


Section in which the canal appears interrupted. 
m,m,m,m*. Magnified views of the laminz in relation to the canal. 


a 
ost 
I 
rm 
A 
2 
by 
3 
Coe] 
4 
PY 


bi 


EXPLANATION OF PLATE XXXVI. 


Fig. 
94—96. BELEMNITES EXPLANATUS, n. s. 
94. Young specimens from Aylesbury, in the upper part of the Kimmeridge Clay. 
v', v’, vo’. Aspect of the ventral face, showing the flattening and slight furrow 
toward the apex. 
7. Lateral aspect, showing the longitudinal depression. 
s’. Section across the alveolar region ; s”, section further back on the guard. 


95. Specimens more advanced in growth, from Waterstock and the Railway-cutting 
in Shotover Hill, in upper part of Kimmeridge Clay. 
. Lateral view, showing the longitudinal groove; v", ventral aspect of the 
same, showing the apicial flattening; o, the same view of a somewhat 
larger individual. 


s”. Section across the alveolar cavity; s‘, across the same cavity near 
its apex. 


96. Full-grown individual from Waterstock, upper part of Kimmeridge Clay. 
v. Seen on the ventral aspect ; the apicial depression wide. 
/“. The same seen sideways; the lateral depression very distinct, the apex 
somewhat bent downward. 
d". The same seen dorsally, where no trace of furrow appears. 
s*. Section across alveolar cavity ; s", section behind the alveolar apex ; @, the 
apex seen axially, to show its compression. 


PL. XXXVI. 


P. Lackerbauer lith. - mp Becquet a Paris 


PALAONTOGRAPHICAL SOCIETY. 


INSTITUTED MDCCCXLVIL. 


VOLUME FOR 1869. 


DCCCLXX. 


A MONOGRAPH 


OF 


mae FISHES 


OLD RED SANDSTONE OF BRITAIN. 


BY 


JAMES POWRIE, F.G.S., 


EH. RAY LANKESTER, B.A., 


JUNIOR STUDENT OF CHRIST CHURCH, OXFORD, AND COUTTS’ GHOLOGICAL SCHOLAR OF 
THE UNIVERSITY. 


PART I (concnupEp).—THE CEPHALASPID/. 
BY 


E. RAY LANKESTER. 


Pages 83—62; Plates VI—XIV. 


LONDON : 
PRINTED FOR THE PALZONTOGRAPHICAL SOCIETY. 
1870. 


PRINTED BY 
J. E. ADLARD, BARTHOLOMEW CLOSE. 


CEPHALASPID J. 33 


disposed concentrically round a middle point, as in P#. Crouchii, but this portion of the 
shield is apparently elevated into a slight convex boss. he surface-striations of P¢. 
rostratus differ considerably from those of Pt. Crouchii. They are very fine, the 
groovings being only ,th of an inch apart near the centre of the disc, and leaving a 
rounded dull ridge with crenate margins between them. 

Localities —This species occurs abundantly at Cradley, near Malvern, Herefordshire ; 
and in beds of the same horizon at Whitbach and in the neighbourhood. It is found 
associated with Scaphaspis Lloydit. 


3. Preraspis MircnEnui, Powrie. Pl. V, figs. 1, 2, 6, 10, 11. 


PreraspPis MITcHELLI, Powrie. Geologist, vol. vii, pp. 170—172, 1864. 
— — Lankester, Brit. Assoc. Report, Trans. Sect., p. 58, 1864. 


I have not had specimens of this species sufficiently perfect to enable me to charac-. 
terise it properly. The specimen figured (Pl. V, figs. 6, 10) was briefly noted by Mr. 
Powrie in the ‘ Geologist,’ in 1864, and an outline-sketch was given. he lateral cornua 
are not seen, but the disc, rostrum, and spine are, to a certain extent. The disc appears 
to be intermediate in form between that of P¢. rostratus and Pt. Crouchii. he specimen 
and others even less well preserved were obtained by Mr. Powrie from quarries in 
Forfarshire, where Heterostraci had been discovered for the first time in Scotland by 
the Rev. Hugh Mitchell. 

A few rhomboidal scales also obtained by Mr. Powrie from this locality (Pl. V, fig. 1) 
probably belong to this or an allied species. 


§ IX. Sectio B.—Osrrostract. 


We have now to consider the forms which are represented by the original type 
Cephalaspis Lyellii of Agassiz, which have bone-lacunz as structural elements of their 
shields, a tubercular ornamentation of the surface, and mesially placed orbits. The 
bodies, with the scales and fins, of some of these fishes have heen discovered, and are of 
the greatest interest, but are not sufficiently common or well known to allow of their 
being used in framing generic and specific distinctions. ‘The division into genera of the 
Osteostraci is based upon the presence or absence of plates in addition to the cephalic 
shield, and upon the form and relation of this additional piece with regard to the cephalic 
shield. 

The species are most safely distinguished by the character of the superficial tubercular 
ornamentation ; where that is not possible the general outline of the head must be 

5 


34 OLD RED SANDSTONE FISHES. 


depended on. But this is a very unsafe criterion, since, owing to the fact of its being 
very thin, the shield is lable to the most deceptive distortion. Examples of this are 
familiar to all who know the fish-heads of the Herefordshire Cornstones. So great is the 
uncertainty in this matter, that I have had much doubt as to the distinction between the 
Scotch type-specimen of C. Zyellit in the British Museum and another Scotch specimen 
in the museum at Arbroath, and again between these two and the common Herefordshire 
species included by Agassiz under Cephalaspis Lyellit. 

Before proceeding to the description of genera and species, in which I shall rely more 
on the figures than on the words which can be used to describe them, remembering the 
motto inscribed by the illustrious Della Chiaje in one of his works, “ Res non verba,’ a 
few general remarks may be offered with regard to the variations of the shield in 
structure and form, and also a general description of the body-scales and fins. 

1. Variations in the Intimate Structure—For a general description of the minute 
structure of the test in the Osteostraci I must refer back to page 7 of this Monograph, 
and also to the plate of microscopic drawings and explanation accompanying this part. 
The microscopic structure of Cephalaspis Lyellii and of the large form called Cephalaspis 
asterolepis by Dr. Harley’ has been carefully studied, but of other species proper material 
has not come to hand. In Awchenaspis and Didymaspis I was able to ascertain the 
presence of bone-lacune of an elongated character, with their long axes crossing in the 
various layers, thus giving the appearance of cross-hatching described by Prof. Huxley. 
In the so-called Steganodictyum Carteri to which I called attention in the ‘ Quart. Journal 
of the Geol. Soc.,’ 1868, p. 547, as being really an Osteostracous Cephalaspid (the 
Steganodictyum Cornubicum having been pronounced a Pteraspid by Prof. Huxley and 
Mr. Salter), I have found small bone-lacunee with very numerous fine canaliculi interlacing 
in every direction. The lacunze were not elongated, and were smaller in size than those 
in the lower portion of the shield of Cephalaspis Lyellit. 

2. Variations in the Surface-markings.—The tubercular ornamentation which appears 
to be a constant characteristic of this section of the ancient Sturgeons presents con- 
siderable diversity of arrangement. In some cases each hexagonal area (described at 
page 8, and figured in Pl. XIV, fig. 7), supports a simple tubercle or one large 
tubercle and a smaller one. In other cases there are many of these tubercles, of a 
smaller relative size, placed irregularly on the area; or, again, there may be a central 
tubercle surrounded by six or more smaller ones, like a mimature volcano with its 
secondary cones. At the margins of the shield, and especially on the ‘cornua,’ the 
tubercles become very closely packed and elongated. ‘The tubercular ornamentation is 
continued on to the under surface of the shield, to some extent, along the enlarged 
margin and its posterior expansions (see woodcut, fig. 12), just as in Péeraspis the 
striations of the surface were shown to continue on part of the inferior surface of the 
rostrum, and along the margin formed by the hollow cornua. ‘The meaning of this 


1 This species is not to be distinguished from the Cephalaspis Salweyi of Egerton. 


CEPHALASPID A. 35 


continuation of the superficial marking to the inferior surface is, of course, simply that 
the parts of the inferior aspect of the shield so marked were as anatomically superficial 
as the tuberculated superior surface. The recognition of this fact is important, as being 
inconsistent with the supposition of a mouth reaching along the margin of the fish’s head, 
which involves the notion that the upper jaw is formed, as it were, by the margin. Probably 
no one acquainted with recent forms allied to the Cephalaspide would hazard such a 
conjecture. ‘The woodcut, fig. 15, is interesting to compare in the consideration of this 
matter. The tubercles on the margin of the shield sometimes assume a very spinous or 
even tooth-like form, e.g. in the species C. asper and Hukeraspis pustulifer (see woodcut, 
fig. 23), and hence it is not surprising that detached portions of the margin with its 
tubercles should have been mistaken for bits of fish-jaw. On the inner side of the 
cornua the tubercles are modified; even the common Herefordshire species presents 
the form of long tooth-like excrescences, and in C. asper, one of Mr. Powrie’s disco- 
veries, the scales, which are always tuberculated in the Osfeostraci like the head-shield, 
present very close-set and acute spinous ornamentation (Pl. X, fig. 5). 

The tubercles themselves are generally round, with a bright and smooth surface, 
similar to that of the intervening matter, both resembling in their microscopic structure 
the ‘cosmine’ of Prof. Williamson. In one species discovered by Mr. Lightbody, of 
Ludlow, in the beds of the Tilestone group which have furnished the Auchenaspis 
Salteri, the tubercles have a curious truncated appearance, giving them a really crater- 
like form. In other parts of the same head-shield, or on what are perhaps additional 
posterior plates, the tubercles are intersected by a network of coarse grooves forming 
an irregular crocodiloid pattern. ‘The tubercles of Auchenaspis are large in proportion 
to the size of the shield; those of Hukeraspis very closely set and small, though not so 
small relatively as those of Cephalaspis Powriei and Cephalaspis Agassizit. 

The exceeding rarity of the preservation of the tuberculated surface in specimens of 
Osteostraci must be borne in mind. I have only one specimen of the common Here- 
fordshire species that shows them at all well. It is much more difficult to get this 
surface in a state of preservation than that of the Pteraspids, difficult as that is to obtain, 
Frequently the lower layers of the Osteostracous shield are preserved, and have been 
mistaken for the true superficies by Prof. Agassiz and by many collectors. When the 
matrix is of sucha character as to preserve well the test, the shield is very generally 
crushed and distorted, so that we are almost invariably offered a choice of evils. 

There are no ‘pits’ on the surface of the Osteostracous shields analogous to those 
described by me in the genera Scaphaspis and Pteraspis, which seem to represent the 
lateral line. 

The scales, like the head-shields in Cephalaspide, are to be looked upon as bones or 
parostotic formations, and not as mere scales. hey belong to the aponeurotic region 
spoken of by Mr. Parker in his great work,’ and, the ganoin-layer being absent from them, 

1 «On the Shoulder-girdle,’ &e. (Roy. Soc.), 4to, 1848. 


36 OLD RED SANDSTONE FISHES. 


there may have existed a separate overlying set of scales, though the ornament renders 
this unlikely. 

Of the Form and Construction of the Shield—In the Osteostraci the shield is very 
greatly convex when not distorted by pressure, attaining an almost hemispherical 
curvature in some parts. The test bends over at the margin, and is continued inwards, 
forming a hollow rim to the shield, which at its posterior angles widens out, as seen in 
the woodcut (fig. 12), forming what may be termed the ‘lateral floors’ of the cephalic 
shield. Since these inflections terminate abruptly, there is no general floor to the cephalic 
box as far as calcareous matter goes. It was completed by a tough skin and a mouth, no 
doubt. The test in Cephalaspis is, moreover, reflected from the margin and floor against 
the roof of the shield, forming a lamina of great tenuity subjacent to the upper tuberculated 
lamina which forms the roof of the shield. This secondary ‘roof’! thins out towards the 
centre of the cephalic plate, and probably became membranous (see woodcut, fig. 13, section ; 
and woodcut, fig. 12, inferior aspect). The hollow rim must be compared with the 
hollow cornua of Pferaspis ; and probably like them it had apertures at its posterior angles, 
close to the attachment of the pectoral appendages (see fig. 16). In the genus Cephalaspis 
proper, which may be placed subgenerically as Lucephalaspis, the shield is ellipsoid in 
its anterior portion, and is produced into cornua of considerable length at each posterior 
angle. ‘I'he median portion is also extended in a backward direction, and in the middle 
line presents a short spine. In the subgenus /emicyclaspis the cornua are not developed, 
nor is the median portion markedly produced, the whole shield having a truncated 
hemispherical outline. In the subgenus Zexaspis the cephalic shield has the character 
of that of Eucephalaspis ; but one at least, and perhaps more, scutes placed dorsally in the 
median line succeeded it. This arrangement paves the way to Sir Philip Egerton’s 
genus Auchenaspis, in which the shield is identical with that of Auwcephalaspis, supposmg 
its median posterior portion but slightly developed, and a large neck-plate soldered to it. 
The subgenus Lukeraspis, which I have associated with Auchenaspis, though no neck- 
plate has been yet found attached to the cephalic portion, possessed exceedingly long 
cornua, extending backwards to more than double the length of the shield itself. Other 
characters of importance render it desirable to form this subgenus of duchenaspis. The 
genus Didymaspis had a neck-plate fully as large as the anterior or cephalic plate, which 
appears to have possessed no cornua. The neck-plate may perhaps be regarded as 
representing in those genera which possess it the distal portion of the compound 
shield of Péeraspis, though it is not advisable to trace a very close morphological 
relation between the two groups /eterostraci and Osteostraci in the details of the 
construction of their superficial armature. The remarks which were made (p. 17) with 
regard to the nature of the suturing or fusion of the pieces of the shield in the Pteraspids 
apply to the similar cases among Osfeostraci with equal force, the difference of histological 
composition in the two cases being, however, duly taken into consideration. 

The position of the orbits, and certain very remarkable concavities and eminences 

' Observed by Prof. Huxley. 


CEPHALASPID. 37 


between and in front of them, are very constant in the Osteostracous shields. The 
structures alluded to are seen in the woodcut (fig. 11), which has been carefully constructed 
from a remarkably well-preserved concave cast of the parts in question of a specimen of 
Cephalaspis Agassizii. Between the orbits is a well-marked tubercle, which I call the 
interorbital prominence (1.p.). In front of each orbit the material of the test is also 
raised into a projecting mass, which is to be called ¢he antorbital prominence (a. p.). 
Between the two aztorbital prominences is a very deep depression of the shield, 
divided beneath the superficial lamina into two parts by a narrow septum continued 
from the inferorbital prominence, which is hollow. ‘The two cavities thus formed 
(see also Plates IX, fig. 2, and Pl. XIII, fig. 4 a) are the antorbital fossa (a. f.).' The 
material of the shield forms a somewhat elevated rg round each eye, which may he 
Fie. 11. Fig, 12. 


m. C. 


>~ 
oe 


——— 
Pinon: 


Mite Dp. Ts p. 
Restored outline sketch of the head-shield of Diagrammatic view, from below, of the cephalic shield 
Eucephalaspis Agassizii. of a Eucephalaspis. 


a. f. antorbital fossz, i. p. interorbital prominence, o. r. orbital rings, i. g. interorbital groove, p. 0. v. post-orbital valley, m. c. marginal cells, 
p. c. posterior cornu, p. a. posterior angle, p. 7. posterior ridge, p. s. posterior spine, 7. rim or margin. 


designated the orbital ring (o.r.). Posterior to the interorbital prominence, and more strictly 
placed between the two orbits, which in some specimens it has the appearance of uniting, 
like the nose-saddle of a pair of spectacles, is a deep well-defined impression, which 
may be distinguished as the cnterorbital groove (7. g.). Immediately behind this, and 
reaching backwards towards the posterior spine, as it approaches which it narrows, 


1 These occupy such a position as to suggest a connection with the olfactory organ, which hence may 
be inferred to have been double in these fishes—a fact which makes it certain that, though these are the 
earliest fishes yet found, they are very far indeed from the first Vertebrates, their ancestors or forerunners ; 
they are also, it seems likely, far in advance of the ‘ Protamphirine’ of Haeckel. 


38 OLD RED SANDSTONE FISHES. 


is a flattened oblong area, which is even a very little concave, and may be named 
the postorbital valley (p. o. v.). Quite anteriorly and near the margin or rim of the 
shield two round masses are indicated by dotted lines (mz. ¢.). Like the double antorbital 
hollow, they do not appear superficially, but are very constantly to be observed 
im specimens, since the superficial portion of the test is so rarely preserved. They 
are really areas in which the concavity of the inferior aspect of the shield is not 
maintained in its regularity, but increased between the two lamine. In Hukeraspis 
(Pl. XIII, fig. 12) they are very markedly developed all round the margin of the 
shield, and are very conspicuous modifications of its structure. They may be termed 
marginal cells. It may, perhaps, seem superfluous to designate each of these 
parts by a distinct name, but it is of great importance that they should be clearly 
recognised and distinguished, since they are present or are represented in all 
genera of Osteostraci, and were they better known in all the species would un- 
doubtedly furnish valuable diagnostic characters by their modification and variation. 
A glance through the plates will enable the reader to recognise some of them 


s. superior lamina. z. inferior lamina. m. margin. 


in nearly every specimen figured, whilst in most the crushing or fracture of the 
specimen has obliterated some or other of them. The letters p.c. imdicate the 
posterior cornua, p.r. the posterior median ridge, p. s. the posterior spine, and p. a. the 
posterior angles of the shield. \t would be important to ascertain with what soft parts 
these protuberances and cavities were connected, and what relation they may have to 
similar structures in recent fish; but into this inquiry, as well as that relating to the 
‘central pit’ in front of the disc of Cyathaspis and Pteraspis, I do not propose now to enter, 
such an investigation being unnecessary in a descriptive monograph of fossil remains. 
Scales, Fins, and Form of the Body in Osteostraci—The two specimens of 
Cephalaspis figured by Agassiz, in his ‘Recherches sur les Poissons fossiles,’ gave 
some evidence as to the nature of the parts of the body in these fish, but our 
knowledge on these matters has been very much extended by the discoveries of 
my friend and ccadjutor Mr. Powrie in Forfarshire, and by other ‘finds’ in the 
same district. Ido not at all suppose that the details ascertained with regard to the 
genus Cephalaspis are necessarily true for all the allied forms ; indeed, the existence of 
large dorsal plates in the subgenus Zenaspis clearly shows that they are not; but as yet 
the only Osteostraci (the only Cephalaspide, indeed) of which we have remains of the 
body (excepting a few scales of Pteraspis) are Cephalaspis Lyellii, C. Pagei, and C. 


CEPHALASPID. 39 


Powriei, and others less satisfactory of the same subgenus. What is said here, there- 
fore, must be accepted as based merely on that limited amount of evidence. 

In the first place, it is probably a very constant character that the scales of the body 
present the same tubercular ornamentation as do the head-shields, since it is observed in 
all the specimens discovered, and is parallelled in the Heferostraci by the identity of orna- 
mentation of the scales and shield of P¢eraspis. The body in those forms known is very 
thin, small, and tapering, probably embracing but a small portion of the viscera, which were 
rather covered in by the so-called head-shield. ‘The scales (in Hucephalaspis) are in four 
chief series on either side the median line—a dorsal, of oblong rectangular form ; a /aferal, 
of much longer proportions ; a marginal, projecting from these at an angle ; and a ventral, 
meeting in the median line on the ventral aspect, as do the dorsal on the dorsal aspect. In 
Lucephalaspis the scales are in a single row in each series, with the exception, probably, of 
the ventral series, where they appear to be broken up into four or more (Pl. XI, fig. 2). 


Fie. 14. 


Loricaria platystoma, Giinther. 


The specimen referred to in the Arbroath Museum is the only one which clearly shows 
this remarkable series of scales. Posterior to the dorsal fin the three well-defined rows 
of scales, visible when the fish is in a lateral position, become broken up and confused in 
a continuous armour of rhomboid scales. The manner of the breaking up of the series I 
do not know, since no specimen is well preserved in this part; but it may be conjectured 
from the numerous analogous cases among Siluroids. Prof. Agassiz was led to suppose 
that the dorsal series of scales was double on either side, by the observation of the rhom- 
boid scales belonging to the region behind the dorsal fin, and probably also by Mr. Dinkel’s 
very imaginative representation of the profile specimen (fig. 1, pl. 1, in the ‘ Poissons 
fossiles ; and in ‘Siluria,’ 1867, pl. 36, fig. 3), now in the possession of Sir Philip 


40 OLD RED SANDSTONE FISHES. 


Egerton. I was completely astonished on comparing the specimen kindly lent to 
me by Sir Philip with the figure which has been so widely copied. The specimen 
presents the concave surface of half of the head-shield and the inner surface of the 
body, showing well the large lateral scales; but none of the detail as to dorsal scales 
which the artist has drawn is to be seen, and there is no real foundation for assigning 
two rows to each dorsal series of scales. In Péeraspis undoubtedly there are many 
dorsal rows of nearly equilateral rhomboidal scales, and one may be quite prepared to 
find that such was the case in some Osteostract. ‘he restored outline sketch (woodcut, 
fig. 16) of a Cephalaspis (Lucephalaspis Lyellii) gives a better notion of the relative pro- 
portions of the scales than mere words can, especially when compared with some of the 
plates. But it is necessary to state again that the way in which the series of scales break 


Fie. 15 a, 6. 


= 


Loricaria platystoma, Gunther, a recent fresh-water fish, of the Siluroid group, from Surinam.! 


up at the dorsal fin is not known. ‘The scales are thicker and broader in C. Lyedlii than 
in C. Powriet relatively. They are deeply imbricated in each series, and have a consider- 
able thickness of structure, histologically agreeing, as far as I have been able to examine 
them, with that of the head-shield. The body in vertical transverse section presents a tri- 
angular form as seen in fig. 17. It appears that the scales of the lateral series admitted 
of considerable movement of the body, though so large and strong; for, whereas in the 
specimen drawn in Pl. VHI they are inclined posteriorly, in that drawn in PI. XI they 
present exactly the opposite direction. This is, no doubt, due to muscular contraction, the 


1 I have to thank the Secretary of the Zoological Society for the use of the cuts, figs. 14 and 16, 
which illustrate a paper by Dr. Giinther. 


CEPHALASPIDA. 


scales becoming overlapped to a greater or less extent, accord- 
ing to the angle assumed. Probably each scale indicates a 
sclerotom. 

The fins are two pectorals of very peculiar form and character, 
a dorsal placed posteriorly and a caudal. The dorsal and the 
caudal were known to Prof. Agassiz ; but the pectoral appendage, 
though indicated in Mr. Dinkel’s drawing in the ‘Poissons 
fossiles’ of the Glammis specimen (which has since been chiselled 
so as to obliterate these organs), are not described in the letter- 
press. Mr. Powrie called attention to these strange pectoral 
organs in the ‘ Geologist,’ 1861, p. 137, and they are well seen 
in some of his beautiful specimens illustrated in the plates of 
this work. They differ from the other fins in presenting no 
trace of fin-rays or of the fine scales which extend upwards 
from the base of the dorsal and caudal. They are simply ellip- 
soid expanses, with some calcareous matter in their structure, 
which has caused them to be preserved, and has rendered 
visible, especially in fig. 1, Pl. X, a kind of reticulate or areolate 
markings, quite peculiar to them. ‘The character of these 
fins differing so much from that of the other fins tends to 
suggest that they may have had other functions than that of 
mere locomotion, and it seems not at all improbable from 
their position that they may have been efficient in causing 
currents of water to pass to the branchial organs covered m 
by the great head-shield (whose outlets are indicated by the 
lateral perforations in the shield of Péeraspis), and have thus 
aided respiration as well as locomotion, as is observed in the fry 
of 'Teleostean fishes at the present day with regard to the 
pectoral fin. ‘These pectoral organs are mentioned here espe- 
cially because it seems hardly doubtful that they are charac- 
teristic of the group, possibly also of the Heterostraci ; it 
is not likely that they were developed in but one single genus 
of the closely allied series classed as Osteostraci. 

A very remarkable specimen of shagreen-like structure has 
been discovered by Mr. Powrie in Forfarshire, in beds which 
have furnished Cephalaspis. As I have not been able to 
assign it to Cephalaspidian fishes, though it may possibly 
be connected with them, I only allude to it here. It consists of 
a surface covered with minute spinous tubercles, the whole having 
the appearance of a fossilized piece of shagreen, and its shape 


4] 


Fie. 16. 


iN 
a 


« 


Restoration of Eucephalaspis 
Lyellii. 


6 


A2 OLD RED SANDSTONE FISHES. 


is more or less that of a Cephalaspid with head and body. The spinous tubercles on some 
species of Cephalaspis (Pl. X, fig. 5) suggested the idea that this might belong to 
the lower surface of one of these fishes, covering in the ventral aspect of the head and the 
ventral series of scales ; being detached, as it were, in this part of the animal, but adhering 
closely to the deeper aponeurotic layers of the exoskeleton in the superior parts, and form- 
ing the characteristic tuberculated surface of the shield and scales. There is, however, no 
proof of its connection with these fishes, and it seems more probable that it belongs to some 
early representative of the Sharks and Rays (representative, perhaps, only in the character of 
its dermal ossifications) than that it is the ventral covering of a Cephalaspid. Its mention 
here may incite investigation and lead to the discovery of specimens showing clearly a 
connection, but as matters at present stand there is not sufficient evidence to justify the 
introduction further of the specimen into these pages. 

Value of Specific and Generic Distinctions —Before proceeding to describe the genera 
and species of Osfeostraci, I would allude to the principles which have determined me in 
forming genera and species. In the first place, as to species. It appears to be better for the 
end of advancing knowledge to signalise a doubtful form by a name, than to pass it over 
as a possible variety or questionable species. The error of associating under one head what 
are really distinct species 1s undeniably as great as that of founding new species on 
specimens which may eventually prove to be imperfect conditions or locally distorted 
states of species already known. ‘The object to be held in view is to attract attention to 
the matter in doubt, and this can be best done by the use of @ name, which has only to 
be rescinded in the event of further discovery rendering it desirable so to do. The 
object of a descriptive work such as this, in which the material is fully placed before the 
reader, is to point out distinctions and peculiarities in the various specimens studied, and 
it seems better in this case to err on the side of division than on that of fusion. 

‘The only use that can be made of generic divisions in the case of fragmentary fossil 
remains is to place those things which are alike in the same group, and those which differ 
in different groups, thus pointing out the gaps in our knowledge of the continuity of 
forms. If our knowledge were more complete there would be greater difficulty in 
indicating groups, a fact. which is true for all groups of organisms. Grouping is 
facilitated by the occurrence of gaps that are not real, but are due to our imperfect 
knowledge. 


List of Species of Ostnostract, arranged in Order of their Occurrence. 


Upper Siturtan. Thyestes verrucosus, Eichwald; Cephalaspis (?) Schrenkii, Pander. 

Downton Sanvstone. Lukeraspis pustuliferus. 

PassaGnBuDs. Didymaspis Grindrodii, Auchenaspis Salteri, Av. Egertoni, Cephalaspis (Hemicyclaspis) 
Murchisoni, C. Lightbodii. : 

Lower Drvontan (of Devonshire and Cornwall). Cephalaspis (?) Carteri. 

Cornstones (West of England). Eucephalaspis Agassizii, Zenaspis Salweyt. 

Lower OLp Rep Sanpstone (Scotland). Hucephalaspis Lyellii, E. Powriei, Z. Paget, EB. asper. 


CEPHALASPID/, 43 


Descriptions of Genera and Species. 
Genus 1.—Crrnatasris, Agassiz. Poissons fossiles, vol. 1, p. 149, 1835. 


Derivation.—xepadn, the head ; aszic, a shield. 
Characters.—Scutum cephalicum simplex semicirculare. 


The genus Cephalaspis, as here indicated, may be conveniently split into three 
subgenera. 


1. Evcrrnaaspis.'—Scutum postice cornibus lateralibus instructum, in medio 
aliquantd productum. 


2. Hemicyciaspis.2—Scutum sine cornibus lateralibus: postice subtruncatum. 


3. Zenaspis..—Scutum Eucephalaspidis scuto simile: scutellum dorsale (vel scutella) 
post scutum cephalicum positum. 


It seems to be a clearer representation of the relations of these fishes to dissociate the 
form destitute of cornua, and that with body-plates as well as a cephalic shield, from the 
other species (which agree in possessing cornua and in other matters very closely), than 
to leave them all in one group, each being equally designated Cephalaspis. It is quite 
possible that further research will render it desirable to consider what I have here called 
sub-genera as genera. Fragmentary remains indicating species belonging to the genus 
Cephalaspis, or only sufficient to pomt to the section Osfeostraci, and not furnishing 
evidence of sub-generic detail, will be spoken of here as Cephalaspis. 

Such remains occur in the Tilestones; in the Ledbury passage-beds we have /emi- 
eyclaspis, and in the Cornstones of Herefordshire Zenaspis. Hucephalaspis is found in 
the Cornstones and the Scotch Lower Old Red. 


1. Evcepnazaspis Lyeiiu, Agassiz. Pl. VIII, fig. 1; Pl. XI, figs. 1 and 2. 


Crernataspis LYELii1, Agassiz. Pois. foss., vol. ii, p. 149, pl. 1, fig. 2, 1835. 


Name.—Named after Sir Charles Lyell, Bart. 
Stratigraphical Position.—Lower Old Red Sandstone (Scotch area). 


| eb; and xegadaoms, Cephalaspis. 
2 jpexueAcos, semicircular ; dozis, a shield. 


3 Zevs, Znrvds, Jove; aovwis, a shield. 


44. OLD RED SANDSTONE FISHES. 


Characters.—I must refer to the figures of the two specimens which are considered 
as belonging to this species for a notion of its characters. It was associated by Agassiz 
with the heads from the English Cornstones, which will here be spoken of as Hucepha- 
laspis Agassizit, since they appear to differ in this poit of structure, viz. that the orbits 
in the Scotch specimens are placed more posteriorly in the shield, and the cornua are 
less produced and less divergent than in the English heads. At the same time it must 
be confessed that there is very close agreement in the outline of the head-shield as exhi- 
bited in the best-preserved specimens. Neither of the Scotch specimens show the surface 
ornament, so that we cannot compare them in this regard. C. Lyellii and C. Agassizii 
were undoubtedly very closely allied, but the evidence does not justify their association 
under one specific name. The figures in the plates give all that is known on the matter. 

General Remarks.—The specimen drawn by Mr. Dinkel in Pl. VIII, fig. 1, is the 
same which he figured more than thirty years ago for Prof. Agassiz. It has since then 
been worked out a little from the matrix, so as to exhibit the marginal series of scales. 
In the process the indications of the remarkable pectoral appendages drawn by Mr. 
Dinkel in Agassiz’s plate, but not referred to in the letter-press, have been destroyed. 
Mr. Dinkel has, however, reintroduced them here, especially on the left side. A false notion 
of the structure of the body of Cephalaspis has been conveyed by the former drawings 
of this specimen and the copies in woodcuts illustrating popular treatises on geology. 
A jointed structure, as though the body were composed of a series of hard rings fitting 
into one another, has been in this way erroneously attributed to the fish (which mistake 
was by no means due to Prof. Agassiz, who knew and described the scales). he fact 
is that the scales of the body are of considerable thickness, and in the celebrated British 
Museum specimen drawn in Pl. VIII, fig. 1, their upper layers are entirely broken away, 
and only a flake of calcareous matter is left coverimg in their impressions, and thus pro- 
ducing the false jomted appearance. The deficiency in the specimen towards the caudal 
extremity marks the position whence the dorsal fin has been broken away. ‘The other 
specimen, drawn in Pl. XI, fig. 1, which is referred to this species, is with, perhaps, the 
exception of Mr. Powrie’s specimen drawn in Pl. X, fig. 1, and referred to C. Powriet, 
the most instructive and beautifully preserved remnant of these curious little old 
Sturgeons which has been found. It belongs to the Museum at Arbroath. The pectoral 
fins are well shown in the specimen, though I have to explain that my over-zealous friend 
Mr. Dinkel has given an appearance of a joint to the right-hand pectoral, which is not 
clearly seen in the fossil itself. The dorsal fin is also well shown, as is the caudal, but 
it is not so perfectly preserved in this as in another specimen (of another species probably) 
drawn on the same plate. The woodcut restoration (fig. 16) may help to elucidate 
the drawing of the fossil. The scales in this Arbroath specimen are unfortunately in the 
same broken condition as in the Glammis (Brit. Mus.) specimen, and no trace of the sur- 
face ornament is to be detected. 

‘The Arbroath specimen has very fortunately been broken in such a way that a piece 


CEPHALASPID. 45 


of the body can be removed, and the view given in fig. 2, Pl. XI, is obtaimed. This 
demonstrates, firstly, a series of ventrally placed scales, which, though rather confused, 
covered in the ventral surface, and appear to have consisted of several rows. The 
great thickness of these scales, and indeed of those of the upper part also, is noticeable. 
The calcified exoskeleton in these fishes bore some resemblance to that of Lophobranchs, 
and, no doubt, formed a rather rigid and tough kind of armature. Secondly, the fractured 
piece of the Arbroath specimen exhibits the body in vertical section, and shows its very 
small calibre and triangular form. 


Fie. 17. 


Ideal vertical transverse section of the body of Eucephalaspis. 


The number of rows in each series of scales is, no doubt, an important matter—pos- 
sibly differing in species. Both specimens of C. Lyellii are unsatisfactory as regards this 
point, since the scales are fractured ; but there appears to have been in front of the dorsal fin 
but one dorsal row of scales on each side the middle line, attached by their lower borders to 
the single series of great flanking or lateral scales: posteriorly to the fin the series became 
broken up into more numerous scales. The number of scales in a row and the relative 
length of head and body are important for specific distinction ; and though it does not 
bear on the question of distinction between C. Lyedlii and C. Agassizii, since the body of 
the latter is unknown, yet a very clear difference in this respect is exhibited between 
C. Lyellii and another Scotch species, C. Powriei. 

The two specimens of C. Zyellit agree, as nearly as their rough condition will 
permit one to estimate, in having about twenty-five scales of the lateral series, placed 
preedorsally, 7. e. which are in front of the anterior origin of the dorsal fin. In the British 
Museum specimen of C. Zyellii the length of the head is 3 inches, of the whole fish 82 
inches, allowing for the tail, which gives a ratio of 1 to 2°4. In the Arbroath specimen 
the head is 2# inches in length, and the whole fish 74 inches, giving a ratio of 1 to 2°6. 
In C. Powriei, on the other hand, the ratio appears to be about 1 to 3:2. Allowance is 
made in these estimates for the parts which are broken. 

The details of structure of the fins shown in the specimen at Arbroath do not allow of 
much being said, since they are not very clear. Small scales appear to extend along the 
fin-rays in both dorsal and caudal fin. It is, however, very important to notice that the 


AG OLD RED SANDSTONE FISHES. 


pectoral fin has quite a different structure, which is well seen in fig. 2, PDX 
There is no evidence of a separation of the calcareous matter imto regular scales, but 
crescent-like areae appear to be marked out by the greater thickness of this matter in 
some parts than in others; neither can any fin-rays be traced in these very curious 
organs. 


2. HucepHataspis Acassizi. Pl. IX, figs. 2, 3, 6. 


CEPHaLAsPis LYELL, Agassiz (in part). Pois. foss., vol. ii, pl. 1 4, fig. 3, 1835. 


Name.—Named after Prof. Louis Agassiz, the author of the ‘ Poissons fossiles.’ 

Stratigraphical Position —The Cornstones of Herefordshire and Worcestershire. 

Characters.—The woodcut, fig. 11, gives what is, I believe, an accurate indication 
of the form of the shield in this species. So many specimens of this are found that 
it is possible to get a more trustworthy idea of the form of the shield when uncrushed, 
and in all its parts, than it is m the case of any other Cephalaspis. Numerous as these 
specimens are, only one has come to hand showing clearly the form of the orbital region 
and the surface-markings. ‘This has been used in drawing these parts in the woodcut, 


Fic. 18 a. Fie. 18 6. 


Ornament of Eucephalaspis Agassizii, magnified 7 diameters. 
‘ 


a, from the orbital region. 6, from the posterior angle. 


fig. 11, and was lent among many others by Dr. Grindrod, of Malvern. It is unnecessary 
to attempt any concise definition of the species in words when the woodcut figures express 
so much more clearly what is meant than a compressed sentence can be made to do. 
The relative position of the orbits in this species and in C. Lye//ii may be seen on com- 
paring the figures in Pl. VIII and Pl. IX. 

General kemarks.—The greatest variation in form is exhibited by the abundant head- 
shields of this species which have been exhumed in the West of England—a variation 
due simply to pressure. Some shields are quite flat, and yet tolerably perfect ; the 
outline in this case becomes very much widened, and the cornua are directed quite away 
from one another, instead of running backwards in nearly parallel directions; others are 
compressed laterally, but many are to be obtained which are regularly convex, and by 
their symmetry lead one to believe that they have not suffered any distortion. No speci- 
meus show the form and proportion of head-shield which is attributed to the following 


CEPHALASPID A. AT 


species (C. Powrie?), and hence we may consider the distinction, so far as it rests on the 
form of head-shield, a good one. Some small specimens, as that drawn in Pl. VIII, 
fig. 5, which probably belong to this species, show an outline rather like some small Scotch 
heads I have seen, one of which is drawn in PI. XII, fig. 1. But the eyes are not so far 
forward in the Scotch specimens as in the English, and we may conclude that they belong 
to C. Lyellii, whilst the English ones belong to C. Agassizii. The Scotch member had ‘a 
longer head’ than the English representative, even among these fishes of the Devonian 
times. 

No bodies of C. Agassizit appear to have been ever found attached to the heads, with 
two exceptions, one of which is obscurely drawn in Pl. VIII, fig. 5, whilst the other shows 
little beyond the fact of a scale-covered body. Both specimens are very small, and, indeed, 
may not belong to this species at all. ‘They are from the Cornstones in the neighbour- 
hood of Ludlow. Small specimens of scales, exhibiting a fine tubercular ornament like 
that of the head-shield of C. Agassiziz, have been found, but give no evidence as to the 
form or relative proportions of the body. It is not a little remarkable that in the Corn- 
stones of Herefordshire and in the slaty beds of Devonshire and Cornwall—which have 
lately (1868) been shown to contain Cephalaspide—the smallest traces of the body, well 
covered in by scales, as we know it to have been, should be so exceedingly rare as com- 
pared with the head-shields, which abound in the West of England, and are packed and 
pressed together in innumerable quantities in Cornwall. The relative size and weight of 
head and body, no doubt, favoured the preservation of the head-shields, which, like the 
Belemnite’s guard, would sink, whilst the body would be broken off from the thus partially 
imbedded head. ‘The large size and freedom from open sutures of the adult head-shields 
would also, no doubt, tend to their preservation, as compared with the smaller and 
therefore more easily scattered scales. A strange instance of the capricious imperfection 
of that very fragmentary document, the ‘ Geological Record,’ is afforded in the absence of 
bodies to English Cephalaspids, and their more frequent occurrence in Scotch specimens. 


3. Evcepnaaspis Powrrer. Pl. X, fig. 1, and Pl. IX, fig. 5. 
Crruataspis Lyeiui, Agassiz (in part). Pois. foss., vol. ii, pl. 1, fig. 1, and pl. 14, fig. 1, 1835. 


Name.—After Mr. Powrie, of Reswallie, Forfarshire, the discoverer of this and other 
Cephalaspids. 

Stratigraphical Position —Lower Old Red Sandstone of Forfarshire. 

Characters.—The form of head-shield distinguishing this species is given in the 
diagram (woodcut, fig. 19). A form of curve which it is difficult to describe in 
words, but which might be expressed mathematically, distinguishes the contour of the 
shield of this form from those of the two preceding. ‘The cornua have a different relative 


48 OLD RED SANDSTONE FISHES. 


proportion, and are more incurved than in Aw. Lyellii or Hu. Agassizii. There appear to 
be about thirty pradorsal scales in’ each series instead of twenty-five, and the scales are 


Ornament of the same from the cornu, 
magnified 7 diameters. 


Diagram outline of the head-shield of Eucephalaspis Powriei. 


narrower in proportion to their length and of less thickness apparently than in Aw. Lyelliv. 
The head measures 1? inch, the whole 53 inches in length, giving a ratio of 1 to 
3°14, instead of 1 to 2°6. The ornament in this species is very finely tubercular. 
In Mr. Powrie’s magnificent specimen, figured in Pl. XI, the ornament can be seen in 
parts with a lens, and may also be traced on many of the scales. 

General Remarks —The grand specimen of fig. 1, Pl. XI, was obtamed by Mr. 
Powrie at Ley’s Mill, near Arbroath. It has been most carefully developed, and shows the 
concave side of the head-shield, and the body twisted so as to bring the dorsal surface to 
the left side, as one looks at the specimen. Other head-shields, detached from the body, 
have come to hand, and one, belonging to Mr. Powrie, is figured in Pl. X, fig. 5. 
A similar specimen is in the collection of the Rev. Hugh Mitchell, of Craig, near 
Montrose. ‘The specimen figured in pl. 1, fig. 1, of the ‘Poissons fossiles, and now 
in Sir Phihp Egerton’s collection, appears to belong to this species. It is most erroneously 
drawn in the figure referred to. The concave surface of half a head is presented with the 
half of the attached body, showing the position of the dorsal and caudal fins; but the 
dorsal series of scales is not preserved as drawn by Mr. Dinkel, the two sets of opposite 
sides being confused in a mass. ‘The lateral series present the same appearance as those 
of Mr. Powrie’s specimen; but I could not count as many as thirty preedorsal. The 
structure of the dorsal and caudal fins is not shown. 


CEPHALASPID/. 49 


4, Evcrpnatasris Pacu. Pl. X, figs. 3 and 4; Pl. XI, fig. 4. 


Name.—After Mr. David Page, of Edinburgh, in accordance with the wish of 
Mr. Powrie, the discoverer of this form. 

Stratigraphical Position Lower Old Red Sandstone of Forfarshire. 

Characters —The form of the head is very similar to that of C. Powriei ; but it is 
difficult to be certain of its exact outline, since all the specimens obtained are much flat- 
tened. Some, however, show this character better than seen in the figured specimens, 
and from these the woodcut outline is drawn (fig. 21). The ornament is notably different 


Fie. 21. 


vuic 
ee 
0 lll 
0° 
SEN 
» vo H\\ 
Ornament of the same, magnified 
7 diameters. 


Diagram outline of head-shield of Eucephalaspis Pagei. 


from that of the several preceding species, for the tubercles are disposed in sets, consisting 
of a central large tubercle, surrounded irregularly, more or less, with smaller tubercles. 
The same character of marking is carried on to the scales. In one specimen (Pl. X, 
fig. 4) I could count about twenty praedorsal scales in a series; but the specimens of 
this species, though appearing very perfect at first sight, are so highly carbonized and 
crushed that it is impossible to ascertain the true arrangement of the scales. Some seem 
to indicate quite a different disposition of the series to that which I have described in 
Lu. Lyellii and Eu. Powriei. The dorsal and caudal fins are well seen in some specimens, 
but have not furnished any special characters; in no specimen of this species have the 
pectoral appendages yet been observed. The size of the majority of specimens is small. 
They may be young individuals; but their great number and the rare occurrence of large 
specimens with them, which seem to belong to another species, leads to the supposition 
that this is a Cephalaspis of small size when adult. At the same time it may prove to be 
a younger form of the succeeding species. 

General Remarks.—Mx. Powrie discovered this form in some abundance in a remarkable 
bed of fine, dark, laminated shale and sandstone, which furnished also several Acanthodians, 


7 


50 OLD RED SANDSTONE FISHES. 


and a few specimens of a larger Cephalaspis. 'The specimens are much carbonized and 
quite black, showing, however, the surface-markings and the scales in parts with a 
distinctness which varies in specimens and is liable to diminish by exposure after the 
exhumation. 

There is a fact of interest with regard to the position assumed by the specimens in 
the rock. They are all very much flattened, but many have apparently also burst 
along the ventral line, so that the fossil presents the two rows of large flank scales spread 
out on each side of the dorsal series. This indicates great toughness in the tegumentary 
skeleton, and implies that the scales were held together by a very strong matrix of fibrous 
tissue, which was less firm along the ventral surface than in other parts. It has been 
suggested, as before noted (page 42), that the ventral part, if not the whole of the Cepha- 
laspids, was covered and firmly connected through the aponeurotic scales with a skin 
containing small shagreen-like ossicles in parts. The continuity presented by the scales 
when crushed, as in the specimens of C. Page, favours this notion. In the Heferostraci 
it is clear, fromthe gland-pits, that there was a considerable development of the parts 
superficial to the calcified layers of the integument. 


5. HucepHaLaspis asper. PI. X, fig. 5. 


Name.—F¥ rom the pointed spinets which exist on the scales and rm of the cephalic 
shield. . 

Characters and General Remarks.—The specimen figured, which is from a nodule in the 
Lower Old Red Sandstone of Perthshire, and another from the bed in Reswalli, Forfar, 
which furnished Mr. Powrie with C. Pagei and others of his valuable discoveries, present 
conoid or dentate spinets (fig. 23) situated on the marginal rim of the head-shield, being 


Fie. 23. 


Spinets from the margin of the shield of Hucephalaspis asper. 


modified tubercles, and also on the scales, as seen in the figure. How far this structure 
is sufficient to characterise a species may very well be held to be doubtful, since such 


CEPHALASPID/. 


or 


denticular processes are very apt to get lost and obscured in these fossils, as, for example, 
those on the inner edge of the cornua of the common Lucephalaspis Agassizit. Perhaps 
this species may be merely a larger growth of C. Pagei, the spinets being preserved in 
the large specimens, but undeveloped or obscured in the smaller. Nevertheless, it is 
right to signalise this characteristic structure by a name. Further search in regard to 
this and other similarly dubious matters indicated in these pages is required, and the 
matter is best pressed on the attention of collectors and others by provisionally naming 
the supposed species. The horror with which the making of a species on small data is 
regarded by some naturalists appears to be a superstition which may lead to evil results, 
for the opposite proceeding of passing over all differences and indications of distinction 
among forms, unless of the most certain and obvious character, is far more injurious to 
the progress of knowledge. The use of a name, whether of genus, species, family, or what 
not, is merely to briefly draw the attention to a supposed speciality of structure separating 
the individuals, to which the name is applied, from others. There is nothing sacred in a 
name, and, if the progress of discovery render it desirable, old names can be changed or 
suppressed. As long as the data on which a species is named are fairly and fully stated, it 
can be no encumbrance to science, even though it should eventually prove to be not 
distinct from another form. 


6. Hemicyczaspis Murcnisoni. Pl. VIII, fig. 6; Pl. IX, fig. 1; Pl. XII, figs. 3 and 4. 


Crpuatasris Murcuisont, Egerton. Quart. Journ. Geol. Soc., vol. xiii, p. 284, pl. ix, 1857. 
— ORNATUS, Id. Ibid. 


Name.—After Sir Roderick Impey Murchison, Bart. 

Stratigraphical Position.—Tilestones and Auchenaspis-grits. 

Characters.—The woodcut (fig. 24) presents the characteristic outline of the sub-genus 
and the ornament which characterises the species. The tubercles ornamenting the surface 
are arranged in distinct groups, marked out as polygonal arez, in this species, more 
distinctly than in C. Pagei, some distance of intertubercular surface intervening between 
the contiguous groups. 

General Remarks.—Sir Philip Egerton (loc. cit.) described C@. Murchisoni and 
C. ornatus originally as two distinct species, suggesting, however, that they might prove 
to be identical should specimens of C. J/urchisoni showing the surface be discovered. 
I have received such a specimen from Dr. Grindrod, and do not doubt that the two specimens 
figured from the Tilestones near Ludlow and the two from the Ledbury Grits are the same 
species. A question was also raised by Sir Philip as to whether C. Murchisoni might 
not belong to his genus Auchenaspis, being thus the anterior portion of a shield divided 
into head- and neck-plates. The presence of a groove running parallel to the edge of 
the semicircular shield seems to me to oppose this notion, for a similar groove runs 


52 OLD RED SANDSTONE FISHES. 


along the posterior margin of the shield of Hucephalaspis, and also along the posterior 
margin of the posterior or neck-plate of Auchenaspis, whilst there is no trace of such a 


Fic. 24. 


Ornament of the same, magnified 
7 diameters. 


Diagram outline of the head-shield of Hemicyclaspis 
Murchisoni. 


grooving along the posterior margin of the anterior piece of the shield in either Auchenaspis 
or Didymasprs. 

I was told by Dr. Anton Fritsch, of Prague, that he had obtained when in England 
a specimen of this species showing parts of the body, but I have no details as to its 
state of preservation, and am not certain that this was the species obtained. Fragments 
of this species are not uncommon in the micaceous sandstone at Ledbury, which has 
furnished a large species of Auchenaspis. ‘There are only two specimens in collections, 
which come from the finer argillaceous Tilestones, where it occurs with another 
Cephalaspis and with the small Auchenaspis described by Sir P. Egerton as Au. Salfere. 
As might be expected, the ornament is rarely preserved in the specimens from the ‘ Grits,’ 
whilst the two from the Ludlow 'Tilestones show it admirably. 


7. Zenaspis Satweryr. Pl. XII, figs. 2, 5, 6; Pl. VIII, figs. 2, 3, 4. 


CrpHALaspis SaLweyI, Lgerton. Quart. Journ. Geol. Soc., vol. xiil, p. 283, pl. x, 1857. 
— ASTEROLEPIS, Harley. Ibid., p. 503, 1859. 
— — Lankester. Brit. Assoc. Reports, 1864. 


Name.—Named in honour of Mr. Humphrey Salwey, of Ludlow. 

Stratigraphical Position.—Cornstones, probably in a special horizon, with Scaphaspis 
rectus and Pteraspis Crouchii. 

Characters (see the woodcut, fig. 26).—The large size of the tubercles, which are 


Fic. 26. 


= ——— 


| ‘ 


iil t 


ie ; i |! al | 

ilar 4 Hr i Hi : | 

¥ Ni lal ly it Mt | | 
t MANTA 


: 


H 
shit 


: 


Nh 


Fig. 26.—Diagram outline of the head-shield of Zenaspis Salweyi, natural size. 
», 27.—Dorsal scute of the same. 
» 28a.—Ornament from near the front of the shield, magnified 7 diameters. 
», 285.—Ornament from near the middle of the shield, magnified 7 diameters. 


54 OLD RED SANDSTONE FISHES. 


largest in the large shields, as may be seen by comparing Pl. XIU, figs. 2 and 6, and 
Pl. VII, fig. 4, and their less frequency, distinguish the ornament of this species from 
that of Hucephalaspis Agassizii, specimens of nearly equal size being compared. The 
thickness of the substance of the shield is relatively great, and it has a bright polished 
surface. The cornua are long, subcylindrical, and solid, differing in this, as does the 
shield-substance in thickness, from Lucephalaspis Agassizii ; the tubercles on the surface 
are oblong and closely packed. ‘The ends of these cornua are not unfrequently found 
detached, having survived the destruction of the rest of the shield, from which they appear 
to have been readily broken. ‘The postorbital valley is shorter and broader relatively in 
this species than in other Cephalaspids. Sir Philip Egerton attached importance to the 
great breadth between the eyes, but the size of the individual and variations in pressure 
are liable to affect this character. A greater interspace between the orbits is, no doubt, 
present than between those of Hw. Agassizii, if orbits of equal size or nearly equal size 
be compared. But the real pomt is that the orbits are much smaller in this species, 
relatively to the size of the whole shield, than in Hucephalaspis. 

General Remarks.—After some hesitation I have decided to associate C. Salweyi and 
C. asterolepis as one species, not being able, on careful examination, to find any character 
which should separate the large specimen described by Dr. Harley from Sir Philip Egerton’s 
original C. Salwey:. A magnificent specimen has been kindly lent to me by Mr. Lee, of 
Caerleon, from which the outline woodcut has been in great measure drawn; two cornua, 
belonging to Dr. MacCullough, and found in the same quarry at Abergavenny, furnishing 
the evidence of these parts which are wanting im Mr. Lee’s specimen. This specimen 
came to hand after my plates had been completed; it has been photographed and pub- 
lished in a recent volume of the ‘ Proceedings of the Woolhope Nat. Field Club.’ Another 
specimen, which is better than either of those drawn in my plate, has been recently obtained 
for the British Museum. It is very similar to Mr. Salwey’s specimen drawn in PI. XII, 
fig. 6, but it has the long cylindrical cornua attached to the angles of the shield, which 
are wanting in this specimen. The ornament is best shown on these two shields, the 
irregular ‘splashed’ character of the tubercular markings being well seen. In parts 
they assume a much more regular character, each tubercle being hemispherical, and all 
nearly of equal size (PI. VIII, fig. 4). Bits showing this character of marking from shields 
of twice the size of that drawn in Pl. XII, fig. 6, and therefore with much larger tubercles, 
are not uncommon, and have been regarded distinctively as C. asterolepis, but I cannot 
believe that the difference is anything but one of size. None of the large specimens 
usually referred to C. asterolepis show the tubercles of the surface of the shield in such a 
way that they can be compared with those of the smaller specimen drawn in fig. 6, 
Pl. XII. On such specimens a few of the larger tubercles are left here and there 
adhering. I do not doubt that if we have had the ornament properly shown, we should 
find oval, circular, and irregular tubercles of various sizes on parts of the shield, as in 
typical C. Salweyi. I say typical C. Sa/wey2, for though Pl. XII, fig. 5, is the figured type 


CEPHALASPID/. 5: 


~'t 


specimen, and shows very little of the surface, fig. 6, Pl. XII, which shows so much more, 
may be taken as more fairly representing it, and was so regarded by Dr. Harley. 

The character of the tubercles varies on different parts of the same shield in this 
species very greatly; in some parts there are fewer tubercles than others, and they are 
round and of equal size; in other parts they are crowded so as to leave no intertubercular 
surface, and are small and large, oval and round, and irregular. ‘This latter arrangement 
is seen near the margin more particularly, which, as we saw in Scaphaspis (P1. I, fig. 4), 
is subject to crowding and irregularity. 

The remarkable scutes drawn in Pl. VIII, figs. 2 and 3, present a very large form of 
ornament, which is regular in size and hemispherical, as seen in Pl. VIII, fig. 4. They 
ocewred in association with the specimen drawn in fig. 2, Pl. XII, and, no doubt, belong 
to the same species of fish, the character of their substance and ornament leaving little 
room for doubt on this pomt. It is the occurrence of these very remarkable scutes which 
has induced me to form the subgenus Zenaspis, since they indicate an arrangement of the 
armature of the body quite different from that of C. Zye/i7. The scutes are symmetrical in 
form, and were therefore probably placed in the median line, probably on the dorsal 
surface. In Pl. XIII, figs. 17 and 18, somewhat similar and smaller scutes are drawn. 
These additional scutes, which thus characterise the subgenus Zenaspis, apparently belonged 
to bigger individuals than that drawn in Pl. XII, fig. 6, for which their ornament is too 
large. All fragments of such additional plates as these should be carefully looked after 
by collectors, since the specimens figured, and others less definite which have been sub- 
mitted to me, indicate a considerable development of such plates, in the place of the 
scales of Hucephalaspis, and may possibly require a generic in place of a subgeneric 
recognition. ‘The flank-scales of individuals as large as Mr. Lee’s specimen must have 
been of considerable strength and size, if they retained merely the relative proportions 
seen in ELucephalaspis ; but if, as is not impossible, they united in parts to form still 
larger plates, we may expect assuredly that they should be detected in the Corn- 
stones. 

‘The ‘rims’ of the cephalic shield of this species are sometimes found alone, being 
solid and of a shape to resist destruction after the breaking away of the expanded dome 


which carries the orbits, &c., as may be observed in other species (see Pl. IX, fig. 4, 
and Pl. XIII, fig. 19). 


8. Cernaxaspis Ligutsopu. Pl. XIII, fig. 19. 


Name.—After Mr. Lightbody, of Ludlow. 

Stratigraphical Position.—Tilestones, near Ludlow ; from the same fine argillaceous 
bed which furnishes Hemicyclaspis Murchisoni and Auchenaspis Salter. 

Characters.—The peculiar form of ornament definitely characterises this species. On 


56 OLD RED SANDSTONE FISHES. 


the margin or rim of the shield are very densely set conical or pyramidal tubercles of 
the size of a pin’s head and larger. The apices of some of these tubercles are curved so as 
to form little hooks. The upper surface of the shield, to judge from a piece seen at the 
lower portion of the figure, which may, however, belong to another plate, was ornamented 
with equally close-set tubercles, so closely set and of such irregular angular form that 
their impress in the matrix has the appearance of a network. But what is chiefly 
remarkable is the presence of a set of grooves, independent of the tubercles, winding 
about amongst them in such a way as to mark out irregular areas of a quarter of an inch 
or less across, giving an appearance which is seen on some Reptilian scutes, and which is 
but feebly rendered in Mr. Dinkel’s drawing in Pl. XIII. 

General Remarks——This species was pointed out to me some time ago by my 
friend Mr. Lightbody, who has a few fragments besides the rim here figured. I have 
great pleasure in naming this very interesting form after one who has done so much in 
the investigation of these matters, and has given me so much assistance in my task. 

It is impossible to say what subgenus this Cephalaspis would fit into, or whether it 
would require a new one. Its large size and very definite peculiar ornamentation render 
it probable that we shall soon hear of it again. Extending inwards from the rim are seen 
in the specimen the ‘lateral floors’ of the shield. The tubercles on their surface are 
much smaller than those on the rim itself. 


Genus 2.—AvucuENaspis, Hyerton. Quart. Journ. Geol. Soc., vol. xin, 
p. 286, 1857. 


Derivation.—-avy nv, the neck ; aczic, a shield. 

Characters.—Scutum in duas partes divisum, anteriorem semicircularem in qua sunt 
oculi positi, cornibus lateralibus magnis instructam majorem, posteriorem oblongam 
minorem. 

a. AvcugeNnasprs.—Cornibus lateralibus modestis. 

6. EuKraspis.\—Cornibus lateralibus longissimis, cellulis marginalibus magnis. 


1, Aucnenaspis Saururi, Lyerton. PI. XIII, figs. 7, 8. 
AUCHENASPIS SALTERI, Hyerton. Quart. Journ. Geol. Soc., vol. xiii, pl. ix, 1857. 


Name.—After the late Mr. J. W. Salter, of the Geological Survey. 


1 ed, intensitive; xépas, cornu; daozis, shield. 


CEPHALASPID®. 5 


~ 


Stratigraphical Position—The Tilestones (the fine-textured deposit occurring near 
Ludlow). 


Diagram outlines of head-shield of 4uchenaspis Salteri and Au. Egertoni. 


Characters.—This species is distinguished by its small size and by the small diver- 
gence of its lateral cornua. The surface-ornament is not known. 

General Remarks.—I have had but few specimens of this species to examine. | 
think that the uniformly small size of the specimens from Ludlow, whence Sir Philip 
Hgerton’s type was obtained, as also the difference in the apparent curvature of the 
anterior portion of the shield and its less divergent cornua, justify the separation of 
this form from that which occurs in some abundance in the micaceous grits of the Led- 
bury Passage-beds. 


2. AucuEnasPis Eerrtoni. Pl. XIII, figs. 3, 4, 5. 


Name,—After Sir Philip de Malpas Grey Kgerton, Bart., the author of the genus. 

Stratigraphical Position.—The Ledbury Passage-beds, or ‘ Auchenaspis-grits.’ 

Characters——Compare the woodcut outlines of the two species, figs. 29 and 30. ‘The 
three specimens of this species drawn in Pl. XIII are variously imbedded in the matrix, 
one showing more of the cornua than another. The specimen which is intended to be repre- 
sented in fig. 3 is the most characteristic, but has not been well drawn on the plate. 
The cornua are largely developed and diverge from the posterior or neck-plate ; the 
shield has more than twice the vertical diameter of the preceding species. The surface 
shows in some specimens tubercular ornament of relatively very large size. 

General Remarks.—This species used to be almost abundant at Ledbury in the grey 
micaceous grits which were exposed in the railway-cutting on that side of the Malvern 
and Hereford Railway-tunnel. Mr. Brookes, well known to many as a clever collector 
and a working-man of great intelligence, sometimes obtained four or five of these heads 
on one hand-specimen of the stone; they occurred associated with fragmentary remains 
of Hemicyclaspis Murchisoni. 


58 OLD RED SANDSTONE FISHES. 


3. EUKERASPIS PUSTULIFERUS, dgassiz. PI. XIII, figs, 9—14. 


ScHLERODUS PUSTULIFERUS, Agass. Sil. Syst., pl. iv, figs. 27, &c.; Siluria, pl. xxxv, 
figs. 9—12. 
PLECTRODUS MIRABILIS, Agass. Sil. Syst., pl. iv, figs. 14, &c. ; Siluria, pl. xxxv, figs. 
3—8. 


Name.—Pustuliferus, bearing pustules, on account of its fine tubercular ornament. 

Stratigraphical Position —The Downton Sandstone of Ludford Lane, near Ludlow. 

Characters.—Posterior portion of the shield not known; cornua twice as long as the 
central or vertical diameter of the anterior portion of which they are processes ; a series of 
six cavities (cellule marginales), woodcut, fig. 31, c. m., on either side excavated in the 


Fie. 31. 


Ornament of the same, magnified 
7 diameters. 


Diagram outline of head-shield of Eukeraspis pustuliferus. 


a. 0. p. orbital prominence ; ¢. m. marginal cavities; p. 0. v. post- 
orbital valley; 7. 0. p. interorbital prominence. 


substance of the shield; a similar pair of cavities (appearmg like orbits m fig. 13, 
Pl. XII) are placed posteriorly to the true orbits and on either side of a very marked 
prominence (i. 0. p. in the woodcut), like the interorbital promimence of Cephalaspis (seen 
in Pl. XIII, fig. 10). The true orbits are deeply sunk, and are seen in figs. 13 and 14; 
the right orbit, which is preserved in the specimen drawn in fig. 13, and lies anteriorly 
to and somewhat to the left of the right hand of the two oblong cavities, there drawn, iS 
not clearly seen. The antorbital prominences (a. 0. p. in the woodcut) are very marked 
and prominent in this form; as to the antorbital fosse I am uncertain. The whole 
surface is covered by a very fine densely set series of circular tubercles of very small size 
and great regularity, extending on to every prominence and furrow of the shield. The 


CEPHALASPID i. 59 


cellular cavities of the margin and of the central part of the shield were only indicated 
superficially by slight undulations of the tuberculated test. The owfer margin of the 
shield, especially of the cornua, contrary to what is seen in Cephalaspis, where the zaner 
border of the cornua is denticulate, is furnished with irregularly tooth-lke processes of 
various length. 

General Remarks.—TVhe remains of this beautiful and well-marked little Cephalaspid 
were described in the ‘ Silurian System,’ and the earlier editions of ‘ Siluria,’ from detached 
specimens of the cornua, as toothed jaws of small ganoid fishes, Pt. (Sclerodus) pustuliferus 
and mirabilis. In the last edition Sir Roderick remarks that Dr. John Harley 
(formerly of Ludlow) had suggested the true nature of these supposed fish-jaws. (See 
description of Pl. XXXV in ‘Siluria,’ 1867.) At page 241 of the same edition it is 
stated that Mr. Salter was inclined to regard these cornua as the jaws of Cephalaspis or 
Pteraspis. ‘The most important feature about Hukeraspis appears to me to be the existence 
of the remarkable cavities between the two lamine of the shield-test, but unfortunately it 
is exceedingly difficult to ascertain definitely their structure and relations, on account of 
the rarity of specimens of the fish, and especially of specimens well preserved. Were it 
possible to describe these structures with accuracy, their peculiarity, though paralleled 
by feebly developed marginal cavities in Cephalaspis, would, I think, justify the erection 
of Hukeraspis into a totally distinct genus. The small vertical breadth of the shield, and 
the outline of its posterior border, as seen in fig. 13, Pl. XIII, renders it improbable that 
we have in these beautiful little specimens a complete cephalic shield comparable to that 
of Lucephalaspis, and leads one to suppose that an additional plate such as is presented 
by Auchenaspis was attached along this border. ‘This is a question which inquiry with 
the hammer may soon decide; provisionally, Hukeraspis may remain as a subgenus of 
Auchenaspis. 

The only specimens yet obtained of this form are from the Downton Sandstone near 
Ludlow. 


Genus 3.—Dipymasptis, Lankester. 


DipymaspPis, Lankester. Geol. Mag., vol. iv, p. 152, 1867. 


Derivation.—6vpoc, twin; aozic, a shield. 
Characters.—Scutum in duas partes subzequales divisum, anteriorem semicircularem 
in qua oculi sunt positi, sine cornibus divergentibus, posteriorem oblongam. 


1. Dipymasris Grinprop1, Zankester. Pl. XIII, figs. 1, 2. 


Dipymaspis Grinvropi, Lankester. Geol. Mag., vol. iv, p. 152, pl. viii, 1867. 


Name.—In honour of Dr. Grindrod, of Malvern. 


60 OLD RED SANDSTONE FISHES. 


Stratigraphical Position —In Red Sandstone, immediately above the ‘ Auchenaspis 
Grits’ of Ledbury. 
Characters and General Remarks.—See woodcut, fig. 33. The fossil is in two 


Fie. 33. 


Diagram outline of the head-shield of 
Didymaspis Grindrodi. 


pieces, one showing the convex cast (Pl. XIII, fig. 2), the other concave surface 
(Pl. XIII, fig. 1), of the cephalic shield. This shield is of an oval form, about an inch in 
length and a littie less in breadth, Anteriorly it is divided by a well-marked junction- 
line into two portions—an anterior semicircular piece and a posterior larger and somewhat 
square piece. ‘The anterior portion exhibits two distinct oval orbits, placed close together 
at its centre. Radiating channels mark the inner surface of this portion of the scute, 
recalling the similar ‘channeling’ in Cephalaspis (also seen in Auchenaspis, being the 
structure which gave rise to the supposition of a fibrous bony skull in Agassiz’s descrip- 
tion of Cephalaspis). The line of junction between the two plates describes a double 
curve; the two produced angles of the semicircular plate embracing the posterior plate, 
but not diverging from it, whilst in the median line the anterior piece is produced into 
the posterior to a small extent: in this way the double curvature of the margins is 
effected. Posteriorly the posterior plate becomes contracted, and its margins tend towards 
describing a dome-like outline when it is abruptly truncated, and the truncated margin 
thickened and inflected. A thickened ridge passes anteriorly along the median line. No 
radiating channels mark its inner surface. Flake-like fragments from the inner substance 
of the scute, soaked in Canada balsam and examined beneath the microscope, show large 
bone-lacunee very densely packed, arranged at right angles in the different lamelle of the 
bony material, so as to produce the appearance of cross-hatching. 

The small specimen figured, which is from Dr. Grindrod’s great collection, is the 
only relic of this form which we at present possess. 


A. In Pl. XIIf one or two fragments of scales are figured which may possibly be 
connected with Cephalaspids, or with the bearers of those ichthyodorulites which not 
unfrequently occur in the Cornstones of Herefordshire and subjacent beds, and have been 
designated Onchus. 

I would particularly draw attention to the beautiful fragments which I have termed 


- 


CEPHALASPID®. 6] 


Kallostrakon podura, on account of the resemblance of their ornament to the well-known 
microscopic markings of the scales of the insect Podwra. The microscopic structure of a 
portion of the scale drawn in fig. 21, Pl. XIII, is given in fig. 6, Pl. XIV, and is well 
worth attention, on account of its very peculiar nature. 

The numerous indeterminable fragments of fish remains, together with the Oxchus 
species, which occur in Siluro-Devonian beds require careful collation and study; they 
appear to indicate the presence of other growps of fish in company with the Cephalaspide 
described in these pages. 

B. Since the plates of this Monograph were completed, and the first part issued, 
a great discovery has been made, or rather a rediscovery relating to its subject-matter. 
Cephalaspide have been found in Devonshire and Cornwall, in beds assigned to the Lower 
Devonian series. Mr. Peach long since (in 1847) described the remains in question as fish, 
and latterly, from specimens belonging to Mr. Pengelly, the Cephalaspidian nature of 
the remains, which had in the interim been considered as Sponges, has been decided by 
the Rev. W. 8S. Symonds, by Prof. Huxley, Mr. Salter, and myself. Innumerable remains 
of the shields of a Scaphaspis, more than a foot long, fragments of Cephalaspis, numerous 
spines, and other indeterminable fragments, besides the splendid scale and spine 
assigned by Mr. Pengelly respectively to Phyllolepis and Ctenacanthus, are the fish 
remains of these beds. Mr. Pengelly, Mr. Peach, and the Royal Geological Society of 
Cornwall, have kindly lent me their specimens for further examination. 

C. Before leaving the subject, and now that the various forms and remains of the 
Cephalaspidian Fishes known have been described and figured to the reader, I should wish 
to revert to the question of their zoological position discussed in the first pages of this 
Monograph. Since they are the earliest remains of Fish presented to us by the geological 
record, we should naturally expect them to exhibit a difference of organization as com- 
pared with living fishes, leading us downwards to some invertebrate group. But it 
cannot be too strongly asserted that these Fishes are, as far as can be seen, by no means of 
a low type, and that, so far from showing any real affinities with lower types of animal 
life, such as the fancied relationship with Crustacea or Cephalopoda, they disclose, upon 
careful examination, points of structure, such as the double olfactory cavities and the pec- 
toral appendages, which place them very far above some living Vertebrata (Lampreys) 
classed as Fishes. At the same time there is nothing in the remains known to us which 
will indicate even approximately their affinities to any one of the large groups recognised 
in the classification of Amphirhine Fishes. So great is the variation in the distribution of 
calcareous matter in the exo- and endo-skeletons of Fishes that in these, as in so many other 
fossil members of the class, no evidence is given by which we can judge of the condition of 
the important respiratory, circulatory, and digestive organs which accompanied the skeleton. 
The calcified skeleton in Cephalaspide, apparently, was entirely confined to the aponeu- 
rotic or ‘splint-system’ of Mr. Parker; and whilst the endo-skeleton was cartilaginous, it 
may have been of high or low development. Apparently, too, like the Lophobranchs and 

9 


62 OLD RED SANDSTONE FISHES. 


Plectognaths, the Cephalaspide may have had the aponeurotic part of their exo-skeleton 
separated wholly or in parts from the upper layers of the ewtis vera, which may or may not 
have existed in a calcified condition. The series of scales or bones along the body of 
Cephalaspis—so strongly recalling the cmetures of Callichthys (which has a complete endo- 
skeleton)—are, probably, morphologically of the same nature as those structures, but ante- 
riorly I have not been able to detect any modification of the flanking ‘ scales’ in Cephalaspis 
in the form of clavicular bones. The very peculiar pectoral fins of Cephalaspis are unlike 
those of Ganoids or Teleosteans ; their broad membranous character is most: nearly repre- 
sented in Sharks; but just as the character of their skeleton does not render it impossible 
that they were Elasmobranchiate, nor point distinctively to either Ganoids, Siluroids, 
Lophobranchs, or Plectognaths as their modern congeners, so do the fins fail to give any 
decisive indication, for have we not classed among Ganoids /o/optychius, Polypterus, 
Amia, and Sturgeon ? 

It is best, then, to let the group Cephalaspide stand alone,’ since they present a 
combination of characters which is not inconsistent with any one of the recent types of 
internal organization presented by Fish, whilst agreeing with no living type of skeleton 
entirely. When their position in time is considered, it becomes very probable that they 
possessed an internal skeleton and viscera of more gezera/ character than any of the higher 
living Fishes now present to us, which would justify their distinct position. The solid 
character of the aponeurotic skeleton in the body of Cephalaspis and its vertical segmen- 
tation, together with the non-calcification or the separation of the more superficial parts of 
the integument and of the cartilaginous endo-skeleton, may be considered as indicating a 
generalised condition of structure, in which the splint-skeleton was so far dominant as to 
take on that self-repetition which is, in other cases, known to us as vertebral segmen- 
tation. 

The Hurrrosrract are associated at present with the Ostrosrract because they are 
found in the same beds, because they have, like Cephadaspis, a large head-shield, and because 
there is nothing else with which to associate them. There is at present no evidence that 
the body and fins of P¢eraspis and its allies were like those of Cephalaspis, and the 
shields are not so closely similar in plan, much less in histological structure, as to warrant 
any inference of similarity in other parts. specially it is to be noted that, unless the 
apertures considered as orbits in Pteraspis represent the antorbital fosse or olfactory 
organ of Cephalaspis, there is a total absence of such fossze in the Pteraspidian shield, 
whilst the alternative gives the absence of orbits, either of whichis an important difference. 

The discovery of the bodies of Heterostraci will be a most welcome addition to 
science, and for this we wait. 

1 Professor Huxley’s last suggestion on this matter (‘ Academy,’ No. 2, p. 42) is that the Cephalaspide 
possibly connect the Monorhina (Lampreys) with the Sturgeons among the Amphirhine; but why should 
their exo-skeleton have so much importance attached to it as to indicate a leaning towards Ganoids rather 


than Sharks? Professor Huxley even speaks of the Cephalaspide as Ganoids simply, which seems to be 
taking too much on ‘faith,’ as regards their unknown viscera. 


Fie. 


— 


to 


cs 


6. 


PLATE VI. 


Pteraspis rostratus, Ag. A very fine specimen; with the surface markings (strie 
and pits) well shown. Cradley. Dr. Grindrod. 


A similar specimen, showing more of the posterior portion of the shield. Cradley 
Dr. Grindrod. 


A natural cast of the surface of part of the shield of P¢. rostratus, showing the 
disposition of the markings round the orbit and on the cornu. Notice in this 
and in fig. 1 the aperture (a) in the cornu, giving the appearance of a detached 
projecting angle to part of the cornu. Author’s cabinet. 

A disc of Pt. Crouchii, showmg clearly the characteristic form and the deep 
incision posteriorly for the spine. 

Cyathaspis Symondsi. Cornstones, Herefordshire. In the Museum of the Geolo- 
gical Survey (Jermyn Street). 

A specimen of P¢. rostratus, showing the superficial markings of the posterior 
portion of the shield. In the collection of Mr. James R. Gregory. 


7,8. Rostra of Péeraspis Crouchii. From the neighbourhood of Ludlow. 


9. 


A fragment of P¢. rostratus, consisting of the ‘ orbital portion.’ 


~ 


¥ Frias LCi 


Fic. 


N 


10, 


18. 


PLATE VII. 


A portion of the test of Scaphaspis Lloydii ; considerably enlarged, to show the form of the surface 
ridges and the coarser structure of the lower layers of the shield. The piece is from near the 
centre of the shield. Observe the flatness of the ridges. 

A similar piece from the shield of Scaphaspis rectus, drawn to the same scale (the lower nacreous and 
cancellated layers not being given). The ridges are of smaller size. 

A similar piece from the shield of Pteraspis rostratus. The ridges are much narrower, and are rounded. 
Drawn to the same scale. 

A similar piece from the shield of Pteraspis Crouchii. The ridges are nearly as broad as in Se. Lloydii, 
and quite as flat. Drawn tothe same scale. 

A portion of the shield of Pé. rostratus, taken nearer to the margin, and including three of the ‘pits’ 
or mucous-gland-sites, Same scale. 


. A fragment, showing the cancellated layer or polygonal cavities. Same scale. 


A portion of the shield of Se. Lloydii, from very close to its margin. The ridges become exceedingly 
narrow, but maintain their relative proportion as compared with ridges from similar parts in P¢. 
rostratus. Compare this with fig. 5, and fig. 1 with fig. 3. 

The microscopic structure of the shield of P¢. rostratus. A section has been taken traversing the 
ridges, four of which are seen in section. The dark matter above these is matrix. After Professor 


Huxley. 
Restoration of the cephalic shield of Pteraspis Crouchii, Salter. 
» 3 Pteraspis rostratus, Agassiz. 


These drawings and those in Pl. VI should be examined with a magnifying glass. The 
striations have been very carefully rendered by Mr. Fielding, but it was not possible for him to 
draw them actually as closely packed as they are in reality. 

The microscopic structure of Pteraspis rostratus. A section has in this case been taken parallel 
to the ridges. The vascular tufts and their connections with the polygonal cavities are seen. 
Original. 

Under surface of the rostrum of Pteraspis Crouchii. Restored. 

5 36 Pteraspis rostratus. Restored. 

Ideal longitudinal section through the shield of Pteraspis rostratus. 


e Ss Pteraspis Crouchii. 
» % Scaphaspis Lloydit. 
Ideal transverse section passing just in front of the insertion of the spine in Pteraspis Crouch. 
‘i Ba more posteriorly. 
* more anteriorly. 


Transverse section of Scaphaspis Lloydit. 


natiun 


F Waller nm lé 


E Belding Det 


6. 


PLATE VIII. 


Eucephalaspis (Cephalaspis) Lyellii ; the specimen figured by Agassiz, ‘ Poissons 
fossiles,’ pl. i, fig. 1, 1835, now in the British Museum. The artist has 
introduced the indications of pectoral appendages which were broken away in 
exposing the series of marginal scales. Glammis, Perthshire. Compare the 
direction of the scales in this and in Pl. XI, fig. 1. 

A specimen of a scute, not forming part of the cephalic shield, and proved by its 
ornamentation and occurrence to belong to Zenaspis (Cephalaspis) Salweyi 
(= asterolepis). | Cornstones, Herefordshire. In the Museum of the 
Geological Survey. 

Another, similar scute. Cornstones, Herefordshire. In the Museum of the 
Geological Survey. 

A somewhat similar fragment of doubtful character, but showing well the orna- 
mentation as an intaglio. Cornstones, Herefordshire. 

A small Cephalaspis from Herefordshire, showing the body in a rather fragmentary 
condition, which the artist has failed to render. In the collection of Mr. 
Morton, of Liverpool. 

Hemicyclaspis (Cephalaspis) Murchisoni ; a fine specimen of the head-shield from 
the Passage-beds of Ledbury, Herefordshire. In the collection of Dr. 
Grindrod. 


Ww 


PLATE IX. 


Head-shield of Memicyclaspis (Cephalaspis) Murchisoni, showing well its posterior 
margin and the angle of the shield on the left side. Passage-beds, Led- 
bury. Dr. Grindrod. (See also Pl. I, fig. 6, and Pl. XII, figs. 3 and 4.) 


Lucephalaspis (Cephalaspis) Agassizit; a head-shield from the Cornstones of 
Herefordshire, which has not been crushed. The part of the cornu which is seen 
is not its upper, but its lower wall. Mr. Lightbody, of Ludlow. 


1. Agassiz ; a head-shield, exhibiting a more complete outline than fig. 2, but a 


little crushed, whence the more ovoid and smooth contour which it presents. 
Cornstones, Cradley. Dr. Grindrod. 


Th 


A rim of a Cephalaspidian shield, perhaps Hemicyclaspis Murchison. Passage-beds 
of Ledbury. Dr. Grindrod. 

A large head-shield of Hucephalaspis Powriet. The very great breadth of the 
shield is due to the flattening by pressure to which the fossil has been exposed; 
making allowance for this, the form of the cornua and the relative size and 
position of parts of the shield may be contrasted with those in the neigh- 
bouring figures. Mr. Powrie. 

Eucephalaspis Agassizii ; a head-shield showing well the relative size of the cornua, 
anterior position of the orbits, and form of the posterior median prolongation of 
the shield. Compare this and figs. 2 and 3; their differences are simply due to 
different pressure and preservation. Cornstones. Dr. Grindrod. 


PLATE IX 


J Dinkel hth W West imf 
C. MURCHISONI, AGASSIZII POWRLEI 


PLATE X. 


Fic. 

1. LEucephalaspis (Cephalaspis) Powriei. This most beautiful specimen shows the 
scales and fins in a marvellous state of preservation. The aspect presented is 
the ventral one, and the body has been twisted so that the marginal series of 
scales is on the right-hand side as one looks at the figure, whilst the dorsal 
series is on the left, as also the dorsal fin, the great lateral scales intervening. 
The form of the head (see also fig. 5, Pl. IX) contrasts forcibly with that of 
C. Lyellii or C. Agassizii, whilst the relative proportion of body and head is 
different from that presented by the specimens of C. Zyel/ii im Pls. VIII and 
XI. This specimen is from near Arbroath, Forfarshire, and is in Mr. Powrie’s 


Collection. 

2. Portion of the head-shield of a Cephalaspid from Forfarshire, showing the 
orbits as seen from below, with indications of blood-vessels passing in from the 
subjacent parts. 

3.  EHucephalaspis (Cephalaspis) Pagei, two thirds the natural size. 


4. Another specimen, drawn of the natural size. ‘These two specimens have been 
selected, as showing better than others in Mr. Powrie’s magnificent series the 
surface markings and the scales. ‘These scales are not well represented in 
fig. 3, in which it is not impossible that the ventral aspect 1s presented. ‘The 
dorsal fin is well seen in fig. 4. The woodcut, fig. 21, gives a faithful 
outline of the head, restored from specimens less crushed than these are. Both 
from the Cephalaspis-bed, Lower Old Red Sandstone, near Reswallie, Forfar, 
In Mr. Powrie’s cabinet. 


4a, 4b. Represent magnified bits, showing the surface markings of some of the scales; 4d 
are, perhaps, ventral scales. 


5.  Eucephalaspis (Cephalaspis) asper. This specimen is in a grey nodule, the fossil 
itself having a red tinge. The polygonal ares are very strongly brought out 
by the great pressure and the infiltration to which the shield has been sub- 
jected. A piece of the ‘rim’ of the shield has become detached, as also in 
the specimen in fig. 3. Mr. Powrie. 


5a, 5b. Represent the under and upper surface of two of the scales of the preceding 
specimen, with some of the characteristic asperities. 


Fic, 


ww 


4A. 


PLATE XI. 


Eucephalaspis (Cephalaspis) Lyell. Lateral view of the specimen from near 
Arbroath, Forfarshire, now in the Arbroath Museum. ‘The head is not well 
preserved, nor is the material of the scales; but their order, and the pectoral, 
dorsal, and caudal fins are admirably displayed. 


The same specimen, with a fragment of the body removed, displaying thus the 
ventral series of scales. The two pectoral appendages are well seen in this 
drawing, but the artist has introduced a joint-like structure in the right-hand 
appendage, which cannot be justified from the specimen itself. Compare the 
pectoral appendages of this specimen with those of Hv. Powriei, Pl. X, fig. 1. 

Caudal fin of a Cephalaspis, from the same bed which contains Hucephalaspis Paget, 
Eu. asper, and perhaps another species. Reswallie, Forfar. Mr. Powrie’s 
collection. 


A small specimen of C. Paget, with the head so crushed as to give the cornua an 
increasedly incurved appearance. 


PLATE XI 


J. Dinke) jith 


CG. LYBLLNT, PAGHI 


Fie. 


(he) 


PLATE XII. 


Head-shield of a small Aucephalaspis from Forfarshire. ‘The outline agrees with 
that of the little specimen drawn in PJ. VIII, fig. 3. 


A portion of the cephalic shield of Zenaspis (Cephalaspis) Salweyi, Egerton 
(= asterolepis, Harley). Cornstones, Herefordshire. In the Museum of the 
Geological Survey. The outline woodcut, fig. 26, has been drawn from a very 
much finer specimen, which I obtained too late to figure in a plate. For the 
dorsal scutes of this species see Pl. VIII, figs. 2 and 3. 

Hemicyclaspis (Cephalaspis) Murchisoni = C. ornatus. From the Passage-beds near 
Ludlow. In Mr. Lightbody’s collection. 

A similar specimen belonging to the Rev. P. B. Brodie. This and the above are 
the specimens figured by Sir Philip Egerton, ‘ Quart. Journ. Geol. Soc.,’ 
vol. xin, pl. IX, 1857. 

Compare these two specimens with those from the Ledbury beds, Pl. VIII, 
fig. 6, and Pl. IX, fig. 1. Their identity is fully proved by the 
ornament. 

Cephalaspis Salweyi. Six Philip Egerton’s figured specimen, /oc. cit. Cornstones 
of Acton Beauchamp, near Ludlow. Mr. Humphrey Salwey. 

A specimen from the same locality, and probably referable to the same species. 
The oblong form and imequality of the tubercles on the surface at one time 
appeared to me to separate C. Salwey from C. asterolepis, but a distinction 
cannot be maintained on these grounds, since the tubercles are equally irregular 
in specimens referred to C. asterolepis. This and the more perfect specimen 
with cornua in the British Museum are rather small specimens of the species. 
The cornua, which are long and rounded, are not preserved in this specimen. 
See the woodcut, fig. 26, page 53. This specimen belongs to Mr. Humphrey 


Salwey. 


PLATE XIII. 


Fie. 
1. Didymaspis Grindrodii. The shield is imbedded with its concave surface upwards. Dr. Grindrod’s 
collection. 
2. Natural cast of the same. From the red micaceous beds above the Auchenaspis Grits of Ledbury. 


The artist has not fairly represented the character of these specimens, which are better seen in 
vol. iv of the ‘ Geological Magazine,’ pl. viii, figs. 4 and 5. 

3. <Auchenaspis Egertoni. 

4. Ap . 

4a. A part of fig. 4 enlarged, showing the orbits, antorbital fossee and their septum, the superficial 
lamina which covers them in having been broken away, and the hexagonal areolation of 
the material of the test, The surface is not preserved; it is tuberculated, as in Cephalaspis. 

5. A part of fig. 4 enlarged. The difference in form and size of the cornua in these specimens is due to 
the more or less covering by matrix, a condition which the artist has not rendered apparent 
in these or the other figures. 

3, 4, and 5 are from the Auchenaspis Grits (Passage-beds), Ledbury, and belong to Dr. Grindrod. 
6,6a. Fragments of scales from beds at the base of the Cornstones, Ledbury. Dr. Grindrod. 

7, 8. Auchenaspis Salteri. The Passage-beds near Ludlow. Mr. Lightbody. 

9—14. Eukeraspis (Auchenaspis?) pustuliferus, from the Downton Sandstone of Ludford Lane, near 

Ludlow. 
9. Central part of the head-shield, showing the orbits and finely tuberculated ornament in intaglio. 
The margin is fractured, and has not the regularity here given by Mr. Lightbody. 

10. Fragment, showing the central part of the shield, enlarged. Dr. Grindrod. 

10a. The tubercular ornament still more enlarged. 

1]. One of the long cornua, showing the denticulations on the outer margin which led to such spe- 
cimens being considered as the jaw of a fish, the Plectrodus pustuliferus of Siluria. Mr. 
Lightbody. 

12. A specimen showing the marginal cavities or cells exposed by the breaking away of the upper 
surface of the shield. Dr. Grindrod. 

13. Aspecimen which shows well the form of the shield and cornua (see woodcut, fig. 31). The 
whole of the superficial lamina of the test is gone, and the subjacent cavities and laminz 
exposed. The part here preserved appears to agree with the anterior moiety of a shield of 
Auchenaspis or Didymaspis. The two bodies which have the appearance of orbits are cavi- 
ties underlying the shield surface, like the marginal cells. The true orbits lie anteriorly to 
this. Compare the figure of these genera. Mr. Lightbody. 

14. A fragment which shows beautifully the impression of one of the long cornua and a part of the 
head. ‘The denticulations of the margin are not drawn finely enough, and the cornu is 
represented as thicker than it is really. Mr. Lightbody. 

15,16. Fragments, the like of which are not uncommon in the lower parts of the Cornstones of Here- 
fordshire, probably the angles or cornua of Cephalaspidian shields which have been rolled and 
washed. Dr. Grindrod. 

17, 18. Mesially placed scutes belonging to some Cephalaspid of the same character and mode of 
occurrence as figs. 15 and 16. A specimen of very much larger size, similar to fig. 17, has come 
under my notice, and appeared to belong to Zenaspis. It is not figured in this work. Compare 
Pl. VIII, figs. 2, 3, and 4, with these. Dr. Grindrod. 

19. Cephalaspis Lightbodii, from the Passage-beds near Ludlow; the under surface of a very large and 
characteristic marginal rim is presented ; in the lower part of the specimen a portion of perhaps 
the dome of the shield is seen in intaglio; more probably this belonged to an additional scute. 
Mr. Lightbody. 

19a. A part of the tubercular ornament enlarged. 

20. Kallostrakon podura. ower Cornstones, Herefordshire. Dr. Grindrod. 

21. Tntaglio of a similar fragment, differing a little in the character of its markings. Dr. Grindrod. 


Lif) 


J Dinikel hth 
AITCHENASPIS., KALLOSTRAKON &° 


PLATE XIV. 


Fic, 


1. Horizontal section about the middle layer of the shield of Zenaspis, magnified 50 
diameters. 

2. The same, magnified 200 diameters. 

3. Horizontal section of the lower layer of the same, magnified 200 diameters. 

|. Horizontal section of tubercles of Zenaspis, magnified 30 diameters. 

5. Thesame, magnified 120 diameters. 

6. Horizontal section of a portion of the scale of Kallostrakon podura, magnified 200 
diameters. 

7. Aportion of the shield of Zenaspis Salweyz, magnified 4 diameters. The substance 
of the shield is worn away from the right down to the left side of the 
specimen, and thus the successive layers are exposed; on the right hand top 
corner the shining tubercular surface; below this the radiating stellate forms 
due to the disposition of the vascular canals ; below this again simple hexagonal 
or multiangular pieces completely emarginated by canals ; and lastly, horizontal 
branching canals, causing the “fibrous structure” spoken of by Agassiz. 
Vertical canals are seen in section as minute dots in all the different 
thicknesses. 

5. Vertical section of a part of the shield of Cephalaspis Lyellii, copied from Prof. 
Huxley’s paper in ‘ Quart. Journ. Geol. Soc.,’ vol. xiv, 1858. 


~*~ 


a 


PALMONTOGRAPHICAL SOCLETY. 


INSTITUTED MDCCCXLVIL. 


VOLUME FOR 1869. 


DCCCLX 


MONOGRAPH 


Or 


THE FOSSIL REPTILIA 


OF THE 


LIASSIC FORMATIONS. 


BY 


PROFESSOR OWEN, F.R.S., D.C.L., 


FOREIGN ASSOCIATE OF THE INSTITUTE OF FRANCE, 
ETC. ETC. 


PART SECOND. 
PER OS AUR TA. 


Pages 41—81; Plates XVII—XX. 


LONDON : 
PRINTED FOR THE PALHONTOGRAPHICAL SOCIETY. 


1870. 


| PRINTED BY 
J. E. ADLARD, BARTHOLOMEW CLOSE. 


MONOGRAPH 


THE FOSSIL REPTILIA 


LIASSIC FORMATIONS. 


OrperR—PTEROSAURIA, Ovwen. 
Genus—DimorPHoDoNn, Owen. 
Species—DIMORPHODON MACRONYX, Buckland. 


Remarns of volant Reptiles (P/erosawria) were later recognised, and, save in the 
instance about to be recorded, in a more fragmentary or scattered condition, in England 
than in Continental localities. 

A single bone or tooth gives value to a slab of Stonesfield Slate, and the evidence of 
a Pterodactyle rarely goes beyond such specimen in that Oolitic deposit. A jaw with 
teeth, or a skull more or less entire, from the Chalk of Kent, or the Upper Green-sand of 
Cambridge, has been welcomed for the fuller information so yielded; and such fossils, 
with a few detached vertebrae and wing-bones, have expanded our conceptions of the 
bulk attained by some of the Flying-dragons at the decline of the Mesozoic period. 

When the waters over which they flitted had a clayey or muddy bottom it afforded a 
quieter resting-place to the dead body of the Pterosaurian therein entombed. So the first 
discovered specimen of one of these in the upraised petrified ocean-bed now forming the 
Liassic cliffs of western Dorsetshire afforded BuckLanp' subjects, in the compass of a slab 
about a foot square, for a description and figures of the leg and wing-bones, with part of the 


1 «On the Discovery of a New Species of Pterodactyle in the Lias at Lyme Regis.” By the Rev. W. 
Buckland, D.D., F.R.S., F.G.S. (Read Feb. 6, 1829.) 


London,’ second series, 4to, vol. iii, 1835, p. 217, pl. xxvii. 


6 


‘Transactions of the Geological Society of 


42 FOSSIL REPTILIA OF THE 


vertebral column, of the species which he called Pterodactylus macronye—the first 
evidence of the genus from deposits so low, or ancient, in the Oolitic series. 

In 1858 I obtained the skull, with a few other parts of the skeleton of the same or a 
closely allied species, from the Lower Lias at Lyme Regis, and communicated a brief 
notice of it to the British Association, which that year met at Leeds.’ 

This specimen confirmed the accuracy of Buckland’s conjecture, which I had doubted, 
viz., that the portion of lower jaw with the series of small lancet-shaped, close-set teeth,” 
ina second slab of Lias, belonged to the same Pterodactyle as the limb-bones he described ; 
but it also showed that these teeth, so like those of some Fishes, were limited to the lower 
jaw, and were associated, in the same mouth, with long, slender, trenchant and sharp- 
pointed laniaries, projecting with wide intervals, and set in advance; which kind of teeth 
had, hitherto, alone been found in the different species of flying Reptiles. 


The chief result of the study of the second discovery of a Pterosaurian in Lias, viz., 
its evidence of a new generic form (Dimorphodon) in the order of volant Reptilia, in 
addition to Ramphorhynchus, von Meyer, and Pterodactylus proper, was noted in the com- 
munication above cited. 

The third specimen about to be described confirms that taxonomic deduction, showing 
a combination of the caudal character, mainly differentiating MRamphorhynchus from 
Pterodactylus, with the dental character above defined. 

I propose first to describe and figure the two specimens yielding the cranial and 
dental characters of Dimorphodon, and then to attempt a restoration of the Liassic species, 
D. macronyx. 

The first specimen with the skull is figured in Pl. XVII. It is on a slab of Lias, mea- 
suring 11 inches by 7 inches. The right side of the head is exposed :* it has been subject 
to pressure and some degree of dislocation. Certain bones of both wings, and a few 
other parts of the skeleton are preserved, pell-mell, in this slab, pressed amongst 
and upon the bones of the head, especially at the back part of the skull. 

The right premaxillary (22), maxillary (21), and nasal (15), are almost in their natural 
positions, give the profile contour of that part of the skull, show most of the teeth of the 
right side upper jaw, and reveal the singular expansion of the nasal (x) and antorbital (a) 
vacuities. The alveolar part of the left maxillary (8’), with its ascending postnarial 
branch has been pushed obliquely downward, with fracture, but without much 
displacement, of the beginning of the alveolar ray, the inner surface of which is 
exposed. 

The mandible (32) has been dislocated and pushed below the place of its articulation 
with the tympanic (gs): the left ramus has also been subject to the same force which has 


1 «On a New Genus (Dimorphodon) of Pterosauria, with Remarks on the Geological Distribution of 
Flying Reptiles ;”’ in ‘ Reports (Sections) of the British Association,’ 1858, p. 97. 

2 Buckland, loc. cit., pl. xxvii, fig. 3. 

3 The specimen has been drawn, in Pl. XVII, without reversing. 


LIASSIC FORMATIONS. 43 


dislocated that side of the upper jaw; the hind part of this ramus is obliquely depressed, 
so as to expose the inner surface (32). 

The anterior entire or undivided part of the premaxillary (22) is about 2 inches in length, 
and 14 inch in vertical height at its back part: it contains four pairs of teeth, which are 
the largest and longest of the series. The foremost tooth (1) is terminal, with a crown 
5 lines in length, rather over 1 line in breadth (fore-and-aft) at its base ; it is subcompressed, 
subrecurved, and sharp-pointed. An interval of 4 lines divides it from the second tooth 
(2), with a crown 5: lines long. After an interval of 7 lines projects the crown (8) of 
the third tooth, 7 lines in length and 2 lines in basal breadth, sharp-pointed like the first, 
but less bent. The socket and base (?) of the fourth tooth appear at an interval of 
6 lines, and below is the entire and displaced homotypal tooth (4’) of the left side, showing 
the cavity on the inner side of its root which would have received the successional laniary. 
This tooth measures 1 inch 2 lines in total length, of which the exposed enamelled crown 
forms two-thirds. In advance of the foremost tooth (1) is seen part of its homotype (1’) of 
the left side, also displaced from the socket, and showing the depression and vacuity on 
the inside of the base, in relation to the succeeding tooth. Beyond the fourth alveolus 
the maxillary (21) appears, underlapping the part of the premaxillary (22") which defines the 
lower and anterior part of the narial vacuity : the maxillary is continued straight backward, 
with feeble indications of two crushed alveoli (5, 6) for 1 inch 9 lines, when the seventh 
laniary (7) projects almost straight downward: the crown of this tooth is 5 Jines long; the 
root, covered with rougher cement, slightly contracts to its implanted end, which has 
slipped a short way out of its socket. An interval of 4 lines divides this from the next 
laniary (8), which shows a crown of but 3 lines in length; this projects opposite the fore 
part of the lateral post-narial branch (21s) of the maxillary. The base of the left 
homotypal tooth (g’) projects from the same part of the dislocated left alveolar branch of the 
maxillary ; and above this, on the inner side of that bone, is exposed the coronal germ of 
asuccessor. In the right maxillary two other straight laniaries (9, 10) of rather decreasing 
length, project with similar or rather lessening intervals: then follows, after an interval of 
3 lines, a pointed compressed crown 14 line in length (11); and, at shorter intervals, 
two smaller pointed compressed teeth (12 and 13). 

These thirteen cuspidate teeth of the upper jaw are included in an extent of the 
alveolar border measuring 5 inches 2 lines. ‘That border is continued backward, straight 
and edentulous, for 9 lines beyond the last tooth, when it is crossed by the large and long 
first phalanx (zr 1) of the wing-finger. his edentulous part of the maxillary forms the 
lower straight border or base of the large triangular antorbital vacuity (a), at the back 
part of which it is overlapped by the fore part of the slender malar (26). | Above this 
vacuity are parts of the nasal (15) and prefrontal (14), both somewhat displaced in this 
crushed part of the skull. The arched part of the frontal forming the upper part of the 
rim of the orbit (0) is recognisable at (11) Pl. XVII. Above its hind part are indications of 
the post-frontal (12) and mastoid (s), with the process of the latter descending external to 


44, FOSSIL REPTILIA OF THE 


its articulation with the tympanic (2s). The metacarpus and dislocated unguiculate digits 
of the wing-limb are confusedly interblended with the crushed and dislocated back part 
of this skull; three phalanges (77 1, 1 2, 7 3) of the wing-finger are determinable. 

‘The two anterior teeth (1’, 2’) of the mandible show longitudinal angular depressions at 
their base, indicating exposure of their inner side, and that they belong to the left ramus. 
The corresponding part of the right ramus may have been broken away : the third laniary 
(3') clearly belongs to this ramus, which is fractured beneath its socket. The point of this 
tooth is broken off: what remains of the body is curved, and is implanted more obliquely 
backward than the two preceding teeth. ‘This at first led me to suspect it might be the 
foremost tooth of the mandible, and that the left ramus had been pushed in advance as well 
as downward: but my doubts on this point have been set at rest by the specimen (PI. 
XVIIT) next to be described, and I view the tooth in question as the third of the mandibular 
series : it is divided from the second by an interval of 6 lines, and the second stands at a 
rather shorter interval behind the first. Five lines behind the third tooth is the base of a 
fourth lamiary (4’), and four lines further back is an indication of a fifth (5’). This is followed 
by the characteristic series of between thirty and forty very small, subcompressed cuspidate 
teeth, each less than a line in length, corresponding in extent with the maxillary part of 
the upper jaw. ‘The entire series of mandibular teeth occupies an extent of alveolar 
border measuring 5 inches | line. 

The depth of the right ramus gradually increases from 5 lines below the last laniary 
to 10 lines below the last denticle. The inner side of the dislocated ramus (32’) shows a 
strong longitudinal ridge projecting inwards about 3 lines above the lower border. ‘The 
outer surface of the ramus seems to have been strengthened near its lower border by a 
similar but lower ridge. 

‘he distal ends of the antibrachial bones (54, 55) overlap the hind part of the mandible: 
that which shows the larger articular surface, opposite the three slender metacarpals, 
should be the radius. The base of the supplementary styloid bone appears near the distal 
end of the ulna, but is better shown in Buckland’s original specimen.’ Indications of 
two carpals intervene between these and the metacarpus. ‘This overlies and conceals the 
articular pedicle of the mandible and contiguous parts (squamosal, malar, &c.) of the 
skull. ‘The metacarpus includes the three slender supports of digits 47,maz, and the 
strong and thick metacarpal of the wing-finger (;,). ‘This bone, being almost con- 
cealed by the first phalanx in Buckland’s specimen, was overlooked, and that phalanx 
was described as the metacarpal of the wing-finger, which, accordingly, in the restoration, 
fig. 2, Pl. 27, of ‘Buckland’s Memoir,’ is made three times the length of the other and 
more slender metacarpals (3'). I have, therefore, had that part of the original specimen, 
now in the British Museum, redrawn (PI. XIX, fig. 1), the true metacarpal being shown at 
m4, It corresponds with the same bone in previously described Péerosauria by surpassing 
in thickness, not in length, the other constituents of the metacarpus. In the specimen, 


1 As in the part re-drawn in fig. 1, Pl. XIX. 


LIASSIC FORMATIONS. 45 


Pl. XVII, the metacarpal of one wing-finger is clearly shown at zm. That of the other, lying 
upon the cranium, is more obscure. ‘The thin compact wall of this pneumatic bone has been 
crushed in upon the wide air-cavity, as with most of the other long bones, so that it looks 
hike two metacarpals. The proximal articular surface of 17 is partly concave and partly 
convex : the distal articulation is trochlear, moderately concave from side to side at the 
middle, convex from behind forward, with a depression behind, above the articulation, for 
securing the olecranoid process of the proximal phalanx. This phalanx (;,,1), in one 
wing, is bent back upon the fore-arm, crosses the dislocated mandible, and has been 
pressed upon it, long and hard enough to leave a channel in the right ramus, where part 
of the phalanx has been removed: its length is 4 inches 6 lines. 

The second, more slender and longer phalanx (jy, 2), is bent at nearly a right angle 
with the first, and lies below and parallel with the mandible: it is nearly 5 inches in 
length. ‘The third phalanx (;;,3) is bent upward in front of both lower and upper jaws: 
43 inches of its length is preserved in the slab: from the analogy of the better pre- 
served specimens (Pl. XVIII, ,, 3), about 1 inch 3 lines are wanting from the 
distal end. 

Of the three unguiculate digits the characteristic large claws are preserved: one (;;) lies 
above the frontal (11) with the penultimate phalanx ; the other two are between the upper 
and lower jaws, with some of the slender phalanges: all these parts of the ramus having 
been dislocated and scattered. 

Parts of the distal ends of the radius and ulna (54) 55’), the metacarpal of the 
wing-finger (am), and the proximal end of its first phalanx (77 ~), of the opposite fore- 
limb, occupy a lower corner of the slab: carpal bones, one of the accessory styloid ossicles 
of the forearm, some of the slender metacarpals of the claw-fingers can be made out 
above these: and there are more obscure indications of vertebrae at that end of the slab, 
curving toward the cranium. 

All the osseous and dental textures are black, as if charred by slow combustion of the 
animal matter. 


DimorpHopon macronyx. Pl. XVIII. 


In August, 1868, I was favoured by the Earl of Enniskillen, then at Lyme Regis, 
Dorsetshire, with a list of parts of a Pterodactyle, in a slab of Lias about 20 inches by 
1] inches, and of other parts in detached portions of Lias, including the entire tail with its 
bone-tendons, which his Lordship had observed at Messrs. James and Henry Marder’s, 
the judicious and persevering collectors of the fossils of that rich locality. 

The result of this valuable and timely information was the securing for the British 
Museum the entire series of these Pterosaurian fossils. 


4.6 FOSSIL REPTILIA OF THE 


‘They proved to be parts of the Dimorphodon macronye, confirmed many of the 
observations made on previously acquired specimens, corrected others, and added almost 
all that was required for the restoration of the skeleton of this remarkable genus and 
species, which I have accordingly attempted in Pl. XX. ; 

The slab of Lias with the second specimen, including the skull of Dimorphodon macronyz, 
is of larger size, shows more of the skeleton and in a more separated and definable state 
than in Pl. XVII. Nine dorsal vertebrze, third to eleventh inclusive, in natural juxta- 
position, with the twelfth slightly dislocated, are preserved at the upper part of the slab 
(Pl. XVIII, pv). The summits of the neural spines (ns) of most of these, and the disposition of 
many of the preserved ribs, show that they lie mainly with the dorsal aspect downward (as the 
specimen is figured). ‘This explains and accords with the position of the parts of the pelvis, 
which lie a little way behind the dorsal vertebrae. The comparatively slender ilium (6, 62) 
is downward ; the broad ischium (;s), and the pair of spatulate pubic bones (,,), are turned, 
like most of the ribs, upward, as I conclude the abdominal or ventral surface of the trunk 
was directed as the fossil lies in the figured slab. The bones of the hind-limb, m connection 
with the acetabulum, are turned outward, with their inner surface exposed. The projections 
of the trochlear terminations of the metatarsals (i, ix 9), show that the sole of the foot 
is turned to view. Accordingly, we have here the bones of the left hind limb. On the 
hypothesis that the femur and tibia are seen from the outside, which at first suggests 
itself, they would belong to the right limb, viewed in profile. But then, the broad thin 
plate of bone contributing to the acetabulum, would represent the ilium, and the indi- 
cations of the pelvis below the acetabulum and head of the femur would represent ischium 
and pubis. ‘This interpretation, however, gives to Dimorphodon proportions of pelvic 
bones very different from those determined by Wagner in Plerodactylus Kochu,' and by 
Quenstedt? in Pterodactylus suevicus; and, besides, it leaves undetermined the pair of 
bones (64, Pl.X VII) which closely resemble in form and proportion the ‘pubic bones’ (w, «) in 
Quenstedt’s instructive plate? ‘In this plate the ilia (s, s) are represented as long slender 
bones, contributing the upper but smaller proportion of the acetabulum, and extending 
horizontally beyond it both forward and backward. The pelvis, in the position in which 
I conclude it to lie in the slab figured in Pl. XVIII, might well afford such indications of 
the pre- and post-acetabular productions of the ilium as are there shown at gg. In 
Pterodactylus suevicus the ischium contributes the lower and major half of the 
acetabulum (¢r, loc. cit.), and expands into a broad thin plate (s, 1b.), having the pro- 
portions to that of the spatulate pubis, which the bone ¢s bears to 4, in Pl. XVIII. The 
portion of the pelvis in the original specimen being preserved in natural connection with 


1 « Ueber Ornithocephalus Kochii,’ in ‘Abhandl. d. math.-phys. Klasse der Bayerischen Akad.,’ ii, 
4to, Miinchen, 1837. 

2 «Ueber Pterodactylus suevicus,’ &c., 4to, Tibingen, 1855. 

3 In the Memoir above cited. 


LIASSIC FORMATIONS. L7 


the sacrum and contiguous vertebra is figured in Pl. XIX, fig. 2; and the constituent 
bones are rightly recognised by Buckland (op. cit., p. 222). 

It is interesting to note, that the pelvis of P/erosauria, so determined, resembles more 
closely that of the existing representatives of the section of Rep/ilia with the 4-chambered 
heart and double-jointed ribs, viz., Crocodilia, than it does the pelvis in Chelonia and 
Lacertia. The ischium in Crocodilia, e. g-, Surpasses the pubis in size, and excludes that 
hemapophysis from the acetabulum.!. The ischium seems to contribute the larger share 
of the acetabulum in Dimorphodon, Pl. XVIII, a. In Birds, as in Lizards, the pubis 
forms part of the acetabular cavity.’ 

In the specimen, Pl. XVIII, a portion, ed, of a long tail, of which the vertebrae were 
surrounded by numerous slender bone-tendons, extends backward and downward beyond 
the pelvis: a better preserved portion with three caudal vertebree (c @) is preserved in a 
detached part of the matrix found in the vicinity of the larger slab. But to this part of 
the vertebral column I shall return in describing the more perfect specimen of the tail of 
Dimorphodon, from another individual (Pl. XIX, fig. 4). 

Behind the skull are four cervical vertebree (Pl. XVIII, c), and part of a fifth in natural 
juxtaposition, or perhaps a little separated at the articular surfaces. ‘The under surface of 
the centrums and articular processes of the neural arches are exposed. ‘The sides of the 
centruims show a slight concavity, but their crushed state obscures the natural contour of 
the under surface. The hind part of the under surface, in the last two of these 
vertebrae, shows a pair of low obtuse processes, with an indication of a convex terminal 
articular surface. ‘The centrum expands in breadth as it advances, and sends out a short 
thick process (parapophysis) from each side of the fore part; to which, in the last three 
vertebrae, are indications of attachment, or parts, of a backwardly produced styliform mb. 
At the midline of the fore part of the last two of these vertebrae a fracture indicates a 
ridge or process there to have been broken off. |The pre-zygapophyses are thick, and project 
far in advance of the concave anterior articular surface of the centrum: the convex 
posterior articular surface of the centrum projects as far beyond the post-zygapophyses. 
Their joints are more vertical than horizontal: the posterior surfaces looking slightly 
outward and downward. 

The superior breadth of the neural arch, as compared with that of the centrum, 
brings its articular processes into view, along each side of the vertebral bodies, in the 
degree shown in Pl. XVIII, c. The character of the articulations indicate less extent and 
freedom of movement of the cervical vertebre than in Birds, and more restriction in. the 
lateral than in the vertical directions. ‘The interlocking joints resulting from the different 
lengths of the fore and hind articular processes add strength to the part of the spine 
supporting the head. 

The cervical vertebrae of Dimorphodon, so far as their structure is exemplified in the 


1 «Anatomy of Vertebrates,’ vol. v, p. 188, fig. 119. 
2 Ib., p. 190. 


48 FOSSIL REPTILIA OF THE 


present specimen, conform to the pterosaurian characteristics of these vertebree, as shown 
in those of Pferodactylus Sedgwickii, described and figured in the ‘ Monograph on the 
Fossil Reptilia of the Cretaceous Formations,’ Supplement No. 1 (1859), pp. 7—10, 
Pl. II, figs. 7—18 ; and in those of Pterodactylus simus, ib. Supplement No. II (1861), 
p. 7, Pl. IL, figs. 1—5. 

The skull preserved in the present specimen agrees in size with that in the slab pre- 
vious!y received (PI.X VII), repeats the characteristics of the genus Dimorphodon, and shows no 
differences of greater degree or value than may be set down to individual modifications. The 
part defective and partly obscured by intrusive bones from other parts of the skeleton is un- 
fortunately that which leaves the precise determination of structure unsatisfactory in the pre- 
viously described specimen. A trace only of tympanic remains at 98, and of the descending 
styloid process of the mastoid at s: the thick metacarpal of the wing-finger (iy, m), intrudes 
into the orbit, and overlaps the upper end of the malar (26). More of the part of the frontal 
forming the superorbital arch (11) is shown than in Pl]. XVII. Part of the concave surface 
of the orbital cavity beneath the superciliary ridge is here seen. The lacrymal (23) or 
descending branch of the prefrontal (14) meets the ascending process from the combined 
malar and maxillary, dividing the orbital from the antorbital cavity. The true size and 
shape of the latter vacuity (0) is here well displayed. The maxillary styloid process (21%) 
rises, at the same angle backward as in Pl. XVII, to join the nasal (15). The medial branch 
or ray of the premaxillary (22’), the end of which is depressed below the prefrontal in 
Pl. XVII, preserves its position in the present specimen, and yields the true arched 
contour of the profile of this remarkable skull. 

The entire vertical extent of the vast narial vacuity, n, is here given, the longitudinal 
one, 3 inches, precisely agreeing with that in the first-described skull. The anterior 
part of the premaxillary (22) shows, also, the same proportions and shape, viewed side- 
ways, as in the first specimen. ‘The conformity is instructively continued in the characters 
of the dental system. The apex of the crown of the lamiary (Pl. XVIII, 1) from the fore end 
of the premaxillary shows the same curvature and proportions as in Pl. XVII; the same 
interval divides it from the second laniary (2); the longer interval, again, occurs between 
the second and the third laniary, with a longer and less curved crown. After an interval 
of seven lines comes the fourth tooth (4), corresponding in size and shape with the one 
which is displaced in P]. XVI, 4’. After an interval of nine lines the apex of the crown of: 
seemingly, the successor of the fifth lamiary (5) appears. It may be, normally, smaller 
than the rest; the socket of this tooth is feebly indicated in the subject of Pl. XVII. The 
sixth laniary (6) shows the same size and relative position as in that subject, and the same 
may be said of the five succeeding teeth, save that the last is rather larger than in Pl. XVII, 
which also shows an additional small hind cuspidate tooth. ‘he suture between the 
premaxillary (22”) and the maxillary (21) is more plainly discernible in the present specimen. 

The extent of alveolar surface of the left upper jaw occupied by the above-described 
dental series is 5 inches 3 lines. 


LIASSIC FORMATIONS. 49 


In the left ramus of the mandible two of the large anterior laniaries are in place ; one, 
answering to the second in Pl. XVII, 2’, projects across the diastema between the second 
and third tooth above ; in size, shape, and curvature, it resembles the second upper laniary, 
close to which it terminates. ‘lhe next mandibular tooth is larger, less curved, and crosses 
the middle of the interval between the third and fourth upper laniaries. The tooth (1) 
displaced beneath the mutilated fore part of the mandible, I take to be the foremost of 
the mandibular series and suppose that its point would naturally project across the interval 
between the first and second of the upper teeth. The fourth laniary appears to be more 
displaced; its base or root, with a lateral depression, is shown behind the fifth tooth of 
the minute serial teeth, and the crown passes obliquely backward on the inner side of 
that of the sixth upper laniary, by which it is concealed. Of the serial teeth, with pointed 
crowns from half a line to a line in length, about thirty may be reckoned occupying an 
alveolar extent of 2 inches, 9 lines. 

At the hind part of the left mandibular ramus, here exposed, three longitudinal 
ridges define two vacuities, of which the inferior may be natural. The upper one seems 
more plainly due to loss of the thin outer plate of bone extended between the upper two 
ridges. ‘The proportions of the ramus closely accord with those of the first-described 
specimen. ‘The fore part of the mandible is too much mutilated for useful comparison. 

The dentition of Dimorphodon, as displayed by the second specimen of skull, consists, 
in the upper jaw, of laniaries with wide intervals, eleven in number on each side; in the 
lower jaw, of four, if not five, laniaries implanted at the fore part of each ramus of the 
mandible at intervals corresponding with three of the four anterior lamiaries above ; then 
follows the long series of close-set and minute pointed teeth. The difference of dentition 
as compared with the first specimen (Pl. XVII) is, in the upper jaw, in the additional 
small laniary or cuspidate tooth at the back part of the series in that specimen. In the 
lower jaw there does not seem to be any noteworthy difference in the number, kinds, and 
position of the teeth. The longest laniaries are included between the second and fifth in 
both jaws : the upper laniaries after the fourth become small and straight. 

At the first view of the framework of the huge head of our Liassic dragon one is 
struck with the economy of bony material and the purposive skill with which it has 
been applied or disposed, so as to give strength where resisting power was most required. 

The lodgment of the poorly developed brain enlists a miserably small proportion of the 
skull: the cranium proper, or brain-case, is relegated to an out-of-the-way corner, so to 
speak, and there it is almost concealed by the projections for joints or muscular 
attachments. The orbits accord with the large eyes given to this volant and swift-moving 
Reptile. 

One can conceive no necessary interdependent relation between the wide external 
bony nostril (”) and the organ of smell, nor be led to conjecture that the tegumentary 
inlets to the nasal chamber were larger than is usual in Reptiles. 

The main purpose of the head is for prehension of prey. ‘The jaws are produced far 


7 


50 FOSSIL REPTILIA OF THE 


forward to form a wide-gaping mouth, and are formidably armed. We may conceive, 
therefore, that the dragon may have occasionally seized an aninial of such size as to 
require considerable force of jaw for overcoming its struggles. ‘The means of resist- 
ance were afforded to the upper or fixed maxilla, not by a continuous wall of bone, but 
by curved columns or abutments. The chief of these is the upper medial arch of bone 
which overspans the skull lengthwise, from the short roof of the cranium to the 
fore part of the premaxillary (22); the frontals (11) and nasals (15) combining with the 
mid-fork or branch of the premaxillary (22’,) to constitute this arched key-ridge of the roof 
of the head. 

From it two piers or buttresses out-span on each side, to give strength and resistance 
to the upper jaw, and especially its alveolar tracts. One, proceeding from the 
nasal, meets the uprising process of the maxillary (21); this abutment, curving from 
above outward and obliquely forward, expands and backs the part of the jaw where the 
second group of large laniaries project. The second buttress is continued from the pre- 
frontal (14), and arches more directly outward to meet the uprising process of the 
malo-maxillary. A third arch, due to the post-frontal (12) and malar (26), expands to abut 
upon the hind end of the maxillary arch, and gives support to the part of the skull which 
the temporal muscles tended to pull downward when they were giving to the mandible 
the power of a strong bite or grip. Finally, comes the strongest of the four piers, due to 
the mastoid (g) and tympanic (28), for giving articular attachments to the rami of the 
lower jaw. 

Thus, four vacuities appear in the side-walls of the skull: the first (,) 1s the largest, 
between the small consolidated or continuous fore part of the skull (22), and the naso- 
maxillary pillar (21x,15). This vacuity answers to the external bony nostril of the same side, 
in the Lizard’s skull (Pl. XX, fig. 3, »), where the nostrils are divided and more or less lateral. 
The second vacuity (q) is somewhat less, of a triangular form, with the base downward : it 
answers to the antorbital vacuity in Zyriocephalus (ib., a) and a few other Lizards, and to that 
in Teleosaurus, where, however, it is very small. The third vacuity (0) still decreasing, is 
oval, with the narrow end or apex downward: it answers to the orbit, but is of large size 
compared with most Saurians ; it is, however, exceeded in relative expanse by the orbit in 
Lyriocephatus (ab., 0). 

The fourth vacuity is the narrowest: it answers to the so-called ‘temporal fossa’ and 
was occupied by the muscles of the same name. Extension of surface, for their origin, and 
additional strengthening of this back part of the skull are gained by laying horizontally 
across the temporal fossa the bony beams called ‘upper and lower zygomata,’ arching 
from the postfronto-malar to the masto-tympanic vertical columns. The heavy phosphate 
of lime, thus singularly economised by the disposition of the bones on mechanical prin- 
ciples plainly to that end, is made to go still further by the arrangement of the osseous 
tissue. Every bone is pneumatic, the abundant, open, cancellous structure being included 
in a very thin layer of compact osteine. 


LIASSIC FORMATIONS. 51 


The bones of the limbs are dislocated and dispersed in the way and degree common 
to the specimens of this animal! hitherto discovered (Buckland, loc. cit., pl. xxvil; and pls. 
XVII and XVIII of the present Monograph). ‘The scapula (Pl. XVII, 51) and coracoid 
(ib., 52) in the same anchylosed condition as in the first-described specimen, are at the end 
of the slab opposite to that with the head. The corresponding humerus (53), preserved in a 
separate portion of the block of Lias, shows the entire contour of the pectoral process (4). 
The right humerus (53') lies below the dorsal vertebrae (p) ; the upper part of the pectoral 
process (%) is wanting, but the obtuse thickening of the end of that remarkable pro- 
duction is well shown. The ridge (e) called ‘ulnar,* descending from the ‘lesser 
tuberosity,’ appears in this view of the ‘palmar’ surface of the bone.’ The sigmoid 
flexure of the shaft is much better marked than in the humerus of Pferodactylus suevicus.® 
The stronger walls of the humerus have resisted the pressure better than those of 
most of the other long bones. 

Of the antibrachial bones parts of the shafts, crushed, are seen at 54, 55, apparently 
of the right wing. With the distal ends of these, the right carpus (56) and metacarpus 
(57) appear to have retained their natural connections. The slender metacarpals of the 
first (1), second (x), and third (1) digits appear emerging from beneath the left hind 
foot which overlies their proximal ends. The phalanges of the first digit (x), two in 
number, preserve their natural articulations. As are also those of the second digit, three 
in number. The metacarpal of this digit is longer by 25 lines than that of the first. The 
additional phalanx would seem to be the proximal one, by its shortness: the second 
phalanx more nearly agrees in length with that supporting the claw-phalanx in the first 
digit; but it is thicker and a little longer. The four phalanges of the third digit (1) are 
dislocated; but the penultimate, which is the longest, retains its connection with the 
ungual phalanx. The proximal phalanx is longer than the second, which resembles in 
length, and seems homotypal with, the proximal phalanx of the second digit. It may be 
concluded, therefore, that the additional phalanx to 1 and mt was developed at the 
attachment of the digit to the metacarpus. ‘The largely and abruptly expanded meta- 
carpal of the fourth digit is in great part covered by the correspondingly thickened and 
much elongated phalanx (;7, 1) therewith articulated. The olecranoid process of this 
phalanx is well shown, and the entire bone is preserved: its length is 4 inches 2 lines: it 
is bent directly and abruptly back upon its metacarpal. .To the distal end is attached 
part of the second phalanx (;;. 2). 

The proximal phalanx of the left wing-finger is preserved in a detached (7, 1) part of the 
slab (Pl. XVIII) containing the major part of the skeleton. The second phalanx (jy, ») of the 
left wing-finger lies in that slab, is entire, and yields a length of 4 inches 9 lines. The third 


1 “Monograph on Fossil Reptilia of Cretaceous Formations,’ Suppl. No. iii, 1860, p. 14, pl. iti, 
fig: 1, e. 

2 Op. cit., 190. 

3 Quenstedt, op. cit., ¢ J, er. 


52 FOSSIL REPTILIA OF THE 


phalanx (jy, ) is 5 inches 6 lines in length; near its distal end is part of the slender terminal 
phalanx of this digit (;4). There is no trace of a fourth unguiculate digit, and I return to 
Cuvier’s view of the structure and homologies of the hand of the Petrodactyle,’ which I 
had abandoned in favour of the seemingly more perfect evidence supporting Professor 
Goldfuss’ restoration of Pterodactylus crassirostris,’ adopted by Buckland* and myself.‘ 

The metacarpal of the left wing-finger (j7.,,, Pl. XVIII) lies beneath the back part of 
the skull, and is over-lapped by the superorbital part of the frontal. Portions of two of 
the unguiculate digits of the same fore-paw (; ;,) are seen in the wide narial vacuity. 

The definition and finish, so to speak, of the joints of the wing-finger are worthy of 
note, especially of that between the metacarpal bone and proximal phalanx. In Reptiles 
generally the articular extremities of the long bones are not very definitely sculptured, and 
do not manifest that reciprocal adaptation of their inequalities which are observed in the 
joints of Mammals and Birds. The difficulty of determining the coadapted extremities of 
detached bones of Reptiles is increased by the great thickness of the cartilage which covers 
them and renders their mutual contact more intimate, and which is always wanting in 
fossil bones. The Pterosaurian modification is, however, purely adaptive ; and the relation 
to Warm-blooded Vertebrates in this respect is one of analogy. An argument in favour 
of avian affinity from the joint-structures could only be propounded by one not gifted with 
the judgment needed to deal with problems of this nature. 

The left femur (65’) preserves its natural articulation with the acetabulum ; the head is 
bent forward from the line of the shaft for an extent like that at which the condyles are 
produced backward ; the shaft is straight, the great trochanter is feebly developed. There 
is no evidence of a modification of the distal condyle for the interlocking articulation with 
the fibula, which in Birds relates to their bipedal station and walk. The length of this 
femur is 3 inches 4 lines 

The left tibia (66’), bent back at an acute angle upon the femur, measures 4 inches 10 
lines in length. There is no trace of patella, nor has this sesamoid bone been found in 
any Pterosaur. The inner side of the bone being exposed, the styliform rudiment of the 
fibula is hidden from view. ‘The trochlear termination of the distal end of the tibia is 
better marked than in Crocodilus, or even than in Scelidosaurus (‘ Monograph on Oolitic 
Reptilia,’ Part TT (1863), p. 16, Pl. X, 66), and consequently approaches more nearly to 
the characteristic form of the joint in Birds. The resemblance to the bicondyloid termina- 
tion of the femur is instructively shown in the distal portion of the Pterosaurian tibia 
figured in Pl. XIX, figs. 8 and 9, and in the distal half of the right tibia of Dimorphodon 


1 «Ossemens Fossiles,’ 4to, v, pt. ii, p. 371. 

2 Beitrage zur Kentniss verschiedener Reptilien der Vorwelt, in ‘Nova Acta Acad. Natur. Curios.,’ 
Leopold Carol., &c., 4ito, tom. xv. ‘‘ Reptilien aus dem lithographischen Schiefer, Pterodactylus crassiros- 
tris, nobis, tabs. VII—X.” 

3 «Bridgewater Treatise,’ 8vo, 1836, pl. 22. 

4 Owen’s ‘ Paleontology,’ 8vo, 1861, fig. 97. 


LIASSIC FORMATIONS. 53 


in the slab, Pl. XVIII, at 66, which crosses the right antibrachium (54, 55). The deflected 
posterior ends of the condyles are here shown, and beneath them three tarsal bones 
(0, 2, 6), with the characteristic short and thick metatarsal of the fifth toe (m, v)." 

The tarsal bone between the tibial trochlea and the three metatarsals (2, 77, 727), 
answers to the astragalus, marked a, in Scelidosaurus and Crocodilus (Monograph and 
Plate above cited); two tarsals, of which the one representing the second row is the 
smallest, intervene between the tibia and the fifth metatarsal; the larger of these ossicles 
answers to the caleaneum (/ in Scelidosaurus and Crocodilus, Monograph, ut supra), the 
smaller and distal one to the cuboides (4, 76.). 

The bony frame-work of the left foot (69’) is instructively preserved; the first four 
metatarsals are, as usual, long and slender, and resemble those in previously described 
Pterosauria ; their under or plantar surface is exposed. ‘The metatarsal of the first or 
innermost toe (2) is the shortest, that of the fourth toe (7v) is next in length ; the third (277) 
is the longest, but there is little difference in this respect ; their distal condyles project 
toward the sole, and are made trochlear by a mid-groove. 

The innermost digit shows the proximal and ungual phalanges in natural connection 
with each other and with the metatarsal: the ungual phalanx (7) is scarcely half the size 
of that of the corresponding digit (7) of the fore-foot. The ungual phalanges of the three 
other toes (é, ai, wv) are preserved, showing the usual uniformity of size in the hind- 
foot of Pferosauria : the number and disposition of the contiguous but scattered phalanges 
best accord with the phalangeal formula (3, 4, 5) presented by the second, third, and fourth 
toes respectively, in better preserved feet of other Pferosauria (P1. XIX, fig. 5). There 
is, however, here unequivocal evidence of a fifth toe, and that not merely rudimental but 
recognisably functional though without a claw. The tarsal bones (%, 4) support a meta- 
tarsal (m, v) directed parallel with the metatarsals (:—7v), but much shorter and also thicker : 
_it is 6 lines in length, and expanded at both ends, the proximal one bemg 25 lines in 
breadth, the distal one 2 lines, and the middle of the shaft 15 line. ‘The under or 
plantar side of the bone is exposed, as in the others, and shows a shallow oblique channel 
passing from the proximal end obliquely to the inner side of the shaft, dividing two 
elevations at that aspect of the proximal end. The distal end is a moderately convex 
condyle, the outer and plantar prominence of which is broken off. I regard this bone as 
the fifth metatarsal. It supports a digit of two phalanges: the first (1, v) is slightly 
dislocated, so as to show the concavity of its proximal joint close to the condyle to which 
it was articulated: it is 1 inch 3 lines in length, and is thicker as well as much longer 
than the corresponding phalanx of the other toes. The second phalanx (2, v) is 1 inch in 
length: it is bent back upon the first, and gradually tapers to a point. Both phalanges, 


1 This throws expository light on the idea, revived by Gegenbaur (‘ Vergleichend-anatomische Bemer- 
kungen iiber das Fussskelet der Vogel, in Reichert’s ‘Archiv fiir Anatomie, Physiologie, und wissensehi. 
Medicin,’ 1863, p. 445), viz., that the distal trochlear epiphysis of the Bird’s tibia represents its proximal 
tarsal series, or astragalus. 


54 FOSSIL REPTILIA OF THE 


in the specimen described, pass obliquely across and beneath the four long metatarsals sup- 
porting the unguiculate claws.’ 

From the position of this exunguiculate long and slender toe, as well as from its 
difference of structure, we may infer its application to a different office from that of the 
other toes. These obviously subserve the purposes of terrestrial locomotion, and 
perhaps of suspension: the fifth toe I infer to have helped to support, like 
the similarly shaped production of the calcaneum in certain Bats, the interfemoral 
expansion of alar mtegument, in the way indicated in the restoration (fig. 2, Pl. XX) of 
Dimorphodon macronye. In the habitual mode of locomotion by vigorous act of flight 
this toe would be in action while the other four were at rest; hence the necessity for 
greater thickness and strength of its bones, and the size of one of the tendons, as indicated 
by the groove in the metatarsal. Interesting, also, is it to note the analogy of this 
‘wing-toe’ with the ‘wing-finger,’ though they be not homotypes, as shown in the 
shortness as well as thickness of the metapodial bone and the length of the pointed, claw- 
less, terminal phalanx. 

The fourth slab of Lias adding to our means of reconstruction of Dimorphodon, was 
observed by the Earl of Enniskillen in the collection of Henry Marder, Esq., M.R.C.S., of 
Lyme Regis. It had been quarried from the same cliff as the preceding specimen 
(Pl. XVIID, and displayed the vertebrae and bone-tendons of a long and stiff tail 
(Pl. XIX, fig. 4). 

Indications of such a tail, in which the vertebree were associated with ossified tendons, 
were apparent, and have been noted in the description in the second specimen with the 
skull (Pl. XVIII, cp); whereby one was able to show that the vertebree in the originally 
described specimen supposed to be cervical (Buckland, loc. cit., pl. xxvii, a, a’) were truly 
caudal, with similarly associated bone-tendons, as, indeed, Von Meyer had recognised 
after the discovery of the caudal structure of his Ramphorhynchus.? The specimen 
now to be described of the entire tail, as represented by its petrifiable parts (Pl. XIX, fig. 
4)? I conclude, from the identity of character of some of its vertebrae with the three 
shown in Pl. XVIII, ¢ d’, and from the discovery of this specimen in the same formation 
and locality, to belong to Dimorphodon macronyz. 

The series of caudal vertebree, to judge from the size of the anterior ones, comes from an 
individual as large as that represented by the fossils in Pls. XVII and XVIII, and, no donbt, 
from an adult or full grown one. This series is 1 foot 9 inches in length, following the curve, 


1 “Cuvier, Wagler, und Goldfuss lassen den Fuss aus fiinf ausgebildeten Zehen bestehen ; in allen 
Pterodactyln habe ich aber nie mehr als vier solehen Zeben und héchstens noch einen Stummel vorgefunden.” 
Von Meyer, op. cit., p. 20. But see ‘Ossemens fossiles,’ 4to, tom. v, pt. ii, p. 374—“ Le cinquiéme réduit 
ai un léger vestige,” &c. 

2 « Beitrage zur naheren Kenntniss fossiler Reptilien,” in Leonhard und Bronn’s ‘ Neues Jahrbuch fiir 
Mineralogie,’ &c., 8vo, 1857, p. 536. 

3 Tt has been drawn with the neural aspect downward. 


LIASSIC FORMATIONS. 55 


which is single and slight ; and it includes upwards of thirty vertebrae. These vertebra, 3 
lines in length of centrum in the first five, progressively increase to a length of 1 inch at 
the twelfth, begin to shorten gradually after the fifteenth, the twenty-first being 11 lines, 
the twenty-fourth 9 lines, the twenty-eighth 6 lines, and the thirtieth 5 lines in length. In 
breadth or thickness the vertebrae decrease from the first to the tenth; and then again 
gradually from the fifteenth to the last, which is filiform. 

The first caudal, or the first of the series here preserved, has the anterior articular 
surface of the centrum subconcave. The inferior surface describes a slight concavity 
lengthwise; the upper part of the anterior half projects as a parapophysis, the end of 
which has been broken off, showing the open cancellous structure. A ridge from its upper 
part was continued to the fore part of the anchylosed neural arch. This arch developed 
zygapophyses, of which the anterior extend beyond the centrum; but they are better 
shown in succeeding vertebra. 

In the second caudal the base of the parapophysis has receded and now projects from 
the upper part of the side of the centrum, occupying more than its middle third. Part 
of a quadrate spinous process is here preserved, projecting above the centrum as far as 
the vertical diameter of that element. 

In the third caudal the base of the parapophysis, reduced in vertical thickness, 
occupies the same positions and longitudinal extent. The postzygapophysis, after a deep 
hind notch of the neurapophysis, curves over the prezygapophysis of ihe succeeding 
vertebra, which enters that notch. 

In the fourth caudal the base of the parapophysis has lost in longitudinal as well as 
vertical extent, and is more posterior in position. ‘The subconvexity of the hind articu- 
lation of the centrum is here well shown. The confluent neural arch is low, attached to 
rather more than the fore half of the centrum. The postzygapophysis does not extend 
back beyond the centrum ; the prezygapophysis is continued beyond the front or concave 
surface of the centrum into the neural notch of the preceding vertebra. 

In the fifth caudal the parapophysis is smaller and more posterior. The neurapo- 
physis rises from the anterior half of the side of the centrum and continues to show the 

zygapophysis, though reduced in size. Between the fifth and sixth caudals a small, 
slender hamapophysis (4) has been articulated to the under part of the intervertebral 
space. 

The reduced parapophysis is continued from the sixth caudal; this vertebra shows a 
much reduced indication of neurapophysis. The base of a hemapophysis crosses the lower 
part of the space or joint between it and the seventh caudal, then expands both forward 
and backward, and more so in the latter direction; the inferior border of this expansion 
is straight. 

In the seventh caudal the prezygapophysis is still indicated, though much reduced in 
size. The hemapophysis, similar in shape to the preceding one, is longer ; and three bone- 
tendons rise from the side of the hind projection of this hemal arch. 


56 FOSSIL REPTILIA OF THE 


In the eighth caudal the base of a reduced parapophysis projects from the side of the 
centrum behind its middle; a low prezygapophysis projects from the neural arch: but 
beyond this vertebra all trace of that arch disappears, or is indicated by fecble prominences 
in the fasciculus of bone-tendons which seem to be attached to neural processes of the 
non-elongated centrums. Six or seven filamentary bone-tendons, one thicker than the 
rest, extend lengthwise above the centrum. Some of these may be traced over two 
centrums, then end in a point, their place being taken by another bone-tendon beginning 
by a similar pointed end. The parapophysis disappears in the tenth vertebra. 

The caudal vertebree in the first discovered specimen of Dimorphodon* answer to the 
eighth—eleventh in the present series. The elongate centrums of the tenth and succeeding 
caudals, usually more or less uncovered by the bone-tendons, show a low lateral ridge, 
and a slight expansion at the ends. The haemapophyses are traceable, much reduced in 
size, to the fifteenth—sixteenth vertebra. 'The bone-tendons are in two fasciculi, one 
neural, the other heemal, in position. rom five to eight may be counted in the side view 
given of each of these fasciculi. The seeming increase of thickness of some, usually the 
more peripheral of the filaments, may be due to this flattened form, and to more or less 
of the side coming into view, instead of the edge. Five or six may be counted in each 
fascicule, even beyond the twentieth caudal; the number varying at parts through the 
formation of the bundle by successive tendons, as above mentioned. ‘They are reduced to 
two or three at the thirtieth vertebra. ‘The terminal joints of the elongate centrums 
appear to be flattened and closely adapted, allowing of very little motion. It is evident 
that, as in Ramphorhynchus, the tail was stiff as well as long, and doubtless served as a 
sustaining ray of the parachute of membrane continued backwards from the wings and 
hind limbs. 

The vertical diameter of the second caudal showing its neural spine is five lines. 
The diameter of the ninth vertebra, including the neural and heemal fasciculi of bone- 
tendons, is the same; and beyond this the vertebrae and their surroundings gradually 
diminish to the pointed end of the tail. 


§ Restoration or DimorrpHopon. Plate XX. 


The several parts of the skeleton of Dzmorphodon preserved in the slabs of Lias 
described and referred to in the foregoing pages have ultimately yielded the desired result 
of their scrutiny and comparison, viz., a restoration of the extinct animal, such as I have 
endeavoured to exemplify in Plate XX ; and I propose to apply that plate in illustration of 
a summary of the osteology and dentition of Dimorphodon, comparing therewith the 


1 Buckland, loc. cit., pl. 29, a, a. I have had these vertebra carefully redrawn, from the specimen, 
in Pl. XIX, fig. 3. 


LIASSIC FORMATIONS. 


or 
~ 


previously known Péerosauria, and adding such deductions as to the status and affinities 
of the order as seem legitimately to flow from the facts. 

The first distinguishing feature of Dimorphodon, or of the present liassic type of the 
genus, is the disproportionate magnitude of the head—the more strangely dispro- 
portionate, as it seems, in an animal of flight. 

The head is large in proportion to the trunk, not only in respect of length but of 
depth, and probably, also, breadth; nevertheless, the shape and disposition of the con- 
stituent bones are such that, perhaps, no other known skull of a vertebrate is constructed 
with more economy of material—with an arrangement and connection of bones more 
completely adapted to combine lightness with strength. 

So far as the skulls of Pferosauria have been sufficiently entire to show the shape of 
the head, no other known species resembles Dimorphodon. The cranial part is singularly 
small: the rest is mainly devoted to the formation of the large, long, and powerful 
prehensile and manducatory jaws. Among the débris of the cranial bones, in specimens 
Pls. XVII and XVIII, the mastoid (g), parts of the occipital (paroccipital, 4), the parietal (7), 
post-frontal (12), frontal (11), prefrontal (14), and nasal (15), are recognisable: the last two 
bones, however, are concerned more with the scaffolding or buttressing of the upper jaw 
than with the protection of the brain or formation of its ease. ‘Though contributing their 
shares to the otccrane, the chief developments of the paroccipital (Pl. XX, 4) and mastoid 
(ib. 8) relate to the muscular connections of the head with the trunk: the mastoid joins the 
postfrontal to form an upper zygoma, giving origin to part of the temporal muscles ; it also 
affords a fixed articulation to the tympanic, and sends down a pointed process external to the 
masto-tympanic articulation. ‘The parietals (Pl. XX, 7), confluent at the mid line, where they 
develop a low crest, swell out slightly at the temporal fossa, indicative of the size and saurian 
position of the mesencephalon. ‘The frontal (11) is narrow and flat between the orbits, of 
which it contributes most of the upper part of the rim. This is continued by the postfrontal 
(12) behind, which sends down a long pointed process to unite with the malar (26), and a 
shorter and thicker one to join the mastoid (g). ‘The prefrontal (14), of a triangular form, 
contributes to the upper and fore part of the orbit, and, either directly or by a connate 
lacrymal, unites with the ascending malo-maxillary process (21, 26), and the base of the pre- 
frontal articulates with the frontal and the nasal. The nasals (15), to the usual con- 
nections with the frontal, prefrontal, and medial process of the premaxillary (22'), superadd 
a union with the lateral ascending process of the maxillary (21x), completing the bar 
between the nostril (z) and the antorbital vacuity (a). The nasal bone forms the upper part 
of the nostril; the rest of the boundary of that singularly wide aperture is formed by the 
premaxillary and maxillary. Of the basis cranii and palate there do not appear to be 
any recognisable parts preserved. ‘The maxillary is overlapped by the hind alveolar part 
of the premaxillary, and unites therewith by a long oblique suture (21”). ‘The maxil- 
lary, receding, expands and sends upward a long slender pointed process to articulate 


8 


58 FOSSIL REPTILIA OF THE 


with the nasal; it then joins the malar and the prefronto-lacrymal, and descends internal 
to the mandible to join the palatine.’ Each maxillary (21, 21’) affords alveoli for eight or 
nine teeth. 

The premaxillary is the largest of the bones of the head. The pair, by confluence or 
connation, constitute the fore part of the upper jaw (22), expanding from a sub-obtuse 
apex as it recedes, and preserving its entireness for an extent of about two inches. This 
tract seems to be arched above transversely, with a slightly convex upper longitudinal 
contour continued along the medial ray or process (22’). Of the configuration of the 
palatal surface the specimens give no evidence. From the analogy of Pteroductylus 
Cuviert and Pt. Sedgwickii,” we may infer that this (premaxillary) part of the 
bony roof of the mouth was entire, and strengthened by a median ridge. The lateral or 
alveolar borders formed alveoli for four teeth on each side. Thus the hind expansion 
of the premaxillary divides into three rays or processes. ‘The upper medial or nasal ray 
is the longest: it is continued backward, continuing the initial curve of the upper 
contour of the face as far as the nasals, the mid suture or confluence of which bones it 
overlaps, and joins suturally to an extent precluding any movement of the upper jaw on 
that part of the head. The length of this ray is about 3} inches. The pair of lower or 
alveolar rays extend back for about 14 inches. 

The malar (26) forms the lower narrower end of the oval orbit, sending up one pointed 
process (united with that of the maxillary ?) toward the prefrontal, and a longer and stronger 
one to join the postfrontal. ‘The squamosal (27) continues the zygomatic bar backward to 
abut against the tympanic. Its precise position and direction are left doubtful in the 
specimens hitherto obtained, but it is unquestionably present, and contributes to the fixa- 
tion of the tympanic. 

This (28) isa moderately long and strong pedicle, immovably articulated to the mastoid, 
paroccipital, and squamosal ; thickest posteriorly, where it is strengthened by an 
outer marginal ridge, sending forward and inward a process which may articulate with 
the pterygoid (but of this I could not get clear evidence), expanding at its distal end to 
receive the abutment of the squamosal or lower zygoma, and to form the convex condyle 
for the articular element of the mandible. 

The dentary parts of the mandible are confluent at the symphysis, which is as long as 
the undivided fore part of the premaxillary. The ramal part of the dentary is compressed, 
and gains a depth of about 10 lines before it bifurcates. The alveolar border of the dentary 
extends as far as that of the maxillary, viz. about 5 inches, beyond which the upper prong 
(Pl. XX, 32’) is continued above the mandibular vacuity, underlapping the surangular (29) 
and terminating ina point. The lower prong (ib. 32”) terminates in a point before attaining 
the vacuity ; it is underlapped by the fore part of the angular (30), with which it articulates. 


1 This description is on a homological hypothesis, subsequently discussed (p. 64). 
* «Monograph on Cretaceous Pterosauria,’ Suppl. 1, 4to, 1859, Pl. I, fig. 1, 6. 


LIASSIC FORMATIONS. 59 


The divergence of the hinder prongs of the dentary exposes a small part of the splenial 
(31). The vacuity, if it be natural and not due to abrasion of a thin outer wall, is a long 
and narrow oval, 1 inch 8 lines in length, 6 lines in breadth. It is circumscribed behind 
by the confluent angular and surangular elements (29). ‘The angular (30) forms a slight 
projection behind the articular concavity ; it expands vertically, and contracts transversely 
as it advances, contributing a small share to the lower border of the vacuity, and con- 
tracting to a point below the dentary, about 5 inches from the angular process. 

The range of variety shown by the skull is considerable in the order Péerosauria. In 
relative size, as in the expanse of the antorbital vacuity, P¢erodactylus crassirostris ‘ 
comes nearest to Dimorphodon ; but the orbit is relatively larger, and the nostril much 
smaller. In Rhamphorhynchus Gemmingi the nostril and antorbital vacuity are of equal size, 
and each is about one eighth the size of the orbit, which is proportionally larger than in 
Dimorphodon. In Pterodactylus longirostris” the nostril is larger than the orbit; the 
antorbital vacuity is. not half the size of the orbit. In P¢erodactylus suevicus* the antorbital 
vacuity is still smaller. In Pterodactylus Kochii* that vacuity is limited, as in Chlamydo- 
saurus, to the upper part of the boundary between the large orbit and the long and large 
nostril. In Pterodactylus longicollum’ it appears to be wanting. 

The shape of the skull offers many modifications in the several species, from the long 
and slender type of that of Pterodactylus scolopaciceps and Pt. longirostris to the shorter 
and deeper cone indicated by P¢. conirostris,® and to the inflated and more or less 
anteriorly obtuse form exhibited by Dimorphodon and the more gigantic Pterodactylus 
simus." 

The position of the tympanic pedicle varies from the almost vertical one in Dimorpho- 
don to the almost horizontal one in Pterodactylus longirostris and Pt. Kochit. In Pt. 
crassirostris it shows an intermediate slope or position. 

The mandible, conforming in relative depth and length to the general shape of the 
skull, has the symphysis longest in those species with long and slender jaws. In 
Pterodactylus suevicus the symphysis extends along the anterior third part of the mandible. 
In Pt. crassirostris it is shorter, and still shorter in Dimorphodon. he depth of the rami 
decreases behind the dentigerous part in Plerodactylus longirostris. 

The generic dental character of Dimorphodon has been given in detail in the special 
descriptions of the specimens figured in Pls. XVII and XVIII. The range of variety mani- 


1 «Monograph on Fossil Reptilia of the Cretaceous Formations’ (1851), Péerosauria, Pl. XXVII, 
figs. 2—4. 

# Ibi, ibry tips be 3 QUENSTEDT, op. cit. 

4 Von MEYER, op. cit., tab. i, fig. 2. 5 Ib., ib., tab. vii, figs. 1—4. 

® Dixon’s ‘Geology and Fossils of the Tertiary and Cretaceous Formations of Sussex, dto, 1846, 
Pl. 38. 


7 “Monograph on Fossil Reptilia of the Cretaceous Formations’ (Péferosauria), Suppl. No. 3 (1861), 
Pl. I, figs. 1—3. 


60 FOSSIL REPTILIA OF THE 


fested in this character is considerable in the present order, although in no species has any 
departure been observed from the predatory zoophagous condition. The teeth, always 
simple and pointed, vary in shape, in number, in position, in relative size. Pterodactylus 
crassirostris exemplifies the laniariform type of teeth, more or less elongate, and separated 
by intervals of varying extent. In this not uncommon condition the teeth are longest in 
the upper jaw, as offering more resistance than does the lower jaw in aid of the weapons 
most deeply implanted in the struggling prey. 

In Pterodactylus longirostris the teeth are rather small, subequal, with short intervals, 
a little widening toward the hind end of the series, which is restricted to the anterior half 
of the jaw, both above and below. 

In some Pferosauria a certain extent of the fore part of both upper and under jaws is 
edentulous, and from its shape has been inferred to have supported a horny sheath. The 
teeth are long slender canines, with wide intervals. They number from about 8 to 10 
on each side of the upper jaw, and from 7 to 8 in each ramus of the mandible. Von 
Mryer proposed for this modification of mouth the generic name Rhamphorhynchus. 

Dimorphodon shows the combination of scattered laniaries, with small, more closely 
set serial teeth in the lower jaw; it has more numerous teeth, occupying a greater extent 
of the alveolar margins of the jaws, than in any other Pterosaurian. 

The very small teeth which have been observed in the short jaws of the little Ptero- 
dactylus brevirostris' are most probably characters of immaturity, not of species. 

In regard to the bony structure of the head and the dentition, the general result of 
observation and comparison of Pterosaurian fossils, and common consent of competent 
investigators, having excluded the volant Mammals from the claim of affinity, the question 
becomes narrowed to whether the skull in Péerosauria more resembles that in the cold- 
blooded or the warm-blooded oviparous air-breathing Vertebrates. 

Hzrmann von Meyer, who has contributed a great and valuable share to our know- 
ledge of the Pterosaurian order,” quoting Oken’s opinion, “that the skull is intermediate 
in character between that of the Chameleon and Crocodile,” sums up his own conclusions 
on that head in the following terms :—“'The skull of P/erodactylus is essentially comparable 
only with that of Birds and Saurians. The preponderating resemblance with the Bird’s 
skull cannot be contested. Against this, however, is a remarkable dissimilarity in certain 
parts which, on the other hand, approximates it to the type of Saurians.” ° 

The term Sauria is here used in the sense of Broneniart and Cuvisr, and it is open 


1 Gotpruss, loc. cit., tab. x, fig. 2. 

2 Especially in the admirable summary of his own and others’ researches, in the part of his great work, 
‘Zur Fauna der Vorwelt’ relating to “ Reptilien aus dem lithographischen Schiefer,” &c., fol., 1860. 

3 «Der Schadel der Pterodactyln, der nach Oken zwischen Chamileon und Crocodil stehen wiirde, 
lasst sich eigentlich nur mit den Végeln und den Sauriern vergleichen ; die iberwiegende Aehnlichkeit mit 
dem Vogelkopfe kann nicht bestritten werden; ihr gegeniiber steht aber eine auffallende Unahnlichkeit in 
gewissen Theilen, die dafiir zum Zypus der Saurier hinneigen.”—Op. cit., p. 15. 


LIASSIC FORMATIONS. 61 


to the unbiassed investigator, and, indeed, becomes plainly his business, to determine, not 
merely whether Avian or Saurian characters predominate in the Pterosaurian skull, but to 
define the degree of affinity or correspondence of cranial structure therein traceable to 
such structures in Lnaliosauria, Dinosauria, Dicynodontia, Crocodilia, Lacertilia, each of 
which may be a group, organically, of co-ordinate value with Aves. 

Greater respect to the memory of so unbiassed a seeker after truth cannot be shown 
than by weighing with due care and what judgment one may be able to bring to the task 
the value and significance of each well-determined evidence of the cranial structure which 
Von Meyer has described and reasoned upon. 

It is to be regretted that not in any of the numerous figures of the skull of Péero- 
sauria, original or copied, has Von Meyer indicated the bones which he describes. When 
he writes—“ ‘The temporal bone lies external to the parietal and principal frontal bones, 
one much wishes he had indicated his ‘ Schlafen- 


27] 


and mainly forms the temporal fossa, 
bein’ in the skull of Rhamphorhynchus Gemmingi, pl. ii, fig. 4; pl. ix; pl. x, fig. 1; 
or in the more instructive example of cranial structure which he has borrowed from 
Goldfuss for the subject of his pl. v (Péerodactylus crassirostris). 

By ‘ Schlafenbein’ Von Mryrer may mean that element of the compound ‘temporal 
bone’ of anthropotomy which I have called ‘squamosal.’ No doubt in Man and 
most Mammals the squamosal does contribute a notable share to the formation of the 
temporal fossa, whence the name ‘temporal’ given to the incongruous group of cranial 
elements coalescing in such warm-blooded Vertebrates with the squamosal, so exceptionally 
expanded in the Mammalia. But as to the value of the bed of the temporal muscles in 
determining the homology of the bones forming it, I would refer to the remarks in my 
work on the ‘ Homologies of the Vertebrate Skeleton.’ ’ 

Some clue to the bone signified by Von Mryer may be got from the following remarks— 
“ Anteriorly it seems not to take, as in Birds, a share in the formation of the orbital rim ; 
here, much more as in Saurians, it is pushed aside or supplanted by the postfrontal.” * 

The term ‘temporal bone’ (Schlifenbem) has been used in various senses, but 
whether it be applied to that element which I, with Cuvier, call ‘mastoid’ in Repfilia, 
or to that which others,* with Cuvier, call ‘temporal’ (meaning squamosal) in Birds, 
there is no bone that Von Muyur can be supposed to mean by ‘Schlafenbein’ which 
forms any part of the rim of the orbit in Birds. 

Von Meyer recognises a ‘ postfrontal’ (‘ Hinterstirnbein ’) in P/erosauria, and states 
that it pushes away his temporal (Schlifenbein) from the orbit. In Péerosauria the post- 


1 « Das Schlifenbein liegt aussen an dem Scheitelbein und Hauptstirnbein, und bildet hauptsachlich 
die Schlifengrube.”’—Op. cit., p. 15. 

2 8vo, 1848, p. 33. 

8 “Vorn scheint es nicht wie in den Végeln an der Bildung des Augenhohlenrandes Theil zu nelmen, 
hier vielmehr wie in den Sauriern durch das Hinterstirnbein verdringt zu werden.’’—Op. cit. p. 19. 

4 Hatiman, “ Die vergleichende Osteologie des Schlifenbeins,” p. 8, pl. 1. 


62 FOSSIL REPTILIA OF THE 


frontal (PI. XX, 12) is undoubtedly interposed between the bone I determine as ‘ mastoid ’ 
(ib. 8) and the orbit (ib. 0) ; and my ‘mastoid’ in Péferosauria answers to Cuvier’s and 
Hallman’s ‘temporal, 7z.e. squamosal, in Birds. We may conclude, therefore, that Von 
Meyer's ‘Schlafenbein’ in Péerosauria is that marked 8 in the skull of Pterodactylus 
crassirostris.’ 

Certain it is that no bone answering to 8 in Pls. XVII, XVIII, XX of the present 
Monograph contributes to the formation of the orbitin any Bird. In the great majority of 
that class, as is well known, the rim of the orbit is mcomplete below; it is formed above 
by the frontal, before by the prefrontal and lacrymal (‘ antorbital’ of ornithotomists), 
behind by the postfrontal (‘ postorbital,’ ib.). Where, as in some Psittacide,’ the orbital 
rim (‘ Augenhohlenrandes ’) is complete, the lower complement is formed by an extension 
of ossification from the antorbital to the postorbital processes, independently of either 
Cuvier’s temporal (8) or my squamosal (27) in Birds. 

I confess that the foregoing result of the analysis of a main ground of Von Meyzr’s 
assertion as to the “ incontestable similarity between the Pterosaurian and Avian types of 
cranial structure ” has not a little tended to shake my confidence in the grounds on which 
he has pronounced definite judgment on the matter. So far as we have yet got evidence 
of the structure of the skull in Péerosauria, it seems that, contrary to the rule in Birds, 
the orbital rim is entire; and that its lower border is completed by the zygomatic arch, and 
chiefly, if not exclusively, by the malar element; whereas, such arch passes freely beneath 
the orbital rim in the few Birds with that rim entire. Now, in this part of the cranial 
structure the Pterosauria agree with the Crocodilia: as in them the malar (26) sends up a 
process to unite with one descending from the postfrontal (12) to complete the orbital 
rim behind. 

In the small species of Pterodactyles (P¢. longirostris, Pt. scolopaciceps, and in the 
perhaps immature animal represented by Pt. drevirostris) the hind convexity of the cranial 
wall is not marked by the apophysiary developments of paroccipital and mastoid, and 
accordingly resembles that part of the cranium in Birds, especially the smaller Galle ; but 
before this similarity of shape can be pressed into the argument for the Avian affinity of 
the Plerosauria, it should be shown to be common to or constant in the extinct volant 
order. 

But this is far from being the case. When a Pterosaur has gained the size of 
Pterodactylus crassirostris*® or Pter. suevicus,' the back of the skull shows no cerebral 
swelling, but only the crests and processes for muscular attachments, as in other Reptilia 


1 *Monograph on Fossil Reptilia of the Cretaceous Formations’ (Pterosauria) (1851), Pl. XXVII, 
figs. 3 and 4. 

* «On the Archetype and Homologies of the Vertebrate Skeleton,’ 8vo, 1848, pl. i, fig. 1 (Calypto- 
rhynchus) ; ‘Anatomy of Vertebrates,’ 8vo, vol. ii (1866), p. 51, fig. 30 (Psittacus), also p. 63. 

3 Goldfuss, op. cit., pl. vii. 

* Quenstedt, op. cit. 


LIASSIC FORMATIONS. 63 


of similar size. Even in Rhamphorhynchus Gemmingi the cranial convexity is not posterior, but 
is limited to the temporal fossee behind the orbit, as in the specimen figured by Von Meyer 
in pl. ix, op. cit.; and this indication of the optic lobes is less conspicuous in the subject of 
pl. x, fig. 1. In Démorphodon there is still less trace of this alleged Avian characteristic. 

The bone which, in the Bird, as in the Pterosaur, forms part of the otocrane, articulates 
with the ex- and par-occipitals behind, with the alisphenoid in front, with the parietal above, 
and with the petrosal within, which contributes the articular surface to the tympanic and 
the upper rim to the meatus auditorius, also articulates in the Pterosaur, as in the Crocodile, 
with the postfrontal : and this character appears to be constant in the P/erosauria as in the 
Crocodilia, while it is exceptional in Aves. In the particulars in which the bone 8 differs 
in the Péerosaurian from that in the Bird, it agrees with 8 in Crocodilia ; as e.g. in its high 
position in the cranium, owing to the low development of the cranial chamber ; its 
greater degree of projection from the true cranial walls ; the extensive and suturally fixed 
character of its articulation with the tympanic as compared with the more definite and 
restricted glenoidal movable articulation which the mastoid (8) affords to 28 in Birds. In 
all these circumstances, whether the bone 8 (PI. XX, fig. 1) be called mastoid or squamosal, 
it is Reptilian, not Avian, in the Pterosaur. 

Herr Von Meyer states, in another of his comparisons, that in the Monitor, Iguana, and 
Stellio, the prefrontal (‘ Vorderstirnbein ’) enters into the formation of the periphery of the 
external nostril (Nasenloch).’ This is the case with Varanus,’? not with true Monitors.’ 
In Zejus nigropunctatus some extent of the suture between the nasal and the maxillary 
intervenes between the prefrontal and the nostril. ‘The non-extension of the prefrontal to 
the external nostril shows no Avian affinity in Pferosauria; rather an agreement with the 
majority of Reptilia, as, for example, with the whole order of Crocodilia. 

In some Crocodilia (Teleosaurus) and Lacertilia (Chlamydosaurus, Lyriocephatus) 
there is an antorbital vacuity, which, in the latter Lizard (Pl. XX, fig. 3, a), 1s equal in size 
with the nostril (ib., z) and intermediate in position between that cavity and the orbit (ib., 0), 
which is large. A process of the maxillary rises obliquely backward to join the nasal, and 
to separate the intermediate vacuity from the external nostril. The lacrymal and pre- 
frontal form the bar dividing the intermediate cavities from the orbit. In most Birds a 
small intermediate vacuity is partitioned off from the nostril by a process of the maxillary 
rising to join the nasal, and is similarly separated from the orbit by the lacrymal, which 
descends to join the malar. The great range of variety in the development of this 
‘intermediate ’ or ‘ antorbital vacuity,’ in Péerosauria, has already been pointed out; but 


' «Zur Fauna der Vorwelt,’ fol., 1860, p. 16. 

2 See Cuvier, ‘Ossemens fossiles,’ v, pt. 2, pl. xvi, fig. 1 (‘grand Monitor du Nil, Lacerta nilotica’), 
p. 259, the Varanus Dracena of Merrem, Varanus niloticus of most modern erpetologists ; also in pl. xvi, 
fig. 7, ‘ Monitor du Java,’ p. 260; the Varanus bivittatus, of Merrem. 

3 As e.g. Tupinambis teguixin, ‘Sauve-garde d’Amérique.’ Cuvier, vol. cit., pl. xvi, figs. 10, 11, and 
Thorictes Dracena, ib., figs. 12, 13; ‘La Dragone,’ ib., p. 263. 


64 FOSSIL REPTILIA OF THE 


the comparable structure is by no means peculiar, as Von Meyer would lead one to infer, 
to the skulls of Birds." 

In no Pterosaurian has any obvious and unmistakeable suture been seen indicative of 
the respective shares taken by maxillary (21) and premaxillary (22) in the formation of the 
dentigerous part of the upper jaw: both bones combine to support the array of teeth; they 
have coalesced, at least at their external or faci-alveolar plates ; as, likewise, have the right 
and left premaxillary portions forming the fore end of the upper jaw. The suture between 
this premaxillo-maxillary bone and the suborbital portion of the zygomatic arch remains. 
Accordingly, there is a choice of analogies in the interpretation of the observed facts: a 
proportion of the compound bone may be assigned to the premaxillary, according to the 
analogy of the Crocodile and Lizard; or the whole may be called premaxillary, according 
to the analogy of the Ichthyosaur. 

Goxpruss, guided by the Lacertian analogy, limits the premaxillary to the anterior part 
of the upper jaw, and to the upper part of the external bony nostril (z) ; and he illustrates 
this view by a dotted line representing the assumed suture in his restoration of Ptero- 
dactylus crassirostris, in pl. ix (op. cit.).” Von Meyer assumes, as arbitrarily, the Ichthyo- 
saurian analogy, but views it as a specially Avian one, and ascribes to the Plerosauria a 
bird-like premaxillary,® and this determination is indicated by the numerals on the restora- 
tion of the skull of Pterodactylus compressirostris in my Monograph of 1851, quoted below, 
Pl. XXVII, fig. 5. 

Of the maxillary bone (my 21) Von Meyer merely remarks that ‘it does not follow the 
type of Birds” (“folgen nicht dem Typus der Vogel,” ib., p. 15). And yet, if the Pterosau- 
rian premaxillary be interpreted according to that type, forming so large a proportion of the 
upper jaw as to include all the teeth, the edentulous maxillary must have had a correspond- 
ingly Avian proportion and position. Only, whereas in most Birds the small and slender 
maxillary sends up a process helping to define the back part of the nostril and fore part of 
the antorbital vacuity, the corresponding process in Plerosawria would be (as indicated in 
my Pl. XVIII, 22°), part of the premaxillary. 

I incline to believe, however, that it may prove to belong to the maxillary ; that 
the dentigerous part of the upper jaw is due, in Pferosauria, to the combined maxil- 
laries and premaxillaries, but that the latter take a larger share in the formation of 
the alveolar tract than Goxpruss conjectures. One ground of such opinion is this: 
the portion of upper jaw with six pairs of laniary teeth in the huge Pterodactylus 
Sedgwickii, in which the palatal surface could be clearly worked out,* showed that the 
anterior expansion, with the group of three pairs of teeth, could hardly have been 


1 ««Zwischen Nasenloch und Augenhohle liegt eine dritte Oeffuung, die wiederum an den Vogel- 
schidel erinnert.””—Op. cit., p. 16. 

* Copied in Pl. XXVII, fig. 4, of my Monograph above cited of 1851. 

8 “Vin Vogeln-aihnlichen Zwischenkiefers,”’ v, p. 15, op. cit. 

4 Monograph, Suppl. No. 1 (1859), Pl. I, figs. 1, a, 6. 


LIASSIC FORMATIONS. 65 


separated by a suture, at the slight constriction suggesting that structure in Pé. crassi- 
rostris, without leaving some indication of its original existence, especially on the palate. 

In the anterior confluence of right and left premaxillaries, and the backward produc- 
tion from their upper part of a bony bar uniting with the nasals and dividing the nostrils, 
we have a character of the Dicynodonts and of some Lacertians (Varanus) as well as of 
Birds, and the Saurian affinity is shown to be the truer one by the firmness of the naso-pre- 
maxillary union and the absence of any power of, or provision for, that hinge-like movement 
of the upper mandible upon the cranium which is peculiar to, though not constant in, the 
Avian class. Moreover, the outer surface of the premaxillary shows none of that spongy 
porosity and rugosity which relates to the sheath or horny covering of the beak character- 
istic of the Bird. Such structure has not even been detected in the feeble trace of eden- 
tulous anterior production of the upper jaw in Rhamphorhynchus, Von Meyer. I cannot, 
therefore, see, with Von Meyer, the beak of the Bird in an animal with a fixed and toothed 
upper jaw ;” for on every hypothesis of its bony structure it finds a closer resemblance 
among the toothed Reptiles than in the class of Birds. 

The mandible, or lower jaw, is supported, as in all Vertebrates below Mammals, by the 
tympanic, viz. the bone (28, Pls. XVIII and XX) which is shown by its osseous connec- 
tions, its relations to the ‘ facial nerve,’* or its equivalent the ‘ramus opercularis,’* and 
by its mode of formation, to answer to that which in Mammals is mainly reduced to the 
function of supporting the ear-drum. In air-breathing Ovipara it superadds this function 
to its more constant and essential use in non-mammalian Vertebrates, of supporting the 
lower jaw. 

In reference to the question of affinity before us, the tympanic gives valuable evidence by 
reason of the moveable articulation and peculiar connections with the upper mandible 
essentially correlated to a covering of feathers. In Péerosauria the tympanic at its 
proximal end resembles that of Lizards by its fixed sutural mode of union with the 
cranium, and it furthermore resembles that in Crocodiles by the abutment of the zygoma 
against its distal end, to which it is suturally attached. 

In Birds the tympanic enjoys a synovial moveable articulation by a single or double 
condyle at its proximal or cranial end, and presents a synovial cavity to a condyloid con- 
vexity of the hind part of the zygoma. By this test, therefore, the Péerosauria 
are shown to be not only ‘Saurian,’ but to be nearest akin to the existing orders 
which possess double-jointed ribs and the correlated cardiac structure. ‘The difference 
of shape between the tympanic of the Pterodactyle and that of the Bird is too strongly 
marked not to have attracted attention; but I do not find in that of the Chameleon the 


1 Goldfuss, loc. cit. 


2 “Wir sehen also hier die Schnautze der Vogel auf ein Thier mit unbeweglicher und mit Zilnen 
bewaffneten Schnautze angewendet.’’—Op. cit., p. 15. 

3 * Anatomy of Vertebrates,’ vol. ii, 8vo, 1866, p. 124, vol. ili, p. 155. 

* Ib., volii, p: 303. 


wy) 


66 FOSSIL REPTILIA OF THE 


most resemblance to the Pterosaurian tympanic.’ For, besides the Lacertian freedom of 
the bone from zygomatic abutment, the tympanic in the Chameleon has not the 
longitudinal strengthening ridges, nor the process turned toward the pterygoid. 

The dentigerous mandible, like the maxilla, speaks for the Reptilian affinity of 
Pterosauria ; the distinct sockets for the teeth ally them to the higher forms of Sawria. 
In reference to the generic modification of dentition in Dimorphodon, it has been remarked 
that this early form of flying dragon seemed to have derived one feature or modification 
from the Fish, and the other from the Crocodile or Plesiosaur.2 

The length of the neck, which is not always equal to that of the head, is due, in 
Pterosauria, rather to the length than the number of the vertebrae. Counting the axis 
with the small coalesced atlas * as one, I give seven cervical vertebrae to the Dimorphodon 
macronye (Pl. XX, fig. 1, c). Of these a series of four are preserved in the specimen (PI. 
XVIII, c), showing, as described, the characteristics of the Pterosaurian cervical vertebra 
which had been determined and illustrated in a former Monograph.* 

Cuvier,’ in his searching analysis of the evidence at his command of the osseous struc- 
ture of the Pterodactylus longirostris, concluded that the cervical vertebrae were not fewer 
than seven, as in Crocodilia and Mammalia, or not more than eight, as in Chelonia. 

Go.pruss was able to demonstrate the vertebral formula in his famous specimen of 
Pterodactylus crassirostris.© The number, ‘ seven,’ was, however, obtained by reckoning 
the atlas distinct from the axis, and the last cervical may have been relegated to the 
dorsal series. 

QueENstED'’ shows seven cervicals in his instructive example of Pferodactylus suevicus, 
reckoning the atlas and axis as one vertebra; and this analogy I have followed im the 
restoration of Dimorphodon. 

Rhamphorhynchus Gemmingi has six cervicals, counting the coalesced atlas and axis as 
one ; but in the specimen figured by Von Meyer in his pl. ix,* there seems to be the centrum 
of a short ‘seventh’ cervical between the longer ‘sixth’ and the first (dorsal) vertebra 
supporting a long free pointed rib. It is certain that the number of cervicals does not 
exceed the latter reckoning or fall short of the first. Thus it is plain that the Pferosauria 
exemplify the Crocodilian affinity in the cervical region of the vertebral column. Lacer- 

1 “Dieser Knochen ist nicht wie in den Vogeln quadratisch, sondern cylindrisch stielformig 
beschaffem.—Hierin, so wie in einingen andern Theilen, zeigt das Thier die meiste Aehnlichkeit mit Cha- 
maeleon.”’—Von Meyer, op. cit., p. 16. 

2 «Report (Sections) of the British Association for the Advancement of Science,’ 8vo, 1858, p. 98. 

3 «Monograph on the Fossil Reptilia of the Cretaceous Formations,’ Supplement, No. I, Pterosauria 
(1859), pp. 7—8, Pl. I, figs. 11—14. 

4 Tb., p. 9—11, Pl. II. ‘Monograph,’ &c., Supplement, No. III (1860), p. 7, Pl. II, figs. 1, 
2 and 4. 

5 “Ossemens fossiles,’ tom. cit., p. 367. 

6 « Man zihlt 7 Halswirbel, 15 Rippenwirbel, 2 Lenden, and 2 Kreuzbeinwirbel,” loc. cit., p. 79. 

7 Op. cit., figs. 1—7. 

85Op, cit. 


LIASSIC FORMATIONS. 67 


tians have fewer definite cervicals ; Birds have more. I have not seen any Bird with fewer 
than eleven cervicals.’ The length and flexibility of the neck is correlated with the covering 
necessitated by the high temperature of the Bird.’ The cold-blooded flying Reptiles 
have a comparatively short and rigid neck, but of a thickness and strength proportionate 
to the size of the head, and adequate to the work to be performed by the jaws in over- 
coming and bearing away the prey they may have seized. 

The chief variety manifested by the Péerosauria in the cervical region is in the relative 
length of the last six vertebra; this is greatest in Plerodactylus longicollum and Pt. 
longirostris ; it is least in Pf. crassirostris and Dimorphodon macronyz, and apparently 
also in Pterodactylus simus, if we may judge by the breadth, compared with the length, of 
the vertebra figured in PI. X VIII, (figs. 1 and 2) of my Monograph, above cited, of 1860. 

There seems to have prevailed a greater range of variety in the number of vertebre 
between the cervical series and the sacrum. In Plerodactylus longirostris, Cuvier esti- 
mated at least twelve which supported moveable ribs,’ and nineteen or twenty in the 
dorso-lumbar series. Von Meyer concluded that the number of dorsal vertebree fell not 
below twelve in any species, nor exceeded fifteen or sixteenin Plerosauria, Pterodactylus 
Kochii shows fourteen dorsal vertebrae ; P¢. crassirostris not more than twelve, reckoned 
by the number of pairs of free ribs, which can be satisfactorily discerned. 

I have seen no specimen of Dimorphodon yielding definitely the number of the dorso- 
lumbar vertebree, 2. e. of the vertebrae between the cervical and sacral ; it is from the best 
considerations I have been able to give to the analogies of these vertebral formule, in better 
preserved examples of other species of ,Péerosauria, that I assign thirteen to this series in 
my restoration of Dimorphodon macronyx (Pl. XX); and I conclude that the thirteenth 
was a true lumbar vertebra or without connection with a free pair of ribs. If there 
should prove to be error in this estimate I cannot think it will extend beyond one vertebra, 
or at most two, in excess of twelve dorsals. 

The nine dorsal vertebra, which have kept together, in almost a straight line, in the 
specimen (Pl. XVIII, p), testify to the strength and closeness of their reciprocal articu- 
lations, under disturbing influences which have affected so great and general a degree of 
dislocation of most other parts of the skeleton. 

Buckuanp seems first to have observed the convexity of one of the terminal articular 
surfaces of the centrum of a dorsal vertebra, and to have deduced an affinity therefrom ; 

1 The Sparrow (Pyrgita domestica) has twelve (‘ Osteol. Catal. Coll. of Surgeons, No. 1571, vol. i, 
p: 297). 

* « As the prehensile functions of the hand are transferred to the beak, so those of the arm are per- 
formed by the neck of the Bird; that portion of the spine is, therefore, composed of numerous, elongated, 
and freely moveable vertebree, and is never so short or so rigid but that it can be made to apply the beak to 
the coccygeal oil-gland, and to every part of the body, for the purpose of oiling and cleansing the plumage.”’ 
—‘ Anat. of Vertebrates,’ ii, p. 39. 


5 Vol. cit., p. 368 :—Il semble qu’il en est resté au moins douze en place du coté gauche.’ The 
specimen figured by Von Meyer, op. cit. in pl. i, fig. 1, shows thirteen ribs on the left side of the trunk. 


68 FOSSIL REPTILIA OF THE 


(the specimen is marked d in the Plate 27 of his Memoir, loc. cit.), and is described 
‘as the body of a vertebra showing a convex articulating surface, as in the Crocodile” 
(p. 221). Quenstept’s Pterodactylus suevicus showed similar detached dorsals, in one 
of which it appeared that “the articular surfaces of the body were convex at the back end, 
and concave at the fore part.”! Buckland’s specimen serves to dissipate any doubt on the 
point so important in reference to the Crocodilian affinity. It might be assumed that the 
Author viewed the convexity as posterior by the expression “as in the Crocodile ;” and in 
the last of the dorso-lumbar series, which I regard, with Buckland, as ‘ probably lumbar,’ 
in the sense of not being costigerous, the position of “ its concave articulating surface ” is 
demonstrated by those of the articular processes (zygapophyses) at the same end of the 
vertebra, which prove them to be the anterior pair, slightly prominent, looking upward 
and inward. Buckuanp notes these as “two anterior spinous processes, an obvious 
typographical error for ‘oblique’ or ‘articular,’ venial in one not professedly an 
anatomist.” 

With regard to the Crocodilian affinity inferred from this structure, it must be remem- 
bered that the proccelian stucture, though it has been observed in Crocodiles from the 
Greensand of New Jersey,’ is characteristic of the Tertiary and existing species, rather 
than of the order at large, which had more abundant and diversified (amphiccelian and 
opisthoccelian) representatives in the Secondary ages of Geology. Moreover, the anterior 
concavity and posterior convexity of the vertebral body obtain in most recent, Tertiary, 
and Cretaceous Lacertilia ; and finally, the cup- and ball-joints of the centrum appear in 
the dorsal vertebree of at least one genus of Birds, though with the ball in front.* 

In the series of nine dorsals, preserved in the subject of Pl. XVIII, pb, the centrums 
slightly lose length as they recede in position from the neck; the anterior ones measure 
0:009 mm. = 44 lines ; the posterior ones measure 0:008 mm. = 4 lines ; the transverse 
diameter of the articular ends is 0°007 mm. = 3 lines. The dorsal vertebra in Buck- 
land’s specimen presents the same dimensions. These dimensions increase as the two or 
three anterior dorsals approach the neck, but the greater enlargement of the last cervical 
is somewhat abrupt. 

For the shape and proportions of the ribs (in the Restoration, Pl. XX), I have those 
marked 4, ¢ in the original specimen,’ and the more numerous and better preserved ones 


' «Die Gelenkflache der Wirbelkérper war auf der Hinterseite convex, wie beim Crokodil, vorn dagegen 
concav. So scheint es wenigstens.”—QuENSTEDT, Ueber Pterodactylus suevicus im lithographischen 
Schiefer Wiirtembergs. 4to, 1855, p. 45. 

2 Buckland, loc. cit., pl. 27. [This vertebra is shown in Pl. III, fig. 2, of the present Mono- 
graph. | 

3 «Notes on Remains of Fossil Reptiles discovered in the Greensand Formations of New Jersey,” 
‘Quarterly Journal of the Geological Society,’ vol. v, 1849, p. 388. 

4 As in Aptenodytes ; “On the Vertebral Characters of the Order Pterosauria,” ‘ Phil. Trans.,’ 1849, 
pl. x, fig. 22, p. 163. 

> Buckland, loc. cit., pl. 27. 


LIASSIC FORMATIONS. 69 


in the specimen figured in Pl. XVIII. Their articulations with the vertebra have already 
been noticed. The ribs increase in length to the fifth or sixth, with some diminution of 
breadth after the third, and acquire a characteristic tenuity beyond the sixth pair. On 
the outer surface a groove extends from the neck, or interspace between the head and 
tubercle downward ; the front border of the groove being somewhat prominent, but sub- 
siding in the hinder ribs. Epipleural appendages are indicated in some specimens ; 
but the indications are feeble, and, if rightly so interpreted, these appendages seem to 
have been but partially ossified. 

The sternal ribs, beyond the sternum, unite below with the free ends of the abdominal 
V-shaped, intermuscular styles. 

The irregular elongate mass (marked 18 in pl. xxviii of Buckland’s Memoir) and 
conjectured to be “ sternum—much broken, and its form indistinct ” (loc. cit., p. 221) in- 
cludes two crushed cervical vertebrae, and part of a third. Of the sternum I have not been 
able to discern a satisfactory trace in any of the specimens of Dimorphodon ; its propor- 
tions and position are, therefore, indicated in the ‘restoration’ (Pl. XX) according to the 
analogy of that in Pterodactylus suevicus,’ Pt. simus,’ and in Rhamphorhynchus. 

In the main, as regards breadth of the hind part and depth of the fore part, the breast- 
bone of Péerosauria is formed on the Ornithic pattern ; 7. e. it is shield-shaped, and it 
has a keel. But the keel does not descend from the expanded portion ; it is formed, as 
shown in a former Monograph (Suppl. No. III, p. 8), by the vertical development of the 
anterior production answering to the ossified sternum of Crocodiles and to the episternum 
of Lizards. I would recommend a comparison of the figures of the sternum in /ywana and 
Notornis, given at p. 21, vol. ii, of my ‘Anatomy of Vertebrates,’ to whosoever may 
desire to form an opinion of the evidence of affinity to Birds or to Reptiles, respectively, 


Fre. 1. Fie. 2. 


59 = 


ee 
), 


Pterosaur. Crocodile. 


afforded by the Pterosaurian sternum, especially as this is illustrated in figures 7 to 12 of 
Pl. II of the Supplement No. III, above cited. No one desirous of simply getting at the 


1 Quenstedt, loc. cit. (1855). 
? Monograph,’ Suppl., No. III (1860), Pl. II, figs. 7—12. 
3 Von Meyer, op. cit. (1860), pl. vii, figs. 1 and 3, and pl. ix, fig. 1. 


70 FOSSIL REPTILIA OF THE 


truth of the matter can put aside the ‘ post-coracoid lateral emarginations,’ and other 
modifications defined in that Monograph as ‘distinctive Pterosaurian characters.’ No 
Bird has shown any approach to them. What modifications of the Pterosaurian sternum 
Dimorphodon may have presented, we have yet to learn. 

In all cases in which it has been observed, the sternum in Pterosauria (fig. 1) resem- 
bles in essential characters that of Crocodilia (fig. 2); its chief part is a longitudinal, com- 
pressed, deep bar (59), expanding laterally, some way from the fore-end, for the articulation 
of the coracoids (51),! and having the posterior expansion (60), which remains cartilaginous 
in the Crocodilia, more or less ossified, in the form of a thin semicircular plate: but the 
whole bone, though adaptively modified for attachment of muscles of flight, preserves the 
characteristic shortness compared with the trunk, and offers a striking contrast to the long 
and large subabdominal plastron in most birds of flight. There is no distinct ‘T-shaped 
episternum, such as exists in most Lacertia, and no trace of clavicles as in Lizards and 
Birds. Distinct lateral elements for articulation with sternal ribs I have not satisfactorily 
made out in any specimen. 

The abdominal heemal arches consist of slender heemapophyses and of chevron-shaped 
heemal spines. 

There is evidence of one lumbar or ribless vertebra anterior to the sacrum, in Dimor- 
phodon ; and no Pterosaurian appears to have shown more than two such vertebree: in this 
character we are again directed to the true Reptilian relation of Péerosauria, and warned 
off the beguiling marks of Avian affinity. 

The indications of epipleural appendages of ribs, more or less bony, if rightly mter- 
preted, answer to the gristly ones in Crocodiliaand some Lacertia.” The restoration of the 
bony cage of the thoracic-abdominal cavity of Dimorphodon (Pl. XX) is based on the 
analogy of better preserved specimens of Péerosauria in regard to this part of the skeleton. 
Scattered elements of the hzmal arches, ‘ abdominal ribs,’ &c., have alone been met with 
in the specimens of Dimorphodon hitherto obtained. 

The sacrum, on the probable hypothesis of retention of the length of centrum shown 
in the lumbar vertebra, would include at least four vertebree ; if, as by the analogy of the 
sacrum (figured in Pl. II, fig. 26, of the Monograph, &c., Supplement No. 1, 1859), the 
vertebrae lost length at this confluent tract, there might be five or six sacrals articulating 
with the iliac bones in Dimorphodon. Von Mxyzr figures 5—6 anchylosed sacral vertebrae 
in his Pterodactylus dubius ;* and the sacrum appears to consist of at least six confluent 
vertebrae in Lhamphorhynchus grandipelvis, Von Meyer.* 

With all the evidence that the Péerosauria, like the Dinosauria and Dicynodontia, 


? “Monograph on Cretaceous Reptilia,’ Supplement, No. III (1860), Pl. II, figs. 7—12. 

? As in Hatteria, see Giinther’s excellent Memoir, in ‘ Philos. Trans.,’ Part II, 1867, p. 13, pl. ii, 
figs. 17, 24. 

3" Op. cit.,ps l7, plevi, ae.1- 

4 Op; cit.) p. 03, pl. vill, figs, 


LIASSIC FORMATIONS. 71 


exceeded the sacral formula prevailing in existing Crocodilia and Lacertilia, we should 
gain no firm ground therefrom for predicating Avian affinity or for building thereon a 
derivative hypothesis of the class of Birds. Many existing Chelonian Reptiles have a 
sacrum composed of more than two vertebree.t 

The perfect specimen of tail-vertebree and associated bone-tendons in the specimen 
(Pl. XIX, fig. 4) completes satisfactorily the restoration of this part of the vertebral 
column in Dimorphodon. Before the discovery of Rhamphorhynchus, the order Plerosauria 
was known only through species having the tail very short. Not only were the vertebra 
comparatively few, estimated at twelve or thirteen in Pterodactylus longirostris,? at 
fourteen in Pt. spectabilis, at fifteen in Pé. scolopaciceps,’ and as low as ten in Pt. 
Meyeri,* but they were very small and short. ‘The great advocate of the Avian affinity 
of the Pterosaurs, Sozmmerrine, based his chief argument in this character. But 
Cuvier was able to adduce instances of Reptilia with tails as short; and he might now 
have cited a Bird with a tail-skeleton as long, as slender, and as many-jointed as in divers 
Saurians.” ‘The earliest indication of a range of variety in this part of the bony frame- 
work of a Pterosaur was deduced, with his usual sagacity, by Buckianp. 

In the original specimen of Dimorphodon are three caudal vertebre at the base of the 
tail, marked K, in pl. xxvii of his Memoir, from the size of which vertebra, together with 
the larger and longer legs, as compared with Pterodactylus longirostris, BucKLAND 
inferred that the entire “tail was probably longer, and may have co-operated with the 
legs in expanding the membrane for flight.”° “A long and powerful tail,’ he proceeds 
to remark, “is in strict conformity with the character of a Lizard” (ib.).7 

Buckianp would have had further direct confirmation of the length and strength of the 
tail of his Lias Pterosaur, if he had recognised the series preserved at a, a’, in his pl. xxvii, 
as caudal vertebrae; but they were conceived to belong to the neck, notwithstanding 
their slenderness and length, and that around them were ‘small cylindrical bony 
tendons, resembling the soft tendons that run parallel to the vertebrae in the tails of 
Rats.”* When the evidences of caudal structure were first recognised by Von Meyer, in 
Rhamphorhynchus Gemmingi, he detected the homologous structures in pl. xxvii of 


1 © Anat. of Vertebrates,’ vol. i, p. 65. 

2 By Cuvier, vol. cit., p. 368. 

5 Von Meyer, op. cit., p. 17. 

4 Ib., p. 17. 

5 Owen “On the Archeopteryz,” ‘ Philos. Trans.,’ 1863, p. 33, pls. i—iv. 

6 Buckland, loc. cit., p. 221. 

1 Archeopteryx lad not then been discovered ; else, it might have been objected to the above hint of 
affinity, not only that there had been short-tailed Pterodactyles, but also long-tailed Birds. 

8 «Mr, Clift and Mr. Broderip have discovered that the remaining cervical vertebrae are surrounded 
with small cylindrical bony tendons of the size of a thread. These run parallel to the vertebra, like the 
tendons that surround the tails of rats, and resemble the bony tendons that run along the back of the pigmy 
musk and of many birds’’ (loc. cit., p. 218). 


72 FOSSIL REPTILIA OF THE 


Buckland’s Memoir, and suggested that its subject might belong to the same section or 
genus.’ ‘I'he subsequent discovery of the skull and dentition has, however, shown that 
another generic section of Pferosauria, or at least one species thereof, had a similar long 
and stiff tail. The modification involving that quality does not, however, extend 
throughout; the anterior caudal vertebra retain the more normal character, and the 
appendage would be most moveable at its base. No doubt a small degree of yielding at 
the many persistent vertebral joints—for complete anchylosis has not been observed— 
would allow a slight curvature to the extent to which the tail is represented as yielding 
to a lateral force in the restored figure (Pl. XX, fig. 2). The number of the caudal 
vertebrae in Dimorphodon macronye was at least thirty; the termination of the specimen 
figured in Pl. XIX, fig. 4, does not indicate a loss there of as many centrums as would 
bring the number up to thirty-eight, which are assigned by Von Meyer to his Rhampho- 
rhynchus Gemming?. 

As we cannot, therefore, with Soemmerring, insist on the shortness of the tail in some 
Pterosauria as proof that they were Birds, so neither can we conclude from the length of 
the tail in other Péerosauria that they were Reptiles. The legitimate taxonomic deduction 
from such caudal modifications is, that they are not of sufficient importance for determi- 
nation of a class, and that they do not exclusively characterise the genus. They 
indicate adaptations in an extreme and variable part or appendage of the body to special 
powers or ways of movement, or sustentation, in air of the present group of volant animals. 

So, likewise, it cannot be, as it has been, inferred from the length of tail in Archeopterye, 
that it was a Reptile.” What we learn from that Avian fossil is akin to what we 
have learnt from Pterosaurian remains, viz., that the tailis aseat of extreme modification, in 
respect of length and number of joints, within the limits of the feathered class. Mamma- 
logists, with a like drift, could add instructive evidence of corresponding caudal variability 
within the limits of the order, as in the volant C/eiroptera, and even within the bounds of 
the family (Bradypus and Megatherium, e.g.). 

The value of the discovery of Archeopterye, in relation to Plerosauria, is enhanced by 
the peculiar nature of the matrix, conservative of cutaneous as well as of osseous 
characters ; showing casts of down and feathers,’ impressions of the fine foldings or 
wrinkles of thin expansions of naked skin, as well as delicate tendons surrounding, 
working, strengthening, and stiffening the caudal framework. 

With these parts the fine lithographic lime-marl should have preserved the plumose 
appendages of the long tail of RAamphorhynchus, if that flying Reptile had possessed such ; 
and, along with caudal plumes and vertebree, should have been preserved the bone-tendons 
of the tail, if Archeopteryx had possessed that structure. 

It is probable, from the constancy with which caudal vertebra of long-tailed 

1 In ‘Leonhard und Bronn’s Neues Jahrbuch fiir Mineralogie,’ &c., Jahrgang, 1857, p. 536. 

2 E.g., as the Gryphosaurus of Andreas Wagner. 


3 A few of the delicate, downy body-feathers of Archeopteryzx are clearly indicated near one side of the 
trunk in the slab with most of the bones of the specimen of drcheopteryz in the British Museum. 


LIASSIC FORMATIONS. 73 


Pterosaurs have been found associated with their tendons,' that detached caudal vertebrae 
of Archeopterye might be recognised through the want of them. 

We may confidently conclude that the Oolitic mud which has entombed the greatest 
number and variety of the flying reptiles of its period would have shown us, when petrified 
into lithographic slate, their feathers, if, as warm-blooded animals, they had needed such 
heat-conserving a covering. ‘The plumose clothing of the long-tailed bird of the period 
proves its heematothermal character, as the want of it shows the long-tailed pterosaur to 
have been cold-blooded. 

The tyro, fresh from the lecture-room of his physiological teacher, ambitious of soaring 
into higher regions of biology than were opened to him at the medical school, impressed 
with the relations of active locomotion to generation of animal heat, may be pardoned for 
inferring that the amount of work involved in sustaining a Pterodactyle in the air would 
make it, physiologically, highly probable that it was a hot-blooded animal. But a competent 
friend, finding him bent on rushing with such show of knowledge into print, would counsel 
him to provide himself with a thermometer adapted to the delicate testing of the internal 
heat of small animals. So provided, if he should chance to beat down a chafer in full flight, 
the experiment, made with due care and defence of the fingers guiding the instrument, 
would teach him how fallacious would be the inference that, because an animal can fly, it 
must, therefore, be hot-blooded. Unless he happen, in introducing the bulb by the 
widened vent into the abdomen, to plunge it into a mass of ova, he will find the heat of 
the beetle, notwithstanding the amount of work involved in sustaining and propelling 
itself in air, not to exceed by more than one degree that of the atmosphere. If he has 
knocked down a female cockchafer prior to oviposition, the ovarian masses may indicate half 
a degree, or even one degree, higher of temperature (Fahr.). With the cooling of the air 
in the summer night the temperature of the JJelolontha concurrently falls. So, likewise, 
would that of the flying reptile, whatever “amount of oxidation and evolution of waste 
products in the form of carbonic acid” * might have attended their exercise of flight. The 
constant correlative structure with hot-bloodedness is a non-conducting covering of the 
body. We may with certainty infer that Archeopterye was hot-blooded, because it had 
feathers, not because it could fly. 

There is no ground, from observation of the Sharks and Porpoises that accompany 
swift-sailing vessels, maintaining themselves near the surface, exercising their several and 
characteristic evolutions in quest or capture of prey, for inferring that the amount or the 
energy of muscular action is very different in the two surface-swimmers. 

Sharks have and, no doubt, work a greater proportion of muscle than Cetaceans ; a 
less proportion of their body is excavated into visceral cavities. Yet the Shark is cold- 
blooded; its temperature rises and falls with that of its medium ; it has no provision, by 


_ 1 As seen in Pl. II, at cd, and in PI. III, figs. 3, 4, 5. 
2 *Proceedings of the Zoological Society,’ April, 1867, p.417, Prof. Huxley ‘* On the Classification 
of Birds.” 


10 


74 FOSSIL REPTILIA OF THE 


a blanket of blubber or other superficial modification, in aid of the maintenance of a fixed 
and high degree of blood-heat. 

There are conditions, it is true, in which a Reptile generates a higher degree of heat 
than is usual, but they are not those accompanying any unusual or excessive muscular 
work and waste ; they are attended with rest, not locomotion. The incubating Boa gives 
to the hand that may be insinuated between the coils surrounding the eggs the sensation 
of a warm-blooded animal. Va .encrennes' found, in the Reptile-house at the Jardin 
des Plantes, when its temperature, in the month of May, was 23° (Centigrade), that the 
heat of the Python, between the folds and upon the eggs, was 41°5° (ib.); so also the 
heat of the incubating surface of the Bird may rise to 10 degrees (Centigr.) above the 
ordinary temperature—higher in this passive state than it ever reaches during flight. 

The organic condition which determines the hot-blooded or cold-blooded nature of a 
volant Vertebrate is the separation or the commingling of the arterial and venous bloods 
in the course of their respective circulations. From the demonstrated absence of any 
heat-retaining covering of the skin in Pferosawria—the kind and amount of negative 
evidence hereon being decisive—I infer that the black and red sanguineous streams were 
mixed by intercommunication of the aortic trunks of the right and left ventricles, as in 
the Crocodile.” The plumose integument of Archaeopteryx bespeaks the separation, not 
only of the pulmonic and systemic ventricles, but of the arterial trunks thence arising ; it 
was, consequently, hot-blooded, not because it could exert the muscular force required to 
sustain itself in the air. The all-important condition of the circulating system has wide 
correlations, not only with the extensive superficies acting upon the surrounding medium, 
and being reacted upon thereby, but with a rapid and uninterrupted respiration, with an 
advanced status of the nervous system, especially the brain, involving higher intelligence 
and more lively and varied instincts, especially the parental. In the organic character 
determining temperature, breathing, and higher phenomena of life, Birds agree with Mam- 
mals and differ from Reptiles. 

Birds agree with Implacental Mammals (Zyencephala) in the development, by the 
embryo, of a vascular allantois devoid of villi for placental connection.” They agree with 
the same Mammals and differ from Reptiles in the transversely and deeply folded cere- 
bellum, and in the larger proportion of that and of the cerebrum to the optic lobes. Birds 
resemble Reptiles in the absence, not only of a corpus callosum, but of a fornix and 
hippocampal commissure. The Lyencephala have the hippocampal commissure, but no 


1 «Paites pendant l’incubation d’une femelle du Python a deux raies (Python bivittatus, Kuhl),” &c. 
“Comptes rendus de ]’Acad. des Sciences,’ Paris, 19 Juillet, 1841. 

2 « Anat. of Vertebrates,’ 7, pp. 510—512, figs. 339, 340. 

3 This character is affirmed to be ‘of extreme importance, and to define Birds and Reptiles, as a whole, 
very sharply from Mammals.’’—Prof. Huxley ‘On the Classification of Birds,’ loc. cit., p. 416. But, then, 
the emphatic assertion comes from a writer on Elementary Physiology, who infers the blood of the Péero- 
sauria to have been hot because they were able to sustain themselves in air ! 


LIASSIC FORMATIONS. 75 


corpus callosum; this characterises the Placental Mammalia. Birds differ from other 
Oviparous Vertebrates in the chalaziferous ovum. ‘The particulars in which Birds differ 
from all Mammals and agree with Reptiles are comparatively unimportant ones of the 
skeleton. The occipital condyles (¢.y.) are more completely blended or unified than in 
Cetacea. ‘The tympanic is interposed between the mandible and the mastoid, as in 
Reptiles.’ 

Two genera of Lyencephalous Mammals retain the osteological character common to 
Birds and Reptiles of the connection of the scapula with the sternum by the intermedi- 
ation of a fully developed coracoid, and it is one of several and more important characters 
disproving any sharp definition of the higher warm-blooded Ovipara, at least, from the 
Ovo-viviparous or Implacental Mammalia. 

The scapular arch retains, in P/erosauria, its crocodilian simplicity, modified in shape 
and in the angle at which the scapula meets the coracoid adaptively for the function of 
flight in the limb suspended thereto. ‘There is, consequently, a close similarity to the 
same elements in Birds of Flight,’ but without any trace of the superadded furculum. The 
articular grooves on the sternum for the coracoids communicate or run into each other at 
the mid line. The articulation of the corresponding end of the coracoid must be as secure, 
and yet with as easy a motion, due to a well-turned synovial joint (shown first in 
Pterodactylus Woodwardi and Pt. simus),*? as in any Bird. ‘The confluence of the 
scapula with the coracoid seems not to be constant in the order Péerosauria ; and where 
it has been found, as in Dimorphodon and Pterodactylus Fittoni, traces of the original 
suture are present, as represented in the large Neocomian Pferosauria in my Monograph 
of 1859.* 

In some specimens of Ramphorhynchus Gemmingi and in Ramphorhynchus longicaudus 
the scapula and coracoid seemed not to have coalesced.? The coalescence is complete and 
constant (so far as may be inferred from two specimens) in Dimorphodon. 

For the analysis of the characters of the humerus in Péerosauria, 1 may refer to my 
Monograph, Suppl. No. III (1861), pp. 13—17, Pl. III. The chief seat of variety is the 
“radial crest ” (Pl. XVIII, 53, 4, of present Monograph). In the shape and proportions 
of this extraordinary process Dimorphodon resembles Pterodactylus more than it does 
Ramphorhynchus. n the proportions of the humerus to the body there is little diversity 
in the several species. 

The antibrachium is commonly two sevenths longer than the humerus. It consists 


1 Asa taxonomic character—whatever degree of value may be adjudged to it—this mode of connection 
of the lower jaw with the skull gains nothing by calling the tympanic ‘quadrate boue,’ or by affirming 
it to represent the ‘izews’ or the ‘malleus’ of Mammalia, whichever may happen to be the favourite 
fancy of the day. , 

2 ‘Monograph,’ Suppl. No. I (1859), p. 13. 

3 ‘Monograph,’ Suppl. No. III (1861), p. 12, pl. ii, figs. 7—12. 

* Suppl. No. I, Pl. II, figs. 1—5. 

5 Von MEYER, op. cit., p. 18. 


76 FOSSIL REPTILIA OF THE 


of two equal-sized, closely and extensively united bones, with one or two slender styliform 
ossicles attached lengthwise, having the base a little below the distal ends of the radius 
and ulna. The latter bone shows no pits for the attachment of quill-feathers, as in the 
hot-blooded volant Ovipara. A carpus with one large and one small bone in a proximal 
row, and with a second large and at least one smaller bone in a distal row, is another 
character by which the Péerosauria manifest their closer affinity to Reptiles than to 
Birds. The remains of the gigantic species from the Cambridge Greensands have yielded 
the characters of the two larger carpal ossicles." 

Variation, as usual, begins to assert its sway as the segments of the limb recede from 
the trunk. ‘This is mainly shown in the relative length of the metacarpus. In Rampho- 
rhynchus Gemmingi it is to the antibrachium as 2 to 7, and to the first phalanx of the 
wing-finger as 1 to 5, or rather less. In Dimorphodon the metacarpus is to the 
antibrachium rather more than 2 to 6, and is little less than one half the length of the 
first phalanx of the wing-finger. In Pterodactylus longirostris the metacarpus is 
two thirds the length of the first phalanx. In Pterodactylus longicollum the metacarpus 
is almost four fifths the length of the first phalanx of the wing-finger. In Pt. suevicus 
the metacarpus is one eighth longer than the antibrachium. 

There are diversities also in the relative length of the phalanges of the wing-finger. 
In Dimorphodon they increase in length from the first to the third. In Ramphorynchus 
Gemmingt the first and second phalanges are of equal length, and the third is shorter. In 
Pterodactylus longirostris, Pt. scolopaciceps, Pt. Kochi, they decrease in length from the 
first to the third, and in a greater degree in Pt. suevicus. 

The most marked variety, however, if the structure has been rightly determined or 
be not due to some accidental mutilation of the individual, is that on which Von Meyer?” 
has founded his genus Ornithopterus, viz. a reduction in the number of phalanges of the 
wing-finger from four to two, and the articulation of the proximal one to two large 
metacarpals. The last pointed phalanx of the wing-finger in Ramphorhynchus is rather 
longer than the penultimate one; in Ornithopterus Lavateri it is only one third the 
length of the penultimate phalanx. 

The evidences of pelvic structure in other Péerosauria, already referred to, leaves no 
doubt as to that in Dimorphodon, as restored at s, 62, 63, 64, in Pl. XX. The 
expansion of the ischial and pubic elements and the direction of the latter are strong 
evidences of Reptilian affinity, and decisive differences in the comparison with Birds. 
Given the greatest number of vertebrae grasped by the ilia, it falls short of the least 
number presented in the class of Birds, as by certain JVatatores, which concomitantly 
manifest a vacillating or waddling gait. Nothing in the structure, proportions, and con- 
nections of the pelvic arch squares with the notion of bipedal progression or erect 
sustentation of the body and wings of the P¢erosaur. The share taken by the hind limbs 


1 “ Monograph,’ Suppl. No. III (1861), p. 17, PL. II, fig. 6; Pl. IV, figs. 5—9. 
2 Op. cit., p. 25, pl. vi, fig. 5. 


LIASSIC FORMATIONS. 77 


in resting or moving on dry ground was that indicated in the restoration of the skeleton 
in PE YY: 

The hind limbs of Dimorphodon are, nevertheless, larger and stronger in proportion 
than in other Péterosauria. The femur, in most species, equals the humerus in length, and, 
in Dimorphodon, also in thickness. In Pterodactylus longirostris and Pt. Kochii the femur 
is the more slender bone; in Ramphorhynchus it is likewise shorter than the humerus. 

The tibia, more slender than the antibrachial bones, in Pterodactylus longirostris and 
Pt. Kochii, is of equal length therewith. In Dimorphodon the tibia is less slender in pro- 
portion to the antibrachium, and is longer by one seventh. In Ramphorhynchus it is 
much more slender than the antibrachium, and is nearly one third shorter. The ankle- 
joint works between the tibia and tarsus, which, as in other Reptiles and Mammals, is 
distinct from the metatarsus. There is no calcaneal prominence, and the foot admits of 
easy rotation, as in the ‘ Restoration,’ Pl. XX, fig. 2, where the inner toe is turned out- 
ward and the sole presented to view, to show the application of the wing-toe in flight to 
the interfemoral web. 

Whether the trochlear terminal joint of the tibia be ossified from a separate centre in 
the Pterodactyle as in the Bird requires a specimen of the requisite immaturity for deter- 
mining. If the hind limbs and pelvis presented the structure for sustaining and moving 
the animal erect on land, an epiphysial state of the articular ends of the long bones might 
be physiologically inferred. I conclude, from the absence of the modifications essential to 
bipedal station and progression in Péerosauria, that the articular ends of both femur and 
tibia, including the distal condyles of the latter bone, were co-ossified with the shaft as in 
other Saurians. 

When in warm-blooded Vertebrates,whether Birds or Mammals, the metapodial elements 
of different toes coalesce, the epiphyses of such coalesced series, or 
‘cannon bone,’ are usually connate, forming a single bone. As, e. g., at 
the proximal end of the Cow’s and Bird’s metatarsus (figs. 3 and 4, ¢)," 
and also even at the distal end of the cannon-bone in Ruminants 
(fig. 3, Z). I demonstrated the fact in both the metacarpus and meta- : 
tarsus of a young Giraffe, in my ‘ Hunterian Lectures’ of 1851. The © = 
specimens are Nos. 3631 and 3635 in the Osteological Collection of 
the Royal College of Surgeons (‘ Catal.’ 4to, 1853, p. 601). 

The distal trochlear end of the Bird’s tibia, in its epiphysial state 
(fig, 4, d), answers to the distal trochlear epiphysis of the Ruminant’s ae) 
tibia (fig. 3, a). In its anchylosed state the distal bicondylar troch- ‘* 
lear jomt or end of the Bird’s tibia answers to the distal bicondylar 
trochlear joint or end of the Pterosaur’s tibia. The proximal 


Fie. 3. Fig. 4. 
tale 


Win ” 


Ruminant. Bird. 


1 «The upper articular surface is formed by a single broad piece. The original separation of the 
metatarsal bone below into three pieces is plainly indicated.’’—‘ On the Anatomy of the Southern Apteryx,”” 
‘Trans. Zool. Soc.,’ ii (1838), p. 293. 


78 FOSSIL REPTILIA OF THE 


epiphysis of the Bird’s metatarsus (fig. 4, ce) answers to the proximal epiphysis of the 
Ruminant’s metatarsus (fig. 3, ¢). 

The interspace between the leg and foot is the seat of variable and inconstant centres 
of ossification, from zero, as in Proteus, Amphiuma, Aves, to the four ossicles in Crocodilus, 
and the seven ossicles in Che/one. 

‘The functions of the hind leg in Birds require peculiarly strong, firm, close-fitting, 
interlocking joints. hus, the fibula articulates directly with the femur, and the meta- 
tarsus as directly with the tibia, No interposed ossicles are permitted to affect the simple 
efficiency of this tibio-metatarsal joint in the long-footed feathered bipeds. In quadrupeds 
and in the short- and broad-footed mana tarsal ossicles, interposed at the space 4 (fig. 
3), have their use. But whether the tarsus exist or not, in the Hematotherma the articular 
ends of the long bones begin as ‘epiphyses ;? and when two or more metacarpals are to 
become massed into one bone, the epiphysis (c) is single—a very significant developmental 
guide to the homology in question. 

The strangest aberrations in homological aims have arisen from a non-recognition of 
the distinction between teleological and homological centres of ossification.!. Not only is 
a tibial epiphysis made into a tarsal bone—and why other epiphyses, such as the proximal 
one of the tibia, or the distal one of the femur, should be differently treated is not obvious— 
but new bones by the score are added to the cranial series. ‘ Basitemporals,’ ‘ prevomers,’ 
‘antorbitals,’ ‘ perpendicular ethmoids,’ ‘ali-ethmoids, &c. &., have been heaped up to 
obstruct the comprehension of the plain and intelligible nature of the bird’s skull. 

The four unguiculate digits of the foot are of nearly equal length, but present a slight 
difference in their proportions” in the Péerosauria. Cuvier having determined the Lacertian 
character of the phalangial formula of these digits, viz. 2, 3, 4, 5, adds that, apparently, 
the fifth digit was reduced to a.slight vestige of two pieces in Pferodactylus longirostris.’ 
Subsequently discovered species have offered a like indication, to which Von Meyer alludes 
as a rudiment or stump (‘ stummel’) of the fifth toe.* No other specimens, to my know- 
ledge, save the third of Dimorphodon (Pl. XVII) and the Ramphorhynchus (P|. XIX, fig. 5) 
have shown the condition of the fifth digit as of three pieces, viz. a metatarsal (m, v) and 
two phalanges (v, 1 and 2). 

The metatarsal of this toe shows an interesting affinity to that in the Crocodilia by its 
greater breadth and shortness in comparison to the other metatarsals. The two phalanges 
have proportions and forms which clearly show their adaptive relations as aids in sustaining 
the interfemoral or caudo-femoral parachute (‘ Restoration,’ fig. 2, Pl. XX). 

Owen, “ Lectures on the Comp. Anat. of Vertebrate Animals,’”’ 8vo, 1846, p. 38. 


See Monograph, pl. xi, fig. 3. 
“Tl paroit qu’ici Je cinquieme étoit réduit a un leger vestige de deux pitces.”’—‘ Oss. Foss.,’ vol. 


eo 3 _ 


cil., p. 374. 

* “Cuvier, Wagler, und Goldfuss lassen den Fuss aus fiinf ausgebilteten Zehen bestehen; in allen 
Pterodactyln habe ich aber nie mehr als vier solcher Zehen, und héchstens noch einen Stummel vorgefunden.”’ 
—Op. cit., p. 20. 


a ee ee ee 


= 


a 


LIASSIC FORMATIONS. 79 


The crushed condition of most of the long bones in the specimens of Dimorphodon 
show the wall of the shaft to have been compact and thin, the cavity large. Although | 
have failed to detect such clear evidence of the foramen pneumaticum in these crushed 
bones as in some of the vertebre, I cannot resist the inference from the structure of the 
long bones that they were filled with air in the living animal, as has been demonstrated 
in remains of the larger Pferosauria of the Cretaceous series.’ 

This general osteological character of the Pferosauria leads me to offer a few remarks 
on its relation to their peculiar power of locomotion among Reptilia, and to the affinity it 
may indicate to other groups of volant Vertebrates. 

Weight is, of course, indispensable to directed motion through the air; but, given 
the weight requisite for the action against gravity resulting in flight, whatever structure 
tends to dispense with additional burthen enables the force to act with more avail—with 
less unnecessary resistance to overcome. 

Where provision is made for unusual flying force, as by the enormous pectoral muscles 
and concomitant shape of wing in the Swift, the required weight of body called for heavier 
bones ; hence the non-pneumaticity of the skeleton. Diminished flying force, especially 
with increased bulk of body, is attended with modifications of bony structure obviously 
adapted, and which have always been recognised in relation, to reduction of weight in the 
mass to be moved through the air. It is true that the mere quantity of air contained in 
bones would have an effect inappreciable in aid of the force raising a weight of 5 Ib. or 
10 lb. from the ground ;? but the true view of the question is—given a bone of 1 foot in 
length and 3 inches in circumference, whether the restriction of bony matter to a thin- 
ness of 3 a line at the circumference, and a substitution of air for the rest of the diameter 
throughout the shaft, be not a provision for diminution of weight and conservation of 
strength which does relate to facilitate locomotion through air ? 

If the humerus of the Ostrich (No. 1373, Osteological Collection in the Museum 
of the College of Surgeons, London, ‘Catalogue’ of do., 4to, 1853, p. 265) be compared, 
as to weight, with the similarly sized humerus of the Argala Crane (No. 1107, ib., 
‘Catal.,’ p. 214), the difference is striking and suggestive; the latter bone being 
“remarkable for its lightness, as compared with its bulk and seeming solidity ” (ib., 
‘Catal.’ ib.). I demonstrated the cause of the difference by a longitudinal section of 


1 «Monograph on the Fossil Reptilia of the Cretaceous Formations (Order Pterosauria),’ (‘ Pal. Soc. 
Mon.’, vol. v), 4to, 1851, pp. 80, 98, 101. 

? A writer impugning the physiological inferences of Hunter and Camper, the discoverers of the 
pneumaticity of the bird’s skeleton, remarks :—“ A living bird weighing 10 lb. weighs the same when dead, 
plus a very few grains; and all know what effect a few grains of heated air would have in raising a weight 
of 10 lbs. from the ground. The quantity of air imprisoned is, to begin with, so infinitesimally small, and 
the difference in weight which it experiences by increase of temperature so inappreciable, that it ought 
not to be taken into account by any one endeavouring to solve the difficult and important problem 
of flight.’"—Prtricrew, ‘On the Mechanism of Flight,” ‘Linnean Transactions,’ vol. xxvi, 
p. 218, 1868. 


80 FOSSIL REPTILIA OF THE 


the two bones. In the Bird incapable of flight the humerus is solid; in the Bird remark- 
able for the long-continued power of soaring in upper regions of the air the shaft of the 
bone is a ‘thin shell of compact osseous tissue.’ ‘The relation of the weight of the volume 
of air occupying the capacious cavity of the Argala’s wing-bone to the total weight of 
its body need not be taken into account in considering the problem of flight, but the 
relation of a hollow instead of a solid humerus is a legitimate element in the endeavour 
to solve that complex kind of animal locomotion. To say that a certain amount of weight 
in the bird is essential to the momentum of flight is no argument against the reduction 
to such requisite weight of the body to be upborne. Every structure so tending to 
lighten the body of a volant animal within the required limit is, and ought to be, recog- 
nisable as physiologically related to flight. 

By the pneumaticity of the bones of the Pterodactyle, it might be inferred, from a 
single bone or portion of bone, to have been an animal of flight. For, although certain 
volant Vertebrates, e.g. the Bat and the Swift, may not have air-bones, no Vertebrate save 
a volant kind has air admitted into the limb-bones. But the effect of such admission, 
of such substitution of a lighter for a heavier material, is to diminish the weight without 
impairing the strength of the bone; the legitimate, if not sole, inference, therefore, is 
that it contributes to perfect the mechanism of flight. 

It is a purely adaptive character, and the insignificant, barely appreciable, difference 
of weight due to difference of temperature in a given bulk of air makes the pneumaticity 
of the skeleton as available and advantageous to a cold-blooded as to a warm-blooded 
volant Vertebrate. 


In concluding the description of the subjects of the present Monograph I am moved 
again to express my sense of acknowledgment for the most instructive of the evidences 
of Dimorphodon macronyx due to my friend from the beginning of our paleontological 
pursuits, the Earn or EnniskiLuen, F.R.S.; and, whilst fulfillmg this pleasurable duty, I 
would add a testimony to one whose loss Palzeontology has much reason to deplore,—to 
the unwearied and undaunted explorations of the precipitous cliffs of Lyme-Regis by 
Mary AnninG, to which, and to her singular tact of discernment of the feeblest evidence 
of a fossil in that dark matrix, science is indebted for the discovery of the first evidence 
of a Pterosaur in ‘ Lias’ of the locality, which has since yielded the grounds for the 
reconstruction of the strangest representative of the order. 


Ramphorhynchus Meyeri, Pl. XIX, fig. 5.—In further illustration of the characters 
of Dimorphodon macronyz 1 have added to Pl. XIX a figure of a long-tailed Pterosaur 
from the lithographic slate of Pappenheim, which, in the feebleness of its hind-limbs and 
the general proportions of the tail, resembles Ramphorhynchus Gemmingi, V. M. 


1 See Von Meyer, op. cit., pl. ix, fig. 1. 


LIASSIC FORMATIONS. 81 


The present specimen, from Dr. Hiberlein’s collection, now in the British Museum, 
shows the fifth or ‘wing-toe’ of the foot, ib. fig. 5, v, which had not been preserved in pre- 
viously described specimens of the genus.’ At least thirty-four vertebrae extend beyond 
the sacrum ; thirty-eight caudals are the reckoned by Von Meyer in the specimen of his 
R. Gemmingi with the best preserved tail ;” but this difference would not have yielded 
sufficient ground for specific distinction. ‘There are, however, differences in the length 
of the parts of the hind limb which indicate this to have been longer in proportion to the 
tail and the ‘symphysis mandibule’ than in 2. Gemmingi. 


R. Meyeri. R. Gemmingi (pl. ix, V. M., op cit.). 
in. i ke 
Length of femur 

tibia . 

ss Ist toe 
A 2nd toe 
- 3rd toe 
$5 4th toe 
m 5th toe 


to)e 


— 


1 

] 8 
1 2 
1 4 
1 6 
] 3 


lH ole 


woOoH ORD OWnN = 


oO EFS = — — LOK 


In both specimens the number of phalanges of the toes increases from the first or 
innermost to the fourth, in the usual saurian ratio, 2, 3, 4,5. In Ramphorhynchus 
Meyeri the fifth toe consists, as in Dimorphodon, of two phalanges, the first being six lines 
in length, the second three lines, and ending in a point. The metatarsal of this ‘wing-toe’ 
is short, broad, and flattened, with a convex outer border at its basal half. 

The bones of the left hind limb are well preserved in the specimen figured. 

The caudal vertebree are surrounded by the bone-tendons. ‘Their proportions, as 
shown in the figure, accord with those in Dimorphodon, Pl. XIX, fig. 4.2 The posterior 
dorsal vertebree of Ramphorhynchus Meyeri show the broad diapophyses supporting the 
ribs, which are more slender than those at the fore part of the chest. The symphysis 
mandibulz is one inch six lines in length, including the edentulous pointed end. Four 
pairs of long slender laniary teeth are preserved at the fore half of the symphysis. The 
teeth at that part of the lower jaw in Ramphorhynchus Gemmingi are fewer in number 
and less closely arranged.’ 


1 “Bin Stummel wur nicht mit Sicherheit zu ermitteln.’—Von Meyer, op. cit., p. 72. 
2 Opetit:, p. 69. . 

3 This figure has the neural surface downward in the Plate. 

4 Compare with Von Meyer, op. cit., pl. ix, fig. 1, and pl. x, fig. 1. 


11 


i a Rl 


PLATE XVII. 


Dimorphodon macronyz. 


Skull and parts of the skeleton : nat. size. 


From the Lower Lias of Lyme Regis. In the British Museum. 


L. XVI. 


TI SAP 007 ecnaneet 


on 


,by J. Erzlebe 


om Hat on Ston 


PLATE XVIII. 
Dimorphodon macronya. a 
Skull and parts of the skeleton: nat. size. 


From the Lower Lias of Lyme Regis. In the British Museum. 


T XVII 


PLATE XIX. 


Dimorphodon macronyx. 


Bones of fore foot and part of wing-finger. 

Pelvis. 

Anterior caudal vertebre. 

Vertebree of entire tail, with surrounding bone-tendons. 


From the Lower Lias of Lyme Regis, Dorsetshire. 


Rhamphorhynchus Meyeri. 


. Dorso-lumbar, sacral, and caudal vertebre, part of pelvis, with bones of the 


pelvic limbs. 
Fore part of mandible and teeth. 


From the Lithographic Slate, Pappenheim, Bavaria. 


. Second phalanx of wing-finger of Péerodactylus validus. 


Distal end, and section of shaft, of tibia of Pterodactylus curtus. 
Ib. ib., side view, of do. do. 
Section of second phalanx of wing-finger of Pterodactylus nobilis. 


From the Wealden of Sussex. 


All the specimens of the natural size. In the British Museum. 


7 
$s | 
oz 


6 


PLATE XX. 


Dimorphodon macronye. 


Fic. 


1. Restoration of the skeleton : nat. size. 
2. Restoration of entire animal : reduced (see ‘ Scales’ at foot of plate). 


3. Side view of the skull of a recent Saurian (Lyriocephalus). 


PALAONTOGRAPHICAL SOCIETY. 


[INSTITUTED MDCCCXLVII. 


VOLUME FOR 1869 


DCCCLXX. 


ry nAa 
es 


MONOGRAPH 


ON THE 


BRITISH FOSSIL 


CEKETA CEA 


FROM THE 


RED CRAG. 


BY 
PROFESSOR OWEN, E.R.S., D.C.L.. 
FOREIGN ASSOCIATE OF THE INSTITUTE OF FRANCE, ETC ETC. 
No. I, 
CONTAINING 


Gunus ZIPHIUS. 


Pages 1—40; Plates I—V. 


LONDON : 
PRINTED FOR THE PALAONTOGRAPHICAL SOCIETY. 


1870. 


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a 4 lea) 
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> ee as 


2 = 


PRINTED Bi! 


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, id DTA TA CL ‘ 
J. BE ADLARD, BARTHOLOMEW cross. ‘ 


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MONOGERAY i 


OF 


BRITISH FOSSIL CETACEA 


OF THE 


"RED CRAG? 


Genus—Zivuivs, Cuvier. 


Tne more abundant evidences of Cetacea from the ‘Red Crag’ of Suffolk are teeth 
and ear-bones, by which, in 1842 and 1843, remains of the order were determined in 
that formation and locality.” Portions of cranium, chiefly rostral, referable to the Ziphioid 
family, are rare; and these are always more or less rolled and worn,’ a condition which, 
with the break-up of the cranial parts of the skull, and the scattering of its densest bony 
parts, with detached teeth, indicates the long-continued operation of sea-waves, breakers, 
and currents, on the deposits of a Tertiary period, which, in England, occupies a very 
limited area. Nevertheless, there are grounds for estimating the amount of these 
deposits, which must have been broken up and transported in order to furnish the 
Cetacean nodules of the ‘Red Crag,’ at thousands of cubic acres. The remaining 
débris of older Pliocene with probably Upper Miocene beds, known as ‘ Red Crag,’ 
occur in patches from Walton-on-Naze, Essex, to Aldborough, Suffolk, extending 
from five to fifteen miles inland. The thickness of the Red Crag is variable, but 
does not now average more than 10 feet; its greatest observed thickness is 40 feet, 
including some sand-beds at the top, which have no shells. Broken-up septarian nodules, 
and other so-called ‘rough stones,’ the débris of washed-off ‘London Clay,’ form in 
some places a rude flooring to the Red Crag, and the Cetacean with other phosphatic 

1 «Recherches sur les Ossemens fossiles,’ 4to, 1823, tom. v, pt. 1, p. 352. 

2 « Reports on British Fossil Mammalia,” ‘Trans. Brit. Assoc.,’ 1842. At this date I was misinformed 
as to the formation in which the ‘ physeteroid tooth’ described in that ‘ Report’ had been discovered. Mr. 
Charlesworth traced the origin of the then unique fossil to the Red Crag at Felixstow. In the following 
year (1843) Prof. Henslow submitted for my determination and description a number of ‘concretions’ 
from the same formation and locality, which are described in the ‘Appendix’ to Prof. Henslow’s papers 
“ On Concretions in the Red Crag,” &c., ‘ Proceed. Geol. Soc. London,’ vol. iv (Dec., 1843), p. 283. 

3 As in the specimen, closely resembling Ziphius longirostris, Cuv., described and figured in my 


“Description of Mammalian Fossils of the Red Crag,” ‘Quart. Journ. Geol. Soc.,’ vol. xii, 1856, p. 228, 
fig. 24. 


1 


2 BRITISH FOSSIL CETACEA 


‘nodules,’ ‘concretions,’ or ‘cops’ of the diggers! are most abundant immediately 
over the ‘rough stones.’ 

It will be understood that I use here, as in my ‘ Memoir’ of 1856, the term ‘ Red 
Crag’ in the sense in which it was first applied by Edward Charlesworth, Esq., F.G.S., 
as “characterised by its ferruginous colour ;”” fully recognising, with the same perse- 
vering and accurate explorer of the Eastern Counties’ Crag-beds, that the rolled and frag- 
mentary Cetacean remains belong to a deposit older than those which, by their testaceous 
fossils, may be truly or strictly defined as ‘ Red Crag :’ that the older deposit in question— 
more or less destroyed and broken up in Suffolk—answers, in time, to the better preserved 
Belgian ‘ Sable noir’ of the ‘Systeme Diestien’ of Nyst and von Koenen ;* and also that, 
though I have received Cetotolites from the London Clay of Essex, I hold the same opinion, 
as does Prof. Van Beneden in regard to his ‘ Placoziphius,’ from the ‘ Rupelian Clay’ of 
Edeghem,* that they have gravitated into such older deposits in the course of their 
agitation and rollmg by the surf-waves. : 

By the term Zp/ius I understand, with Cuvier, a genus having close relations with 
Physeter, Lann., and still closer with Hyperoodon, Lacép.° (if it really merit generic distinction 
therefrom), characterised by a more or less elongate, slender, edentulous, beak-shaped upper 
jaw, or ‘rostrum, varying in form, abruptly and considerably expanding between the 
orbits, behind which both maxillaries and premaxillaries rise to build, with the frontals, a 
boundary wall concave upward and forward, the middle part of which, formed by the 
premaxillaries and nasals, arches forward, so as more or less to overhang the nostrils.7_ In the 
mandible or lower jaw the teeth are reduced usually to a pair, which are subterminal or 
terminal in position, and are most conspicuous, or may only be visible, in the male sex.® 

In the existing species of this genus, discovered since the date of the classical 


! An abbreviation, according to our Saxon proclivity, of ‘coprolites,’ which these nodules were generally 
supposed to be prior to my discovery, in 1840, of the nature of the coprolitoid fossilin Mr. Joun 
Brown’s collection, afterwards figured in ‘ British Fossil Mammalia,’ p. 536, figs. 219, 226, and 227. 

2 «Proceedings of the Geological Society of London,’ vol. ii (May, 1835), p. 196. 

3 «Geological Magazine,’ 1867, vol. iv, p. 501. 

4 See the excellent Memoir by Mr. Gopwin-AvstEN, ‘Quart. Journ. Geol. Soc.,’ vol. xxii, 1866, p. 228. 

° Ziphius—“a de grands rapports avec le cachalot et encore de plus grands avee l’hyperoodon.” 
—‘Ossem. Foss.,’ tom. cit., p. 351. 

6 « La partie du museau formée comme a l’ordinaire par les maxillaires et les intermaxillaires, est une 
espéce de cylindre ou de prisme quadrangulaire dont les angles sont arrondies.’’—Ibid., tom. cit., p. 354. 

7 * Ces intermaxillaires—remontent le long des cétés des narines, et se recourbent en avant pour former 
avec les deux os du nez, , ”, qui sont encastrés entre eux, une espéce d’auvent sur le dessus des narines.” 
—Ib., p. 351. 

* The taxonomic applications of the teeth in the species of Ziphius is affected by the singular arrest of 
development of the dental system. Rudiments of teeth may be found hidden in the gum, filling the alveolar 
groove of the premaxillaries, from which one small pair may slightly project, according to the age of the 
individual, Small teeth, or concealed rudiments, may precede or follow the pair, or two pairs, which are 
better developed in the lower jaw. As generic characters, I deem these dental conditions to be valueless. 


OF THE RED CRAG. 3 


work cited, the blow-hole is single, transversely crescentic, with the horns forward ; 
there is a small but rather high falcate dorsal fin, the pectorals are small and placed 
low, the caudal broadly and terminally emarginate.' 

The rostrum in Ziphius, as demonstrated in the skulls of the existing species above 
described, is mainly composed of the maxillaries and premaxillaries; but it likewise 
includes at the medial line, along a greater or less extent from the base, the prefrontal 
and the vomer, while at its base are parts of the palatines and pterygoids. 

For aright use of a generic name it is requisite, in case of doubt, to refer to the speci- 
men or specimens of species on which the genus has been founded, called ‘type- 
specimens’ by those naturalists who use the word ‘type’ in that sense. The genus 
Ziphius was defined by Cuvier from characters afforded by more or less mutilated 
skulls of three or more species. They were mainly derived from the least mutilated 
specimen, which had been found by a peasant on the sea-shore (‘sur le bord 
de la plage”) between the village of Hos and the embouchure of Galégeon, in the 
Department of the “ Bouches-du-Rhone,” near the canal leading from the marsh of 
‘! Estomac’ to the sea.” The occiput and most of the 
cranial cavity were wanting; but the specimen showed Fre. 1. 
the temporal fossee with the orbits and interorbital 
expanse of bone, including the nostrils and the whole 
of the rostrum, which was edentulous. It is figured 
im the ‘Ossemens Fossiles,’ tom. v, pt. 1, pl. xxvii, 
fig. 3 (7 the nat. size). I reproduce in the cut, fig. 1, 
the original figure of part of this specimen ona larger 
scale, given by Prof. Gurvats, i his ‘ Zoologie et 
Paléontologie frangaises,’ 4to, pl.xxxviu, fig.2. These 
figures show that the premaxillaries (g2) forming the 
end and upper part of the sides of the rostrum 
expand as they rise (’22), curving outward and up- 
ward, and inclining forward at their summits (22°) 


to support a pair of small massive nasals (15) 
wedged in grooves between them: the expanding 
nasal plates of the premaxillaries bound or form the 
lateral walls of the cavity (prenasal fossa), into which opens posteriorly the upper outlet of 
the nostrils, the septum of which (14) extends forward, bisecting in part the prenasal fossa. 
The premaxillary wall or ridge dividing this fossa (29’) from the maxillary (g1’) is represented 
on the left side of Cuvier’s fig. at 0, 0 ; the rostral part of the premaxillary (99, fig. 1) is marked 
g' 0 the same figure. The maxillaries (91, fig. 1, pl. xxvii, fig. 3, ¢, ¢, f, inthe ‘ Ossem. Foss.’) 


' As in Ziphius micropterus, ‘Mémoires de lAcadémie Royale de Bruxelles,’ tome xii, pl. i; and in 
Ziphius patachonichus, ‘ Anales del Museo Publico de Buenos Ayres,’ tom. i, pl. xv. 


5 


* «Ossem. Foss.,’ tom. cit., p. 150. 


Ziphius cavirostris, Cuv. 


4 BRITISH FOSSIL CETACEA 


greatly extend transversely the interorbital platform, of which they constitute the two large 
lateral wings or concavities ; they are continued forward along the under part of the sides 
of the rostrum, and for some distance in the form of an outstanding ridge (fig. 1, e, e). 
The upper mid-tract of the rostrum (‘vomer,’ 4, of Cuvier) is formed by a production of the 
‘Jamina perpendicularis «thmoidei,’ answering to my prefrontal,’ of singular thickness (fig. 
1,14’). The lower mid-tract of the rostrum is, in part, formed by the veritable vomer. This 
rostrum was edentulous. 

From the longitudinal extent of the prenasal fossa, through interruption of ossification 
of the prefrontal, between 14 and 14, in fig. 1 (4 and 4 of fig. 3, pl. xxvii, ‘Oss. Foss.’), 
Cuvier gave to this original type of his new genus the specific name Ziphius cavirostris. He 
notes the cetacean character of want of symmetry in the twist of the nasal bones to the left ; 
and that the contiguous plate of the right premaxillary is the largest, while, in the fossa (ib., 
0), the left premaxillary is the largest (‘dans la grande fosse c’est le gauche qui reprend de la 
largeur et qui rejette vers la droite la suture qui la sépare de l'autre” (tom. cit., p. 351). The 
posterior wall of the nasal chamber is perforated on each side the septum by a single orifice ; 
Cuvier writes—“pour la communication du nerf olfactif avec les cavités nasals:’’ it more pro- 
bably transmitted, as in Delphinide and Hyperoodon, the nasal branch of the first division 
of the trigeminal nerve, with accompanying branches from the anterior meningeal artery. 

Cuvier adds that this skull was very heavy and very dense (‘‘cette téte est tres pesante, trés 
dure,” ib., p. 352), by which he may be recording another character of his genus, viz. the 
singular petrosal density of much of the osseous texture; this character, however, he 
believed to be due to posthumous petrifaction of the specimen, and so deemed it to 
belong to a fossil Cetacean. The specimen, now in the Museum of Comparative Anatomy 
at the Jardin des Plantes, though not ‘“complétement pétrifiée en calcaire,” ib., does 
appear to have imbibed calcareous matter, probably by long imbedding in the superficial 
deposit from which it had been dislodged. 

The new and more enlarged view of the expanded interorbital part of Cuvier’s original 
or type-specimen of Ziphius cavirostris, which Prof. Gervais has given,” is chiefly for the 
purpose of comparing it with a similar view of the skull of a Zpdzus found in 1850, on the 
beach of Aresquiers, Department of Hérault, which Zphius, still existing in the adjoining 
seas, he refers to the same species—Z. cavirostris.2 In this instance the mandible was 
obtained, showing one of the series of germs of teeth in the alveolar groove of each ramus to 
have been developed at the end of the symphysis. The small size of this protruding tooth 
may relate to sex, and indicate the stranded Zphius to have been a female. M. Gervais notes 
that the prenasal fossa (his ‘excavation conchoide’) is deeper and less expanded (‘‘ moins 
évasée et plus considerable,” op. cit., p. 8) than in Cuvier’s specimen, owing to differences in 
the direction and development of the premaxillary walls of the fossa. But M. Gervais is 
inclined to include these differences within the range of sexual or individual varieties of 

* «On the Archetype and Homologies of the Vertebrate Skeleton,’ 8vo, 1848, p. 58. 


2 «Zoologie et Paléontologie francaises,’ 4to, pl. xxviii, fig. 2; woodcut fig. 1 of present Monograph. 


3 Op. cit., p. 154, pl. xxxviii, fig. 8, and pl. xxxix, figs. 2—7. 4 Op. cit., pl. xxxix, figs. 4, 4a. 


OF THE RED CRAG. 5 


the same species of Ziphius. ‘There seems, however, to be another difference, if we may 
judge from the figure of the upper surface of the cranium from Aresquiers, viz. the 
continuation of a canal more or less unoccupied by the prefrontal (or so-called ‘ vomer’) 
along the upper part of the rostrum to its termination. The text of M. Gervais supplies 
nothing to correct the plain inference from his figure. Moreover, the new and valuable 
figures from the original type-skull of Zphius cavirostris, in the Museum at the Jardin 
des Plantes (pls. xxxviii, fig. 2, and pl. xxxix, fig. 1, Gervais, op. cit.), confirm the 
accuracy of Cuvier’s account of the proportion contributed to the rostrum by the pre- 
frontal (fig. 1, 14’: ‘vomer, h, “ d’une singuliere épaisseur,” Cuv., tom. cit., p. 351) ; whereas 
the appearance of the homologous tract in the Zphius from Aresquiers (Gervais, pl. xxxviii, 
fig. 1) plainly shows it to form a narrower and less elevated tract at the upper part of the 
rostrum. If this should signify something more than sexual or individual difference, I never- 
theless concur with Prof. Gervais in rejecting its interpretation as a generic distinction. 
Returning to Cuvier’s illustrations of his (supposed extinct) genus Zphius, we find it, 
next, illustrated by veritable fossils, from what is now known as the ‘ Middle Crag’ at 
Antwerp. Ziphius planirostris, Cuvier, is founded on specimens (figs. 2 and 3) which 
include the rostrum, and so much of the expanded bases of the maxillaries and premaxil- 
laries, with the palatines and pterygoids, as give the characters of the nasal passages 
and base of the rostrum. The posterior part: of 
the basal or interorbital expansions of the maxillaries 
and premaxillaries are, with the overarching nasals, 
broken away. In the second of the two fossils in the 
above general condition, to which Cuvier applied the 
names Ziphius planirostris, he recognised and points 
out differences which might be of specific value. 
Regarding the first as the type of Cuvier’s Ziphius 
Plamirostris, its more perfect condition permits both 
upper and lower apertures of the nasal passages to 
be seen 


a condition of value in appreciating the 
generic osteological characters of Zphius. In the 
upper view (fig. 2) the prefrontal (14) forms the 
posterior wall and septum of the vertical respira- 
tory canals, the septum being continued into a sud- 
denly expanded tract (14'), which advances with an un- 
symmetrical bend to the right, as in Ziphius Layardi, 
Plate I of this Monograph. Any further definite tracing 7 
of this mid-tract is not afforded by Cuvier’s reduced Ziphius planirostris, Cuv. 
figure, beyond the canals (d, 2) which terminate the pre- 


' «J,'autre morceau—plus mutilé que le premier, offre des différences, qui pourroient passer pour 
spécifiques.””—Op. cit., p. 355. 


6 BRITISH FOSSIL CETACEA 


nasal fossee (22, 29’) anteriorly, and partially reappear at », », fig. 2. The premaxil- 
laries expanding at the sides of the nostrils develope each a ridge, extending toward 
the nasals and dividing the prenasal fossze from the maxillary ones (21', 21’). 

‘The specimen figured (‘ Oss. Foss.,’ loc. cit., pl. xxvii, figs. 4, 5, 6; fig. 5 being copied 
in my figure 2) shows a rostrum of similar proportions to that in the type-species (fig. 1, 
and ‘Oss. Foss.,’ loc. cit., fig. 3); but the prenasal fossze are more shallow, the premaxil- 
lary walls being less elevated; and a more marked difference is seen in the continued 
ossification from the ‘septum narium’ or ‘lamina perpendicularis’ forward (fig. 2, 14’), 
expanding to form a broad and dense mid-tract along the’ upper surface of the rostrum 
(as at f in Cuvier’s figures 4 and 5, “créte plate, qui me paroit appartenir au vomer 
par sa partié inférieure et élargie,” ib., p. 354). I shall subsequently show that this mid- 
tract does in the present and some other species of Zphius coalesce by its lower 
expanding surface with the canaliculate vomer, and in that respect, but in that only, may 
be said to belong thereto. 

The third specimen, also fossil (fig. 3), and from the same Mid-tertiary formation and 
locality, resembles the second in the proportions of the rostrum, and in the continuous 
ossification (14') of the cartilage extending forward from the ‘septum narium’ (14), and 
expanding as shown at f in Cuvier’s pl. xxvii, fig. 7. 
Only, whereas in fig. 2 (‘Oss. Foss.,’ ib., fig. 5) 


Meh, BY 


the prenasal fossee terminate anteriorly, each in 
a foramen leading to a canal which reappears 
further forward on the surface of the snout; in fig. 
3 (Oss. Foss.,’ ib., fig. 7) the fossze are continued by 
open canals (d, d) to the same part of the snout, 
gradually contracting forward to such canals. More- 
over, in the present species, which may be noted as 
Ziphius Cuviert, the upper surface of the maxillaries, 
from the anterior third of the rostrum to the sides of 
the prenasal fossve (d, @), is roughened by irregular tu- 
bercles and ridges. Thischaracter appears indifferent 
degrees in other fossil Zphit, e. g. Z. gibbus, t. 1, fig. 
2,e; Z. (Ziphiopsis, Du Bus.) phymatodes, &c. 
From the expanded superorbital platform of 
the maxillaries (¢, c, figs. 2 and 3), these bones 
rapidly contract, with a concave outline to the base 


; of the rostrum, along the sides of which a ridge (¢) 
Ziphius Cuvieri, Ow.; Z. planirostris, No. 2, Cu. is eontinued for some way, gradually subsiding. 
The third species of Ziphius, Cuv., is represented by a petrified edentulous rostrum (‘Oss. 
Foss.,’ loc. cit., pl. xxvii, figs. 9, 10), with so much of the base as shows it to be of similar 
composition with that in the preceding Zipizi, but in shape longer and more slender. As 


OF THE RED CRAG. . i 


the upper mid-tract of the rostrum shows a mesial linear groove, Cuvier attributes its 
formation to the premaxillaries ; but in the previous species the anterior continuation of 
the so-called ‘ vomer’ is similarly grooved. The veritable premaxillaries are perforated at 
their basal expansion, as in Ziphius planirostris, by the incisive foramina (marked d, d in the 
illustrations of the present Monograph). 'The maxillaries, where, tracing them backwards, 
they begin to expand, show the lateral ridge marked e, and the outlets of the suborbital 
canal. On this fossil rostrum Cuvier founds his species Ziphius longirostris. 


I now proceed to the inquiry, whether any existing species of Cetacea manifest 
characters which can legitimately be interpreted as generically those of the Zphius of 
Cuvier, and whether they manifest corresponding modifications in the construction of 
the rostrum with those interpreted by Cuvier as of specific value. 

I shall commence with the description of one of the series of Cetacean skulls transmitted 
from the Cape of Good Hope, for description and comparison, to the British Museum. 
Two figures of the instructive specimen in question have been published by my colleague 
Dr. Gray, F.R.S. ; one is a copy of a drawing by Mr. Trimen, of Cape Town, under the name 
Hyperoodon capensis ;* the other is an original woodcut from the specimen itself, as exem- 
plifying the genus Petrorhynchus, Gray.” It would be an ill return for Mr. Layard’s 
liberal labour of transmission if the British Museum did not furnish him with such 
anatomical accounts of his rarities as might serve or help to determine their genus and its 
true affinities. 


Zrpuius inpicus, Van Beneden ;> PutRORHYNCHUS CAPENSIS, Gray ; (figs. 4, 5, 6, and 7). 


This species exhibits the interesting and instructive condition of a partial ossification 
of the prefrontal or cranio-facial cartilage—so-called ‘vomer’ (fig. 4, 14, 14’). In this 
skull the cranio-vertebral elements continued by or into that cartilage can be completely 
traced. The prefrontals form, as usual, the posterior concave wall of the nasal passages, 
where each is perforated by one larger foramen, for the trigeminal olfactory (?), and a few 
smaller vascular foramina; they coalesce at the mesial line, and rise as a low buttress- 
hike ridge abutting obliquely and unsymmetrically against the nasal bones; the coalesced 
parts then extend forward, as the thin ‘ septum narium,’ with a sharp, free, superior concave 
border ; the plate slightly thickens as it descends to be wedged into the vomer, expands 
more as it advances in front of the nasal apertures, but subsides to the bottom of 
the prenasal fossa, sinking almost to the level of the vomer, with which it has 
coalesced, and there, at 14, shows a surface from 2 to 3 inches across and 9 inches 
in length, concave both lengthwise and transversely, in which concavity was lodged the 


1 “Proc. Zool. Soc.,’ April 11, 1865, p. 359. 
2 Ib., June 27, 1865, p. 527. ‘ 


3 «Mémoires couronnes et autres Mémoires de |’Académie Royale de Belgique,’ t. xvi, 1863. 


8 BRITISH FOSSIL CETACEA 


remaining prefrontal cartilage, into which had extended a few spicule of bone from 
the lateral premaxillary walls (22', 22’) of the deep cavity left between them by reason of 
the defect or arrest of ossification of this part, as in Zphius cavirostris (fig. 1). It is 
most interesting to find indications of a similar structure in the original figure taken from 
Cuvier’s type-specimen of Ziphius cavirostris, in pl. xxxviii, fig. 2, of Prof. Gervais’ 
excellent work above cited. 


Fic. 4. 


f, 


Ny 
i 
Dae 


Ziphius indicus, V. B.; 1-8th nat. size. 


In Ziphius indicus, at the distance of 9 inches from the upper nostrils, ossifica- 


' Dr. Gray supposes this figure to be from a specimen distinct from that figured by Gervais, in his 
plate xxxix ; he observes, ‘The skull of this genus (viz. Petrorhynchus) resembles in several particulars 
the skull of Ziphius cavirostris figured by Gervais (‘ Zool. et Paleont. frang.,’ t. 39); but the cavity on 
the crown of that species is only slightly developed, though it is apparently rather more developed in the 
other specimens figured on the plate, t. 38 (figs. 1, 2) of that work, and the vomer is sunk in a groove, as 
in the other Zipbioid genera’? (‘Catal. of Whales, &c.,’ p. 346). But the figure 1 of pl. xxxviii, Gervais, 


OF THE RED CRAG. 9 


tion has abruptly proceeded to convert the prefrontal cartilage into a dense mid- 
tract of bone (fig. 4, 14’) of a petrosal texture, and of unusual breadth, viz. 2 inches 
9 lines, at about three inches in advance of the part where this ossification gains the level 
of the premaxillaries. Prof. Gervais’s figure shows the ossified prefrontal (his ‘ vomerine’) 
tract to rise and expand with similar abruptness in the type Ziphius cavirostris (cut, 
fig. 1, 14’). From its broadest part, in Zphius indicus, the mid-tract contracts, at first 
gradually, then more suddenly and unsymmetrically ; the sutural fissures between it and 
the premaxillaries (22) becoming shallower and widening into grooves, which demonstrate 
at their bottom the complete coalescence of the prefrontal and premaxillary elements of the 
rostrum. The grooves are continued to the end of the rostrum, and indicate the share 
contributed there by those bones respectively. 

The side walls of the rostrum, formed by the premaxillaries (22), rise almost vertically 
from the sutures with the maxillaries (21), and terminate above in a border, which is 
obtuse for the first six inches, from the end of the snout backward; it then becomes a 
sharp margin bent inward, so as partly to overlap the mid-tract (14’). ‘The inward 
inflection becomes greater, and the border is again thickened and also roughened where 


Ere: 5. 


Ziphius indicus, V. B. 1-8th nat. size. 


it arches over the part of the cavity left by the unossified part of the prefrontal (14). The 
same modification characterises the type-specimen of Zphius cavirostris (fig. 1, 22%). 


is an enlarged upper view of part of the same skull (‘de la plage des Aresquiers’), as the upper view of the 
entire skull given in figure 2 of pl. xxxix; while figure 2 of plate xxxviii is a view of the upper surface of 
part of Cuvier’s specimen (‘de la plage de Fos.’), of which a side view of the whole specimen is given in 
fig. 3, pl. xxxix, of the ‘ Zool. et Paleont. frangaises.’ 


2 


10 BRITISH FOSSIL CETACEA 


The premaxillaries then expand and diverge (22’), curving outward and upward to articu- 
late with the sides of the basal half of the short and massive nasals (15). These are 
rather turned to the left. The nasal plate of the right premaxillary (22’) is broader and 
less vertical than that of the left; it is, as it were, pushed somewhat outward and down- 
ward. ‘The kind and degree of this symmetry closely resemble those described and figured 
by Cuvier in his type-specimen of Ziphius cavirostris.' 

The cavity, so circumscribed or bounded externally by the premaxillary plates, answers 
to that marked d in figs. 4, 5 and 6, of pl. xxvu, op. cit., of Cuvier’s Ziphius planirostris ; 
we shall find it common to all the Zpz, with varying proportions [such as are exhibited 
by the recent Zphius Layardi, Pl. I, fig. 2, of the present Monograph]. The maxillaries, 
forming, in Ziphius indicus, the lower and lateral parts of the rostrum to within about 
three inches of its free end (fig. 5, 21), gradually expand vertically and transversely as 
they pass backward, bending inward below, along the palato-premaxillary and palato- 
vomerine sutures, until the right meets the left maxillary at the mid-line (fig. 7, 21), in 
advance of the palatine bones (ib., 20). The suture between the maxillaries is about 
5 inches long. ‘This palatine part of the maxillary is convex transversely and smooth. 
It is bounded above for the first five inches by a narrow (ecto-maxillary) groove, the upper 
border of which projects, at first slightly, and then extends outward, forming behind the 
groove a rough (ecto-maxillary) ridge (fig. 5, e), gaining both in transverse and vertical 
extent or thickness until it reaches the middle of the naso-premaxillary plate, where it 
swells into a convex tuberosity (fig. 5, y), at the part answering to that in which the 
vertical walls rise in [yperoodon. Beyond the tuberosity the maxillary extends outward, 
articulating first with the malar (26), then with the superorbital tract (11) of the frontal, 
sweeping upward, in a graceful curve, with that bone to join the base of the nasals. This 
broad interorbital plate of the maxillary forms, with the similarly expanded nasal plate of 
the premaxillary, a large and moderately deep semilunar cavity, perforated by the 
(antorbital) canal and foramen, transmitting a branch of the second division of the 
trigeminal (a). The cavity contracts and deepens forward, answering to that so marked in 
the canal between the tuberosity (7) and the beginning of the premaxillary expanse, and 
there opens the second canal (fig. 4, 4), continued from the antorbital one, for the chief 
branch of the second division of the trigeminal, mainly answering to the suborbital or 
antorbital nerve in land mammals, as in Physeter (Euphysetes) simus (‘“ Indian Cetacea,” 
‘Trans. Zool. Soc.,’ vol. vi, pl. xiii; Phocena brevirostris (ib., pl.ix). The ectomaxillary 
ridge (fig, 5, e) is grooved along the thick margin of its basal or hinder half. A narrower 
groove, commencing three inches in advance of the foramen (4, fig. 4), extends forward 
along the line of the maxillo-premaxillary suture to the anterior termination of the 
maxillary. ‘The whole of the rostral part of the maxillary extends outward, beyond the 
subvertical plane of the side-wall formed by the premaxillary, and in a degree augmenting 


* Oss. Foss.,’ tom. cit., pl. xxvii, fig. 3. 


OF THE RED CRAG. 1] 


as it approaches the base of the rostrum. The transverse section of the middle of the 
rostrum (at the part marked 13 in fig. 7) thus gives the figure shown in outline, by woodcut, 
fig. 6, in which 14 marks the petrosal production of the ‘prefrontals,’ 22 the premaxil- 
laries, 21 the maxillaries, and 13 the vomer. As far as can be judged from the figure 
of the rostrum in Cuvier’s type-specimen of Zphius cavirostris, it would yield a very 
similar form of transverse section. 

On the palatal surface of the rostrum of Ziphius indicus (fig. 7) the premaxillaries 
(22) alone form the anterior 43 inches ; the vomer (13) then appears at the mid-line for an 


FIG. 7. 


Section of rostrum, Ziphius indicus. 
1-4th nat. size. 


Ziphius indicus. 1-8th nat. size. 


extent of 11 inches as a convex ridge, not exceeding 5 lines across at its broadest part. 
The premaxillaries unite with the first six inches of the exposed tract of the vomer, the 
maxillaries (21) with the succeeding five inches. About three inches behind the vomerine 
tract the palatines (20) meet at the mid-line along a suture 2} inches in extent; then 
follow the broad and deep pterygoids (24), extending downward with their lower margins 


12 BRITISH FOSSIL CETACEA 


bending outward from the line of their mid-palatal suture, which is 6 inches in longi- 
tudinal extent. The free lower borders of the pterygoids approximate and thicken as 
they pass backward, diminishing but not obliterating their interval, at the hind end of 
which the thick border of the pterygoid bends abruptly outward for about three inches, 
terminating in a sharp angle answering to a hamular process. The expanded fore part 
of the malar (figs. 5 and 7, 26) shows the beginning of the styliform backward 
continuation. The squamosal (ib., 27) has a 
Fie. 8. prominent, flat, oval facet at the fore part of 
ia the ‘glenoid cavity,’ the concave articular sur- 
face of which is defined at the back part. Other 
anatomical particulars not specially concerning 
the subject of the present Monograph I here 
omit. 

An upper view of the mandible is given in 
cut, fig. 8. The sex of the mdividual affording 
the above-described skull is not known. From 
the size of the alveoli indicating that the ter- 
minal pair of mandibular tooth-germs were 
developed, it was probably a female. In Prof. 
Van Beneden’s specimen the developed teeth 
were but 23 inches (0°065 mm.) in length, 
and consisted chiefly of root, thickly coated with 
cement. 

From the correspondence of structure of the 
upper jaw of the present Cetacean with that in 
the specimens affording to Cuvier the characters 
Se of his genus Zphius, I refer it thereto; the de- 

Mandible of Ziphius indicus. 1-8th nat. size. gree of ossification of the prefrontal or cranio- 

facial cartilaginous constituent of the rostrum, 
with the proportions of the rostrum, I interpret as specific, and adopt the xomen triviale 
by which this existing species of Zphius has been designated by Prof. Van Beneden 
(loc. cit.). 


Zrpmius Layarvi (Doxicnopon, Gray). Plate I. 


Having shown, in Ziphius indicus, a partial ossification of the cartilage continued 
along the groove of the vomer from the septum narium, or coalesced prefrontals, as in 
Z. cavirostris, Cuvier, I now proceed to demonstrate, in another existing species, the 


OF THE RED CRAG. 13 


condition of that upper mid-tract of the rostrum which Cuvier made distinctive, with 
other specific characters, of his Zphius planirostris and Ziphius longirostris. 

I find this character in the composition of the rostrum of the skull of an adult male of 
Ziphius Layard: (Dolichodon, Gray) liberally transmitted, like that of Z. indicus, for 
description, to the British Museum, by the excellent naturalist, Mr. Edgar L. Layard, 
F.L.S., &., now at the Cape of Good Hope, after whom the species is named.' 

In this skull (Pl. I) the premaxillaries (fig. 1, 22’) retain, at their hind part, the sutures 
connecting them with the nasals (15) and maxillaries (21’), where they bound the upper 
apertures of the nostrils; the sutures connecting them with the prefrontal (figs. 1 and 2, 
14) and maxillaries are traceable a short way along the base of the rostrum, and then 
become obliterated, that with the prefrontal being the first to disappear. Upon the fore 
part of the palate (ib., fig. 3) the sutures remain between the premaxillaries (22) and the 
vomer (13), and between the premaxillaries and the contiguous palatine parts of the 
maxillaries (21). 

I may here recall the remark made in discussing the homology of the ‘ prefronrtals,” 
viz., that the toothed Cetacea afford welcome and favorable grounds for determining the 
nature of the mammalian ethmoid through the absence of the olfactory sense-capsules which 
obscure the homologies of the prefrontals in the rest of their class. The ‘os en ceinture, 
Cuv., of Batrachians, and the similarly conspicuous rhomboid tract of the ‘ ethmoide,’ Cuv., 
on the upper and middle part of the base of the rostrum in Struthious Birds, exhibit the 
partially exposed condition of the prefrontals characterising certain species of Z/phius. In 
this, as in other Cetacea, the prefrontals, prior to their coalescence as ‘lamina perpendicu- 
laris, diverge and contribute a small share to the anterior wall of the cranium and a larger 
one to the posterior walls of the nasal passages, of which their produced and coalesced 
parts constitute the partition; they are connate posteriorly with the orbito-sphenoids, and 
usually coalesce with the vomer inferiorly. I have observed the coalesced ‘lamina per- 
pendicularis’ to be cartilaginous in a young whale’s skull where the rest of the walls of 
the nasal passages were ossified. ‘The forward continuation of the ‘lamina perpendicularis 
eethmoidei’ rests upon the groove of the vomer, in a cartilaginous state, in most Cetacea, 
leaving the vacancy in the dry skull along the upper medial line which suggested to 
Cuvier the term ‘cavirostris’ for the Zphius which he first made known to zoologists.’ 
In the present species (7. Layardz) it is ossified, and, becoming superficial and conspicuous 
between the premaxillary nasal processes, expands as it advances, and rises as a smooth, 

1 The want of definitions of bones or sutures in the descriptions and figures by Dr. J. E. Gray, F.R.S., 
of this instructive specimen, in the ‘ Proceedings of the Zoological Society,’ April 11, 1865, p. 358, and in 
his very useful ‘ Catalogue of Seals and Whales in the British Museum,’ 8vo, 1866, p. 354, has not enabled 
me to use or reproduce them for my present purpose. 

2 «On the Homologies of the Vertebrate Skeleton,” ‘Report of the British Association,” 1846, 
p. 226. 


3 The section of the skull of Zuphysetes simus in my paper “On Indian Cetacea,”’ ‘ Zool. Trans.,’ vol. vi, 
pl. xiv, figs. 1, 39, 34,13, Will aid the student of homologies in following the above remarks. 


14 BRITISH FOSSIL CETACEA 


dense, convex ridge (fig. 2, 14) an inch and a quarter across at its broadest part, gradually 
contracting to a breadth of half an inch when it has traversed one third of the length of 
the rostrum. At about the terminal third of this part the outer margin of what seems 
to be the suture between the prefrontal (14’) and premaxillary (22) rises and forms a free, 
thin, inwardly or medially bent margin of bone, which soon appears as the upper and 
outer border of a longitudinal canal, grooving the upper and medial surface of the 
premaxillary, and gradually gaining vertical extent as it passes forward. ‘The under and 
inner margin of this groove is the, here, persistent medial suture or harmonia between the 
premaxillaries, which suture becomes obliterated at about one fourth of the way backward 
from the anterior end of the rostrum; and, thus, any definition of the boundary between 
prefrontals and premaxillaries becomes impossible. The solid terminal fourth of the 
rostrum, in advance of the vomer (13, fig. 3, Pl. I), is formed by the welded prefrontal 
and premaxillaries ; behind this both bones, together with the vomer and the rostral parts 
of the maxillary (ib., 21), combine to form the dense beak-like production of the upper 
jaw, at the base of which are the palatines (ib., 20) and pterygoids (24). 

On the palatal surface of the rostrum (PI. I, fig. 3) the maxillo-premaxillary suture is 
distinct, or linearly traceable, and the vomer (13) intervenes between the contiguous 
palatal portions of the premaxillaries and maxillaries for an extent of one foot, gaining at 
the middle of this extent a breadth of 8 lines, and having a smooth transversely convex 
surface toward the palate. The degree in which the vomer thus appears upon the 
bony palate exemplifies, with other characters, a specific difference in Zphius as in 
Delphinus. 

In Phocena brevirostris? and Euphysetes simus* the limits of the prefrontals and of the 
vomer can be defined; the latter, in these and other Cefacea, is a long spout-shaped 
bone, its canal looking upward, and this receives, posteriorly, the anterior coalesced parts 
of the prefrontals, and, in advance thereof, the cartilage continued therefrom forward. 
This cartilaginous prolongation of the ‘septum narium,’ formed by the coalesced portions 
of the prefrontals, is ossified in the fossil kinds of Zphius to be described, and projects 
conspicuously between the premaxillaries at the upper surface of the rostrum, as it does 
in Z. Layardi. 

In this species the ectomaxillary groove (Pl. I, fig. 1, ge) commences posteriorly 
between the antorbital plate of the maxillary (21’) and the pterygoid (g4’), and is continued 
forward, diminishing in breadth and depth, upon the upper and outer border of the 
maxillary to the fore end of the rostrum, or of so much as remains in the present specimen, 
the extreme tip having been broken off. The lower boundary of the beginning of the 
ectomaxillary groove formed by the pterygoid and maxillary extends outward as a thick 


" Compare, e.g., the skull of Delphinus (Steno) Gadamur with that of D. (Steno) frontatus; or 
the skull of Delphinus (Lagenorhynchus) fusiformis with that of D. (Lagenorhynchus) Pomegra. 

2 Owen, ‘‘On Indian Cetacea,’’ ‘Trans. Zool. Soc., 

8 Tbid., pl. xii. 


* vol. vi; pl? ix. 


OF THE RED CRAG. 15 


convex buttress (), some way beyond the upper boundary, subsiding, as it advances, at 
about the basal fourth of the rostrum. At about the middle of the rostrum the lower 
border of the ectomaxillary groove again projects, but as a sharp, somewhat jagged ridge; 
at the anterior third of the rostrum the upper part of the groove similarly projects, the 
lower one having subsided ; such ridge or ridges I indicate as ‘ectomaxillary,’ but they 
evidently represent anteriorly the alveolar groove of toothed Delphinide. The interorbital 
plate of the maxillary is perforated by the two large apertures (a, 2) for the transmission of 
nerves and vessels, answering to those marked @ and 4 in Pl. IX (Phocena brevirostris), 
Pl. XIII (Luphysetes simus) of my Memoir, above cited, on Indian Cetacea, homologous 
with the suborbital or antorbital division of the fifth nerve in land mammals. The fore 
part of the nasal processes of the premaxillary are also perforated by a smaller canal, 
whence a groove is continued some way forward along with the suture between the nasal 
plate of the premaxillary and the medial (prefrontal) part. 

The outlets of the bony nostrils (Pl. I, fig. 2, 7,7) are slightly twisted from behind 
forward and to the right, the right outlet (7) being the widest; the intervening septum 
shows a corresponding departure from symmetry. The posterior part of the premaxillaries 
(ib., fig. 2, 22) diverge, expand, and rise to define the cavity around the nostrils, or ‘prenasal 
fossa,’ as in all Ziphit. 

The pterygoid is a broad, vertically extended, triangular plate of bone, widely and 
rather deeply excavated in the major part of its extent (fig. 1, 24’), the non-excavated 
anterior apex (24) being wedged between the maxillary and palatine at the lower and outer part 
of the base of the rostrum. 'The inferior thin border of the excavated part is slightly everted, 
and is applied to the cerresponding part of the opposite pterygoid, leaving a deep fissure 
intervening and contracting posteriorly (ib., fig. 3, 24”). The breadth and depth of this 
‘interpterygoid fissure’ varies in the species of Zphius.. The conchoidal part of the 
petrotympanic is bilobed posteriorly, but less deeply indented than in Delphinus ; it is 
rather widely open or unfolded anteriorly, with a thin, compact, involuted wall ; the petrosal 
part articulates by a posterior process to the mastoid. 

Being unable to draw the line of generic distinction at any of the gradations of length 
and slenderness of snout occupying, in this respect, the interval between Ziphius indicus, 
figs. 4—7, and Zphius Layardi, Pl. 1, and meeting the same difficulty in the degrees in 
which the prefrontal cartilage of the rostrum becomes ossified, I hold by the well-defined 
characters of the Cuvierian genus; and, premising the above definitions of recognisable 
parts of the Ziphial rostrum, I proceed to apply them to the specific definition of the 
petrified snouts from the ‘Red Crag.’ 


1 It is wide, e. gy. in Ziphius patachonicus, pl. xvii, fig. 2, 2, of Burmeister, ‘ Anales del Museo publico 
de Buenos Ayres,’ 4to, 1868. 


16 BRITISH FOSSIL CETACEA 


Genus—Ziputvus, Cuvier.’ 


Species—ZIPH1US PLANUS, Owen. Plate LL, fig. 1. 


This species is represented by the basal part (probably third) of the rostrum, including 
the fore part of the right (-) and left (7) prenasal channels or fossz, leading from the 
nostrils to the ‘premaxillary’ grooves or canals (d,d). These concavities present the 
same unsymmetrical twist as in Ziphiuvs Layardi (P\. I, fig. 2, 7 and 7), Z. plani- 
rostris (fig. 2, 22’, 29’), and Z, Cuviert (fig. 3). The disproportionate breadth of the 
right prenarial channel (PI. II, fig. 1, 22’, 7) is greater in the fossil. The correspondingly 
bent prefrontal septum (14, 14) is relatively thicker than in Z Zayardi, and is more 
directly continued into the mid-tract (14’), which is flatter above, than in the recent 
species, resembling in that respect Zphius Cuvieri (fig. 3). The premaxillary grooves 
(d, d) continued forward from the fore ends of the prenasal fosse (r, 7) show, at their 
commencement, the orifice of a small canal, as in Ziphins Layardi, \eading into the 
osseous substance of the rostrum. The prenasal grooves soon subside. ‘The interme- 
diate prefrontal tract is continued on flattened above, not convex, as in Z. Layardi, but 
the generic resemblance is well marked. The posterior part of the maxillaries include the 
large anterior nervo-vascular (antorbital) foramen (4), and show the posterior beginning of 
the ectomaxillary groove (g,g), the upper border of which is developed to form the ecto- 
maxillary ridge (e, e); this speedily subsides. The superiorly flattened mid-tract (14', 14’) 
slightly expands toward the fractured fore end of the rostrum, where its lateral boundaries 
sink into wide and shallow longitudinal channels. The worn surface of the under part of 
this rostral fragment yields no satisfactory character. 

The upper surface of the fossil is represented of the natural size (Pl. II, fig. 1), and 
indicates a species of Ziphius larger than the Z. Layardi. The vertical diameter of the 
rostrum, at 7, is 4 inches. 

‘The specimen was found in the Red Crag at Shotley, Suffolk, and had been split, 
lengthwise and vertically, by a workman, the fissure extending from the right prenasal 
fossa to the middle of the mid-tract (14’, 14’). 


! Synonyms :—E£piodon, Diodon, Heterodon, Dioplodon, Alaima, Delphinorhynchus, Petrorhynchus, 
Dolichodon, Micropterus and Micropteron, Mesoplodon, Mesodiodon, Ziphiopsis, Placoziphius, Aporotus, 
Ziphirostrum, Rhinostodes, Berardius, Hoplocetus, Eucetus, Ziphiorhynchus, Choenodelphinus, Choenocetus, 
Belemnoziphius, &c. (the mob is included in the family Rhynchoceti, Eschricht). 


OF THE RED CRAG. 17 


It is completely petrified, and yields on percussion the clear metallic ringing sound 
characteristic of the Cetacean fossils from this formation; but it is less deeply stained by 


ferrugineous salts than is commonly seen in the fossil snouts from other Suffolk Red 
Crag localities. 


Species—Ziputus cissus, Owen. Plate II, fig. 2; and Plate III, fig. 3. 


Amongst the fossil, as in existing, ZpAii are some species with an excessive develop- 
ment of the ‘mid-tract’ in the upper surface of the rostrum. 

I propose the specific term ‘giddus’ for one of the ‘Red Crag’ snouts, because the 
mid-tract is not only convex transversely (Pl. II, fig. 2, 14’, 14’), but rises by a more 
defined and elevated longitudinal convexity (Pl. III, fig. 8, 14’) than in the other fossil 
species. 

In the specimen exemplifying this character—the subject of the figures cited—a smaller 
portion of the fore part of the prenasal fossee (Pl. I, fig. 2, 29’) is included than in Zphius 
planus. Sufficient of the fore part of the ‘septum narium’ (ib., fig. 2, 14) is preserved to indi- 
cate an unsymmetrical twist to the left, and also to demonstrate that such part of the bony 
septum is absolutely thicker than in the larger existing species, Zphius Layardi, and that 
it is thinner both absolutely and relatively than in Zphius angustus (Pl. II, fig. 2). It 
expands, in Zphius gibbus, immediately beyond the nostrils, and rapidly, into the ‘ mid- 
tract’ (14’), which gains, with some excess on the right side, a breadth of one inch at twice 
that distance from the nostrils, then gradually narrows to three fourths of an inch, and 
again expands both transversely and vertically to the extent shown in the figures cited. 
The subsidence, longitudinally, to the fractured fore part of this rostrum is more rapid 
than the rise, and that, as it appears, without any mechanical influence of posthumous 
abrasion (Pl. III, fig. 3). The proportion of the upper surface of the rostrum contri- 
buted by the narrower part of the mid-tract (14’), at the point marked by the star im fig. 3, 
is shown in the outline of the circumference at that part (drawn with the upper part 
downward) in PI. III, 14’; the much larger proportion which, through its own expansion 
and the subsidence of the side tracts of the premaxillaries (22), the mid-tract (14) forms at 
the part of the rostrum marked by the star in Pl. II, fig. 2, is indicated at 14’, 14’ in 
the superadded outline of the transverse section at that part of the rostrum. 

The nasal processes of the premaxillaries (22', 22’, fig. 2, Pl. II) are perforated, each 
by an orifice (d) relatively wider than in Zphius Layard: (Pl. I, fig. 2, d); and no 
groove is continued forward from the ‘foramen naso-premaxillare’ (¢) in Ziphius gibbus, 
as is the case in Ziphius planus and Ziphius Layardi. 

The commencement of the interorbital expanse of the maxillary (21, fig. 2, Pl. I) in 

3 


18 BRITISH FOSSIL CETACEA 


Ziphius gibbus shows the nervo-vascular foramen (g, 4) on nearly the same transverse line 
as d,d; the downward and backward direction of the canal leading therefrom to the 
common entorbital foramen’ is shown at 4d, fig. 3, Pl. III. 

The pterygoid ale (Pl. III, fig. 38, 24) would seem to be directly applied to the under 
part of these maxillary expansions, and not to be separated therefrom by the fissure seen 
in Ziphius Layardi (P\. I, fig. 1, between 21’ and 24’); but on this poimt the abraded 
condition of the hinder end of the rostrum makes it unsafe to insist. It appears certain 
that the representative of the ectomaxillary ridge (Pl. II, fig. 2, 21...e, and Pl. III, 
fig. 3, e, eg) is much thinner vertically than in Zphius Layardi, and more rapidly sub- 
sides as it advances forward upon the side of the base of the rostrum. A narrow ecto- 
maxillary groove (Pl. III, fig. 3, ey) is continued forward from the point of subsidence 
to the broken end of the snout, gradually sinking toward the lower part of the snout, as 
in Zphius Layard. No sutural evidence of the proportions of the maxillary and premax- 
illary bones remains upon the sides of the rostrum under description. Neither, though 
there be an indication of the structure, can it be certainly affirmed that the veritable 
vomer contributes any proportion of the smooth, transversely convex under surface of the 
rostrum, the curves of which, in transverse section, are indicated in the outlines taken at 
the two points marked in Pls. II and III. 

A medial linear suture marks the meeting of the palatal plates of the maxillaries, 
anterior to the palatine bones, as in Zphius Layardi (P1. I, fig. 3, 20), and this line can 
be traced inclining to the right side, as in the night vomero-maxillary suture in Z. Layardi, 
but the left one is obliterated, if it existed, in Zphius gibbus. ‘The medial plates of the 
pterygoids have met and been in contact for a greater extent in Zphius gibbus than in 
Ziphius Layard, descending to form a deep median angular ridge, to the fore part of 
which the palatines seem to have contributed some share. But here, again, the 
state of abrasion affects the utility of further attempts to specify structural characters. 

Sufficient, however, of the present rostrum has defied the effects of long ages of surf- 
movements, through the degree of petrifaction undergone before dislodgment from its 
original burial-bed, to justify the specific distinction attempted to be pointed out, and, at 
all events, to afford means of comparison which may lead to a reference of other specimens 
of Crag-whale-snouts to the Ziphius gibbus. 


1 Shown at d, fig. 2, pl. 13, ‘Indian Cetacea,’ loc. cit., in Huphysetes simus. 


OF THE RED CRAG. 19 


Species—ZIPH1IUS ANGUSTUS. 


ZiPHIUS ANGUSTUS, Owen. Plate III, figs. 1 and 2. 


The snout on which a species of Ziphius is indicated by the above name exhibits a 
development transversely and vertically of the prefrontal mid-tract (14’, 14’) similar to that 
in the foregoing species, but it is not identical; the convex mid-tract is relatively narrower, 
as is the rostrum itself, in Ziphius angustus, and the narrower part of the tract, after its 
slight hinder expansion, is of greater longitudinal extent before its second or anterior 
expansion. ‘The whole rostrum is more slender, as is indicated by the contour of the 
transverse section given in Pl. III, at the part marked with the star in figs. 1 and 2, 
which corresponds with the part of the rostrum similarly marked in Zphius gibbus (ib., 
fig. 3). Moreover, the ectolateral maxillary ridge (ib., fig. 1,¢,¢’) is longer, extending 
further forward upon the side of the rostrum, and subsiding more gradually. 

The premaxillary foramina (q, d, fig. 2) are as large as in Zphius gibbus. Only 
the imner walls of the maxillary foramina (4, 4, 21, fig. 2) remain; they show them to be 
on the same transverse line as d, d, but to be nearer thereto, indicative of a continuance of 
the slender character of the snout to this commencement of the maxillary platform. The 
ectomaxillary groove (ib., fig. 1, ey) is narrow, and extends forward, descending, as in 
Ziphius gibbus ; but its lower border begins to swell outward (¢’, 21, fig. 1) as it gets toward 
the fore end of the present fragment, as in Ziphius Layardi, indicating that one half or 
more of the rostrum may be wanting in the present specimen. The sides of the pterygo- 
palatine ridge are slightly concave in Zphius angustus, at the part where they show a 
moderate convexity in Zphius gibbus, and it is improbable that this difference can have 
been caused by posthumous abrasion. 

The specimen described is figured of the natural size in the two views given in 
Pl. II, figs. 1 and 2. 

The density of the bony texture of the fractured fore part of the snout closely accords 
with that of the same part in the recent Zphius Layardi, and suggests an application, by 
the living animal, of the snout analogous to that of the similar rapier-like rostral weapon m 
the Sword-fish (Hestiophorus). 


20 BRITISH FOSSIL CETACEA 


Species—ZIPH1US ANGULATUS. 


ZIPHIUS ANGULATUS, Owen. Plate IV, figs. 1 and 2. 


In this well-marked species the prefrontal mid-tract rises in an angular roof-like 
manner, the sides of the upper surface of the rostrum so formed sloping from a median 
ridge at the angle shown in cut, fig. 9, and in the section below fig. 1 in PI. IV. In this 
specimen the maxillo-premaxillary suture (sin fig. 1) is traceable at the expanded basal part of 

Rats: the rostrum, showing the proximity thereto of the (antor- 
bital) nervo-vascular canal (6) in the beginning of the 
expanded part of the maxillary bone. The premaxillary 
foramen (d) is midway between that suture and the 
prefronto-premaxillary suture ; this foramen is smaller 
than in the two preceding fossil species, and in the 
same degree approaches to the character afforded by 
the foramina (4, Pl. I, fig. 2) in the recent Ziphius 
Layardi. The septum narium (14) rapidly expands, 
anterior to the nostrils, to the breadth of the mid-tract 
shown in fig. 1, at two inches in advance of the nostrils. 
From this part the mid-tract advances, maintaining the 
same breadth to the middle of the rostrum, and thence 
eradually contracts, with subsidence of the roof, to the 
broken anterior end. The right premaxillary shows a 
superficial groove (v,») about four inches long, near 
the margin of the mid-tract, due to a branch of 
the main canal (g). The corresponding canal, exposed by the fracture at 2, in the left 
premaxillary, ran deeper, and its superficial branch emerged later or further forward 
upon the surface. The ectomaxillary ridge of the snout (fig. 2, e), commencing at the 
outer and lower wall of the canal (4), extends forward, gradually subsiding, but not to 
effacement, in the preserved extent of the present rostrum, at least on the right side ; on 
the left side it is interrupted at its middle part by the emergence of a branch canal 
from 4, forming an open narrow groove for two or three inches, beyond which the ridge, 
though lower, is resumed. 

The sides of the pterygo-palatine keel-like ridge (fig. 2, 24) is almost flat ; anterior to this 
the lower and lateral parts of the snout show two successive slight swellings, as at y and z, 


Section of rostrum, Ziphius angulatus. 


OF THE RED CRAG. 91 


a 


fig. 2; and the lower or palatal contour rises toward the upper line of the snout, and this in a 
way that seems not to be due to friction. The post-palatal ridge, due to the confluence 
of the medial plates of the pterygoids (fig. 2, 24’), forms a deep and sharp carina. In 
advance of this the vomer appears upon the palate, forming the keel of the ridge there ; the 
composition of the rostrum at this part is shown in fig. 9, where 14 is prefrontal, 13 
vomer, 21 maxillary, 22 premaxillary. 


a) 
i) 


BRITISH FOSSIL CETACEA 


Species—ZIPuivs MEDILINEATUS. 


ZIPHIUS MEDILINEATUS, Owen. Plate IV, fig. 3. 


In the by no means easy task of choosing a name from specific characters shown by 
this class of fossils, I have been led to the one above given by the extent to which such 
mid-line, indicative of a bipartition of the mid- 

Fie. 10. tract (14', 14’), extends along that tract in the 

present specimen. It is present in a feebler 
degree and for a less extent in another specimen, 
afterwards to be described, which may indicate a 
variety of the present species, but such character 
of the mid-tract of this species is not the only 
one by which it differs from the foregomg Zphit. 
With reference to the character of the largest 
and first-described specimen, suggesting for it 
the name planus, it might be objected that a 
convexity of the mid-tract, as shown in Z. gibdus, 
had been worn down by posthumous abrasion. 
So, likewise, it may be said of the present 
species, that the persistence of the median line 
is due to the nonage of the dividual. But, 
if so, it coexists with a flatness of the part of the 
prefrontal mid-tract along which it extends that cannot be the result of abrasion, for the 
sides of the longitudinal fissure are convex ; and yet the degree of convexity is so slight 
on each side of the fissure that the mid-tract gives as flattened a character of the upper 
posterior part of the snout as in Zphius planus. I conclude, therefore, that the difference 
in the character of the mid-tract between the present species and Zphius angustus, Z. 
angulatus, Z. gibbus, to be due to an original and inherent structural specific character of 
the skull in Aphius medilineatus. From Ziphius planus it differs not only in size, but in 
the greater degree of transverse convexity of the upper part of the snout between the fore 
parts of the ectolateral ridges e, ¢ (as, e.g., at the place, *, of the section figured below fig. 3, 
Pl. IV), and in the greater relative longitudinal extent of those ridges ; there is not, besides, 
any trace in Z. planus of the median linear groove characteristic of the prefrontal tract (14’, 14’, 


Section of rostrum, Ziphius medilineatus. 


fig. 3) in the present species. The short oblique irregular fissure on the outside of 14, in 
fig. 3, simulates a suture defining the prefrontal from the mid-tract, as the mid-line along that 


OF THE RED CRAG. 23 


tract simulates a sutural division of the tract itself; but the one is due to fracture, and the 
other to a linear groove, which groove gradually becomes shallower, and is obliterated at six 
and a half inches from the internarial part of the prefrontal. The linear grooves (22, 22, 
fig. 3) dividing the mid-tract from the lateral parts of the premaxillaries disappear about 
the same part of the snout with such mid-groove. The ectomaxillary ridge (e, e) rather sud- 
denly decreases about three inches from its basal beginning, but is continued forward four 
inches before it finally subsides ; a narrow groove extends forward in the same direction to 
the fractured fore end of the snout. The median fissure between the descending pterygoid 
plates is conspicuous in the present species ; they seem not to have coalesced so completely 
as in the palatal carina so formed in previous species. An inferior median ‘ vomerine’ 
tract is indicated by parallel longitudinal grooves along the anterior two inches of the 
palatal surface of the present specimen. ‘The constituents of the rostrum at this part are 
marked in fig. 10 by the same numerals as in fig. 9. 


24 BRITISH FOSSIL CETACEA 


Species—ZIPHIUS TENUIROSTRIS, Owen. Plate V, figs. 1 and 2. 


The first, or basal, five inches of the prefrontal mid-tract in this specimen is more 
convex than in Ziphius planus or Z. medilineatus, but is less convex than in Z. gibdus or 
Z. angustus.. Beyond that point (*, fig. 1) the mid-tract rises with a well-marked trans- 
versely convex surface, and with a slight longitudinal convexity, after which it is con- 
tinued straight to the fractured fore end of the snout, preserving its transverse breadth and 
convexity, as well as the lateral linear grooves or sutural indications of distinction from 
the side-plates of the premaxillaries (22). These plates descend from those sutures almost 
vertically, with a slight outward expansion, to join or coalesce with the maxillaries (21) 
in the extent of the snout preserved anterior to the ectomaxillary ridges. ‘lhese ridges 
(Pl. V, figs. 1 and 2, e, e), when in advance of the intraorbital expansion of the maxil- 
laries, are continued forward for ten inches before final subsidence, the decrease of breadth 
being very gradual. At about eight inches from their origin the nervo-vascular canal, of 
which the ridge e at first forms the outer wall, emerges, the ridge itself being then continued 
onward as the lower border of that canal. The high position of the basal half of the ridges 
makes them seem to terminate laterally the upper surface of that part of the rostrum, the 
transverse contour of which upper surface is sinuous, moderately convex at the middle, and 
concave on each side, as shown in the section at 14’, 14’, and e,e, Pl. V, but in a minor 
degree than at the more basal parts of the rostrum. ‘The side of the rostrum below the 
ectomaxillary ridges swells into a longitudinal convex tract (m, fig. 1, Pl. V). The 
vomer, emerging upon the palate, about half way from the inner end, of the present specimen 
(fig. 1, 13) contributes about half an inch of its transverse extent to the fractured fore part, 
indicating, according to the analogy of Zphius Layardi (P\. I, fig. 3) that this long and 
slender form of snout must have extended about six inches beyond the fractured fore part 
of the specimen above described. 


! Compare the section at the point *, in fig. 1, Pl. V, in regard to the convexity of 1¥,14, with that 
part in the section below fig. 3, Pl. IV, or the sections given in Pl. III. 


OF THE RED CRAG. 25 


Species—ZiPuius compressus, Owen. Plate V, fig. 3. 


The differential character of the present species is shown by the predominance of the 
dimension of depth over that of breadth at every part of the extent of the specimen 
figured. ‘The prefrontal mid-tract (14’, 14’) is transversely convex from its beginning, the 
convexity increasing as it advances; and, from the low position of the ectomaxillary 
ridges (¢, e) and the steep slope thereto of the premaxillaries (22), the mid-tract seems, of 
itself, to constitute the upper surface of the rostrum, almost in the degree shown at the 
part *, fig. 3, the transverse section of which is given below that figure. ‘The upper part 
of the mid-tract is impressed by a median linear groove, filled up about four and a half 
inches from the base of this rostral fragment. But the difference in form and proportion 
from Ziphius medilineatus is strongly marked, and is exemplified in the section outlined 
in the two species in Pls. [V and V. 

On both sides of the rostrum of Zphius compressus the nervo-vascular canal continuous 
with the ectomaxillary ridge (Pl. V, fig. 3, e) is exposed, its more prominent lower 
border representing the continuation of the ridge. The surfaces below the ridges con- 
verging to the pterygoid carina are almost flat. The figure between the pterygoid plates 
is unobliterated, though narrow. The palatal surface is transversely convex in advance 
of the carina, expanding to the breadth shown at the part whence the section is taken 
below (fig. 3). In thisrespect Zphius compressus markedly differs from Z. medilineatus 
and Z. angulatus. 'Vhe proportion contributed by the vomer to the palatal surface is 
much the same as in Zphius tenuirostris ; the disposition and proportions of the other 
constituents of the rostrum, here, are similar to those denoted by the usual numeral 
symbols in the woodcut, fig. 6. The upper and fore part of the present specimen of 
Ziphius compressus lias been subject to an abrading action, which seems to have shaved 
off, obliquely downward and forward, a deep line from that surface: the sloping contour 
(14’, 22’) shown in fig. 3, Pl. V, is due to that accident. 


Remarks ON REcENT AND Fossin ZipHio1D CETACEA. 


Doticnopon, Gray.—According to the practice—principle I have in vain endeavoured 
to discover—whereby genera have been founded on, or generic names given to, recent and 
fossil Ziphioid Cetaceans, as many genera as species might have been made out of the fossil 
snouts selected from the fruits of many years’ gathering and observation of such, in order 
to exemplify what, on the assumption of transmissibility of such characters, I take to be 


specific departures from a primitive ziphioid type. 
4 


26 BRITISH FOSSIL CETACEA 


The claims of Zphius Layardt, for example, to the generic distinction conferred on it, 
under the name Dolichodon, appear to rest on modifications of the mandibular pair of 
teeth, which are described as “very long, produced, arched, and truncated, with a conical 
process in front.” ? 

I have figured the obtusely rounded extremity of the left of these teeth of the natural 
size, in Pl. I, fig. 5. The first calcified part of the tooth-pulp, which still retains a thin 
coat of enamel, must have appeared as the ‘germ’ of the tooth in the young Zphius, 
and was probably one of a series in the alveolar groove. Its formation in the present tooth 
has been followed by a rapid growth of matrix in the antero-posterior direction, which 
became calcified without enamel, and was progressively added to from below until, finally, 
a tooth of 1 foot in length, 13 inch to 2 inches in fore-and-aft breadth, and from 3 lines 
to 6 lines in thickness, was completed, and most probably, as a sexual character. In the 
present specimen about three inches of the base of the tooth is obliquely inserted into a 
cleft of the alveolar border of the mandible (ce, ¢, fig. 4, Pl. I); the base of the tooth 
terminates in a solid jagged border, showing exhaustion of the pulp and cessation of 
further growth, but abrasion of the fore part of the base of the exposed crown of the 
tooth has exposed a shallow remnant of the pulp-cavity. ‘This abrasion seems due, as 
Dr. Gray observes, to “ friction of the upper jaw.”’® The dense rostrum rested in part 
thereon when the mouth is closed, and the teeth curved backward and inward so as to 
embrace the rostrum, as shown in Mr. Trimen’s drawing, * reproduced by Dr. Gray in his 
‘Catalogue’ above cited. 

Many teeth of Cetacea from the Red Crag have reached me showing abrasion of 
the dentinal part or body of the tooth below an apical or coronal part, on which a thin 
layer of enamel is traceable. The analogy of the structure of the mandibular teeth of 
Ziphius Layardi supports a reference of such Crag fossils to the same genus, and I 
have observed nothing in the anterior laniariform teeth of Zeuglodon (or Squalodon) to 
affect such view. 

It is probable that the mandibular teeth, developed as in Zphius Layardi, ave peculiar 
to the male, but the proof is wanting. It is more probable that they exhibit an abnormal 
direction and state of growth in the specimen figured in Pl. I. In any case I cannot regard 
such modifications of form or size of the pair of laniariform mandibular teeth, character- 
istic of Zphius, Cuv., as grounds for further splitting up this well-marked and intelligibly 
defined genus. 

Prrroruyncuus, Gray.—lIt will be seen that I have availed myself of the various degrees 


! Gray, ‘ Catalogue of Seals and Whales in the British Museum,’ 8vo, 1866, p. 353. 

2 Ib., p. 355. Dr. Gray’s expression, “The edges of the front lower teeth are absorbed or worn 
away,’ &c., would leave the inference that there were other teeth behind, which is not the case; he doubt- 
less meant to say “the front edge of the lower teeth,’ but it is a small part only of that edge which is so 
affected. 

5 «Proc. Zool. Soc.,’ April 11, 1865, p. 358, fig. e. 


OF THE RED CRAG. 27 


and forms in which the cartilage continued forward from the prefrontal ‘septum narium,’ and 
resting upon the groove of the spout-shaped vomer, has been ossified in our Red Crag Zphi, 
to indicate their ‘specific’ distinction. Dr. Gray uses this character in the case of 
Ziphius indicus, in which the cartilage called by its definer, Van Beneden, “cartilage vomé- 
rienne ” is ossified, and in the nearly allied, if not identical, species, showing, also, ossifica- 
tion with superior convexity of the ‘ mid-tract’ (figs. 4, 5, 14’), to support the imposition 
of the generic name ‘ Petrorhynchus.’ Jam disinclined, however, to refer my Ziphius gibbus, 
with the like modification of Van Beneden’s ‘cartilage vomérienne,’ to that nominal 
genus.’ 


Having given careful and impartial consideration to the characters which have been 
proposed, or allowed to be inferred, as those of generic value in other instances, I crave 


leave to continue, in reference thereto, to submit the reasons for a different estimate of 
such characters. 


Epropon, Rafinesque (Schmaltz).—One of the characters assigned to this genus by its 
founder appears to have been due either to anomaly or to accidental mutilation of the in- 
dividual :— Sous-famille. Point de nageoire dorsale. G.1, Catodon, Lac. 2, Notaphrum, 
R. do., sp. 3, Hpiodon, R.,” &c. The dorsal fin is characteristically small in Ziphivs, but 
is never, normally, wanting. Rafinesque’s genus has been subsequently accepted with 
rectification of the fin-character, and the addition of a dental one,’ founded on the position 
of the characteristic mandibular pair of teeth, viz. ‘in front of the lower jaw,’ or near the 
anterior end of the symphysis, and on some modification of the shape of those teeth, 


1 “Tn Ziphius ossification extends along the cartilaginous continuation of the prefrontals forward to 
near the end of the premaxillaries.”’"—“ Report on the Archetype and Homologies of the Vertebrate 
Skeleton,” ‘Report of British Association,’ 8vo, 1846, p. 226. I have been gratified by seeing, at length, 
this homology recognised, and trust we shall hear no more of the vomer appearing upon, or occupying 
any part of, the upper face of the cetacean rostrum. “Les fosses nasales ctant refoulées a la base du 
crane, les maxillaires avec les intermaxillaires et le vomer forment une masse compacte de trois os emboités, 
au centre desquels ou trouve ordinairement la partie cartilagineuse de l’eethmoide qui termine en avant la 
colonne vertebrale.”” ‘‘ Rarement ce cartilage est ossifié: nous n’en connaissons des exemples que dans les 
ziphioides.’—Van Beneden and Gervais, ‘Osteographie des Cétacés, vivants et fossiles,’ 4to (texte), fol. 
(planches), p. 4: no date; but the first Part, with pls. i, 11, xxi, xxiii, was received by me Nov. 2, 1868. 

2 «Analyse de la Nature,’ &c., 12mo, Palerme, 1815, p. 60. 

3 Gray, ‘Catalogue of Seals and Whales,’ 8vo, 1866, p. 340. The Author quotes “ Rafinesque, Précis 
Somiol. 13, 1814 (no character),” op. cit., p. 341; but the latter remark would apply more truly to the 
genus Kogia proposed in the ‘ Zoology of the Erebus and Terror,’ “Cetacea,’’ 4to, 1846, p. 22; with the 
additional remark by Rafinesque, ‘‘les noms trop barbares doivent étre modifiés,”’ ‘Principes fondamen- 
taux de Somiologie,’ &c., 8vo, 1814, ‘ Régles Génériques, No. 41, p. 30. It may not be out of place 
to quote the following from the same judicious Naturalist of Palermo :—“ I] est absurde d’indiquer ou établir 
un Genre, sans lui assigner des caractéres, puisque ces caractéres en sont les bases genériques et que sans 
differences charactéristiques il ne pourrait exister réellement; on doit en conséquence les exprimer par une 
définition, toutes es fois que l’on indique un Genre nouveau, sans quoi autant vaudrait ne pas tantaliser la 
curiosité en mentionnant son vain nom.”’—Op. cit., p. 19. I have that confidence in the common-sense and 


28 BRITISH FOSSIL CETACKA 


respecting which I may first remark that each of such teeth is lodged in an elongate 
fissure at the fore part of the alveolar border, as in Zphius Layardi. The mid-cartilage, 
supported by the vomer and continued from the prefrontal septum, does not become ossified, 
and consequently the bony rostrum appears to be excavated above. ‘his character is 
signified, according to its true value, by the specific term cavirostris, applied by Cuvier 
to a form or species of Zphius still represented in European seas. Nor does the dental 
modification appear to me in any measure to justify generic distinction. ‘'l'o the trouble 
of cetologists, however, the Zphius cavirostris of Cuvier has to sustain, not only the 
generic term Lpiodon, put upon it by Rafinesque, but also those of //eterodon (by Lesson), 
of Dioplodon (by Gervais), of A/aima (by Gray), &e. 

Detenroruyncnves, De Blainville.—This name was imposed upon a Cetacean, 
15 feet in length, stranded at Havre, September, 1825, having a rostriform termination 
of the upper jaw, without conspicuous teeth, and with one pair of teeth at the symphy- 
sial part of the lower jaw. ‘The rostrum was composed of vomer, maxillaries, and pre- 
maxillaries, the latter apart at their upper margins, and enclosing a cartilaginous 
prolongation of the prefrontal, resting upon a canaliculate vomer; the basal parts of 
the premaxillaries, diverging and expanding posteriorly, rose to define and include a 
prenasal fossa, bisected by an ossified prefrontal septum, and at the back part of which 
the nostrils opened. Palatines and pterygoids contributed to form the under part of the 
base of the rostrum.’ 

All the characters of Zphius (Cuvier, 1823), with the main specific modifications of 
the cavirostral species, are here manifested. The real gain to zoology was the opportunity 
of defining the external characters of the entire recent representative of the genus which 
the Founder of Palzontology believed to have become extinct. This specimen was 
unfortunately applied to multiply useless names and divert from the completion of a 
knowledge of a most interesting generic form, ably and sagaciously indicated by Cuvier ; 
it will again be referred to in reference to the claims for acceptance of the genus 
Mesodiodon. 


good judgment of my fellow countrymen and labourers in philosophical zoology which leads me to antici- 
pate a tacit burial and oblivion of the barbarous and undefined generic names with which the fair edifice 
begun by Linnzeus has been defaced. 

1 «Bulletin de la Sociéte Philomathique,’ tom. iv (1825), p. 139. 

* F. Cuvier, ‘ Hist. des Cétacés,’ 8vo, p. 116, pls. vii and viii, fig. 1. 


OF THE RED CRAG. 29 


Genus—Brrarnius, Duvernoy. 


Skull of Ziphius (Berardius) Arnouxii, Duy. 


30 BRITISH FOSSIL CETACEA 


This genus was founded by Duvernoy' on the head or skull of a Zphius brought 
from the ‘mers de la Nouvelle Zélande’ by M. Arnoux, chirurgien-major of the corvette 
Le Rhin, commanded by Capt. Bérard, and presented by him to the Museum of Natural 
History in the Garden of Plants in 1846. ‘The animal is stated to have been stranded 
at Port Akaroa, Banks’ Peninsula, New Zealand. It was about 32 feet in length, 
and the skull (fig. 11) measured 4 feet. The cramium is rather more symmetrical 
than usual, and shows, as in Zphius indicus, a beginning of the vertical productions 
of the maxillaries (e) which distinguish, by their full development, the genus 
HHyperoodon. 

The premaxillaries (22, 22’) are divided along the upper part of the rostrum by a 
channel (originally occupied by the wunossified basis of the prefrontals), exposing 
the upper groove of the vomer, and giving to the transverse section of the 
rostrum a deep upper emargination (“la forme demi-cylindrique,” Duvernoy, ib., 
p- 53). 

The prenasal fossze (fig. 11, 22’) contract forward to the entry of the incisive or naso- 
premaxillary canals (4), asin Ziphius planirostris, Cuv. (fig.2, 4, d, p. 5). In the non- 
ossification of the prefrontal cartilage, Ziphius Arnouati participates with Z. Gervaisii, Z. 
micropterus, and Z. patachonichus. The anterior (¢) of the three outlets (a,4,c) of the 
sub-orbital or antorbital canal opens into a better defined depression of the maxillary (at 
ce fig. 11) than usual, but this cannot be interpreted as generic. The ecto-maxillary 
ridges (e) have a rugous exterior,” as in Ziphius Cuvieri (fig. 3, p. 6). The vomer ‘ puts in 
an appearance ’ at the under surface of the rostrum, between the maxillaries and pre- 
maxillaries, for an extent of about 1 foot 3 inches. Herein Z. Arnouaii resembles 
Z. Layardi, but combines the character with a non-ossification of the prefrontal rostral 
cartilage. The mid-pterygoid keel shows a fissure due to non-confluence of the descending 
plates. In all these characters we have specific modifications of an essentially generic 
ziphial type. ‘The value of the mandibular dental character will be discussed in connection 
with the next nominal genus. 

Mersopiopon, Duvernoy—The chief character of this genus is the position of the 
developed pair of mandibular teeth (figs. 12 and 13). On this character I would remark 
that more than one pair of teeth or of gingival tooth-germs are formed in the alveolar 


1 “Annales des Sciences Nat.,’ 3e série, ‘‘ Zoologie,”’ tom. xv, pl. 1. 


’ 


2 Are these the parts defined as ‘‘intermavzillaries, rather swollen on side of blowers,” in Gray, ‘ Catal. 
of Seals and Whales,’ 8vo, 1866, p. 327? 

3 “TIT, Le genre Mesodiodon, nob., caracterisé par l’existence de deux dents développes et alvéo- 
laires 4 la machoire inférieure, qui sont implantées bien en arriére de celles du genre précédent, & peu prés 
au commencement du second tiers de la longueur de chaque branche mandibulaire. Ce genre, distingué 
dailleurs par d’autres caractéres importants, se compose de quatre espéces, dont trois vivantes et une 


fossile.’’—Duvernoy, loc. cit., p. 41. 


OF THE RED CRAG. bl 


groove of the immature Zphii in both upper and lower jaws. But of these rudimental 
teeth the mandibular alveolar pair, calcified and developed, in sexual or other relations, is 
not the same in‘every species. ‘Then arises the question, Is the difference of place—in 
one a little further forward (figs. 8 and 11), in another a little further backward (fig. 12, 
and Pl. I, fig. 4)—significant of those general or wider modifications of structure 
which alone justify, alone make usefully applicable, a generic section of mammals 
with its distinctive appellation? As a physiologist, or guided by common-sense, I 
cannot admit such interpretation of value of an almost functionally insignificant character, 
yielded by parts also which are the seat of such great and singular variability in the 
Cetaceous order. 

Sometimes two pairs of this series of rudimental mandibular teeth are developed, as 
in FHyperoodon, Lacép. In Hunter’s specimen (from 30 to 40 feet long, No. 2479, 
Mus. Coll. Chir.), as in that figured in my ‘Odontography,’ pl. Ixxxviii, fig. 1, 1, 2, 
the two teeth m each ramus are approximate, and the anterior one is much the 
smallest ; in Lacépéde’s specimen (a male, 22 feet long) they are further apart, the 
foremost answering to the second more developed tooth in No. 2480, Mus. Coll. 
Chir. 

In the Aphius from New Zealand (fig. 11) two teeth are developed in the same 
position in each ramus as in LZyperoodon Butzkopf, Laccép. (H. Baussardi, ¥. Cuv.). 
Thereupon is proposed the genus Berardius. Duvernoy remarks—“ Tl a, entre autres, 
pour caractére quatre fortes dents triangulaires comprimées a l’extremité de la méachoire 
inférieure ” (loc. cit., p. 41). But the term ‘a |’extremité’ would apply more accurately 
to Hunter’s Ziphioid than to Arnoux’s. 

As to the inadequacy of this dental character of two pairs attaining calcification and a 
certain degree of growth, the same remark applies as to the generic importance of the 
accident—for it can hardly be called otherwise—whether the fore or the hind pair of 
such teeth rest rudimentary. 

Of any other generic characters (‘entre autres’) of his Berardius Duvernoy is 
silent. 

Assuming the minor degree of symmetry in the naso-maxillary part of the skull to be 
constant in the species, and not an individual variety, on what intelligible ground can 
such small modification of form and direction in bones, confessedly presenting in all 
essential respects the characters of other ZpAiz, sustain the imposition of a generic 
name ? 

The intermediate step, shown by Zphius Arnouwii, in the development of the parts 
of the maxillaries (fig. 11, e, e), which forms, perhaps, nits extreme degree, the best reason 
for accepting Hyperoodon as a nominal indication of the species of Ziphioids with such 
fully developed processes, is significant of the derivative bond uniting all the Ziphioid 
family. 


BRITISH FOSSIL CETACEA 


Fig. 12 represents the skull of a Zphius' stranded near Havre, at the mouth of the 


Ziphius microplerus. 


(Mesodiodon micropterum, Duv.) 


Seine, which is said by Suriray to have shown four 
parallel Jongitudinal furrows in the skin of the throat, 
about half an inch deep. 

It would be well to look for this structure in other 
recently stranded Zphii.2 It may be a distinctive 
character, but would then be cutaneous and specific— 
nothing more. 

Compared with Zphius Sowerbit, Z. micropterus is 
longer and more slender. The base of the premaxillary 
(22') shows the infundibular expansion of the begin- 
ning of the naso-premaxillary canal (d) (as in Zphius 
planirostris, Cuv.).° 

The rostral parts of the premaxillaries are separate 
above, the ossification of the prefrontal gristle not ex- 
tending beyond the part marked 14, in advance of 
which is exposed, as usual, the deeply situated cana- 
liculate surface of the vomer which supported the 
cartilage. 

The maxillaries (21) show one large (4) and several 
small nervo-vascular outlets (a—c) and the lateral ridge (e). 


1 This skull, ascribed to a genus Delphinorhynchus, is described 
as the type of the Delphinide in the ‘Histoire Naturelle des 
Cetacés’ of Fr. Cuvier, 8vo, p. 75 et seq. 

2 Dr. Gray ascribes to the Ziphius (Delphinorhynchus) micro- 
pterus stranded at Ostend in 1835 the character of “ throat with 
four parallel slits beneath ” (‘Catalogue of Seals and Whales,’ 8vo, 
1866, p. 352); but I do not find this noticed in the memoir by 
Dumortier, ‘Mémoires de l’Académie Royale de Bruxelles,’ tome 
xii, containing the only description of this recent Ziphius with which 
I am acquainted. The incidents ascribed by Gray to “the female 
caught at Havre, on the 22nd August, 1828” (‘Catal. of Seals and 
Whales,’ p. 352), are suspiciously like those narrated by Dumortier 
of his Ostend specimen :—“ Sa longueur totale, depuis l’extremite du 
museau jusqu’a celle de la queue, était 3 metres 45 centimetres, ou 
environ 11 pieds.’? “Le Delphinorhynque d’Ostende fut conservé 
vivant hors de ]’eau pendant deux jours, mais sans rien vouloir manger. 
En vain voulut on lui offrir du pain humecte et d’autres substances 
alimentaires, il les refusa constamment. Souvent il poussait de forts 
mugissements,” &c.—Dumortier, loc. cit., pp. 5 and 6. 

3 «T’entré d’un canal, légérement creusée en entonnoir.”’— 


Cuv., ‘ Oss. Foss.,’ p. 58. 


—" 


OF THE RED CRAG. 33 


The sex of this individual is not noted ; it differs from Sowerby’s species (a male, fig. 
13) in the much smaller size of the mandibular pair 
of teeth, which are similarly situated ; but the speci- 
men was rather larger. The smallness of the dorsal 
fin in both is indicated by the term ‘ mzcropterus,’ 
applied to the Havre Ziphial. This character 
may be more widely or generally manifested in the 
genus Ziphius.’ 

In the Anatomical Museum of Christchurch, 
Oxford, is preserved the portion of skull (fig. 14) of 
the species of Zphius on which Sowerby founded 
his Physeter bidens, with a clear apprehension of 
its close and intimate affinities to the Cachalots. 
The specimen had been stranded on the west of 
Scotland (Elginshire ?). In the skull of this species, 


as in Ziphius cavirostris, Cuv., the premaxillaries 
(22) are separated from each other above for nearly 
the whole length of the rostrum, exposing (at 13) 


the upper canaliculate surface of the vomer, but 


including the proportion of the prefrontals which 
have been ossified into a dense convex tract at 14. 
The departure from symmetry in the naso-maxil- 
lary part of the skull is rather greater than in 
Ziphius Arnoux, rather less than in Z. Layard. 


' See, e.g., Dumortier, loc. cit., plate i, and the figure of 
Ziphius (Ziphiorhynchus) patachonicus in Burmeister’s ex- 
cellent ‘Anales del Museo Publico de Buenos Ayres,’ 4to, 
entrega quinta, 1868, pl. xv. On the 26th of May I was 
favoured by WiLL1AM ANDREws, Esq., M.R.I.A., with a copy 
of his Memoir, from vol. xxiv (1869) of the ‘Trans. of the 
Royal Irish Academy,’ on Ziphius Sowerbii, “ cast ashore in 
Brandon Bay, County of Kerry,” Ireland. The specimen, 
like that found on the Elginshire coast, was a male; it ‘ was 
about fifteen feet in length” (p. 7); the blow-hole was 
crescentic, with the horns turned forward; the mandibular 
tooth entered a notch of a thick padding of fibro-cartilaginous 
substance covering the alveolar border of that part of the 
upper jaw; it showed the usual sexual development and 


Ziphius Sowerbii. 
specific position; the summit of the crown was visible exter- Physeter bidens, Sow. 


nally when the mouth was shut. Unfortunately the skin of (Mesodiodon Sowerbii, Duv.) 


(Dioplodon Sowerbiensis, Gervais.) 
: ane ; (Diodon Sowerbii, Jardine and Bell.) 
fins, as to proportion, form, and position, were not subjects (Mesoplodon Sowerbiensis, Gervais.) 


of observation. 5 


the throat, as to the presence or otherwise of folds, and the 


34 


BRITISH FOSSIL CETACEA 


Prof. Van Benepen figures a mandible of Ziphius Sowerbii' after a photograph 


Ziphius densirostris. 
Mesodiodon densirostre, Duy. 


of the original in the Museum of Christiania, trans- 
mitted to him by Prof. Boeck.’ The teeth are small, 
and situated a little nearer the hind part of the 
symphysis than in the Havre specimen (fig. 12), 
but it may well be a mere individual variety. ‘The 
specimen was found on the coast of Norway, the sex 
unknown—may be inferred to have been female from 
the size of the teeth’ in the male, fig. 13, from our 
own coast, and as contrasted with the size of the teeth 
in the female specimen stranded at Ostend, and de- 
scribed by M. B. Dumortier.? 

Of the Cctacean representing the Mesodiodon 
densirostris of Duvernoy, the sex is unknown; the 
skeleton was sent by M. Leduc from the Séchelles 
Islands to the Garden of Plants, Paris, in 1839; it 
was most probably a male, showing a greater relative 
size of the developed pair of mandibular teeth than in 
Ziphius Sowerbii, with concomitant depth of the part 
of the jaw supporting them, chiefly due to growth of 
the alveolar border. But all this testifies to no more 
than specific value, if, indeed, it truly means that. 
The position of the teeth, more remote from the sym- 
physis, tells better for specific distinction than their size. 

A further degree of departure from symmetry 
than in Ziphius Sowerbii and Z. micropterus is mani- 
fested in the naso-maxillary region (fig. 14, 29’). 
The prefrontal (14) is ossified throughout its rostral 
extent, forming the summit of the ridge, a form which 
the compression and upward convergence of the pre- 
maxillaries (22) give to the upper surface of the ros- 


1 «Note sur un JMesoplodon Sowerbiensis de la céte de 
Norwege,” in ‘Bulletins de l’Acad. R. de Belgique,’ t. xxii, 
1866. Dr. Gray, in the ‘Cetacea of the Erebus and Terror,’ 
pl. v, figs. 1 and 2, also follows De Blainville (‘Nouvelle Dict. 
d’Hist. Nat.,’ t. ix, p. 177) in giving a termination usually 
significative of locality —“ Sowerbiensis ;’ I retain the common 
form, indicative of the individual—Sowerbi. 

2 «Mémoires de l’Académie Royale de Bruxelles,’ 4to, tom. 
xii, 1839. 

3 Ib., pl. ii, fig. 4. 


OF THE RED CRAG. 35 


trum. The upper mesial margins of the premaxillaries, toward the end of the rostrum, rise 
and converge above the prefrontal as in Ziphius Layardi, to which the present species is 
closely allied. ‘The degree to which the end of the prefrontal subsides, the state of con- 
fluence therewith of the premaxillaries, and the extent to which they are naturally or actually 
apart at the end of the rostrum, can be determined only by inspection of the specimen 
itself. ‘lhe rostrum shows the usual ivory-like or petrosal hardness indicated by the 
specific name.’ ‘I'he under part of the rostrum is convex transversely, and part of the 
vomer is there exposed. ‘The antorbital foramen (e) is near the premaxillaries; the entry 
to the premaxillary canals (2) has the usual infundibular shape. Duvurnoy approximates 
Ziphius longirostris, Cuv., to Mesodiodon densirostris, because it shows a similar ossification 
of the prefrontals (which he calls ‘vomer’) along the upper mid-line of the rostrum. 
But we see, as in Ziphius indicus (fig. 8), that this condition may be associated with 
a comparatively short and thick rostrum. 

The Mesoplodon of Gervais is the Mesodiodon of Duvernoy. I may add that, at 
p- 60 of his Memoir (loe: cit.), Duvernoy cites, as one of the generic characters of 
Mesodiodon—‘ Les intermaxillaires, élargis a la base du rostre ont le trou, en entonnoir 
que distinguent les espéces de ce genre ;’ but on this ground Chonezipiius must be trans- 
ferred to Mesodiodon. In truth, however, the funnel-like expansion inte which the 
premaxillary foramina (@) open characterises, with a certain range of variety, the Ziphials 
generally. 

Cuoneztpuius, Duvernoy.—Of Cuvier’s Ziphius planirostris Duvernoy makes his 
genus Choneziphius, acknowledging, however, that the premaxillaries ‘‘ sont creusés d’une 
cavité en forme d’entonnoir” (p. 61, as, mdeed, Cuvier had expressly pointed out), 
which character Duvernoy had previously made a distinction of the species of his genus 
Mesodiodon, which includes, apparently, on that account, Cuvier’s Zphius longirostris. 

The dense petrous ossification of the mid upper tract of the rostrum, due to the 
prefrontals, Duvernoy attributes to the ‘ premaxillaries’ (p. 61), as in Zphius longirostris 
he ascribes it to the ‘ vomer’ (p. 60). 

Puacozipnius, Van Beneden.,—This genus is represented by a fragment including the 
base of a fossil rostrum discovered by Colonel Lu Hon in an old deposit (‘le crag noir” 
or stage of the Neozoic series, at Edeghem, near Antwerp. In the breadth and superior 
flatness of the preserved back part of the maxillaries this fossil resembles Zphius planirostris 
and Z. planus. The prefrontal cartilage has not been ossified ; consequently there is, as in 
Ziphius Sowerbit, fig. 13, and Z. Arnowaii, fig. 11, a vacant channel (“le canal vomérien,” 
V. Ben.) between the premaxillaries, the bottom of which canal is formed by the superior 
vomerine groove. ‘The expanded prenasal parts of the premaxillaries show the foramina 


' Originally imposed on it by De Blainville. 
2 Van Beneden, “Sur un Nouveau Genre de Ziphioide fossile (Placoziphius),’’ in ‘Mémoires de 


Academie Royale des Sciences, &c., de Belgique,’ tome xxxvii, 1866. 


36 BRITISH FOSSIL CETACEA 


answering to those marked ¢, in the illustrations of this Monograph (Zphius Layardi, 
Pl. I, fig. 2, a, et passim), opening or commencing, as usual in Zphius, at the fore end or 
apex of an infundibular cavity. The right premaxillary is unsymmetrically expanded in 
the narial region. ‘I'he maxillaries are too much mutilated to show the position of the 
antorbital foramina (4, 4, in the figures of the present Monograph), but the outer margin 
of the maxillaries is grooved, as usual, by “un sillon trés-marqué,”’ answering to e, g, of 
the same figures. 

The vomer puts in a carinate appearance on the palate, nearer the palatine bones than 
in Ziphius Layard; but this is a difference of barely specific value when unassociated with 
better characters. ‘The constituent bones of the fossil fragment of rostrum described and 
figured by Prof. Van Beneden (op. cit.) appear to have been in some degree separated 
from each other, indicative, like the separate state of the atlas, of the nonage of the indi- 
vidual. How far the non-ossification of the prefrontal cartilage may bear the same inter- 
pretation may be questioned. 

The rostrum is restored (pl. i, fig. 1, vol. cit.) by continuing forward the lateral 
contours of the maxillary parts of the preserved base of the snout with the same degree 
of convergence to the problematical eud, which, of course, makes the snout abnormally 
short for a Ziphioid. One sees, however, how deceptive such basis of restoration may be, 
by continuing, in like manner, the lateral contours of the expanded parts of the maxilla- 
ries forming the base of the prolonged rostrum in most of the ZpAzi figured in the present 
Monograph. 

This at least may be safely affirmed, that it is somewhat premature to propose a new 
genus of Cetacean on the fragmentary and decomposed fossil representing the P/aco- 
ziphius Duboisii of Van Beneden. 

Zipuiopsis, Du Bus.—The first character assigned to this genus‘ raises the question 
whether ‘proportionate length of rostrum’ be available for generic distinction. ‘The inade- 
quacy, to that end, of ‘ direction of rostrum,’ in view of the slight deviations from straight- 
ness hitherto observed in recent and fossil Ziphioids, has already been mooted. I note, 
also, that examples cited of the genus Zphiopsis do not give the length of the rostrum. 
Z. phymatodes, e. g., is represented by a fossil rostrum with the end broken off.’ 
If the end of the rostrum of Zphius Layardi, e. g. (Pl. I, fig. 3), were broken off 
anterior to the vomer (13), it might seem to have had one of but moderate length. 


1 « Rostre de moyenne longueur, droit, 4 peu prés aussi haut que large. Maxillaires supérieurs trés- 
epais. Canal vomerien petit. Incisifs médiocrement développés, & bords internes soudés ensemble dans 
toute leur partié rostrale, depuis la fosse prénasale jusqu’a la pointe.” —‘ Bulletin de l’Académie Royale des 
Sciences, &c., de Belgique,’ tom. xxv, p. 628. 

» «Le vomer apparait a la surface palatine, vers le milieu de la longueur du museau, et disparait vers 
son extrémité, entre les orifices antérieures des canaux palatins. Un peu au dela la picce est brisée.”— 
p. 628. A second specimen is only “un peu plus complet.”—Ib. My experience warns against inferring 
the length of the rostrum in a Ziphius in a specimen not having that part entire. 


———— 


.) lS a 


OF THE RED CRAG. 37 


As to the character of interconfluence of the premaxillaries at the region of the pre- 
nasal fossa, it may be permitted to doubt whether an intervening ossific production of the 
prefrontals, as at 14, Pl. J, fig. 2, Pl. IV, fig. 3, and in woodcut, fig. 13, 14, has not 
been so interpreted. 

Of the second representative of Zphiopsis (Z. servatus) the author knows but a 
single fragment of rostrum ‘presque entier a sa pointe’ (I. c., p. 629), and broken off in 
front of the prenasal fossa; it shows the same palatal portion of vomer as in Ziphius 
Layard. 

Rainostopes, Du Bus.'—This genus is founded on a less compact or spongy texture 
of the fossil bones forming the mid-part of a much mutilated rostrum. 

Zieuirostrum, V. Ben.°—In this genus the rostrum is straight, or bent slightly 
upward at the point. The maxillaries ere more or less thick. ‘The premaxillaries 
(‘incisifs’) are confluent along their inner borders at the mid-part of the rostrum, but 
distinct at the point. The vomerine canal is open; 7.e¢. the premaxillaries are distinct 
or apart at their upper borders, the interspace not being occupied by an ossified prefrontal 
cartilage. 

It is well that the direction of the rostrum is not seized as a generic character. 
Whether the degree of ossification of the prefrontal gristle be more entitled to serve as 
such is questionable. In some Zphirostra the vomer appears as part of the palate, and, 
as usual, “un peu en avant du point occupé par lextremité antérieure des palatins” (loc. 
cit., p. 624, Zphirostrum marginatum, Du B.); and “ dans le tiers moyen de la longueur 
de celui-ci” (ib., p. 623, in Ziphirostrum Turninense). 

As to the confluence of the inner (mesial) borders of the premaxillaries, this, which 
first affects their lower margins, is an affair of age and a matter of degree, in fossil 
Ziphioids difficult to determine with any degree of certitude. 

Aporotus, Du Bus.—This genus’ has the same characters as Zphirostrum, save that 
the inner borders of the premaxillaries in their rostral extent remain applied to one another 
without ever (?) becoming confluent. 


1 « Rhinostodes Antwerpiensis, Du Bus. II n’existe au Musee qu’un seul fragment de téte de cet 
animal; c’est la partie moyenne d’un rostre extremement mutile.” p. 629.—Recent Ziphii, as well as fossil 
ones, exhibit degrees of difference in the density of the tissue of the bones forming the rostrum. The 
prefrontal production appears to have been ossified in the portion of rostrum representing the genus 
Rhinostodes, Du B. 

* «Genre Ziphirostrum, Van Beneden. Rostre droit ou légerement releve vers la pointe. Maxillaires 
plus au moins épais. Incisifs soudés ensemble par leur bords internes, dans la partie moyenne du museau, 
mais séparé a la pointe. Canal vomérien ouvert,”—“ Sur differents Ziphiides nouveaux du crag d’Anvers,”’ 
par M. le Vicomte Du Bus (‘Bulletin de Academie Royale des Sciences de Belgique,’ 2e serie, tome 
xxv, 1868). 

3 “Le genre Aporotus a les mémes caracteres que le genre Ziphirostrum, sauf que les bords internes des 
incisifs, dans leur partie rostrale, restent appliqués l’un contre l’autre et ne sont jamais soudés ensemble.” 
Loc. cit., p. 626. 


38 BRITISH FOSSIL CETACEA 


The confluence or otherwise of borders in contact, even if known with certainty to be 
unaffected by the age of the individual, is not admissible as a generic character. Vte. Du Bus 
exemplifies his nominal genus exclusively by fossil rostrums. In some of the Zphi 
associated under this head the vomer appears on the palate (4porotus recurvirostris), in 
others not (Ap. afinis). 

ZipuioRuyNcuvus.—Ziphiorynchus, Burmeister, founded on a female Ziphius stranded 
on the shore at Buenos Ayres, with a rostrum resembling in premaxillary and prefrontal 
modifications that of Ziphius Gervaisii, rests on the minute and concealed germs of 
teeth’ for its generic distinction. One (terminal) pair of mandibular teeth are more 
developed, but only in the degree characteristic of the female sex. Burmeister has 
recorded valuable anatomical observations on this Zphius; it accords with Ziphius 
(Delphinorynchus) micropterus’ in the small size and position of the dorsal and pectoral 
fins, the dorsal being rather further back in Ziphius patachonicus than in Z. micropterus. 
The caudal fin is widely emarginate in both. 

BrLemNozipuius.—Professor Huxley contributes his mite to this array of names by 
proposing the genus Belemnoziphius for those species in which “the vomer occupies fully 
a third of the width of the upper face of the rostrum,” the extremity of which “is entire, 
not bifid, but sharply pointed, almost like the end of the guard of a Belemnite, the vomer 


294 


and premaxilla seeming to coalesce into one solid terminal cone.”* Unless the prefrontal 
has been mistaken for the vomer, I have not seen any specimen of Zpdius or Ziphioid 
Cetacean presenting these characters, and can only remark that they appear to exemplify 
rather a phase of anatomical knowledge of the individual than a power of recognising 


a genus. 


A few words may be permitted, in conclusion, in reference to the first-described 
specimens on which our earliest knowledge of Cetacean remains in the Red Crag deposits 
was founded. 

Dr. Bowerbank or Mr. Charlesworth would be able to say how long, and through how 
many hands, the fossil (fig. 226, p. 536, ‘ British Fossil Mammals,’ 1846) in Mr. John 
Brown’s (F.G.S., of Stanway) Collection, now in the British Museum, Register No. 
27,862, had been passing, without any clue having been caught as to its nature, before the 
amorphous waicum came, in 1840, into mine, with permission from the possessor to 
slice and apply to it the microscopic test. If the paleontologist cares to turn to the 


1 «Annals and Mag. of Nat. Hist.,’ 1866, pp. 94 and 303. 

* *Anales del Museo Publico de Buenos Ayres,’ entrega quinta, 4to, 1868, pl. xix, fig. 4; pl. xix, 
fig. 6. 

3 « Mémoires sur le Delphinorhynque microptére échoué 4 Ostende,” par B. C. Dumortier (‘ Mémoires 
de |’Académie Royale de Bruxelles,’ tom. xii, 1839). 

* « Proceedings of the Geological Society,’ May 25th, 1864 (‘Quarterly Journal of the Society,’ vol. xx, 
p- 392). 


OF THE RED CRAG. 39 


page above cited, he will be puzzled to understand the note purporting to afford the 
‘unde derivatur’ of the generic name under which it is there described. ‘The truth is that, 
having detected its nature as a tooth, and the affinity in structure to the physeteroid type 
of Cetacean teeth, I suspected it might have belonged to one of the ziphial divisions of 
Physeteride, and named it in the first instance Ziphiodon, as the note tells, from “Ziphius, 
generic name of a fossil Cetacean, and odove, a tooth.” 

Subsequent acquisitions of fossil petrotympanic bones from the same deposit and 
locality as those to which Mr. Charlesworth had traced the tooth in question, and some of 
the larger of which ‘ Cetotolites’ showed the more simple balznoid type of form as con- 
trasted with the division of the involute convexity into an outer and inner lobe, and the 
non-continuation of the overarching wall around the inner end of the cavity, characteristic of 
‘Cachalots’ and Ziphial ear-bones, induced the suspicion that the fossil tooth in question, from 
its agreement in size with those baleenoid ear-bones, might appertain to some such species, 
more especially as the cetotolites of the Cachalot type from the ‘Red Crag’ were much 
smaller in size; and, at the last hour of going to press, I gave this doubt its benefit ; 
but omitted to erase from the proof-sheet the note explaining the first imposed generic 
name. In this stage of knowledge of detached, fragmentary, and scattered fossils 
one can only suggest a guess, nor did I assume to give more, by the light of the 
analogies detailed in pp. 340 and 341, than “a dim and distant view of the actual 
generic characters of creatures revealed to us by a few fragments of their fossilized skele- 
tons, which have been bruised and worn by ages of elemental turmoil.’”! 

Unremitted attention to the Mammalian fossils of the Red Crag has failed, hitherto 
(1868), to add to the grounds for determining the generic nature of the tooth referred by 
me, in 1840, toa Zphiodon, and subsequently, in 1846, to a Balenodon. 

It is affirmed by E. Ray Lankester, Esq., in a paper on Crag Mammalia, in ‘Quart. 
Journal Geological Society’ for February 8th, 1865 (p. 231), on the authority of 
M. Van Beneden, “that the Balzenodon teeth of the first form (that originally described) 
are, without doubt, the teeth of the bident lower jaws of those Ziphioids whose remains 
occur with them in the Red Crag.” 

The reception of the evidence, when the experienced Cetologist of Louvain may have 
the leisure to publish it, proving beyond doubt that the tooth im question belongs to a 
Ziphius of the Red Crag, will be most acceptable. In the meanwhile, however, I cannot 
help resting in the same state of uncertainty, oscillating, as it were, between Ziphiodon’ 
and Balenodon, as from the years 1840 to 1868. And TI have the greater difficulty in 
extricating myself from this expectant state of mind, because of the number of distinct 
species of Zphii which, in that interval of time, I have been able, I trust, to define to 
the satisfaction of paleocetologists. When M. Van Beneden’s demonstrations reach us, 
we shall know to which of the species (or Grayan genera) of British Red Crag Ziphials 
the tooth of Balenodon belongs. 

1 Op. cit., p. 541. 


40 BRITISH FOSSIL CETACEA OF THE RED CRAG. 


There is a vagueness, unfortunately, in Mr. Ray Lankester’s announcement of 
M. Van Beneden’s discovery that of itself begets hesitation. Mr. Brown’s old fossil cannot 
be one of the “teeth of the bident lower jaws of ‘all’ these Ziphioids whose remains occur 
with them in the Red Crag.” At least, the analogy of the range of variety in size, shape, 
direction, and position, of the teeth in the bident lower jaws of existing species (genera) of 
Ziphials would lead us to infer something of the same kind in the fossil species (genera). 
Then, again, recurs the undoubted fact that Cefacea of another family, so far as cetotolite 
characters teach, have also left their remains in our Red Crag. 

When M. Van Beneden adduces as an argument, removing all doubt of the 
belonging of a Balzenodon tooth to the Red Crag Zphiz, that it “ occurs with them in the 
Red Crag,” does he mean to say, as his reporter would have us to understand, that they 
are so associated in closer and more demonstrative relation with the unquestionable snouts 
of Ziphii than they are with the unquestionable ear-bones of Balenoids? I have not, 
as yet, obtained the requisite evidence of the fact, and I may aver to have had as much 
personal experience in exploring and cellecting from Red Crag localities as either M. Van 
Beneden or Mr. Ray Lankester. 

Again, the latter affirms, on M. Van Beneden’s authority, with respect to the 
second kind of cetacean Red Crag teeth, ‘‘ more elongate and with an emarginate mipple- 
like crown of enamel,” that they are, without doubt, the teeth of a species of Sgualodon." 

The grounds that lead me still to entertain doubts on this point are given at page 26, 
in connection with comparisons with a recent Ziphial tooth (see Pl. I, figs. 4 and 5) 
having unquestionably the character above cited ; and I may add that, of the truly cha- 
racteristic and most numerous compressed, two-fanged, serrated teeth of Zeuglodon (= 
Squalodon, V. B.), I have not as yet seen any specimen from our English Red Crag 
deposits. No doubt it is a telling, as it is a sweeping, conclusion from the on dit of the 
experienced and accomplished Louvain cetologist that ‘ Balenodon physaloides’ must be 
** removed from the list of our British fossil Mammals ;’—hbut it is not science. 


1 «Quart. Journ. Geol. Soc.,’ 1865, p. 231. 


or 


PLATE I. 


Ziphius Layard. 


. Side view of skull. 
. Upper view of ditto. 
. Under view of rostrum, with section of ditto at *, in outline. 


.. Side view of mandible (parts broken restored in outline). 


These figures are 1-6th nat. size. 


. Summit of developed mandibular tooth, showing enamelled crown or portion, a, 


nat. size. 


. Transverse section of mandibular tooth, nat. size. 


A recent species, from the Cape of Good Hope. 


[The letters and figures are explained in the text. | 


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PLATE II. 


Ziphius planus. 


Fic. 
1. Upper view of basal part of rostrum. 


Ziphius gibbus. 
2. Upper view of rostrum, wanting the extremity. 


2*. Outline of transverse section of rostrum at the part marked * in fig. 2. 


Fossils, from the Red Crag of Suffolk. In the British Museum. 


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PLATE III. 


Ziphius angustus. 


Fic. 


1. Side view of basal portion of rostrum. 
2. Upper view of ditto. 


2*. Outline of transverse section of ditto at the part marked * im fig. 2. 


Ziphius gibbus. 
3. Side view of basal portion of rostrum. 


3*.Outline of transverse section at the part marked * in fig. 3 (upside down). 


Vossils, from the Red Crag of Suffolk. In the British Museum. 


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PLATE IV. 


Ziphius angulatus. 


Fic. 


1. Upper view of rostrum, wanting the end. 
2. Side view of ditto. 


2*.Outline of transverse section at the part marked * in fig. 1. 


Ziphius medilineatus. 
3. Upper view of rostrum, wanting the end, 


3* Outline of transverse section of ditto at the part marked * in fig. 3. 


Fossils, from the Red Crag of Suffolk. In the British Museum. 


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PLATE V. F 
Ziphius tenurrostris. 


Fic. 
1. Side view of rostrum, wanting the end. 
2. Upper view of ditto. 
2*,Outline of transverse section of ditto at the part marked * in figs. | 
and 2. 


Ziphius compressus. 


3. Side view of rostrum, wanting the end. 


3*. Outline of transverse section of ditto at the part marked * in fig. 3. 


Fossils, from the Red Crag of Suffolk. In the British Museum. 


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AMNH LIBRARY 


iii 


100126826