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Poeetactid ent Be wees le Sc MENG ie a nts © Vey iW ¥ vdeo 7 8 pte 4d Wide “yl Vie na 4 ; Lee | Ll RORORS eo et | aera yiiay dd : va Weveweveven oc eh vee wie ceo tse i vis Le i | WW ao vey hee oN i ieae : deh ‘ “bare hth Were SUL uuri) Spel th ne Legpent y peer “wtannel ng Wet" ps By MRSC Cec aac OOP i int Plo aed Bee Deal Do y Vivir ivy YA ee ee gg ¥e to Ney ve wwe Wh muy see ahel PEE lets babi ty og al id Ae | os oJS MAA J eu LATA “ah vvwry iy bevel e iad wd bsp EEN pel hd A het ae ss | i ! ye aA | ve i) v Ad rg 2 YUsSy weed? wv bee hela heh yy : Oh aacaeen * y vy Nees ca Ler. | Fwy Ch aheihd EMS A a ete tad Ai vd Wider 20 oo dy awwees WIS dd NO IY ¢ z SOS hdd yer evd dil - ka vy J ro he} wv" vw ‘ Tart 4 riot ah ; . EM . J wd / “¥ / het | . vty } AS Sah had MYM Hts re. Sf Pa ak = Were yOES yd Jie a vu v wwe A pet i d WV sh NEP 3 wv CH Sd fo, - .* . . a © ~4 © Sa eg "% } vue dV vid Udy \ bbe 3) “ew te ath pe oe ‘Vv i ¥ ew ed vice ete be Cel a aa Vey ad Oiitty ye ee Vow PALAONTOGRAPHICAL SOCIETY, INSTITUTED MDCCCXLVIL. LONDON: MYPCCOLXVI—MDCCCLXXII. MONOGRAPH OF THE BRITISH PLEISTOCENE MAMMALIA. ORDER OF BINDING AND DATES OF PUBLICATION. PAGES PLATES aie ea sare PUBLISHED General Title of Vol. I = 1900 December, 1900 i—], 1—28 I—V 1864 April, 1866 29—124 VI—X1IX 1867 June, 1868 125—176 XX—XXITII 1868 February, 1869 177—194 and Table of Contents XXIV, XXV 1871 | June, 1872 A MONOGRAPH OF THE BRITISH PLEISTOCENE MAMMALIA Ol, Te BRITISH PLEISTOCENE FELIDA. W. BOYD DAWKINS, M.A., F.RS., AND W. AYSHFORD SANFORD, F.G.S. LONDON: PRINTED FOR THE PALZONTOGRAPHICAL SOCIETY. 1866—1872. PRINTED BY ADLARD AND SON, MS) ies, BARTHOLOMEW CLOSE, E.C., AND 20, HANOVER SQUARE, W. “ , -- ‘ gee : VOLUME FOR 1864. /eosonian institys y fy LONDON 9 MDCCCLXVI. THE Bek | Tbe PLEISTOCENE MAMMALIA. BY ‘ W. BOYD DAWKINS, M.A., F.GS., W. AYSHFORD SANFORD, F.G:S. PAC a INTRODUCTION. (Paces i—1.) BRITISH PLEISTOCENE FELIDA. FELIS SPELAA, Goupruss. (Paaus 1—28; Prares I—V.) OUND OUNG: PRINTED FOR THE PALAONTOGRAPHICAL SOCIETY. 1866. Bi ‘tut _* pRINTED BY J. B.A PREFACE. Siycn the time when Baron Cuvier published the last edition of his great work, the “Ossemens Fossiles,’ in 1825, the materials for working out, in detail, the British Pleisto- cene Mammalia, of which Professor Owen had given an admirable outline in 1846, have been steadily and rapidly increasing. Our great National Museum has been supple- mented by other public ones in most of the principal towns, in which the past Fauna and Flora of their respective districts are carefully preserved. The zeal of private collectors also has added very largely to the heap of accumulated facts, which cnly have to be com- pared and brought to a focus to enable us to realise, in all their varieties of size and form, the animals that lived upon that portion of the Pleistocene continent which now forms Great Britain and Ireland. Zoology also has made great strides, and, armed with a more perfect knowledge of the present order of things, we are daily becoming more fitted to in- vestigate profitably the past. In undertaking to bring the Mammalogy of the Pleistocene up to the requirements of the day, we are conscions of our own shortcomings and of the magnitude of the task. We propose, by adopting the form of a series of Monographs upon each species, and by not commencing a second species until we have exhausted all the attainable information upon the one we may have in hand, to leave the work in such a state that it may be continued by any successors without alteration of plan. We do not pledge ourselves to bring out the Monographs in zoological order, but just as our materials may admit of the complete description of any Pleistocene genus. This arrangement, as each Monograph will be distinct from the rest, like those composing M. de Blainville’s ‘ Ostéographie et Odontographie,’ will not affect their being bound up in their proper order on the completion of the work. For any information as to the remains of Pleistocene Mammals in private cabinets, or anywhere else in Great Britain or Ireland, we shall be extremely obliged, as we wish to give the distribution and relative numbers of every fossil Pleistocene species. W. BOYD DAWKINS. W. AYSHFORD SANFORD. February, 1866. Py * ; rs 1 i \ 1 ' d \ ' ; , ’ ‘i , ; r; t . i f bp ' ' p 2 aa ’ " 2 — iy : Bt > uel 4 ilo : Ay “ t ! hits ‘ if fave ; f f Bt} : . bo di 5 5. ‘ if f Lovhy Hipree rer a / . bis y pam OO PAP Heo) the er te 4 ‘ , Diet, cas ti my F + Ay és Stet + a ee Yt ; aah Paar Ss) aka : F vy 7 nit ‘ ’ 2 mee. t St ay da? 2). ary SM j He Os a A Rich -t tyiad LARP Ei op (Oe Pasir ity De ea Chey ride LQ) #112 a Hy EU, PR \ y ’ Pa ry , : 4 CONTENTS OF INTRODUCTION. § 1. Definition of Prehistoric Period. § 2. Prehistoric Caverns. A. ' Hi 2 4 SF bo w Kent’s Hole. Paviland. Arnside Knott. Cheddar. Whitcombe’s Hole. Plumley’s Den. Uphill. Heathery Burn Cave. § 3. Prehistoric Alluvia and Peat-Mosses. § 4. Prehistoric Mammalia. § 5. Inferences as to Prehistoric Climate. § 6. Definition of the term Pleistocene. § 7. Pleistocene Caverns and River-Deposits. § 8. Principles of determination of Pleistocene Species. § 9. Pleistocene Mammalia. P Hop Ps Boo oF Bimana. Cheiroptera. Carnivora. Insectivora. Ruminantia. Artio-dactyla. Perisso-dactyla. Proboscidea. Glires. § 10. Groups of Species into which the Pleistocene Mammalia naturally fall. A B Cc. D 30) Extinct Species. Species confined to Northern Climates. Py) 5 Southern ,, BS common to Cold and Tropical Climates. Species still inhabiting the Temperate Zones of Europe. § 11. Inferences as to Pleistocene Climate. § 12. Relation of Pleistocene to Prehistoric Mammals, and those now living in Britain. § 13. Probable causes of extinction or dispersal of many of the Pleistocene Species. ; Vat ite i pals Wis Phen i 1h a f Lissa es why s ih dove eH ‘ tai OLA ai ‘ Hy t i Ay de BRITISH PLEISTOCENE FELIDA. Felis spelea, Goldfuss." CHAPTER I. Lower Jaws . : : 4 oe Pian era II. Forrarm : : F go HIE - III. Os InnominaTuM : ’ : : rar ap IV. Tarsus ; ; : ; pee Ves | eve Hinp Paw. ; i: ane When simply numbers are placed at the commencement of the lines of the Tables of Measurement, they invariably signify— . Extreme length or antero-posterior measurement in inches. . Minimum circumference. . Maximum transverse measurement of proximal articulation. .. Vertical ditto ditto. . Maximum transverse measurement of distal articulation. . Vertical ditto ditto. ao fF WwW dO = ; DESIDERATA FOR THE COMPLETION OF THE MONOGRAPH ON FELIS SPELAEA. British specimens of nasal and internal bones of the cranium; vertebre ; atlas; axius; sacrum. } 5 2 complete humerus, or proximal end of the bone; distal end of ulna. | > % carpal bones, viz., cuneiform, trapezium, trapezoid, magnum. 3 Z complete, or proximal end of femur. oa % complete, or proximal end of fibula. E Bones of any other species of Felis or Machairodus from the British Pleistocene. The species at f present known to exist in Britain are Machairodus latidens, Felis pardus, F. catus. But the lynx and, : perhaps, the serval might be expected. The authors of this Monograph would be glad of the opportunity of examining any bones or teeth of Carnivora; and the utmost care will be taken of bones, addressed either to W. Boyp Dawkrns, Esq., Geological Survey, Jermyn Street, or to W. A. Sanrorp, Esq., Nynehead Court, ‘ Wellington, Somerset. Or should the possessors not wish to send them, the authors would be glad of the . opportunity of simple examination. : " Unfortunately we have been unable, from want of time, to publish the Plates and Descriptions of the Bones of Felis spelea in the order in which they should naturally appear. We trust that this may be rectified in our future Monographs. 7 INTRODUCTION. § 1. In going backward in time from the historical period in Britain, we find ourselves landed in the realms of archeology without any guide to absolute date, and without any connected record of events previous to the first landing of Julius Cesar upon our shores. The investigation of the contents of peat-mosses, of alluvia of rivers, and of a large number of caverns occupies our attention ; and while the remains of man are widely spread, we miss the larger Carnivora, the Pachydermata, and others of the Pleistocene Mammalia. For this period, as embracing the deposits more usually termed recent, and extending from the Pleistocene down to the beginning of history, we adopt the name Prehistoric. It is eminently the field of the archeologist, who subdivides it, according to the traces of man that it contains, into the iron, bronze, polished stone, rude unpolished stone, and flint periods." From his poimt of view we have nothing to do with it; but for the sake of showing the relation of the Pleistocene fauna to that now living in Britain, we are obliged to treat it zoologically. It forms a distinct zoological period, separable from the Pleisto- cene, but passing insensibly into the Historic period. § 2,4. Prehistoric caverns.—Caverns, as affording shelter from the weather, have been the resort of man and wild animals in all times, from the Pleistocene to the present day. Hence, very frequently in the same cave remains of different epochs are found. In Kent’s Hole, for instance, overlying the mass of bones dragged in by hyznas in Pleistocene times, and in parts hermetically sealed by stalagmite, there was a stratum of dark earth containmg the remains of the feasts and fires of some early people—bone implements, chert and flint arrowheads, “a hatchet of syenite,” sandstone spindles, shells of mussel, limpet, and oyster, a palate of Scarvs, and numerous fragments of pottery. This last “ is of the rudest description, made of coarse gritty earth, not turned on a lathe, and sun- baked; on its external margin it bears zigzag indentations, not unlike those from the barrows of Wilts.” Its ornamentation and texture are like those of the rude pottery obtained 1 See Sir John Lubbock’s ‘ Prehistoric Man,’ 8vo, 1865. London. R INTRODUCTION. by the Earl of Enniskillen, F.R.S., from the Bears’ Den of Kiihloch, and that from the Pile- dwellings of Switzerland in the collection of the late Mr. H.Christy. In some places the stalag- mite had been broken through, apparently for purposes of sculpture; and human bones, and flints of all forms, “ from the rounded pebble, as it came out of the chalk, to the struments fabricated from them, as the arrow- and spear-heads, and hatchets, were confusedly dis- seminated through the earth, and the whole agglutinated together by stalagmite. Flint cores were lying by the sides of the flakes struck from them.”! The remains also of the wild boar, red deer, fox, rabbit, and rodentia, were obtained from the same layer, and are in part preserved in the Museums of London and Oxford. The metatarsals and -carpals in the Oxford Museum, obtained from the upper portions of the contents of the cavern in association with charcoal, belong to the small short-horn Bos longifrons. To the absolute date of these remains there is at present no clue; but that the cave was inhabited after, to the disappearance of the characteristic Pleistocene mammalia found in the cavern, by savages closely allied to those whose remains are found in hut- circles and tumuli, there can be no doubt. ‘The careful exploration now being conducted by the British Association will doubtless throw great light upon the relative age of the various layers, and possibly the absolute age of some of the superior ones. - § 2, B. The Paviland cave, described by Dr. Buckland, affords another instance of the mixture of Pleistocene and Prehistoric remains. ‘To the one period belong the elephant, rhinoceros, horse, and hyzena; to the other, the human skeleton (which equals in size the largest male skeleton in the Oxford Museum), the bones of ox and sheep, the whelk, limpet, littorina, and trochus, that had been introduced for food. Certain small ivory ornaments found along with the skeletons Dr. Buckland considers to have been made from the tusks of the mammoth in the same cavern, and he justly remarks—“ As they must have been cut to their present shape at a time when the ivory was hard, and not crumbling to pieces, as it is at present on the slightest touch, we may from this circumstance assume for them a very high antiquity.”* May we not also infer from the fact of the manufactured ivory, and the tusks from which it was cut, being in precisely the same state of decomposition, that the tusks were preserved from decay during the Pleistocene times by precisely the same agency as those now found perfect in the Polar regions—by the intense cold; that long after the mammoth had become extinct the tusks thus preserved were used by some race that has passed away; and that at some time subsequent to the interment of the ornaments with the corpse a great climatal change has taken place, by which the temperature in England, France, and Germany has been raised, and decomposition set in in the organic remains that up to that time had re- mained for ages in their natural condition? The presence of the remains of sheep under- * «Cavern Researches,’ by the late Rev. J. MacHnery, F.G.S., edited by E. Vivian, Esq., 4to, 1859 (the larger edition). * Buckland, ‘ Reliquiz Diluvianz,’ 4to, 2nd edit., 1824, p. 90. INTRODUCTION. Xl neath the bones of elephant, bear, and other animals, coupled with the state of the cave- earth, which had been disturbed anterior to Dr. Buckland’s examination of the cave, would prove that the interment was not of Pleistocene date. No traces of sheep or goat have as yet been afforded by any Pleistocene deposit of Britain, France, or Germany. § 2, c. These two instances of the presence of remains of Pleistocene and Prehistoric age in the same cavern are two out of a large number in which a similar mixture of organic remains are met with. Instances also may be multiplied of caverns containing remains of Prehistoric age alone without others. Thus, Professor Owen quotes a cave at Arnside Knott, near Kendal, that yielded wild boar (Sus scrofa), brown bear (Ursus arctos), and other existing species of Mammalia." § 2, D. During our explorations of caverns in Somersetshire we explored three of Prehistoric age. In 1859 a small cave at the head of Cheddar Pass yielded a large quantity of bones. Prior to our examination, on tts first discovery, some of the remains were deposited in the Museums of Bristol and Oxford. The list of Mammalia comprises, besides man, the wolf, fox, badger, wild boar, goat, roebuck, Bos longifrons, and horse. A human skull from this cave, preserved in the Oxford Museum, is very well developed,’ and may have belonged to a person of considerable capacity. § 2, n. In 1863 we examined a second cave, also in the Mountain Limestone of the Mendip range, in Burrington Combe, and named it from its discoverer Whitcombe’s Hole. It was very nearly blocked up with earth mingled with charcoal, and contained a large quantity of the remains of ox, red deer, goat, wolf, fox, badger, rabbit, and hare. In the lower portion of the cave, where the floor dips downwards, we disinterred fragments of a rude unornamented urn of the coarsest black ware, with the rim turned at right angles, and an angle iron which more closely resembled those found strengthening the angles of wooden chests in the Roman graves on the banks of the Somme than anything else we have seen. The accumulation of bones and charcoal proves that the cave was in- 1 ‘British Fossil Mammals,’ 8vo, 1846, p- 429. 2 We are indebted to Professor Phillips, M.A., LL.D., F.R.S., for the following note upon this skull: —‘The cranium is dolicho-cephalic, elevated in the parietal region, very narrow behind, with a very distinct occipito-parietal slope, narrow and evenly convex in the front ; substance thin ; individual young, _ probably female. ‘This cranium most nearly resembles one from the cave at Llandebie (now in the Oxford Museum), which is filled with stalagmite, and was accompanied by bones of elk, bear, and Bos longifrons. The dimensions are— Length . F : 5 in inches 7:22 Breadth, parietal , : ry 5°32 » frontal ‘ ; - 375 ““The last measurement is taken along the supraciliary line, for it is hardly a crest in this individual. “Do you ask what race of men this belonged to? I answer that I have seen plenty of men and women with such crania in the south of England and South Wales.”—Oszford, Sept. 1, 1865. xl INTRODUCTION. habited by man for some considerable time, like those of Perigord.’ The interment is clearly of a later date than the occupation, because it is made in the mass of earth, bones, and charcoal, which resulted from the latter. The interval between the two is of doubtful length. § 2, . In the same year we explored the cave, Plumley’s Den, like the preceding in Burrington Combe. It consists of two large chambers, connected together by two small passages, not more than a few inches high. The natural entrance, but a little larger than a fox-hole, was in the roof of the first chamber, and through this we had to drop down into the cave. Subsequently we blasted a second entrance. The first chamber was at least half-full of broken rocks, forming a talus immediately below the natural entrance, through which in part they may have been introduced. ‘They were covered with a mortar- like mass of decomposing stalagmite. Underneath them we found a group of four skulls, more or less crushed and fractured. One of these belonged to a small variety of the Bos taurus, probably that variety so abundant in deposits of Prehistoric age, Bos longifrons. Two others belonged to a species of the goat tribe, and approach more closely to the Aigoceros Caucasica’ of Asia than any recent species with which we are acquainted, the ‘ horncores being oval, in section, very nearly parallel, and slightly recurved. We have met with a similar form in a deposit of bones at Richmond, in Yorkshire ; but in the absence of the necessary materials for comparison from the Museums of London, Oxford, and Paris, we do not feel justified in imposing a new specific name. The fourth skull belonged to Sus scrofa, and had a round hole in the-frontals, about the size of a crown-piece, which had the appearance of bemg made by human hands. The presence of the lower jaws by the side of their respective skulls indicates that they were deposited in the cave while the ligaments still bound them together. They were all more or less covered with decay- ing stalagmite. Between the interstices of the stones covering the floor were numerous bones and teeth of wolf, fox, mole, arvicolee, badger, bat, the metacarpal of red deer, the radius of Bos, and the remains of birds. ‘The outer chamber was remarkable for the absence of earth of any kind, except underneath the natural entrance, where there was a thin coating. ‘The lower chamber, on the other hand, running in the same slope as the outer, has its lower end entirely stopped up with a fine red earth, deposited by a stream, traces of the flow of which, during heavy rains, were evident. How the animal remains were introduced—for there were no marks of gnawing upon them, and no fragments of charcoal in the cave—is altogether a matter of conjecture. But the fact of finding the skulls grouped together, coupled with the presence of the hole in the frontals of the Sus scrofa, mclines us to believe that they may have been introduced by the hand of man. The entrance was far too small to admit of an ox falling into the cave by accident, and scarcely large enough to admit of a goat or deer squeezing themselves through. 1 «Revue Archéologique,’ 1864. 2 Gray, ‘Cat. of Bones of Mammals in British Museum,’ 8vo, 1862, p. 249. INTRODUCTION. xl § 2,4. Fromacave in the limestone cliffs at Uphill, near Weston-super-Mare, Mr. James Parker obtained the following remains :—Human crania and bones, accompanied with rude pottery and charcoal, the bones of wild cat, wolf, fox, badger, Bos longifrons, and a second species ox of larger size; the red deer, Sus scrofa, and water-rat. A large percentage of these belong to young animals, and some are gnawed by dogs, wolves, or foxes. This must not be confounded with the cavern at Uphill from which Messrs. Beard and Williams obtained Pleistocene Mammals. § 2, u. The Heathery Burn Cave, in Yorkshire, explored by Mr. John Elliot, yielded, besides the remains of man, those of the otter, badger, goat, roedeer, hog, and water-rat. With reference to the human remains, Professor Huxley observes—‘“ I see no reason for believing them to be of older date than the river-bed skulls,” 7. e. those found in the valley of the Trent, associated with Bos longifrons, goat, red deer, wolf, and dog." We have selected these as examples of Prehistoric caverns, and as representing the fauna of the vague interval between the Pleistocene and our own times. § 3. In the alluvia of rivers and in peat-bogs the remains of animals of Prehistoric age are found in large numbers, and correspond remarkably with those of the caverns. Thus, the Manea Fen, in Cambridgeshire, has yielded Ursus arctos; the peaty mud near Newbury the beaver, wild boar, roedeer, red deer, wolf, goat, horse, otter, water-rat, bear, Bos longifrons, and B. primigenius ; the peat and the marly beds below of Ireland the Megaceros, associated with the Bos longifrons, the red deer, and the reindeer (C. tarandus). The peat of Hilgay, in Norfolk, has furnished the beaver and Irish elk, while that of Ross- shire the traces of the reindeer, in an antler presented by Sir P. Malpas de Grey Egerton, Bart., to the British Museum. ‘The reindeer also is described by Professor Owen? as occurring beneath a peat-moss near Hast Dereham, in Norfolk. The remains of Bos longifrons are most universally found with red deer, roedeer, wild boars, otters, and beavers. The marl underneath the peat of Scotland has also yielded the gigantic skulls of the great Urus, Bos primgenius. From the very recent character of the osseous sub- stances of these, Professor Owen infers that this animal may have maintained its ground longer in Scotland that in England. “Tn the Museum of the Natural History at Newcastle is a remarkably fine shed antler of the true elk, Alces malchis. It was found in Chirdon Burn, near the bottom of the recent peat-formation, resting partially on the coarse gritty marl formed by the weathering of the subjacent strata.”* It measured, when perfect, from tip to tip, 2 feet 10 inches ; from burr to extreme end, 2 feet 10 inches ; round the burr, 10 inches; and round the beam nearly 8 inches. The gisement of the fossil stamps it as being of the same relative “ Geologist,’ 1862. : 2) Op -ncit, ‘Trans. of Tyneside Naturalists Field Club,’ vol. v, part ii, p. 111. Paper by Mr. Richard House, 1861: XIV INTRODUCTION. Prehistoric age as the Urus of Scotland and the Megaceros of Ireland, found in the marl at the bottom of the peat. It is worthy of remark that this animal was eaten by the dwellers in the Lake-villages of Moosedorf, Wauwyl, Meilen, Robenhausen, Concise, and Bieune, according to Professor Riitimeyer. In digging the foundations of the large works at Crossness Point for the southern out- fall of the metropolitan sewage a most interesting collection of Prehistoric Mammalia was made from the peat and silt. We identified red deer, Bos longifrons, goat, beaver, horse ; and among the remains forwarded to the British Museum is a remarkably fine antler of reindeer (Cervus tarandus), which My. Houghton, the engineer, informs us came from the bottom of the peat at a depth of fifteen feet below the surface. A large number of cases of similar discoveries may be quoted, as in the estuarme mud of Selsea, of the hind and fore legs of Bos longifrons, with all the bones, to the smallest sesamoids, in place, or in a silted-up river-bed at Waterbeach Mills, near Cambridge, of the same animal, associated with the red deer, goat, horse, and wolf. § 4. The following list, which probably will be largely increased, represents the mam- malia derived from the Prehistoric deposits, and includes those species that began to live in the Pleistocene, and are living in Britain at the present day, and which therefore must have lived in the Prehistoric Period, although their remains have not yet been discovered init. The latter are marked with an asterisk. It consists of thirty-four species. Homo sapiens, Lin. Arvicola amphibia, Desm. Rhinolophus Ferrum Equinum, Leach. » pratensis,* Bell. Vespertilio noctula, Geoff. y) peerestis,*® Pall. Talpa Europea, Lin. Lepus timidus, Faér. Sorex vulgaris,* Lin. » cuniculus, Lin. Felis catus ferus, Schreb. Equus caballus, Lin. Canis familiaris, Lin. Alces malchis, Gray. » vulpes, Lin. Megaceros Hibernicus, Owen. » lupus, Lin. Cervus Tarandus, Lin. Mustela erminea,* Lin. % elaphus, Lin. ¥ martes,* Lin. » capreolus, Lin. s putorius,* Lin. Ovis aries, Lzn. Lutra vulgaris, Eral. Capra egagrus, Gmel. Meles taxus, Schreb. » hircus, Gmel. Ursus arctos, Lin. Bos longifrons, Owen. Mus musculus, Lin. », Urus, Cesar (= B. primigenius, Boj.) Castor fiber, Zin. Sus scrofa, Lin. The absence of the squirrel and dormouse from this list may, perhaps, be owing to their arboreal habits, which would render the chances of their bones being found in caverns or river-deposits very remote. Both genera and perhaps both the English species occur in the French Pliocene strata. —— INTRODUCTION. XV All these animals are still living in Britain at the present moment, except the Irish elk (Megaceros Hibernicus), which is entirely extinct; the remdeer and moose (A/ces malchis), which are now confined to the colder regions of Northern Europe, Asia, and America ; and the Bos longifrons, and B. primigenius, the beaver, the wolf, the wild boar, and the Ursus arctos, which lived on these into the Historical period. The Bos longifrons —which, in our opinion, will ultimately be found to be specifically identical with Bos taurus—was the variety that supplied the Roman legionaries in Britain with beef;* the Ursus arctos was probably, from an allusion of Martial,” exported to Rome for the sports in the theatre; and the beaver was still living in the river Teivy, in Cardiganshire, when in 1188, Giraldus Cambrensis accompanied Archbishop Baldwin on his tour through Wales to collect volunteers for the First Crusade. This latter animal, according to Boethius, lingered on in Loch Ness till the fifteenth century. The wolves, sufficiently abundant in the Andreads Wold to eat up the corpses of the Saxons left on the field by Duke William’ after the Battle of Hastings, lingered on in England till 1306, in Scotland till 1680, and in Ireland, protected by the uncultivated wilds-and the misrule of the country, until the year 1710. ‘he last wild boar was destroyed in the reign of Charles I. The Irish elk (Megaceros Hibernicus), whose remains are so very abundant in the Pleistocene deposits, is the only species that can be proved to have become extinct in the Prehistoric period. | Whether or no the great Urus and the small short-horn (Bos longifrons) be extinct, or live, the one im the larger domestic cattle of Europe, as the Flemish oxen,* and those of Holstein and Friesland, the other in the smaller breeds, has not yet been satisfactorily decided. In the Prehistoric period the dog, the goat, and the sheep, make their appearance for the first time in the world’s history. Bos /ongifrons also has not yet been proved to have lived in the preceding period, the evidence for its coexistence with the extinct Pleistocene mamimalia being founded on some remains cast up by the sea at Clacton and Walton, which therefore may have been derived from a much later deposit than the Preglacial Forest-bed. The remains of the Irish elk and the reindeer may, perhaps, indicate an earlier division of the Prehistoric period, but upon this point we must be content to wait for more evidence. The occurrence of the latter animal in Prehistoric deposits proves that in Britain, as in the 1 See Boyd Dawkins, ‘Sussex Archeological Collections,’ vol. xvi. 2 «Nuda Caledonio sic pectora prebuit urso’ Non fals4 pendens in cruce Laureolus.” 3 ‘De bello Hastingensi Carmen,’ by Guido, Bishop of Amiens, who died in 1075. * Lustravit campum, tollens et czsa suorum, Corpora, Dux, terre condidit in gremio ; Vermibus atque /upis avibus canibusque voranda, , Deserit Anglorum corpora strata solo.” 4 As Professors Nilsson and Riitimeyer suggest ‘Ann. and Mag. Nat. Hist.,’ 1849; ‘Fauna der Pfahlbauten,’ 4to, Basle. XVi INTRODUCTION. south of France, it lingered on after the disappearance of its Pleistocene congeners—the lion, the hyeena, and the great pachyderms. § 5. Inferences as to Prehistoric Climate in Britain, &c.—The evidence of a gradual increase of temperature in France and Germany during the historical period seems to us perfectly incontrovertible. Firstly, the Rhine and the Danube’, during the first four cen- turies, were frequently frozen over in the winter, so that the barbarians, ‘“‘ who often chose that severe season for their inroads, transported, without apprehension or danger, their numerous armies, their cavalry, and their heavy waggons, over a vast and solid bridge of ice.”” On the banks of the Danube the wine was frequently frozen into great lumps.* Secondly,* Czesar mentions the reindeer as existing in the great Hercynian Forest that overspread Northern Germany, along with the gigantic Urws and the elk. This statement of Cesar is singularly corroborated by the discovery in the peat-bogs of Pomerania’ of the remains of these three animals, so that there can be no reasonable doubt of his accuracy in this particular case. Modern ages afford no instance of like phenomena. From some cause or other, the temperature has increased on the banks of the Rhine; and im the fact that at the present moment the reindeer® cannot live south of the Baltic we may recognise a proof of a diminution of cold in that region since it was inhabited by those lovers of a severe climate. ‘This change of temperature is very generally accounted for by the drain- age of morasses and the cutting down of woods; but may it not with more probability be ascribed to a much deeper cause—to a secular change operating throughout Europe, which began in the Pleistocene, and was going on throughout the Prehistoric, and happened incidentally to be noticed in the Historical period? The presence of the reindeer in the Prehistoric deposits of England, Ireland, and Scotland, affords precisely the same evidence as those mentioned by Cesar. At the time they lived in Britain and Ireland the climate must have been suited for them. On the theory of the gradual modification of climate through the Prehistoric and Historical periods, the fact of the manufactured ivory and the tusk preserved in the cave-earth of Paviland, cited above, being in precisely the same state of preservation, is alone explicable. There may have been oscillations of temperature, but the progress on the whole seems to us to have been gradual from the intense cold of the glacial period to the temperate “insular climate” obtaining in Britain at the present day. § 6. The term Pleistocene is used in the same sense as Professor Forbes and M. 1 Diodorus Siculus, i, 5, p. 340, edit. Wesselling. Herodian, i, 6, p. 221. ? Gibbon’s ‘Decline and Fall, chap. ix. 3 Ovid, ‘ Epist. ex Ponto, Lib. 4, ep. 7, Lin. 9, 10. Virgil, ‘Georgic,’ Lib. 3, Lin. 355. Zenophon’s © Anabasis,’ i, 7. 4 mMartes. Hp agrestis. Lutra vulgaris. »» pratensis. Meles taxus. Lepus timidus. Ursus Arctos. s, cuniculus. Sorex vulgaris. Mus musculus. The Bison priscus, or the aurochs of the Pleistocene, that spread over nearly the whole of the Pleistocene Huropzo-Asiatic continent from the Pyrennees through France and Germany as far as Behrings Straits and Eschscholtz Bay on the American shore of the 1 ‘Zoognosia,” ili, p. 221 (8vo, Mosque, 1814) :—‘“‘Sic dictam unciam a Cl. Steven zstumatissimo nostri musei largitore accepi ex montibus Soongoricis et jugo Altaico, pantheram Africanam ex ditissima donatione Excell. Pauli de Demidoff obtinuimus. Hee profundiore colore gaudebat, uncia vero albidiore, sed macularum nullam vidi differentiam. Individium vivum nunc apud Excell. Principem de Yussupoff con- servatum ex Persia venit, etiam albidiore indutum est veste, et maculee subannulares, in pedibus et cauda magis virgatee sunt. Uncia itaque pelle albidiore, juniores pardos indicare videtur.’’—Cuvier (tom. cit., vol. iv, p. 428) endorses this determination. xliv INTRODUCTION. Arctic Sea, now lingers in a Lithuanian Forest, protected by an imperial decree of the Czar of Russia. Its remains are found associated with those of the mammoth, in the frozen brick-earths and gravels of Eschscholtz Bay, throughout the Asiatic “tundras,” the islands of New Siberia and the Lachow Group, and in the caverns and river-deposits of Northern and Western Europe. It is mentioned im a remarkable List of Graces! of the Abbey of St. Gall, written by Ekkehard the younger (who lived from a.p. 980 till 1036), as an article of food, together with the bear, the urus, the beaver (which they called a fish), the wild horse (Hguus feralis), the marmot, and others. Mr. Wylie considers that the Lquus feralis was the offspring of a domestic breed run wild like that of America; but when we consider the vast number of horses that have left their remains in association with those of bison in various Pleistocene deposits, and that at a far later date they formed the food of the tribes that lived in the pile-dwellings of the Swiss Lakes, and the hut- circles of Berkshire, the probability seems to us that reference is here made to a breed of horses as undoubtedly wild as the mouse-coloured wild horse of Central Asia. The musk-shrew, Sorex moschatus,’ of Pallas, described by Professor Owen from the Preglacial deposits of Bacton, under the name of Paleospalax magnus,is now found only ina 1 The list of animals is so remarkable that we have subjoined the whole passage in which they are mentioned : “Sit benedicta fibr caro piscis uoce salubri.”—line 71 “Sub cruce diuina benedicta sit ista ferina. Sub cruce diuina sapiat bene queeque ferina. Et semel et rursus cruce sit medicabilis wrsws. Hune medici sanum memorant nullique nocivum. Dente timetur (petulcus) aper, cruce tactus sit minus asper. Cerui (vel cerue) curracis caro sit benedictio pacis. Heee satan et Larvee fugiant crustamina cerue. Signet wesontem benedictio cornipotentem. Dextra dei ueri (uel benedicat) comes assit carnibus w77. Sit dos siluanus sub trino nomine (crucis hoc slgnamine) sanus, Sit feralis equi caro dulcis in (sub) hac cruce Christi. Imbellem dammam faciat benedictio summam. Capreus ad saltum benedictus sit celer altum. Sit cibus illesus capree. Sit amabilis esus. Capreoli (us) uescam dent (det) se comedentibus escam. Carnes werbicum nihil attulerint inimicum. Pernix cambissa® (fera alpina) bona sit elixa vel assa. Sub cruce diuina caro dulcis sit leporina Alpinum cassum* faciat benedictio crassum. Sit caro syluana crucis omnis robore sana.’”’—‘ Benedict. ad Mensas’ Ekkehardi Monachi Sangallensis (lines 117—136), ‘ Archeological Journal,’ vol. xxi, pp. 355 and 358. ? Pallas, ‘ Zoographia,’ vol. i, p. 128; ‘Second Travels,’ trans., vol. i, p. 48, London, 1802. Fischer (‘Synopsis Mammalium,’ 8vo, Stut., 1829, p. 250-1) gives the literature of the species. 3 Chamois. 4 Marmot ? INTRODUCTION. xlv living state in Southern Russia, in the area between the Don and the Volga. An animal of aquatic habits, it is especially abundant on the banks of the Soura River, in latitude 55° north, and longitude 47° east, under a temperate continental climate, cold im winter but hot in summer. Dr. Pallas, in his ‘Second Travels,’ describes the country which it inhabits, after an unusually severe winter, as covered with tulips, saffron, and the Star of Bethlehem, and although Spermopiilus citillus, the Alpine marmot, was in the neighbour- hood, there were vineyards close by. The water-shrew of the Pyrennees, Myogale Pyrennaica, Geoffr., is a closely allied species, differing from the Sorex moschatus (Myogale moschata, Fischer), in its smaller size, and long and rounded tail. An analysis, therefore, of the fifty-three Pleistocene species, those about which there is any doubt being omitted, gives 14 as extinct. 8 as confined to northern climates. Dy ye southern climates. 1 as common to temperate and hot climates. 28 as still inhabiting the temperate zones of Europe. 53 § 11. Inferences as to Pleistocene climate——The proportion of fourteen extinct to thirty-nine living species proves that, in the geological sense, the present order of things is separated by a small interval from the Pleistocene ; while, from the fact that twenty-eight species, or half, are still living in the same European area, we may infer that the conditions of existence, the climate and food, and the like were then very similar to those now obtaining in the area in which they live. That, however, some great physical change has taken place in Hurope since the Pleistocene times, is proved by the presence of other groups of mammalia—those confined now to cold and to hot countries. They afford evidence that at first sight appears conflicting, but which upon analysis we shall find to be very conclusive, that the climate in Britain, in those days, was very much more severe than at present. From the conditions under which the surviving Pleistocene herbivores now live, we can infer those under which they lived im Britain in that early period. The northern group of Pleistocene mammalia, living only now im a severe continental climate, consists of species that have very different powers of resisting cold and heat. Thus, the musk-sheep is found now only under the lowest temperatures in the vast treeless “ Barren Grounds” of North America, while the reindeer lives also in the forests, along with the - elk, of the Huropzo-Asiatic and North American continents. The red deer and the bison range up to the edge of the province inhabited by the latter animals. The lemmings live tinder a very severe climate, while the marmots are found in the higher and colder districts in Southern Europe and Central Asia. Each of these northern species is dependent upon the oscillation of the climate for its particular habitat in a given year, 2? xlvi INTRODUCTION. retreating northwards or southwards, according to the temperature that regulates the supply of food necessary for its existence. Thus, in North America, Sir John Franklin writes that the migrations of the animals afford a means of foretelling the severity of the season. If the reindeer retreat far south then a severe winter is to be apprehended ; if, on the contrary, they remain very nearly in their usual winter haunts, the season invariably is a mild one. ‘The reindeer of Northern Russia are equally dependent upon the season for their locality ; and if an unusual season occurs, to put the animals off their accustomed route, the inhabitants of the district at the mouth of the Kolyma, living upon the chase, endure the severity of famime. M. Von Matiuskin, the Lieutenant of Admiral Von Wrangel, had the good fortune to see one of these migratory bodies of reindeer crossing a river, consisting of many thousands, divided into herds of two or three hundred each. By some such oscillation of temperature, which regulates the supply of food for the herbivores, the remains of the animals of two contiguous zoological provinces may be found together in one spot, as in the case of the northward retreat of the musk-sheep which, living in Hearne’s time (a.pD. 1770—72) near Fort Churchill, has left that district to be occupied now by the elk and the waipiti. In this manner the admixture of the remains of animals living at the present day, respectively, in a severe, and in a temperate continental climate, may be accounted for in the Pleistocene caverns and brick-earths. Of the district in America, where the animals inhabiting the high northern latitudes meet with those that live under a comparative temperate climate, Sir John Richardson writes :—‘ The subsoil north of latitude 50° is perpetually frozen, the thaw on the coast not penetrating above three feet, aud at Great Bear Lake, in latitude 64°, not more than twenty inches. The frozen substratum does not of itself destroy vegetation, for forests flourish on the surface at a distance from the coast, and the brief, though warm summer, gives birth to a handsome flora, matures several pleasant fruits, and produces many carices and grasses.’” But in the vast plains of Siberia, extending from the Altai Mountains to the Arctic Sea, we find probably a nearer approach to the Pleistocene climate of Western Europe. Covered by impenetrable forests, for the most part of birch, poplar, larch, and pines, and low creeping dwarf cedars, they present every gradation in climate from the temperate to that in which the cold is too severe to admit of the growth of trees, which decrease in size as the traveller advances northwards, and are replaced by the grey mosses and lichens that cover the low marshy “tundras.” The maximum winter cold, registered by Admiral Von Wrangel,” at Nishne Kolymsk, on the banks of the Kolyma, is—65° in January. “Then breathing becomes difficult; the wild reindeer, that citizen of the Polar region, withdraws to the deepest thicket of the forest, and stands there motionless as if deprived of life; and trees burst asunder from the intensity of the cold. Throughout this area roam elks, black bears, foxes, sables, and wolves, that afford subsistance to the Jakutian 1 Back’s ‘ Journey to the Arctic.Sea,’ 4to, 1836, appendix, p. 479. 4 (Ojo Olt INTRODUCTION. ; xl vit and Tungusian fur-hunters. In the northern part countless herds of reindeer, elks, foxes, and wolverines make up for the poverty of vegetation by the rich abundance of animal life. ‘‘ Enormous flights of swans, geese, and ducks arrive in the spring, and seek deserts where they may moult and build their nests in safety. Ptarmigan run in troops amongst the bushes ; little snipes are busy along the brooks, and in the morasses; the social crows” seek the neighbourhood of new habitations ; and when the sun shines in spring one may even sometimes hear the cheerful note of the finch, and in autumn that of the thrush.” Throughout this region of woods a hardy, middle-sized breed of horses lives under the mastership and care of man, and is eminently adapted to bear the severity of the climate. Like the other northern quadrupeds they change their coats in the midst of summer. “'They perform most laborious journeys, often of three months’ duration, with no other food than the half-withered grass, which they get at by scraping away the snow with their hoofs, and yet they are always in good condition.” ‘The only limit to their northern range is the difficulty of obtaming food. The severity of the winter, through the southern portion of this vast wooded area is almost compensated for by the summer heat and its marvellous effect on vegetation. The hypothesis of a series of conditions obtaining in Pleistocene Western Europe similar to those now found in this portion of Northern Asia, will alone satisfy the evidence afforded by the fauna, and the deposits in which they are found. ‘The contortion of the gravels, and the angular state of the pebbles of which they are often composed, are, as Mr. Prestwich infers, explicable only on the theory of ice having been formed in our rivers im far larger quantities than at the present day; the one being the result of the grounding of miniature bergs, the other of their melting away and depositing their burden of pebbles. The large plateaux of brick-earths are probably the deposit of the floods caused by the sudden melting of the winter snow, similar to that which Admiral Von Wrangel describes in Northern Siberia, and Sir John Franklin in the area north of the Canadian Lakes. The winter cold would be sufficiently intense to allow of the northern group of mammalia living in the winter, and even the musk-sheep (of which the remains are rare) might have been obliged to leave the Pleistocene “ tundras,” and take shelter im the zone of the elk and even the bison, in an unusually severe season. On the other hand, in the summer, the animals that are now found in the temperate zones of Europe might advance even into the country of the elk and the reindeer ; and even carnivora now confined to hot climates find their way into the temperate zone of the day. Thus, the Hyena vulgaris, or common living hyeena, is found fossil in the South of France, without penetrating as far north as Britain, France, or Germany. In fine, the evidence afforded both by the fauna and deposits of the Pleistocene seems to us to prove that the climate in Pleistocene Britain was more severe than it is now; that at a time when Britain formed a portion of the Europzo-Asiatic continent, it more closely resembled that now obtaining in the fur-countries of Northern Asia than xlvin INTRODUCTION. elsewhere ; and lastly, that it was subject to oscillations by which the migrations of the herbivores were directed northward or southward, as the case may have been. The remarkable evidence afforded by the thick woolly covering on the carcases of the mammoth and tichorhine rhinoceros of Siberia, as to the temperature of the countries in which they lived, makes it very probable that the Hippopotamus major was in hke manner defended from the cold; but at the same time we must bear in mind that the aquatic habits of the genus are incompatible with the severity of a climate suited to the reindeer; that it has not been found in Russia, nor in any of the vast deposits in the high northern latitudes ; and that therefore it is rather to be put into the same category with the bison of North America, rather than the reindeer, as an occasional visitant rather than a dweller throughout the year in England, France, and Germany. Its remains are very rare, as com- pared with the other herbivores, the fossil elephants, rhinoceroses, Irish elks, bisons, rem- deer, and the like. Its head-quarters probably were on the shores of the Mediterranean, and the north of Africa, from which latter locality M. Gervaise’ cites 1t as occurring in a fossil state near Constantine, in Algeria. In the caverns on the Huropean shores its remains are extremely abundant, as also in the Italian Phiocenes.” There is another interesting point connected with climate. How it may be asked, can you reconcile the presence of the spelzean hysena and the lion with the climate which the reindeer and the musk-sheep required for their existence? Is not the very fact of their coexistence with the reindeer a proof of their specific distinctness from the African or Asiatic lion, or the hyena of the Cape? If they are identical in species must not the Pleistocene climate have been similar to that of the countries in which they now live? An appeal to the zoological distribution of the carnivora over wide areas, proves that it is not so. While the herbivora are dependant upon the temperature for the vegetation on which 1 Tom. cit., p. 363. 2 The occurrence of hippopotamus may be accounted for in a somewhat different manner. While we may be almost certain that the general climate of Britain, during the Postglacial epoch, has been more severe or, properly speaking, more extreme or continental than at present, a period or periods of some length may have intervened, while England and Ireland formed a portion of the European continent, when the climate may have been less severe, and the rivers free from ice throughout the year. This view of the case is strengthened by the fact that the fossil hippopotamus is frequently, if not generally, accom- panied by forms of elephant and rhinoceros 7. e. Hlephas antiquus and Rhinoceros leptorhinus, which appear both in this hemisphere, and as far as E. antiquus is concerned, in America also, to have had a southern, and even a tropical, rather than a northern range. Among other facts which must be accounted for is the existence of a Lusitanian flora on the west coast of Ireland. This flora, or some member of it at least, with difficulty maintain their ground at the sea-level, and must have been exterminated by the severity of the glacial epoch ; and we cannot suppose but that their migration from a southern land has occurred since, and that along a coast-line. It is true that Professor Forbes assigned the Miocene period as that of this migra- tion, but he appears to have overlooked the great severity of the intervening glacial climate. On these points compare, Forbes’s “ Flora and fauna of the British Isles,’ ‘Mem. Geol. Survey,’ vol. i, p. 336 et seq.; Trimmer, ‘Quart. Journ. of Geol. Soc.,’ vol. ix, p. 13, 1853; Lyell, ‘ Ant. of Man,’ ed. 1863, pp. 273 et 320; Croll, ‘ Nat. Hist. Review,’ No. xx, p. 594; and authors quoted by them.—W. A. S. ae Se eee ee 7s. =a ee ee ee ee ey ee = - er Ee ee ee a INTRODUCTION. xlix they feed, and are restricted in range to those districts where the food most fitted for them is to be found; the only limit of the range of the carnivora is to be found in that of the animals upon which they prey.’ Thus, the tiger preys upon buffaloes, deer, and the herbivores peculiar to each district, throughout the length and breadth of India. On the shores of the Sea of Aral it is the scourge of the horses of the nomad Tartars, and in the district of the Altai it preys upon the wild boars, and further north upon the remdeer ; and yet specifically it is the same, the markings on the skin of that of the Aral being of the same character as that of India.” The fox is another example of the same kind, ranging throughout the old and new worlds, and yet not divisible into species. ‘The wolf, also, and the panther, already quoted, are instances of the same kind. ‘The fact, therefore, of the spelzan hon and hyzna having lived in a climate in the Pleistocene far differing from that im which they now live cannot be quoted in favour of the recent and the fossil belonging to distinct species. ‘The remains of Hyena spelea are found in vast numbers in Britain, France, and Germany; those of #7. syelea being more sparingly found; and those of wolverine being abundant in the caverns of Liége and Gailenreuth, and absent from France, and met with but in three caves in Britain. Neither of the two former species have, as yet, been discovered in Russia or in the high northern latitudes of Europe and Asia, where there are such vast stores of fossil remains. § 12. Relation of Pleistocene to Prehistoric mammals, and those now living in Britain. —The following table of British genera and species of land mammalia, from the Pleisto- cene downwards to the present day, shows at a glance the close relation existing between them ; and it shows, moreover, by the gradual elimination of the Arctic group of mammalia, that the increase in temperature from the Pleistocene to the present day has been gradual : Species now living in Genera. Pleistocene species. Prehistoric species. Builant Castor, Lin. fiber, Lin. fiber, Lin. trogontherium, Fisch. Mus, Lin. musculus, Lin. musculus, Len. musculus, Lin. messorius, Sh. sylvaticus, Lin. rattus, Lin. decumanus, Pall. Arvicola, Lac. = Hypudeeus,|amphibia, Desm. Til, Spermophilus, Fred. Cuv. Sciurus, Lin. Myoxus, Zimm. 1 See M. de Serres, “Sur l’Origine des Animaux et des Vegétaux,” ‘Revue du Midi,’ tom. y livraison. agrestis, Flem. pratensis, Bell. citillus, Pail. erythrogenoides, Fale. 2 See preceding note on Felis spelea. amphibia, Desm. agrestis, lem. pratensis, Bell. amphibia, Desm. agrestis, Flem. pratensis, Bell. vulgaris, Lin. avellanarius, Desm. INTRODUCTION. Species now living in Prehistoric species. Britain, Genera. Pleistocene species. Lagomys, Geofr. Lepus, Lin. Lemmus, Link. Elephas, Lin. Rhinoceros, Lin. Equus, Lin. Sus, Lin. Hippopotamus, Lin. Bison, Gesn. Bos, Lin. Ovibos, Blain. Capra, Lin. Ovis, Lin. Cervus, Lin. Alces, (Ces.), Lin. Megaceros, Hart. Machairodus, Kaup. Felis, Lin. Hyzena, Briss. Canis, Lin. Lutra, Eri. Mustela, Lin. Meles, Lin. Gulo, Stor. Ursus, Lin. Talpa, Lin. Sorex, Lin. Vespertilio, Lin. speleeus, Owen. timidus, Aral. cuniculus, Pall. lerimus, Link. primigenius, Blum. antiquus, Fale. priscus, Gold. meridionalis, Nestz. tichorhinus, Cuv. leptorhinus, Owen. megarhinus, Chris. Etruscus, Fale. fossilis, Owen. scrofa, Lin. major, Desm. priscus, Owen. primigenius, Boj. moschatus, Desm. elaphus, Lin. capreolus, Lin. tarandus, Lin. dicranios, Nest. malchis, Gray. Hibernicus, Owen. latidens, Owen. speleea, Gold. antiqua, Cuv. catus, Lin. speleea Gold. lupus, Lin. vulpes, Lin. vulgaris Eral. martes, Lin. putorius, Lin. erminea, Lin. taxus, Lin. luseus, Sabdne. speleeus, Gold. Arctos, Lin. Europzea, Lin. vulgaris, Lin. moschatus, Pall. 2 14 a2 timidus, Hrzl. cuniculus, Pall. caballus, Lin. scrofa, Lan. ? = Urus, Ces. longifrons, Owen. hircus, Gm. egagrus, Gm. aries, Lin. elaphus, Lan, capreolus, Lin. tarandus, Lin. malchis, Gray. Hibernicus, Owen. y catus, Lin. lupus, Lin. vulpes, Lzn. familiaris, Lin. vulgaris, Hral. martes, Lin. putorius, Lin. erminea, Lin. taxus, Lin. Arctos, Lin. Europea, Lin. vulgaris, Lin. timidus, Hrzl. cuniculus, Pall. variabilis, Pall. = caballus, Zin. scrofa, Lin. =? taurus, Lin. =? taurus, Lin. hircus, Gm. eegagrus, Gm. aries, Lin. elaphus, Lin. capreolus, Lin. dama, Lin. catus, Lin. vulpes, Lin. familiaris, Ln. vulgaris, Hrel. martes, Lin. putorius, Lin. erminea, Lin. vulgaris, Lin. taxus, Lin. Europea, Lin. vulgaris, Lin. fodiens, Pail. remifer, Geof. mystacinus, Leist. Daubentonii, Lest. MONOGRAPH THE BRITISH MAMMALIA PLEISTOCENE PERIOD. Order—CARNIVORA. Faminy—FELID A. Genus—FELIs. Species—Felis spelea, Goldfuss. CHAPTER I. Felis spelea—ULower jaw, Pl. I, figs. 1, 2, 3; Pl. VL" 1, 2. A comparison of the jaws of large Feles from the caves and river-deposits, one with the other, and with those of the existing species nearest them in size, shows us most distinctly that, while there are some fossil jaws which, in form and size, are absolutely indistinguishable from those of the lion, others, which are generally considerably superior to them in some of their dimensions, offer characteristics which appear to prove. that they are so closely allied to those of that animal that the differences between them probably do not amount to more than extreme variety, while they essentially depart by the same characters from those of the tiger. If a large series of lower jaws of recent lions and tigers be compared together we find that, while the individual differences are great, resulting possibly from difference of sex, food, climate, and the like, there is one test of specific value by which the one can be distinguished from the other. In the lion the inferior border of the ramus bears a slight ' Unfortunately we have been unable, from want of time, to publish the Plates and descriptions of the bones of Felis spelea in the order that they should naturally appear. We trust that this may be rectified in future monographs. 3 se 1 2 PLEISTOCENE MAMMALIA. process immediately beneath the last molar, which is developed to a different degree in different individuals; in some it causes the outlme of the imferior border of the lower jaw to present a regular rounded, convex outline, nearly straight from symphysis to angle, while in others it reaches a maximum of development, so that the jaw approaches the doubly arched appearance so manifest in the figures Pl. I, figs. 1, 2. In the tiger, on the other hand, the inferior border of the lower ramus is straight, or rather concave in outline, from the symphysis backwards, the only exception out of the large number of tigrine skulls im the museums of Oxford and London being that afforded by one in the Hunterian collection, killed in India, and presented by General Hardwicke, in which there is the faintest possible approximation to the leonine contour. In this point the rami of Felis spelea agree most remarkably with those of Fels Jeo, and, as far as the larger specimens are concerned, present us with the maximum development of this “amal” process, as it may be called (a of figs. 1, 2, Pl. 1). In these large specimens it is far more strongly marked than in any recent varieties of Felis leo; but, as the latter present great variations in this respect, we cannot consider the stout proportions which the former exhibit a proof of specific difference. The smaller jaws above referred to (figs. 1, 2, Pl. VI) do not present a greater development of the “samal” process than ordinary specimens of the lion; being in this respect, as in all others, indistinguishable from that animal. In the leopard, both of Africa and of India, the inferior outline of the lower jaws resembles that of the lion; but that of the jaguar (Helis onca) is straight or slightly concave, like the tiger. In both the large and small varieties the height of the condyle above the angle coincides remarkably with that of the lion, but differs in the tiger; this dimension being in the latter greater, both proportionally and absolutely. ‘The proportion of length to maximum depth of the condyle varies in different individuals; but in the few imstances we have seen, both large and small varieties have this part proportionally somewhat stouter than in the recent species (Pl. I, figs. 1, 2, 3, B; PI. VI, figs. 1, 2, 3). Baron Cuvier considers that the gradual ascent of the coronoid process from the alveolar border of felis spelea is one point of difference between that animal and the living lion. We cannot see that in this poimt any difference exists between the fossil and _ recent species. The coronoid process rises more gently from the alveolar border in the lions that we have examined than in the tigers, but the difference is very slight. The apparently sharper angle at which this process rises from the alveolar border in the lion we find to be caused by the form of the anterior portion of the ramus in that animal, which tapers slightly forwards through the length of the molar series, while in Felis spelea the alveolar and lower borders are nearly parallel (PI. I, figs. 1, 2,c p, az). The angle in both forms, /. spelea and F. leo, is in reality as near as possible identical, when measured with the alveolar border c p produced backwards, 7. e. about 30° in each case. But when the angle is measured from the lower border of the ramus a B, it is about 40° in the lion and 30° in Felis. syelea. his angle, however, is variable, and therefore FELIS SPELAIA. 3 not to be trusted to as a specific character. Another point appears to deserve more attention ; it is that in Felis spelea the coronoid process projects backwards far more beyond the neck of the condyle (PI. I, fig. 1, F) than in the lion or tiger; in the former animal this excess of projection amounting to nearly an inch, while in the tiger it is barely perceptible; and, as far as we have measured, it does not exceed half an inch in the lion. ‘The different appearance this gives to the bone is very remarkable. The “superior border of the coronoid process is a strong, smooth, externally rounded ridge, somewhat stronger than that of the hon or tiger, but almost totally destitute of the strongly marked ridges which are on this part of the bone in those animals. The angle in the larger jaw resembles that of the lion, except that in the lateral aspect it forms one end of an arch (Pl. I, figs. 1, 2, @), of which the samal/ process (a) forms the other, whereas in the lion and in the small variety this aspect presents a slightly con- cave line (PI. VI, figs. 1, 2,6, a). In the adult tiger the angle descends far lower, and the arch above mentioned extends without interruption to the symphysis. The masseteroid ridge is perhaps proportionally somewhat thicker and stronger in the large jaws, and the upper external border of this portion is massive and rounded (PI. J, fig. 1, H), stead of ending in a sharp and knotted ridge, as in the lion and tiger and smaller cave variety (Pl. VI, fig. 1, nH). The contour of the lower border of the anterior half of the ramus is precisely the same in the smaller cave jaws and the lion, as well as in the large fossil jaws; but it is more concave in the tiger. The contour at the symphysis differs in the large form of Felis spelea from that of either of the others. The angle formed by the front edge of the symphysis and the pro- duced plane of the lower border is much larger in Melis spelea than im either Felis leo or the small cave specimens (Pl. I, figs. 1, 2, x, n, a). This angle amounts to 70° in Felis spelea, while it is 45° in the smaller fossil forms, and 40° only in the lion; im the tiger it is 55°. This difference, though very striking to the eye, appears to be variable ; it is probably of little, if any, specific value. The alveolar border is straight in all the forms (Pls. I, VI, figs. 1, 2, c, v). The mentary foramina are very variable in form and position (Pls. I, VI, fig. 1,D. In _ some of the fossil jaws they appear to be each divided so as to form four on each side; we have seen an approach to this variation in recent specimens of both lion and tiger. We may observe here that the silky smooth surface, which is remarkable on the other more robust feline bones of the caves, is also observable on the larger jaws, while the smaller resemble in this respect those of the recent large Feles. ‘The jaws we have principally used in our descriptions are from Bleadon and Sandford Hill. Of these, nearly perfect specimens of both varieties exist in the Taunton Museum, being a portion of the collection of Mr. Beard. ‘There are also in the same collection many large fragments which confirm the description taken from the more perfect specimens. The fragments of lower jaw, showing the angle, condyle, and coronoid process (PI. I, 4 PLEISTOCENE MAMMALIA. fig. 8), which enables us to complete the illustration of the part missing from the Bleadon jaw was obtained from the brick-earth of Crayford, in the Thames Valley, and is now in the possession of Dr. Spurrell, to whose kindness we are much indebted for the loan of the specimen. It was found in association with the tichorhine, leptorhine (Owen), and megarhine Rhinoceros, Wephas antiquus and H. primigencus, horse, red deer, bison, &c. It probably belongs to the smaller variety. A reference to the table of dimensions will show that the measurement of this bone are not greater than those of the lion and tiger. A second specimen from the same locality gives the extent of the alveolar border occupied by the teeth at 3:20 inches, while PM3 = 0°78 X 0°41 X 0:39, PM4 = 1°12 < 0°56 < 0°66, and Mi = 1:15 x 0°51. A third mutilated ramus from the brick-earth at Ilford, in the possession of Mr. Brady, has a circumference anterior to premolar three of 4°90 inches. A jaw of the large form was found at Fisherton by Dr. Blackmore, and others exist in different parts of the country. Messrs. Schmerling, Marcel de Serres, Dubrueil, and Jean-Jean describe under the name of Felis leo, and Messrs. Croizet and Jobert under the name of /. antigua, various bones of large Feles found in the caves of Belgrum and France, and in the Pleistocene deposits of the latter country, and consider them as distinct from Felis spelea. We have no doubt as to the correctness of the four first-named authors in their deter- mination of these bones as those of Felis feo, confirmed as it is by the remains of that animal in the Taunton Museum, from Sandford Hill and Bleadon. 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Ss ve) ° at, S a o Ss S Tmndarrd “uInasnyy woWNe,T, “najads sya 6 PLEISTOCENE MAMMALIA. CHAPTER II. Fehs spelea—Fore-arm, P\. II. J. Ulna, figs. 5, 6,7, 8, 9. We have unfortunately been unable to procure the distal end of this bone. The smaller specimens correspond so exactly with those of the lion that we are unable to perceive any difference, save a very slight diminution in size of the recent specimens, which, we believe, have been all, or nearly all prepared from animals which have lived most of their lives in cages. The larger, however, present some differences, which, we think, necessary to describe. The line drawn from the proximal end of the humeral articulation to the summit of the olecranon forms a more acute angle with the axis of the bone in Ff. syelea than in F. leo (figs. 5, 6, 7, 8, 9, a, 6). The large humeral articulation (figs. 5, 6, 7, 8, 9, a, a) is precisely alike in both. The proximal radial articulation (¢, ¢’, figs. 5, 6, 8, 9) is more nearly at a right angle with the axis of the bone in /. spelea than in F. /eo, though it still forms an angle of 60°, that in F. /eo being 50°. In F. spelea the articulation is shallowest posteriorly, in 7. eo anteriorly. The coronoid process cisstrongly developed, but not more so proportionallythaninthe lion. The shaft of the bone appears, from the largest fragments we have met with, to have been straighter and deeper, in proportion to the thickness in the larger, than the smaller form. But the most marked difference to the eye is shown by the comparison of figs. 6 and 9, which show the extremes of the variation (fig. 8 being a somewhat intermediate form). Tt will be seen that in fig. 6 a strong ridge runs parallel to the back of the bone on the outer or radial surface (figs. 6, 8, 9, d, @’), forming a broad shallow groove which extends -downwards as far as the specimens we have seen allow us to observe. It will be seen that this ridge terminates or dies away in fig. 9 a little below the radial articulation (fig. 9, d); in the recent specimens of the lion it is still less visible. As, however, the length and prominence of the ridge appears to vary precisely as the size of the specimens, and we have examined above twenty, we cannot look upon it as a characteristic difference. Generally speaking, the surface of the bone in the larger specimens appear to present well-defined ridges with rounded contours, and smooth FELIS SPELAA. ji shallow grooves between; while the smaller and more strictly leonine forms present more angular and less defined ridges, and the grooves between appear to be more cut up by small irregular supplementary ridges. Fragments of this bone are very numerous in collections of Somerset bones from the caves of Hutton, Bleadon, Sandford Hull, and Wookey Hyzena-den, specimens of both varieties having occurred im all. The largest number we know of together is in the Taunton Museum, from the collections of Messrs. Williams and Beard. The only bone of the Pleistocene deposits that is likely to be mistaken for this is the same bone of the bear. ‘The latter is, however, much straighter, and never has the broad external groove of the shaft described above. But the most marked distinction is in the proximal radial articulation, which m /. spelea, as in all the Feles, invariably forms an acute angle with the axis of the bone, while in the bears it is at right angles. The form of the olecranon is also somewhat different, but its irregular form can be more easily recog- nised by a comparison of the plates of the bones than from the most elaborate description. There is a figure of this bone belonging to the smaller variety in Sch merling’s ‘ Oss. foss. de Liége,’ vol. 2, pl. xix, fig. 1, and a very characteristic one figured by Marcel de Serres, &e. ‘Oss. foss. de Lunel-Viel.,’ pl. viu, fig. 3. Felis spelea. Felis leo. Felis tigris. ac Bleadn Sandford) § *, * MrasuremMeEnts or Una. ; Hill. |S & 2 2 5 (In Inches.) a 5 = a iL 2. oe 1. mee S |W.AS) = a Figured| Appa- Figured) p.& 2 = = speci- jrentlythe speci- | 4 § s men. [pair of 1. men. | Oo & io) jo) | lhene thayextremcn nur ote) deo Ht nas ae spe | aller || ea. |) ate) Depth just below radial articu- lation,) proxintallnaee nse) sae. 62. 2°68 | 2°69 | 2°25 | 2°66] 1-3 1:74 | 1:62 1°87 Thickness (transverse) at same OWE. sae se rae mee tee cna: 1:05 | 1:00 | 1:17 pte Heil 0°86 | 0°87 0°75 Circumference at same point...... 6:00 | 6:00) 5:12 | 610) 65 4°39 | 3°24 4:0 Minimum circumference ......... oe wf A) sls 2-40 | 2°0 2°25 Humeral articulation, vertical Cinear)) .). cen eee eee on aS 3°00 | 3°40 on 2:0 2:80 | 1:37 1°75 Transverse humeral articulation...| .., Bi) | PIO WN sae 7 150 | 1°62 1:87 Radial articulation:pro- wera aN 0:70 | 0°70 a 0°57 | 0:50 | 0:5 0-62 ximal, vertical ...... posterior | .., nye a8 ten aise 0°60 Ditto ditto ditto, transverse...| ... 2°00 | 1:70 cae 1:6 1°63 | 1:5 1:37 Ditto distal articulation, vertical| .., nD ie 4s Big 0°60 | 0:5 0°5 Ditto ditto ditto, transverse...| .., aes oe nae a 0°60 | 0°6 0°62 Carpal articulation, vertical ...... Bi, : a “ef Bee 0°75 | 0°62 1:0 Ditto ditto, transverse ...| ... a Sob Boe Bee 0:48 | 0°37 0:5 From radial to carpal articulation, S DOtMMIMGIUBIVEs.......,0..-5.cacansee nae sae ae Bar a 165 | 1:25 2:0 * For these measurements we are indebted to Mr. Charles Robertson, the able Demonstrator of Anatomy at Oxford. § PLEISTOCENE MAMMALIA. 2. Radius, Pl. II, figs. 1, 2, 3, 4. The general form of this bone is like that of many other carnivora; the lower half is straight, while the upper is bent gently backwards in a slight curve; immediately below the proximal epiphysis it is bent sharply forwards, so that the humeral articulation which terminates the bone is set at an angle of about 70° to the axis of the bone (figs. 1, 2, 4, a). The section of the bone at the distal epiphysis is nearly triangular, with the outer surface convex, while the inner is nearly flat or slightly concave, and the posterior deeply so. ‘This section gradually passes into one which is a flat oval, rather wider on the after edge. The tuberosities (figs. 1, 4, e) give the bone, where they occur, a triangular section, the oval recurring just below the proximal epiphysis. The humeral articulation is roughly oval (figs. 1, 2, 4, a), being produced into a blunt point on the anterior edge (figs. 1, 2, e). It is concave with the exception of the anterior edge, where it is vertically convex. There is a slight eminence on the external edge (figs. 1, 2, 4, 7) which falls gradually away into the concavity of the articulation. The proximal ulnar articulation commences just under this eminence (figs. 1, 2, 4, /), and is continued posteriorly round three fourths of the head of the bone to the anterior point (figs. 1, 2, 4, e), where it is deepest. The distal ulnar articulation extends nearly the whole width of the bone, and forms a slightly concave oval, wider transversely than vertically, on the upper posterior edge of the distal epiphysis (fig. 3, g, where the edge alone is shown). ‘This is set at an angle of 45° to the axis of the bone. The carpal articulation (fig. 3) is roughly trapezoidal in form. ‘The imner or ulnar edge of the articulation is bounded by a semicircular arc, and the surface is concave, the outer boundary is convex, and the inner is straight when viewed from below, while on the lateral aspect it is boldly curved vertically. ‘The anterior edge of the styloid process is nearly semicircular; the surface of this part of the articulation is concave vertically, following the under surface of the styloid process, while it is convex transversely. The styloid process (2, 7, figs. 1, 3) is set slightly and obliquely inwards, while the process on the upper anterior edge of the epiphysis is set nearly in the plane of the greatest depth of the bone. Some of these bones from the Somerset caves are, as far as considerable fragments allow us to judge, indistinguishable from those of the lion (fig. 4). Others, of which we fizure one perfect specimen, are, as Cuvier remarks, generally far stouter in proportion to their length (figs. 1, 2, 3). The only differences in form which we can discover in this or in numerous fragments that we have examined appear to be due to this excess in stoutness alone, and can hardly be regarded as of specific importance. FELIS SPELAA. 9 All the specimens that we have seen are from the caves of Bleadon, Sandford Hill, and Hutton, and are in the Taunton Museum. This bone may be distinguished from that. of the bear by the greater prominence of the eminence of the humeral articulation in the latter animal (figs. 1, 2, 4, 7). This articulation is more circular in Felis than in Ursus. In the bear the ulnar proximal articulation is less pronounced and more rounded vertically; the tuberosities below this articulation are far smaller, and the bone is altogether more angular in section, more slender and flatter; the distal ulnar articulation is set nearly parallel to the axis of the bone on a small process. In the carpal articulation the strong upward curve on the internal edge altogether disappears, and the articulation is much less concave. Schmerling, in his ‘ Oss. foss. de Liége,’ gives good rough figures of this bone, both of the large variety (vol. ui, pl. xv, fig. 3) and of the smaller with the ulna (pl. xix, fig. 1). Felis spelea. Felis leo. a tigris. MEASUREMENTS OF pee eee g g Ravivs. is 3 Z 2 2 (In Inches.) a f = Large form. Small form. : z zi av ie} S) s) z ; x 6 2 3 = Ee 2 = 1 4 5 a Extreme length ............... olen ree cel eae bie [eset Hes S24 OOF ieee @) Minimum circumference ... | 3°50| 3°50] 3°85| 3°60] ... ... | 3°10} 3°10} 2°66) 2:5 yay hs) Transverse proximal humeral anticnlationiees. ease eee: AAS ee SO esi csc fi scee | 1:00)) epee OO) ie ose Vertical proximal humeral articulations ea.ee eos. ZnOde 2 OOe vy | L:95i|"... ooo (L250) pe ean he Tees 1°62 Vertical proximal ulnar arti- Guilation: yAkeen eee es: 0:55 0-54} 0°80) ... ae waz |) OF42 ee Orton Ons 0°5 Transverse distal ulnar arti- culation. Aenea Sl tONerra re yoga 1 LOO) oc eee OSo LOLS Tien Oo Vertical distal ulnar arti- Culation §..25-0-e eames 0°60] 0°67) ... ee soot OREO) son | O70 Transverse carpal articula- BIO RE! .- sa cooaee eee TAO) eS Ghee Wh ees toatl) UA Olin. aoe | Oot 0 Mee Vertical carpal articulation .. | 2°48) 2°40] ... | ... | 2°75] ... das Pane | SIRE IEG2NEA TIS 7, From proximal articulation ; to tuberosity, both inclu- SING shah ae een ree cae 2°30) 2:35| 2°60! ... ses poo | LIP aoe | APIO 8. ie 10 PLEISTOCENE MAMMALIA. CHAPTER IIL Felis spelea—Os innominatum, PI. IIT, fig. 1. We have met with but four specimens of this bone from the British Pleistocene deposits, all more or less imperfect. Of these, three were obtained by Messrs. Williams and Beard out of the bone-caverns of the Mendip Hills, and are preserved in the Taunton Museum, while the fourth and largest, which we figure, is derived from the brick-earth on the south side of the Thames Valley, at Slade Green, near Erith, in Kent, and is in the national collection. It consists of the iium and ischium, tolerably perfect, and a portion of the os pubis of the left side. The ilrum may be described as resembling the blade of an oar in form, slightly con- cave on the external, and nearly flat on the internal or sacral surface ; the parts answering to the crest (fig. 1, a, a’) and spinous processes (4, 0’, 6”) forming the strong rounded and raised border of the blade. The external surface is traversed by a longitudinal ridge (c, c’) that strengthens the attachments of the glutei muscles. his ridge is very slightly developed in the specimens from the Mendip caverns, while in that figured it reaches a maximum development. The symphysis of the sacrum extends slightly along the upper edge of the shaft or body, and takes the form of a small spine or process at 6. This shaft or body (c, d) is massive, and of great strength, flattened internally and rounded externally, and ending inferiorily in a sharp free edge (@). The anterior inferior spinous process (¢) varies in form and size in the different individuals, but bears a general resemblance to that of the lion. The portion (/) of the os innominatum that composes the apex of the sacro-ischiatic arch immediately above the acetabulum is convex and rounded, and without any ridge. It forms the connection between the shafts of the ilium and ischium. ‘That of the latter (g, 4) is prismatic in form, tapering posteriorly and expanding downwards and transversely mto a broad triangular blade, the anterior edge of which forms the posterior boundary (7) of the pubic arch. The ischial spine (4) is a small pyramidal process immediately above the posterior edge of the acetabulum. FELIS SPELAIA. Wi The shaft of the pubis (/, m) is convex externally and flat internally, and in some of the fossil specimens is proportionally more massive than in those of the lion and tiger, _ with which they have been compared. The acetabulum is proportionally larger, and the -cotyloid notch (z) shorter in the spelzan than in the recent lion. ' With these two exceptions, a close comparison of the speleean ossa innominata with those of the recent large carnivora proves that the only difference between the pelvis of F.. spelea and F. leo and F. tigris is one of merely size. We are unable to detect any specific differences between the pelves of the two latter animals. We have given a figure (fig. 2) of the pelvis of /. tigris in the British Museum, three fourths of the natural size, to show the far greater size of that of the Felis spelea. PLEISTOCENE MAMMALIA. 12 68-€ GV. 1¢.2 “sny “WU *40]09U00 SHOT 06-¢ 05-0 60-1 p9-T 02-2 “sn “IU *snpindoay SHAT 06-7 “SO “IN “DaUuo sya UOT] 94} UL SB oes oY} Ajfe12UeS oq pnom uoyJAOdo.d sity, 60-7 00-9 GEV 6-¢ 09.1 GLe1 08-0 09.1 08-¢ 6:61 “sn “Ig ‘TPIL *s2.060g sya 0¢-G 'S°V-M 09) syag 99.¢ “sn “Ag “TCIl "00) SYayT [3-€ G3-E1 “snyl UoyUNeT ‘IH ptoypues), ‘pajads ‘goojtoduit yyoq are eI asaug, y 0¢-Z 00-2 40G-€ 0¢-V1 "SHI “AG 9219) BpeIS Age A somMey y, SYOT JOJOUIVIP IVI[I-TUL9G Jajawip siqndimag OTA] dIAOg WINIYOST JO LaWVIPIUaG wniyost jo yydaqg WNITIST JO pues 0} WH[NQe0V " WN[NQey20¥ Jo TajawWVIP [BUIpNySuOrT “"" tunyNgezs08 JO JOJOWIVIP ISTOASUBIT, UNIS! JO ssoUyoy) WNUIXEL 2S cee uniyostro qys1aq WNW, WINIE JO FSI9Y [BOTA A “"* 4 8uay [vO], (-seyouy uz) ‘WALVYNIWONN]T SQ dO SUNTWNAUOS VA] ee Ne a ee Se ee See ue ut FELIS SPELAA. 13 CHAPTER IV. Felis spelea—Yarsus, Pl. IV. Astragalus, fig. 1. Tue astragalus of Felis spelea is precisely like that of the lion and tiger in form, but, with the exceptions of certain specimens which we shall notice, is much superior im size to those of either of the latter two animals we have met with. The tibial or proximal articulation (fig. 1, a, a’) is of the usual pulley-like form ; it is inclined to the antero-posterior diameter, at an angle of about 30°; at the back of this are set the two articulations for the calcaneum, the outer being concave and somewhat broader at the top than the bottom, and this matches the great sigmoid articulation ; the other is subtriangular and matches the small lateral circular articulation (¢ of fig. 2) of the calcaneum. ‘These three articulations are the surfaces of the head or proximal portion of the bone. This is connected with the navicular or distal articulation (fig. 1, 4), by a short shaft or neck, as in all the other digitate carnivora. This articulation is suboval in form, and bent vertically and diagonally, so that it is highly convex, and matches well the deep concave articulation of the navicular bone. The only cave fossil that is likely to be taken for this bone is that of the bear. But the latter is easily distinguished by the extreme shortness of the-neck or shaft joining the proximal and distal portions of the bone. The pulley-groove of the tibial articulation is also flatter in the bear, and on the inner posterior edge this articulation is terminated by a sort of spur rising at right angles to the surface, which does not exist in the Feles. Among the many specimens of this bone that have occurred to us there are some in the Taunton collection which are much smaller than the rest, and very little exceed those of a large lion in size. hey do not, however, offer any other difference. 14 PLEISTOCENE MAMMALTA. Merasurements.! Felis spelea. No Bion (aon | me ACO. Sandford Wo Bo Br. Mus.|Br. Mus. Hill. Bis tha TR. 2. Ib 21, IL} Bex | srily | SOO | Be | 270 | BeO | B35 | sem | MoH | aed) | aes) |) oPO2 |} aril | Bead | Be7H | Sl) || SeOsH | So 3 | Use |) lee ew} Meal) |) heals eB yO) |] WAL | ew A) SSS | So |] SW | 279 || Beals | BUH | PSO | Bog | geae Hi wezO | WeOs | eG | desi | WS | esO | RO | 1eA@ | O@e GH Wes |) News | Tee oN) PO | CO | OOO) Teng | ey The 4th and 5th specimens in the above list are of the small variety above mentioned. The specimen figured is the first in the above list of measurements. It is with many others in the T'aunton collection, from Bleadon cavern. Others have occurred to us from Sandford Hill and Oreston. Calcaneum, fig. 2. The general form of the caleaneum of Felis spelea, like that of all the Feles, and of most, if not all, the digitate carnivora, is nearly straight from the bulb which forms the attachment of the tendo Achillis (fig. 2, a) to the cuboidal articulation (fig. 2, 6). From this bulb the bone slightly increases in size to the upper part of the outer astragaline articulation, at which point is the maximum vertical measurement (fig. 2 ¢’). The posterior boundary is generally a gentle convex sweep through the whole length of the bone ; in some specimens this becomes nearly straight or even very slightly concave as it approaches the bulb for the tendo Achillis. The outer astragaline articulation (fig. 2, c, c’) is a broad sigmoidal surface, bent backwards vertically on itself. Near the level of the middle of this surface a stout process rises laterally, at right angles to the body of the bone (fig. 2, d, e), and supports at an angle of about 30° to the axis, a nearly circular surface, which is the inner astragaline articulation (fig. 2,e). In the form of this articula- tion it resembles the lion, that of the tiger being transversely oval. The mass of bone which connects these with the cuboidal articulation is nearly cubical in form, and has on the external surface the prolongation of the cuboidal groove for the tendon of the peroneus ‘In this, as in all cases throughout these monographs, when the numbers alone are given, the measure- ments are—l, extreme length; 2, minimum circumference; 3, virtual measurement at the proximal articulation; 4, transverse ditto ditto; 5, vertical measurement of distal articulation; 6, transverse ditto ditto. i aa FELIS SPELAA. 15 longus (fig. 2, f). The cuboidal articulation (fig. 2, 4) terminating the bone distally at right angles to the axis forms a nearly circular surface, a portion of the circle being cut off by a chord, which forms the mner posterior boundary of the articulation. If this cord were slightly bent, it then would approximate very closely to the recent leonine form, which is nearly an oval. The attachment of the tendo Achillis is formed by the process rising on the inner edge of the bone at right angles to the rounded terminal surface (fig. 2, 7). In this it appears to differ slightly from that of the lion, for in that animal the groove formed by the two surfaces is replaced by a nearly flat or slightly rounded surface at an angle of about 45° to the axis. | The only differences which we can observe between the corresponding bones of this animal and the tiger are that mentioned above, and the rounded or oval form of the cuboidal articulation in the former, while in the latter it forms a quarter circle. Among the large number of calcanea we have examined, we find three or four which do not appear to differ in any respect from those of lion, with the exception of bemg very slightly larger, the difference of size in general being very great. ‘That figured probably belonged to an aged animal, the outer portion of the bone being roughened by exostosis. Trelis spelea. Lion. Tiger. MEASUREMENTS OF CALCANEUM. Bean Sand- BSH Ne ae (In Inches.) a W.A.S.| Mus. | Mus. ie 9 9 il 19 12 13 112 L.| 114 L, Ieee Lotalplenothmermrrrr ss. och ecsc: 5°60 | 5°24 | 5°34 | 5:00 | 5-01 | 4°52 | 4-48 | 3:90 | 4-00 | 4-30 2. Minimum circumference ......,.. 4°73 | 4°24 | 4-30 | 4°40 | 4°30 | 3°90 | 3°70 | 3°30 | 3°15 | 3°45 3. Maximum vertica] measurement | 2°36 | 2:10 | 2:08 | 2°27 | 1-82 | 1°87 | 1°80] 1:59 | 1:60 | 1°77 4, Maximum transverse ditto PON PenOOr 2 Og) 1-97 2-03} 1°80) e706 7a eA Sa eG! 5. From inner articulation to the outer end of the bone, articu- lationeinclad cca 3°95 | 3°70 | 3°90 | 3°68 | 3°77 | 3°36 | 3°45 | 3°13 | 2°90 | 2-74 6. Sigmoidal articulation, transverse measurement ......... SES catch See 0:92 | 0°82 | 0°82 | 0°85 | 0°75 | 0°74 | 0°72 | 0°60 | 0°55 | 0-64 7. Cuboidal articulation, transverse MLEASUTEMeENt 7 eee eee ee 16329) 1-30) 1-45 | 13381) 1-305) 104) 1:25. )0:908 1502)0-92 SS Ditto: “ditto, verticals... 1717 | 1:13 | 1°26 | 1°15 | 1°17 | 0°95 | 0:93 | 1:02 | 0:82 | 0:85 9. Inner astragaline articulation ELAMSVETSE! hecccos ct acc eee ee 0°81 | 0°80 | 0°84 | 0°88 | 0°79 | 0°68 | 0°68 | 0°71 | 0°70 | 0:73 10. Inner astragaline articulation, WET iss eer eB 0°74 | 0°70 | 0°72 | 0°74 | 0:55 | 0°70 | 0°57 | 0°67 | 0:69 | 0°55 The specimen figured is from Bleadon, and evidently belongs to the same individuals * Specimen figured. 16 PLEISTOCENE MAMMALIA. as the cuboid, astragalus, &c., which we have figured ; it is at Taunton, with many others, from the collection of Mr. Beard. ‘The adult bones are from Bleadon and Sandford Hill, and the same bone of young animals from Bleadon and Hutton. Naviculare, fig. 3. This bone, as its name denotes, is somewhat boat-shaped, the upper concave and nearly circular surface forming the articulation for the astragalus (fig. 3, a), and the two flat inferior surfaces, slightly inclined to each other (fig. 3, 4, c), forming those for the ecto- and meso-cuneiforms (figs. 5, 6), while a slightly rounded surface on the inner side forms the proximal attachment for the endocuneiform (fig. 3, d). The hinder portion of the astragaline articulation is curved sharply upwards, and this portion is supported by a process (forming, as far as this bone is concerned, the upper border of the great tarsal groove for the peronews longus), which is generally much slighter in Felis spelea than in lion or tiger, it being proportionately the largest in the latter of these three animals. This forms the only appreciable difference observable in this bone except size between them. It is distinguished from the corresponding bone of the bear by the somewhat greater thickness and by the greater development of the upward prolongation of the astragaline articulation, and also by the more decidedly circular form, as contrasted with the oblong and somewhat angular plan of that of the latter animal. MEASUREMENTS. lege ' Felis spelea. : Lion, , Tiger, : come te Br. Mus. pace Br. Mus. Gal So Bleadon. BAe Te ) 114 L. ] 0°75 0°66 0°66 0°65, 0°63 0°39 0°55 0°56 2, 5°45 5°00 4°77 4:03 4°20 3°98 4:90 4°35 3 1-24 1°20 NE) 0:98 0°91 0:90 0°91 0:97 4 1:79 la4 1:50 1:38 117 1:10 1:25 1°20 5 1°33 1:27 1°26 1:02 1:02 1:00 1:06 1-00 6 1:94 1°66 151 1:00 1:10 0:91 1:00 0°83 The bone figured is from Bleadon, and is at Taunton ; it probably belongs to the same animal as the astragalus, calcaneum, &c.; but it is from the right paw, and for the sake of uniformity is reversed in the plate. Schmerling gives two good figures of this bone, showing the proximal and internal surfaces (‘ Oss. foss. de Liége,’ t. ii, pl. xvii, fig. 4). FELIS SPELAA. | 17 Cuboid, figs. 4, 4’, 4”. The general form of this bone is well expressed by the name, the bounding surfaces being roughly at right angles to each other; the distal surface, however, being larger than the proximal. The outer and posterior or lower surfaces are traversed, diagonally and downwards, by a deep groove for the tendon of the peroneus longus (figs. 4, 4’, 4”, 6, 6); a continuation of the groove being formed on the outer and distal surface of the caleaneum in an upward direction, and by the hook of the ectocuneiform and the process on the hinder part of the naviculare in the opposite and transverse direction. The proximal or calcaneal articulation (fig. 4, 4’, c) 1s nearly flat and subcircular. The distal or metatarsal articulation (fig. 4, 4”, d) is concave, and may be described as an oval, the outer side being bent inwards so as slightly to resemble a B in outline, while that of the ectocuneiform is trapezoidal and nearly flat (fig. 4, 4’, a). The difference in size is the only difference we can remark between this bone and that of the recent lions and tigers. In some specimens of tigers we have observed a dis- position in the tarsal articulations generally to separate up into small surfaces, whereas im the lion and Felis spelea they appear to coalesce to a greater extent, and this is especially the case in the cuboid. MEASUREMENTS. Felis spelea, Bleadon, Lion Lion, Br. Museum, Lion, Tiger, { Taunton. 112 L. Col. Surg. Br. Mus., 114 L. 1 1:39 1:04 1:00 1:05 110 2; 4°22 3°00 3°25 3°80 3°27 3 1°36 0:70 0°86 0:90 0:90 4 1°36 1:00 0°61 0°68 0-78 | 5 1:20 1:00 0°81 0°82 0°76 6 1:24 0:98 0°86 0°82 0°94 | The only specimen we know of is from Bleadon. It evidently belongs to the same individual as the calcaneum and several other bones of the tarsus, which we have figured. Schmerling gives rough figures of this bone, apparently taken from an imperfect speci- men. ‘They appear to be the external and posterior views, but one of them is reversed in position (‘ Oss, foss. de Liége,’ t. ii, pl. xvii, fig. 5). 18 PLEISTOCENE MAMMALIA. Ectocuneiform, figs. 5, 5’. This bone has a wedge-shaped body, the superior surface (fig. 5) forming the head of the wedge, and the inferior or plantar the vertical edge. The flat navicular proximal (figs. 5, 5’, a) and slightly concave distal metatarsal (figs. 5, 5’, 4) articulations are both nearly isosceles triangles. From the inferior surface of the bone a very stout hook-shaped. process is developed (fig. 5’ c), that advances forwards to terminate in a rounded boss. ‘This hamular process affords attachment to the plantar ligaments that extend to the cuboid and meso- and ectocuneiform, and to the tidczalis posticus and flexor pollicis muscles. In all the British specimens of the bone that we have seen, the hamular process advances forward within a short distance, from 0°10 to 0:20 inch of the plane of the distal surface, and in no instance as far as the distal plane. This also is borne out by the examples from the cave of Gailenreuth, in the collection of the Earl of Enniskillen, F.R.S., to whose courtesy we are indebted for the examination of his museum. A large series of ectocuneiforms belonging to lions and tigers, in the museums of London and Oxford, shows, as one might naturally expect, considerable variations in the development of the hamular process ; and while we find many ectocuneiforms of lion that cannot, in this or any other respect, be differentiated from those of tiger, yet, on the whole, it has a smaller distal extension in the former than in the latter (fig. 5’’, c). The maxi- mum distal extension is seen in the specimen of tiger in the British Museum figured fie. 5’”, c, in which it extends as far down as the plane of the distal articulation. We must admit, therefore, that while the development of the hamular process in the ecto- cuneiform of Felis spelea is no absolute test of leonine or tigrine affinities, yet that it points rather in the direction of the former than the latter. M&ASUREMENTS. Felis spelea. Lion. Tiger. Bleadon. Sandie Col. Surg. a ae W.A.S. ae Col. Surg. 1 0°83 0:81 0:96 0°64 0°70 0°70 0:80 0°74 2 3°95 3°52 4:00 PPUT 2°90 3°10 3°25 3°05 3 1°05 0°87 1:00 0°82 0:62 0:90 0°70 0:80 4 1:05 1:02 1:04 0:98 0:95 0°93 0:94. 112 5 1:08 0:98 1:04 0°85 0:80 0°93 0:90 0°80 6 1°36 1°19 1°45 1:00 1:03 111 115 1:14 The specimen figured is the first in the list of measurements, reversed in order to get the most perfect representation on the same side as the rest of the tarsus. FELIS SPELAIA. 19 The three specimens measured formed part of the collection of Mr. Beard, and are now at Taunton. ‘There is a good rough figure of the distal articulation of this bone in Schmerling (‘ Oss. foss. de Liége,’ t. 1, pl. xvii, fig. 6). Mesocuneiform, fig. 6. This small bone has the proximal and distal articulations slightly inclined to each other, and wider apart internally than externally. The proximal or navicular articula- tion is roughly oval, but slightly pointed anteriorly, and is slightly concave transversely and convex vertically, while the metatarsal or distal is wider anteriorly than posteriorly, and is very slightly concave vertically. The inclination of the articulations one to the other, and other minute points in the form of the bone, appear to differ in different individuals of lion and tiger; consequently the slight difference we observe between the only specimen we know of this bone in Feks spelea and the above two animals is probably not of specific value. Generally speaking, however, it appears to be a shorter and thicker bone altogether in the fossil than in either of the - recent large species. It may be easily distinguished from the corresponding bone in the bear by the greater squareness and angularity of the anterior or upper face of the bone in that animal, as con- trasted with the rounded and oval form in Felis spelaa. Both distal and proximal articulations in the bear are also much more concave, and the articulation for the endocuneiform is well developed, whereas scarcely a trace of it exists in Felis spelea. MEASUREMENTS. Lion. Tiger. Felis spelea, Bleadon, Taunton Col. Br. Mus. Col. Br. Mus. Col. 112 L. Surg. 1141. Surg. 1 0°55 0°39 0°35 0°61 0°51 2 2°90 2°10 1°89 2°00 2°40 3 0°54 0°40 0°35 0°43 0°42 4 0°91 0°66 0°70 0°68 0°68 5 0:50 0°46 0°42 0:54 0°52 6 1:02 0°72 0°65 0°69 0°79 The specimen figured is from Bleadon, and probably belonged to the same individual as the astragalus, calcaneum, naviculare, and cuboid, which we have figured. It is at Taunton. We know of no other figure of this bone. 20 PLEISTOCENE MAMMALIA. Endocuneiform, fig. 7. It is with considerable hesitation that we give a figure of this bone. The general form and surface resemble the same bone in the lion and tiger, but there are differences which make us doubtful whether we have assigned the bone its right place. It has lost the proximal epiphysis. If this is restored as a feline endocuneiform (fig. 7, a), it would then be very similar to the same bone im lion and tiger; but the anterior portion of the distal articulation points downwards in an acute angle, whereas in lion it points nearly directly forwards in a right angle. The articulation also is slightly convex in our bone, whereas it is slightly concave in that of the lion. Unfortunately, the corresponding articulation of metatarsal 1 is broken in the only specimen we know of, so that we cannot say whether this bone showed a corresponding variation in form. We think, however, that attention should be drawn to the bone, in the hope that some one, more fortunate than ourselves, may discover a perfect specimen, and decide whether we are right or wrong in our determination. The only specimen we have being imperfect, we give no measurements. It is from Bleadon, and is in the Taunton Museum. We know of no other figure of this bone. F : FELIS SPELAA. 21 CHAPTER V. Hinp Paw, PI. V. Tne metatarsals of the carnivora are so well known, and so much resemble each other in the digitate forms, that comparative anatomists are generally content to refer to the figures of those bones rather than to attempt to discriminate them by descriptions, which must all very closely resemble each other. If we attempt the course which, in nearly all other portions of the skeleton, is held to be absolutely requisite, it is with the view of doing what we can to render the descriptive portion of our work as perfect as we can, rather than to trust to bare figures. Metatarsal 1, fig. 1. It is well known that the first metatarsal of the genus Felis is rudimentary. In fact, it would be difficult for one not acquainted with the bone in question to recognise it as a metatarsal at all. In Felis spelea it is a small wedge-shaped bone, the wedge thinning off anteriorly to an edge, and distally to a blunt pot. The internal surface is slightly rounded; but the external is flat, and the posterior is irregular. The articulation for the endocuneiform is slightly concave transversely and convex vertically; and, as we suppose, from the corresponding bone in the lion, it would be furnished with a small hook-shaped process on the external’ and posterior edge. ‘This part is mutilated in our only specimen (fig. 1, a). In other respects it does not differ “appreciably from that of the lion, except that in that animal the articulation is flatter. The specimen figured is from Bleadon, and is in the Taunton collection, and is reversed from the left paw. We have to repeat what we have written respecting other specimens of various 1 Internal and external, in these descriptions, are invariably used in reference to the position of the whole limb in the skeleton, and not to the position of the bone as to the joint. 22 PLEISTOCENE MAMMALIA. parts of large Feles from the Somerset caves, that there occur metatarsals which, both in size and form, so exactly resemble those of the recent lion, that they cannot be distin- guished from them. (We have observed, however, a tendency in those of the tiger to a ‘greater proportional development in the antero-posterior direction of the proximal articu- lation, which appears generally to distinguish the metatarsals of that animal from the other two large feline animals.) But im all the Pleistocene deposits there occur, but not numerously, except in the Somerset caves, bones of larger size. The set which we have figured evidently belonged to the same animal, though metatarsal 3 is reversed from the left paw. They were found by Mr. Beard, with many other parts of the same skeleton, m Sandford Hill Cave, m the Mendip, and are now in the Taunton collection. Some larger specimens from Crayford, in the valley of the Thames, are figured in PI. VIII. They belong to Dr. Spurrel, and are nearly, if not quite, equal to the largest German specimens. They differ from those we have figured in Pl. V by having the distal articulations comparatively much smaller, as well as by their tapermg more and being more bent. We have observed similar variations in some of the cave specimens, as well as in the metatarsals of recent lions and tigers. Metatarsal 2, fig. 2. The shaft of the second metatarsal is somewhat triangular in section, the sides being flatter towards the proximal and becoming more convex towards the distal articulation, so that the bone then becomes almost cylindrical. The shaft is slightly curved backwards, the outer boundary, or that facing metatarsal 3, being nearly straight, and the inner, or that facing metatarsal 1, being curved slightly inwards, so that the bone appears to bend slightly in that direction. The proximal articulation for the mesocuneiform (fig. 2, a, 4) is nearly at right angles to the axis of the bone. It forms a roughly triangular surface, of slightly double curvature from front to back, where it ends in a small spur, curving sharply upwards (3, fig. 2). The surface is concave, in a transverse direction. The point of contact with metatarsal 1 can hardly be called an articulation ; it is slightly smoothed, and is supported by a small process (¢, fig. 2), a short distance below the proximal articulation, on the inner front edge of the bone. The ectocuneiform articulations, together with those for the third metatarsal, form two very slightly concave oval surfaces at right angles to the proximal articulation, and con- tinuous in direction with the outer surface of the bone. ach of these surfaces is divided by an horizontal ridge, very slightly marked ; the upper part of each belongs to the ecto- cuneiform, the lower to the third metatarsal. The anterior edge of the anterior of these is bent outwards, so that the front of the bone presents the angular projection (fig. 2, d, e), and when the metatarsals are applied to each other the point d rests on the point a (fig. 3). FELIS SPELAA. 23 The distal or phalangeal articulation is, unfortunately, mutilated in all the specimens of this bone that we have seen; but it probably resembles that of the lion, excepting that the lateral development of the internal process (g, fig. 3) appears to be much less in the large cave form than in the recent. It probably, like that of the lion, had the outer surface of the bulb much flatter and more deeply indented than in the other metatarsals. Metatarsal 3, fig. 3; and Pl. VIII. The shaft of this bone is cylindrical, slightly flattened on the front surface; it is very slightly bent backwards, and expands slightly laterally at the distal epiphysis. The proxi- mal or ectocuneiform articulation is very flat (fig. 8, a, 6), and inclined at an angle of about 60° to the axis of the bone downwards and inwardly, being at right angles to the front surface. It is bounded anteriorly by the curved front edge of the bone, inwardly by a waved line, posteriorly by a very small spur, rising from the surface (fig. 8, c), and exter- nally by a waved line like the internal boundary, but set at a right angle to the front, while the internal is at an acute angle with the same. The articulations for metatarsal 2 are the small polished heads of two small eminences, corresponding with the summit of the internal waves above mentioned; while those for metatarsal 4 are two larger oval sur- faces, on the opposite side of the bone, the anterior lining the surface of a hollow facing diagonally backwards and downwards, the anterior edge of which forms the boundary of the front external expansion of the proximal extremity of the bone (fig. 3, 4, d), while the posterior articulation is a flat surface, facing directly outwards and upwards. This, with the posterior articulation for metatarsal 2, and the posterior portion of the proximal articulation, are supported by a large process, the lower part of which gradually slopes into the body of the bone. The bulb-shaped phalangeal or distal articulation is proportionally wider than that of the recent lion, but is in other respects precisely similar. Metatarsal 4, fig. 4. This bone is somewhat more bent than metatarsal 3, and the section is somewhat more angular, giving a square section just below the proximal epiphysis. It is rather more bent backwards than metatarsal 3, and in the slightest degree outwards. The proximal or cuboidal articulation (fig. 4, a) is rhomboidal and convex, both in transverse and vertical directions, and is bounded posteriorly by a shallow groove and small spur. Of the internal articulations for metatarsal 3, the anterior (fig. 4, 4) is a convex surface passing into a concave towards the ridge which separates it from the cuboidal articulation ; it rests posteriorly on a slight process, and faces inwards, upwards, and towards the front. The posterior of these articulations (¢) is slightly convex and oval in form, and faces 24 PLEISTOCENE MAMMALIA. backwards, inwards, and downwards, ma is, like the corresponding iolgubvi on meta- tarsal 3, supported on a massive posterior process. The articulations for metatarsal 5 much resemble those corresponding with them on metatarsal 3, but they are more concave; and the posterior, instead of facing slightly upwards, faces directly outwards, or rather a little downwards. ‘The distal articulation corresponds with that of the lion, except that it is slightly wider in proportion to the depth. Metatarsal 5, fig. 5. This bone is cylindrical, passing upwards into a triangular section, and downwards into an oval, wider transversely than in a vertical direction. The cuboidal or proximal articu- lation is a small oval surface, inclined to the axis of the bone (a, 8, fig. 5), and slightly concave transversely, and convex in an antero-posterior direction. It is bounded, externally, by a high, strong ridge or crest, which passes from the front, round the outside to the back of the bone, where it terminates just above the posterior articulation for metatarsal 4 (e, f). The anterior articulation for metatarsal 4 (fig. 5, c) much resembles the corresponding articulation already described in that bone; but the posterior (fig. 5, d) is a small, uregularly formed surface, the facing of which is generally backwards, inwards, and upwards. The form of the distal articulation corresponds exactly with that of the lion, being flattened internally, and extended laterally in an outward direction, but it shows the same slight difference in the greater transverse diameter. This bone in the specimens we have examined is more bent outwards and backwards than in the recent lion and tiger. In general, these bones may be easily distinguished, when perfect, from those of the bear, by the smaller proportional length, as well as by the far greater proportional size of the proximal end of the bone in Ursus. This gives these bones in the bear a taper shape, quite different to the solid, strong-looking bones of Felis spelea. For the details of difference we refer to our descriptions, measurements, and figures— FELIS SPELAIA. 25 MEASUREMENTS. Metatarsal 1 bemg mutilated, we cannot give measurements. Specimen Felis Felis Relenice f figured. Bl. B. tigris, tigris, | Coll ege of Felis leo, S. H. B. | Taunton. Brit. Mus. | College of | ¢ ceons. W.A.S. Taunton. 114L. | Surgeons. 2 Metatarsal 2. Inch. Inch. Inch. Inch. Inch. Inch. | Rt ccna ete 4°92 dd ie 4°04 4°25 4:08 4:66 FT SUB 2°18 2°36 Bue 1°75 1:60 1°45 We72, Pa eE a-ion co eaner 0-91 0:70 ot 0°60 0°59 0°56 0°63 eS SG aeCneae 1°30 1:50 th 115 0:98 "| 0°78 1:00 Bs eR ick was ne nee 0°84 071 0°65 0°85 6) es Sern Sais 1:80 an ta 1:60 1°45 1°37 1-60 4 5 | Metatarsal 3. BI. B. | Bl. B. Taunton. | Taunton. eer es sane .| 5°50 5°40 5°94) 4:97 4°88 4:53 5°20 2 Ae 2°40 2°20 2°55 1:98 1°78 1°60 1°83 PEED cocduce nes 1:10 1:05 127 0:98 0:98 0°83 0:94 4y he 1:50 1:37 dae 1:32 1°32 1°17 1°24 5 pesca 1:00 0°90 0°86 0°84 0°78 0°70 0°80 areas auaoeee 2°20 1°80 1:80 1°60 1°15 1°40 1°70 Metatarsal 4 Vis eRe arise 5°60 ads baie 4°95 4°55 4°54 5°30 4 Dh) crepe 2°30 eae 238 1:80 158 151 1°65 hs Fk ne 0°84 NG sie 0°62 0°68 0°67 0°70 A) Sue cites 1:60 Mee eigie 1:23 1°20 0:94 1°30 Ot eeeSbasadueNaee 1:98 a ee 1°78 0°61 0°65 0°78 Qiiremiteniy yo 2°00 aah Sele 1°55 1:50 1:50 1°75 2 Metatarsal 5. Bl. B. Taunton. |e Foie: Beer eee 5°20 here ae 4°33 4:05 4:20 4°60 DUNES EE tierce ol 1°87 1:26 ste 19337 1:53 1°40 1°37 Oi aoe osa te 1:00? 1:30 i 0-72 0°83 0:57 0.75 4 PERT ee 0:908 1:10 teh 0°74 0:90 0°83 0°82 Di ace ERR 0:80 a sue 0°64 0°72 0°63 0°63 Os eases: 1:80 Sat ace 1°38 1°53 1:26 1°50 Phalanges, Pl. V., figs. 6 to 14. We have had in the Taunton Museum, and in other places, abundant means of ex- amining the phalanges of Felis spelea. The well-known difficulties of assigning to each ' The largest Somerset specimen. 2 Both 4th metatarsal articulations inclusive. 3 Across cuboidal and metatarsal articulations. 26 PLEISTOCENE MAMMALIA. first phalange the correct digit have been felt by us; and, though we give figures of specimens in the order in which we believe them to occur, we are by no means confident that we are right in all, particularly as the great variation in size has added to our diffi- culty in that respect. Generally speaking, the first phalanges of Felis spelea present the usual character- istics of the genus. The deeply concave proximal articulation, the border of which is broken by a deep depression posteriorly (figs. 6, 7,8, 9, a, 4), the pulley-like distal articula- tion, and the strongly marked muscular attachment on the lower surface, are common alike to the Felidee and to the other carnivora ; whereas, the less taper form of the shaft, swelling out laterally and frontally on the anterior surface towards the distal epiphysis (figs. 6, 7, 8, 9, ¢, d), the deeper proximal articulation, and the less strongly marked muscular attachments below, distinguish the feline phalanges from those of the bear's, which are, as far as this species is concerned, the only phalanges that are likely to be con- founded with them. As we state above, the great variety in size added much to the difficulty of determining the proper place of each phalange, but we came to the determination to trust to form alone, and to figure the largest of each form which we found most to resemble the corre- sponding bone in the lion. Most of the phalanges we have seen are from that great depository of feline remains, Bleadon Cavern, in the Mendip, though we have good specimens from Sandford Hill, Oreston, Caldy, and other places. The phalanges of the hind paw may easily be distinguished from those of the front by the superior stoutness in proportion to their length; the flexure also is not so great. There is but little difference, except in size, between those of digits 3 and 4; but we have always found that of digit 2 curved at least shghtly, but sometimes considerably, outwards towards the centre of the paw, and to have the distal articulation set at an inward angle of about 60° to the shaft of the bone (fig. 6, e). This angle is sometimes, but not always, followed in the corresponding phalange of digit 3 (fig. 7, e); that of digit 4 being always at right angles (fig. 8, e), and that of digit 5 being set inwardly also at an angle of about 60° (fig. 9, e). This last digit is always strongly curved inwards. The outer may be distinguished from the iner side, in all the digits, by comparing the form of the sides of the posterior depression in the border of the proximal articulation. The angle at the summit of the outer side is nearly a right angle (figs. 6, 7, 8, 9, a); the inner side is more or less sloped, so that this angle at the summit is obtuse (figs. 6, 7, 8, 2), &). All those we figure are from Bleadon, and are nearly, if not quite, equal in size to the largest we have seen elsewhere, and are vastly superior in size to those of any living Féles ; but there are many fossil specimens of phalanges which do not differ in size or proportion from those of the existing lion and tiger. FELIS SPELA(A. 27 Second Phalanges, Pl. XI, figs. 10 to 13. The second row of phalanges of Felis may, as is well-known, be easily distinguished from those of other carnivora, by the peculiar outward turn of the distal articulation, which is so formed that the last or ungual phalange may fall back on the outside of the second so as to allow the claw to point upwards and protect its point from imjury- when retracted. We are unable to give any rule for placing the phalanges of each digit in order, except that the distal articulation of digit 2, and sometimes also that of digit 3, is not at right angles to the axis of the bone, but at an angle of from 15° to 30°, as represented in fig. 10 a. That of digit 5 is generally shorter and stouter, in proportion, than the others. We have figured the largest of each form that we have met with. They are all from Bleadon; they occur also from the Sandford Hill Cave, and from Oreston, Caldy, Ilford, and a great many other localities. The bone may be represented as triangular in section, from the distal to the proximal epiphysis, gradually expanding from the former to the latter. The proximal articulation is triangular in form, the inner and outer sides being convex, and the posterior deeply concave, having a deep pit within the concavity. It is divided by a ridge into two lateral concave portions, corresponding with the convexities of the distal articulation of the first phalange, and ends anteriorly in a truncated spur, pointing forwards and upwards. The distal articulation is, as is well known, a cylindrical roll, set transversely on the end of the bone, narrower in front than posteriorly, so that one end projects outwards considerably, and at a varying angle to the axis. The same rule may be given for the determination of those of the hind paws, by their superior stoutness, in proportion to their length, as we have given for the first phalanges. Third Phalanges, Pl. XI, fig. 14. The third or terminal phalange is a bone of rare occurrence. Five examples have occurred to us from English deposits; they are all from Bleadon. But one of them is in a sufficiently perfect state to figure and describe. It is probably that of the smaller hind toe. The lateral aspect may be gathered from the fig. 14, a being the articulation, the point of attachment of the tendons of the flexor, and ¢ that for those of the extensor or retractile muscles. The large triangular portion which extended beyond these points is a hollow sheath, which contains within it at d a strong core, upon which the claw is fixed, so that it is kept firm im its place by the sheath. The form of the whole of these phalanges is alike, and we can find no difference between those of either the fore or hind paw, except size, that of the thumb of the fore paw being enormous. One of these from Gailenreuth is in the possession of Sir P. De Grey Malpas Egerton, Bart., M.P., F.R.S., and measures 2°5 inches in antero-posterior length, and 0°8 inch in width. 28 PLEISTOCENE MAMMALIA. Ist Phalanges : MEASUREMENTS. Figured speci- mens. ates 44, Digit 1 Bl. B., Taunton. Inch. Lie eB 1°86 DARN eee Go oan 2°20 3) aan craanaee 0°97 Alc A een 0°67 Di, | Bele merical 0:73 6 een 1°25 we (l). Digit 2 BI. B., Taunton. ae cole ta 2°36 | 2) TEER ee en 2°36 | Oe ee ue anes 1:10 AGA aro 1°65 OVE LER A eeeneae 0:90 Gi ene oe 1:45 hohe 32. | Digit 3 BI. B., Taunton. AD ib ena aoe 2°01 Deere 2°34 SMEAR AN a. 3 1:02 2, Reset A 0°65 Br eeu a8 0°83 (Cr meen 1:13 Rive 48. Digit 4 Bl. B., Taunton. aes nes ee 151 Dar ee mea 1:97 Sek ees 0°83 A icc EAR 0°62 Ok ahaa eRe 0:68 py tee ary eee 1°10 Smallest Set. At Taunton. Felis tigris, Felis leo, 51. Brit. Mus. 114 L. W.A.S. BI. B Inch. Inch. Inch. 1:70 1°80 1:70 1:80 1:60 e7t 0°80 0°80 0°80 0:48 0:54 0°52 0°67 0°63 0°58 0:70 0°84 0°80 23. Bl. B 1:90 2°15 1:90 1°80 1:70 1:74 0°82 0°86 0:80 0°50 0°59 0°50 0°67 0°67 0°65 0°72 0°52 0°61 24 Bl. B 1°85 1:99 1:80 1:78 1:61 151 0°78 0:80 0:78 0°50 0:49 0°48 0:59 0°62 0:60 0:70 0:51 0°80 26. Bl. B 1°40 1:64 1:60 1°83 1°42 1°46 0°74 0°68 0°72 0°55 0°52 0-48 0:60 0°51 0:60 0:94 0:49 0°75 - PALMONTOGRAPHICAL SOCIETY. : yg INSTITUTED MDCCCXLVIL. LONDON: MDCCCLXVIII. THE beet TiS H PLEISTOCENE MAMMALIA. BY W. BOYD DAWKINS, M.A., F.B.S., GS., W. AYSHFORD SANFORD, F.G:S. 12 Tee BRITISH PLEISTOCENE FELIDA. FELIS SPELAA, GoupEuss. (Paeus 29—124; Prares VI—XIX.) LONDON: PRINTED FOR THE PALHONTOGRAPHICAL SOCIETY. 1868. FELIS SPELAIA. 29 MEASUREMENTS. 2nd Phalanges. bite 14, Felis tigris, Felis leo, Digit 2. Bl. B., Taunton. Brit. Mus. 114 L. Mio ZN SE Inch. Inch. Inch. Da) Re a oh 1°76 1:26 1:38 74, OE A a 2°00 1:45 1°42 Bae o cris Sane 0°78 0°68 0°60 Aaa anata ae : 0°60 0°43 0°50 Oi Seosaseeeee 0:78 0°64 0°60 OMEre etc cc:. 0°91 0:50 0°54 Digit 3 Bins LO eae 1°80 1:38 1:47 DA eee SS na 1°87 131 1225 By son ns ecomee 0°85 0°61 0°58 (a Als he ee 0°58 0:44 0°50 Ra oc 5p ean 0°72 0°62 0°60 Ope 0°75 0°42 0°50 Digit 4 a DSS a a ca 1°65 1°50 1:28 pains ieee Fae me 1:75 1:36 1°31 ON eg 0°83 0°56 0°66 Aiaih en e 0°56 0°43 0-51 De hotest aa: 0°75 0°60 0°54 One Ces 0:69 0-48 0°58 Digit 5 ane (is? 9, pe kD ee 1:57 120 1°20 Dare as OES 1975) 12 151 3 0°82 0°59 0°60 A eS ae a ae 0°53 0:04 0°50 Di pees asae a 0°72 053d 0°50 (i! ee ees 0°65 0°45 0°50 Sesamoids; Pl. V, figs. 15, 16. The sesamoids figured are from Bleadon. They certainly belong to a large carnivore, and are precisely similar to those of a lion in form, though of course much larger in size ; and as they are from the Bleadon Cave, where Felis spelea abounds, and bear is extremely scarce, we do not doubt that we are right in ascribing them to the former animal. : 5) 30 PLEISTOCENE MAMMALIA. CHAPTER VI. SKULEe els, Vil Vill, Vill x, XG Xd. CONTENTS. § 1. Introduction. Skulls of Felis spelea § 11. Squamosal. Sound in Europe. § 12. Malar or Jugal. § 2. Basi-occipital bone. § 13. Lachrymal. § 3. Exoccipital ; Supra-oceipital. § 14. Ethmoid. § 4. Basi-sphenoid. § 15. Wormian. § 5. Ali-sphenoid ; Pterygoid. § 16. Parietal. § 6. Pre-sphenoid ; Orbito-sphenoid. § 17. Frontal. § 7. Palatine. § 18. Nasal. § 8. Maxillary. § 19. Measurements. § 9. Intermazillary. § 20. Summary. § 10. Petrosal; Mastoid ; Tympanic; Malleus. § 1. Lntroduction. Skulls of Felis speleaa discovered in Hurope.—Perfect skulls of Felis spelea are so very rare that we have had the opportunity of studying no more than three which at all approach completeness—that from Sundwig, in Westphalia, now in the British, Museum, of which Professor Owen has given excellent figures in his ‘Memoir on Thylacoleo,”! and the two which we figure from the Mendip Caves, now in the Taunton Museum. ‘The typical skull figured by Goldfuss, and copied by Cuvier,” has altogether eluded our search. It was from the cavern of Gailenreuth from which Lord Enniskillen and Sir Philip de Grey Malpas Egerton have obtained vertebrae and many other bones of Felis, as well as large quantities of the remains of bears. There are, however, several figures of the skull. M. de Blainville® figures a fine and ' «Philosophical Transactions,’ 1859, pt. i, pls. xii, xv. 2 “Nov. Act. Nat. Cur.,’ tom. ix, p. 476, pl. Ixv; ‘Oss. Foss.” 1825, 4to, tom. iv, pl. xxxvi, fig. vi. 3 “Ostéographie, Felis,’ pl. xv. FELIS SPELAIA. 31 apparently perfect specimen, which M. Pictet reproduces in his ‘ Paléontologie.* From the text of the former writer we gather that the figure was taken from a plaster cast in the possession of Count Munster, the original having béen found in a cavern in Fran- conia. D’Alton? also figures a perfect or nearly perfect skull from Muggendorf. Other naturalists’ who have turned their attention to the Pleistocene Fauna describe and figure fragments only of the spelzean skull; for the species, though widely spread through central and western Europe, is nowhere abundant, nor are the remains generally found in a perfect state. The largest English fragment hitherto figured is that found in Kent’s Hole Cavern by Mr. McEnery, and drawn on a slightly reduced scale by Mr. Scharf, and published by Mr. Vivian. It is also figured in a woodcut in Professor Owen’s ‘ British Fossil Mammalia.’ It represents only a portion of the right maxillary and intermaxillary, with the dentition of an animal of the average size. Of the two skulls from the caverns of Mendip which we figure, the more perfect and smaller lay for many years after its purchase from the Rev. D. Williams, broken and unseen in a box in the Taunton Museum. ‘The fragments were put together by the pre- sent able curator, Mr. Bidgood, and the teeth were afterwards found among a quantity of those of hyzena; and thus a fair specimen of the skull of the British spelzean lion was obtained. Mr. Beard, the explorer of bone-caverns in Somersetshire, on his collection of bones being bought for the Somerset Archeeological and Natural History Society, told us of the presence of a skull of a lion from Hutton Cavern, among the bones some time before purchased from his old rival the Rev. D. Williams. As the skull agrees in its condition - and colour with the remains from that cave, we have no hesitation in affirming that the original of Pls. VI, VII, VIII, and IX, is the specimen alluded to. When Mr. Beard’s collection was brought to Taunton, the small pair of nearly perfect lower jaws figured in Pl. VI was found to correspond exactly with the skull in respect to age, size, and colour. We know that it was the practice of Messrs. Williams and Beard to work at the same cave at the same time, and to share the contents. In this way very frequently a fine specimen was divided between them, even in the case of the long bones,—femora, humeri, and the like. These, now that both collections are in the hands of the Archeological and Natural History Society, are in many cases reunited, and form perfect bones. We have therefore every reason for believing that the lower jaws in Mr. Beard’s collection belong 1 «Paleontologie,’ 4to, 1853—7, pl. vi. 2 «Raubthiere,’ pl. vii, fig. a, 4, ¢, d. § Leibnitz, ‘ Protogea,’ pl. xi; Buckland, ‘Reliquis Diluviane,’ pp. 17, 62, 261; Schmerling, ‘ Oss. foss. de Lidge,’ tom. ii, p. 14; Marcel de Serres, Dubreuil et Jean-Jean, ‘Oss. foss. de Lunel-Viel,’ pp. 101, 107, pl. vii, fig. 1; Rev. W. Vernon, ‘Phil. Mag.,’ 1829, p. 225; McHnery, ‘Cavern Researches,’ edit. G. E. Vivian, Esq., 1859; Owen, ‘Brit. Foss. Mam.,’ p. 161; ‘Rep. Brit. Assoc.” 1842; Falconer, ‘Quart. Journ. Geol. Soc. Lond.,’ vol. xvi, p. 490; Blackmore, ‘Cat. of Fossils in Salisbury Museum,’ p. 101; Boyd Dawkins, ‘Quart. Journ. Geol. Soe.,’ xviii, p. 115; Ed. Lartet, ibid., vol. xvi, p- 475; Falconer, ibid., pp. 99, 104; Baron Anca, ibid., p. 460. The last two notices may refer to a dis- tinct species, as the animal is described merely as a large Felis. 32 PLEISTOCENE MAMMALIA. to the same animal as the skull in that of the Rev. D. Williams. ‘They were, however, among the bones from Sandford Hill, and were therefore described as from that cave in our first chapter. A closer examination has shown us that bones from different » caverns in the Mendip can be recognised with much probability by their condition and the colour of the matrix. In both these respects the lower jaws strongly resemble the remains from Hutton. They exactly fit the spelean skull from Hutton. We therefore suppose that they must have been accidentally misplaced either by Mr. Beard himself, or in the removal of his collection to Taunton, and that they really belong to the same animal that furnished the skull in the Hutton Cave to the Rev. D. Williams. The skull in question is that of an adult rather past the prime of life. The teeth are decidedly worn, and the alveolus of the right upper tubercular molar is partially removed by absorption, which proves the loss of the tooth during the lifetime of the animal. Its state of preservation is shown by the following list of its component bones. A minute portion of the right nasal is present im the angle of the frontal suture, also small portions of the palatines adjoining the maxillary suture, and also that with the pre-sphenoid. The maxillaries with their dentition are nearly perfect, the palatine process bemg excepted. From the (otherwise perfect) inter-maxillaries the incisors have gone. The left third incisor was diseased, and probably lost during life. The right malar and squamosal are absolutely perfect, and the left nearlyso. The posterior or cribriform plate of the ethmoid, and a part of its central plate, are present, so that the beautiful tracery with which it fills the anterior end of the cranial cavity may be seen by looking through the foramen magnum. The greater part, however, of the bone has disappeared. ‘he vomer is entirely wanting. ‘The pre-sphenoid and orbito-sphenoid are nearly perfect. The superior parts of both frontals are nearly perfect, but the orbital portions are much broken. The right tympanic bulla is perfect. The articular portion of the squamosal (“corsal” of Straus- Durkheim) is preserved on both sides, as also are the lower jaws, with the exception of the coronoid processes and a small portion of one of the condyles. ‘The basi-sphenoids, ali-sphenoids, parietals, mastoids, basi-occipitals, exoccipitals, supra-occipitals, paro- occipitals, and Wormian, are nearly absolutely perfect. A small part of the lachrymals is attached to the frontals and maxillaries. The petrosals appear to be perfect, though of course they are but slightly visible. The second skull (Pl. X) is from Mr. Beard’s collection, and was found in Sandford Hill Cave. Along with it were found the lower jaws described in Chapter I and PI. I of our work. They were accidentally labelled as coming from Bleadon, and the mistake transferred to our pages was not discovered until the chapter had gone to press. Both ‘skull and lower jaws belong to a young adult. Several bones of an animal exactly cor- responding im size and age were found along with them; and as those adjoining each other in the skeleton exactly fit, we have reason to believe that we have a considerable portion of the same individual. Unfortunately it was the practice of Mr. Beard “to restore,” not very skilfully, the missing parts of crania and other fossils with hard plaster, FELIS SPELAVA. 33 and in this case the result has been very great difficulty and risk in taking his work to pieces and articulating the skull for scientific purposes. After such rough usage the exactness of fit of the component parts and the symmetry have been to some extent lost. We have figured the skull as it stands now free from plaster, without attempting a restoration which very possibly might have been erroneous, and which certainly could have served no scientific end. The specimen retains a small piece of each nasal zm sit, and a large part of the right palatine. The right maxillary is all but perfect, with a small portion of the left. The right intermaxillary also is in part present. The only teeth remaining are the large premolars (four), and a portion of the right canine. Both premolars are nearly perfect, together with the left squamosal and a large part of the right, so that we can form an adequate idea of the size and form of the zygoma ticarch. The frontal bones are present, but their supra-orbital processes are much abraded. ‘The left tympanic bulla is much broken, and the right is almost entirely gone. The basi-sphenoid is all but gone, and only the lower and pesterior portions of the ali-sphenoid are left attached to the lower part of the ali-sphenoids and the squamosals. Both mastoids are imperfect. The basi-occrpital is present, but the exoccipitals are abraded, and the supra- occipital is gone; and of the par-occipitals, only the left fossa remains. In addition, we figure in Pl. XI the maxillaries and intermaxillaries of another specimen from Sandford Hill Cave, which is also from Mr. Beard’s collection. It is of very large size, and exhibits the perfect adult dentition, with the exception of the small tubercular upper true molars, the small premolars (two), and one first incisor. We also give a figure of the articular portion of the squamosal of a gigantic animal from Bleadon Cave (Pl. IX, figs. 2 and 3). Besides the skulls we figure, we have examined a large number of fragments, which are for the most part in the Taunton Museum, as well as the nearly perfect specimen from the Sundwig Cave, in Westphalia, now in the British Museum; from it are absent the greater part of the zygomatic arches, a portion of the left palatine, the pterygoid pro- cesses, the upper part of the supra-occipital, and two thirds of the nasals, the left orbito- sphenoid, together with the adjoining part of the frontal, the ethmoid, and the vomer. We have also examined the specimen figured by Professor Owen and Mr. Scharf from Kent’s Hole, and a similar fragment from Muggendorf, now in the British Museum, and another like it from Ravenscliff in Gower, in the possession of Colonel Wood. ‘These constitute the materials which we have at hand for writing this chapter on the skull of Felis spelea. We shall compare the spelean skull bone by bone with that of the living species most closely allied to it, that is, lion and tiger, beginning with the basi- occipital. § 2. Basi-occipital. (Pls. VIII, IX. No. 1).—The basi-occipital forms the posterior portion of the base of the skull, and is regarded as the centrum of the occipital vertebra by all who hold the “vertebral theory.” From the slight bulging of the sides it 34 PLEISTOCENE MAMMALIA. is somewhat hexagonal in form. It is rather longer than wide, and is slightly longer than the basi-sphenoid, to which if is firmly attached by a straight transverse suture. It forms a strong plate of bone of nearly uniform thickness, articulated behind to the exoccipitals in the whelp by a suture, of which the lateral portions are transverse. In the median line, however, it sends back a square process, the free end of which is the lower and anterior border of the foramen magnum. ‘The lateral portions of this suture are interrupted by the “foramen condyloide,” which passes from the posterior edge of the “foramen lacerum posterius”’ (a) to the interior of the cranium near the anterior border of the foramen magnum. ‘This transmits the large motor hypoglossal nerve. The sides of the bone are in contact with rather than articulated to the tympanic, and above that to the petrosal, the junction between them being interrupted posteriorly by the large 9 irregular ‘foramen lacerum posterius”’ (a) or “foramen jugulare”’ for the passage of the eighth pair of nerves and a large vein connected with the internal jugular. The medullary surface is somewhat concave, forming a lodgment for the “ medulla oblongata.” The flatness of the lower surface is broken by two large rough depressions on each side close to the tympanics for the insertion of the “recti antici majores ” of the head, which take their rise in several roots on the pleurapophyses of the cervical vertebree. Immediately behind them are the smoother but larger impressions of the “recti antic1 mimores,” which have their origin in the “atlas.” These impressions are represented in Pl. VIII, in front of the “foramen lacerum posterius.” In the median line we sometimes find in Lion the commencement of the tubercle for the attachment of the “constrictor pharyngis superior,” which is, however, mainly attached to the basi-sphenoid. This does not occur, as far as we know, in Felis spelea. With the exception, perhaps, of a slight tendency to greater width in the spelzean as compared with the leonine and tigrine basi-occipitals, there is no difference worthy of note. § 3. Eeroccipitals and Supra-occipitals. (Pls. VI, VII, VIII, IX, X. Nos. 2,3, 4).— It is more convenient for purposes of description to treat these as one bone rather than in accordance with their centres of ossification, because they are never found separate except in the very youngest individuals, and because, firmly anchylosed together, they form the main surface of connection between the head and the trunk. ‘They compose a strong plate of bone, triangular in outline, firmly articulated to the basi-occipital (Pl. VIII, No. 1) at right angles, and with it circumscribmg the foramen magnum (Pls. VIII, IX). On either side of the latter are two short thick pedicles of bone which point downwards and backwards, and support the condyles by which the head is articulated to the atlas. These point in their upper portion upwards, in the middle backwards, and in the lower downwards. ‘Their edges project over the sides of the pedicle, forming a fossa, which is called the “condylian fossa” (4, Pls. Vi and X). These are the portions termed by Professor Owen the exoccipitals (No. 2). ‘The two inferior angles of the bone are composed of the paroccipitals, or paramastoids, FELIS SPELAA. | 35 as they are sometimes called, which form on either side a wide and deep cavity on the outer surface, which may be called the paramastoid fossa (c, Pls. VIII and IX), receiving the projecting sides of the glenoid cavity of the atlas, and thus combining great firmness of articulation with freedom of lateral motion. It runs as far downwards as the origin of the massive bony pedicle which projects downwards, and ends in the paramastoid tubercle (Pls. VI, VIIL, IX, d), which is homologous with the jugular tubercle in man. The par- occipital articulates with the mastoid in front, and inferiorly with the tympanic; if we hold to the ‘vertebral theory” of the skull, as propounded by Professor Owen, it is homo- logous with the parapophysis of the basi-occipital vertebra. The portion of the occipital which composes the apex of the triangle, and together with the exoccipitals complete the arch over the spinal cord, is the supra-occipital (Pls. VI, VII, VIII, IX, No. 3), which would be the neural spine of the vertebra. Its sides are decidedly convex, while the continuation on the paroccipitals is concave, so that the whole side of the triangle is distinctly sigmoid ; the interior and inner surface forms the back wall of the cranial cavity, and is entirely in contact with the cerebellum, for the convolutions of which it is deeply grooved and waved. The upper edges of the superior and outer surface are covered with high radiating ridges, of great sharpness and strength circumscribing depressions of various depth, which may be called the splenial fossz (Pl. IX, #), from their being the points of insertion for the tendon of the great splenius muscle. That descending from the apex, remarkable for its size, serves for the attachment of the cervical ligament (ligamentum nuchee). The articulation of the exoccipitals with the basi-occipital has already been described in our account of that bone. Tach of the paroccipitals (paramastoids) covers by a broad overlap the posterior end of the tympanic bulla. Above this it is firmly articulated to the posterior border of the mastoid through the whole length of that bone. Above this, in some individuals among the larger Feles, the supra-occipital articulates with the parietal, but generally the descending process of the Wormian or interparietal passes down- wards so as to join the upper part of the mastoid and prevent the connection of the parietal with both the ex- and the supra-occipital. The suture with the Wormian is of great depth in the aged animals of the larger Feles, owing to the great height of the lambdoid or occipital crest. In advanced age the whole of the lambdoid suture is obliterated, and its position is marked by a very sharp and massive crest, the lambdoid or supra- occipital. . Ligaments and Muscles of Occiput.—A very detailed account of the muscles and liga- ments of Felis are given in Straus-Durckheim’s great work,’ to which we would refer those who wish to become acquainted with the details of this part of the subject. We shall content ourselves with giving a list of the principal ligaments and muscles, with their 1 «Anatomie, descriptive et comparative, du chat, par Hercule Straus-Durckheim,’ 2 vols. 4to, plates folio, Paris, 1845. This work is perhaps the most perfect monograph on the comparative anatomy of a single animal that exists in any language. 36 PLEISTOCENE MAMMALIA. points of origin and insertion, so far as they are connected with the part of the skull under consideration. ) The ligaments which connect the head with the neck are the followimg:—The cervical hgament, or “ligamentum nuchz,” which is comparatively small in the genus Felis, springs from the neural spme of the first dorsal vertebra, and passes among the muscles of the neck to its insertion on the summit of the occipital crest. The “atlo-cephalic capsular,” the representative of the capsular of the head in man, occupies the position expressed by its name. It connects the skull, not only with the atlas, but also with the odontoid process of the axis. The “anterior superficial atlo-cephalic” is the equivalent of the “ anterior cervical” in man. It springs from the anterior and upper border of the hypapophasis of the atlas, and is inserted into the posterior border of the basi-occipital. The “median superficial posterior atlo-cephalic,” the posterior superficial of the atlas in man, fills the space between the upper part of the foramen magnum and the corresponding part of the atlas. It also extends down the sides of the posterior portion of the paroccipital fossa, which it connects with the exterior of the glenoid cavity of the atlas. The “rectus posterior capitis” has its origin in this ligament. The “deeper” ligament of the same name as the last appears to be simply the fibrous envelope of the spinalcord. The “anterior lateral atlo-cephalic,” or ligament of the first vertebra in human anatomy, springs from the border of the glenoid cavity of the atlas, and is inserted into the basi-occipital on the mner border of the condyles. The “lateral atlo-cephalic,” a strong ligament not found in man, has its insertion on the internal border of the paroccipital fossa, whence it passes downwards and backwards, and is attached to the inferior border of the glenoid cavity of the atlas. It hinders the excessive rotation of the head on the atlas. ‘Two other smaller ligaments, having the same function, are called the “ superficial,’ and the “ deep transverse posterior,” “‘atlo-cephalic.” They have the same insertion as the last, but pass upwards and back- wards to their points of attachment on the upper border of the neurapophysis of the atlas. They are not found in man. ‘The “lateral odonto-cephalic,”’ having the same name in man, closes the list; it passes from the end of the odontoid process to the inferior angle of each occipital condyle. In giving a list of the muscles attached to this part of the skull, we will begin with those that serve for the movement of the whole head. The great “ complexus’! of man is represented in Felis by two muscles, that called “‘biventer cervicis” by Eustachius and Albinus, the “intersectus” of Straus-Durkheim, and that to which the latter author confines the name of “complexus.” The insertion of the “intersectus’” is on the inner portion of the occipital arch, over the foramen magnum. Thence it passes backwards, dividing into four principal tendinous roots, which are attached to the transverse processes of the seventh cervical and first three dorsal vertebra. It adheres to the cervical ligament through its whole length. Above this lies the “ com- 1 Op. cit., vol. ii, p. 241 et seq. FELIS SPELAA. 37 plexus” proper, which has the same insertion as the preceding, and passes backwards to the diapophyses of the cervical vertebrae, and thence to the three anterior dorsals. The great, lesser, and middle recti posteriores of the head underlie these two muscles; their insertions extending nearly to the upper edge of the foramen magnum, and their origins being in the atlas and axis. ‘These five muscles serve to lift up, and to a certain extent to rotate, the head. Their points of insertion are therefore necessarily of great strength in the larger and more powerful Feles. In Fels spelea they are not more massive propor- tionally to the size of the animal than in the living tigers and lions. The great splenius muscle springs from the “cervical ligament,” and an aponeurosis which connects it with the last cervical and first five dorsals, and ends in a short strong tendon which is inserted into the occipital bone immediately behind the lambdoid suture. Its enormous size and strength in Felis spelea is seen by the large fosse for its insertion in Pl. IX, fig. 1. It takes part in the same movement as the preceding five muscles. The “trachelo-mastoid ’ of Douglas rises from five tendinous roots attached severally to the last. four cervicals and the first dorsal, and passes in the form of a long thin band along the side of the neck to its insertion on the paroccipital. Near it is inserted the “rectus lateralis,” which has its origin on the ala of the atlas. The “superior obliquus” of the head has nearly the same direction as the last, but passes within it to be inserted on the surface of the paroccipitals in the upper part of the condyloid fossa, at the point where they join the exoccipitals. These three muscles, together with the splenius, bend the head from side to side. The great size of the lateral ale of the atlas stands in direct relation to the development of these muscles, in animals that shake and worry their prey, such as lion, tiger, and Felis spelea. M. de Blainville’ states that the occipital crest is prolonged further backwards in the tiger than in the lion, a point which we have remarked to be by no means of characteristic value, and that the condyles are more detached in the former than in the latter animal. We have frequently found the converse of this latter statement to be true. He also writes that in these two points the plaster cast of Count Miinster’s specimen of Felis spelea’ agrees with tiger and differs from lion. We are unable to lay hold of any character in the occipital bone that would differentiate lion from tiger or from Felis speed. § 4. Bastsphenoid (Pl. VIII, No. 5).—The basisphenoid is articulated to the presphenoid by a transverse suture, which is clearly visible even in animals of considerable age. It is much wider though about the same length as this latter bone. The form of the inferior or guttural surface exposed in the perfect skull is roughly triangular, the apex of the triangle being cut off by the presphenoidal suture, and the base being formed by 1 “Ost. Felis,’ p. 28. 2 “Ost. Felis,” p. 108, 38 PLEISTOCENE MAMMALIA. that with the basioccipital, which in the old animal is obliterated as completely as the frontal suture of human anatomy. The sides are covered to a great extent by the overlap of the anterior portion of the tympanic bulla, and in front by the guttural process of the alisphenoid. ‘The surface in the larger Feles is nearly flat or slightly concave. A small foramen on each side of the basisphenoid’ in the suture between it and the alisphenoid is the posterior opening of the canal which conducts the Vidian nerve and artery to the foramen sphenoidale : from this proceeds a well-marked groove backwards along the above- named suture, to the foramen lacerum medium ; it then passes along the suture between the tympanic and alisphenoid just outside the foramen caroticum, to the small foramen by which the nerve makes its exit from the petrosal proper, within which bone it branches from the facial nerve. When detached the bone in the lion is of the same truncated triangular shape, but it is somewhat wider than was before apparent in consequence of the overlap before described. Its vertical thickness is slightly greater anteriorly than posteriorly, owing to the com- mencement of the upward slope from the bottom of the “sella tureica” to the olivary process within the cranial cavity. The dorsum ephippii rises to a considerable height in all the larger Feles, and is turned much forwards, the inclination from the summit to the posterior edge of the bone being at an angle of 30° to 35° with the horizontal. The sides of the “dorsum” expand into lateral ale, somewhat like the wings of a moth, and homologous with the ‘posterior clioid processes” in man. ‘The arch formed by them, and the anterior clinoid processes is sometimes completed in Felis. We have not, however, met with an instance of this in the speleean skulls: on each side of the “dorsum” is a furrow for the “internal carotid artery.” Sometimes there is a small median foramen on the guttural surface, and two minute foramina on the back of the “dorsum ephippui.” Muscles.—The peristaphyline (Straus-Durckheim) muscle, the representative of the internal muscle of the same name in man, has its origin close to the alisphenoidal suture, at a point where it crosses a line joming the “foramina ovalia.” Its office is to lower the “velum palati.”” The superior constrictor of the pharynx is attached to the posterior por- tion of the bone close to the basi-occipital suture. The only point worthy of note in comparing this bone in Melis spelea with those of lion and tiger is that it has a tendency to be somewhat wider in the two former animals than in the latter. This width, however, is very variable, and cannot be considered characteristic. § 5. Alisphenoid; Pterygoid (Plate VIII, No. 6).—The alisphenoid, usually described as the ala of the sphenoid, and treated by Straus-Durckheim! as a mere process of that bone, lies immediately in front of the squamosal in the surface of the cranium. As, ' Op. cit., vol. 1, p, 395. FELIS SPELAUA. 39 however, it 1s easily separated from the basi- and pre-sphenoid, while the suture between it and the pterygoid is obliterated at a very early age, we treat the alisphenoid and the pterygoid as one bone for descriptive purposes. Nearly the whole of the outer surface of the bone is visible in the perfect skull, as a vertical plate running upwards to form part of the walls of the cranium between the temporal and optic fossze. It also extends horizontally as far back as the petrosal, passing under the anterior part of the tympanic. Inferiorly, the pterygoids extend downwards and backwards on either side of the great guttural groove, ending in the thin, strong, hamular processes in lion and tiger, which in our spelzan skulls are unfortunately broken away. For purposes of description this compound bone may be divided into the horizontal or guttural portion, the supero-vertical or temporo-optic, and the infero-vertical or pterygoid portions. ‘The first of these is a narrow plate, transversely concave, covering the postero-lateral edges of the guttural surface of the presphenoid and the antero-lateral edges of the basisphenoid. In the basisphenoidal suture is the orifice of the Vidian canal, by which the nerve and artery of that name enter the alisphenoid, and pass forwards into the orbit at the external border of the ‘“‘ foramen sphenoidale.” We have already described the groove connected with this canal on the surface of the basisphenoid in our account of that bone (§ 4). The infero-vertical or pterygoidal processes curve downwards from the horizontal portion, and articulate anteriorly with the palatine by a vertical suture. The hamular processes, in which they terminate, are the equivalents of the internal pterygoid plates of human anatomy, the externals being represented by a slight longitudinal ridge immediately in front of the foramen sphenoidale. M. de Blainville’ states that the hamular processes of the tiger are less delicate than those of the lion. The variations, however, in this respect, in both these species, do not enable us to confirm this observation. As might be expected, these parts have not occurred in a fossil state. The horizontal plate expands posteriorly behind the Vidian canal, and articulates with the squamosal just within the boundary of “the glenoid cavity.” At this point it joins the supero-vertical or temporo-orbital process, which is a long, thin, triangular plate, highly convex externally, articulating behind with, and passing under, the squamosal by a highly concave suture, above with the antero-inferior angle of the parietal, in front with the postero-inferior angle of the frontal, and the orbito-sphenoid. At the bottom of this suture is a notch, which, together with a corresponding surface of the latter bone forms the large “foramen sphenoidale,”’ to a certain extent the representative of the sphenoidal fissure or ‘foramen lacerum anterius’* in man, and giving passage to the third and fourth nerves, to the first branch of the fifth pair, or the trigeminal, and to the sixth. Immediately behind this, and rather lower down, is the foramen rotundum, for the trans- 1 «Ost. Felis,’ p. 28. 2 Straus-Durckheim, op. cit., vol. 1, p. 395. 3 Holden, ‘ Human Osteology.,’ 3rd edit., p. 78. 40 PLEISTOCENE MAMMALIA. mission of the maxillary portion of the trigeminal, and more widely separated and exactly opposite the glenoid cavity is the foramen ovale (y), which transmits the infra-maxillary, or mandibular branch of the fifth pair. The latter is called the carotid foramen by M. de Blainville? and some others, in the mistaken belief that it transmits the carotid artery. The foramen caroticum is in all the Feles extremely small; and the external orifice being entirely covered by the Eustachian process of the bulla, and generally, but not always, surrounded by the substance of that portion of the tympanic, it may be said to be within the foramen lacerum medium, immediately on the inner edge of the groove for the Vidian nerve. ‘The internal orifice which is immediately in front of the apex of the petrosal proper leads directly to the groove for the artery described above as on the cerebral surface of the basisphenoid. ‘This is very distinct in the smaller cats, but is less so in the larger. The foramen itself is larger in the jaguar than in any other large Felis that we have examined, admitting a small wire. It is very small in the tiger, and still smaller in a panther, and even in very young lions; in some old animals of both these species it appears to be entirely closed. In Felis spelea it closely resembles the lion. The functions of the carotid artery, as it exists in most other mammalia, appear to be supplemented, or rather replaced, by those of the numerous vessels which accompany the nerves in their passage through the different foramina, and which in this part of the skull - unite an external “rete mirabile” to an internal, for the supply of blood tothe brain. We have observed in many skulls of Felis that the large foramina of the alisphenoid are accompanied by smaller, which appear to be appropriated to the transmission of vessels, though we have not ascertained this to be the case by the actual dissection of those spe- cimens. Muscles—The hamular processes of the pterygoidal portion of the alisphenoid bemg broken away, we can say nothing of the origin of the constrictor superior pharyngis, or of Folian muscle, in Felis syelea. The origin of the circumflexus® (Albinus) is under the foramen ovale (g), whence it passes round the hamular process to its msertion in the velum palati. § 6. Presphenoid and Orbito-sphenoid (PI. VIII, fig. 9).—All that hold the “ vertebral theory” of the skull agree in assuming the homologies of the centrum of a vertebra for some part of the presphenoid, though there are differences of opinion as to the morphological value of the different parts. The bone is firmly anchylosed to the orbito-sphenoid while still foetal, and the sutures have all but disappeared at birth; for this reason we describe them as one bone. ’ Straus-Durckheim, op. cit., vol. i, p. 295. Holden, ‘ Human Osteology,’ p. 395. *Osteol. Felis,’ p. 14. % Straus-Durckheim, op. cit., vol. ii, p. 229. FELIS SPELAA. AJ In the smaller Taunton skull this bone is nearly perfect, and as the palate is broken away the anterior portion is visible. We are consequently able to describe all the free surfaces of the bone. The visible portion of the inferior surface is a narrow strip, widening slightly pos- teriorly and considerably anteriorly, which forms the central portion of the roof of the posterior nares. The posterior portion is covered by a deep overlap of the pterygoid, and the anterior by a similar overlap of the palatine. The posterior end is firmly anchylosed to the basisphenoid by a deep suture convex posteriorly. These sutures are indicated in Pl. VIII, No. 9, by faint lines, as they are nearly obliterated in the skull by age. This surface is nearly flat in Helis spelea; this is also the case in the majority of the lions’ skulls that we have met with; whereas in the majority, if not all of the tigers” skulls, as well as in most of the other Feles, there is a strong longitudinal central ridge, which receives on each side of it the prolonged posterior processes of the vomer. Anteriorly the bone is seen to consist of a thickened central mass, which rises into a thin vertical plate, separating the ethmoidal sinuses, and articulating firmly with the central plate of the ethmoid, a portion of this articulation is seen in the skull we are describing. The central mass expands laterally into two thin plates, covered by the overlap of the palatine before mentioned, which form the floor of the ethmoidal sinuses, for the reception of the infra-lateral processes of the ethmoid, sometimes called the “cornets de Bertin.” The outer walls of these sinuses are formed by thin plates rising from the outer edges of the floor, at first inclining somewhat inwards, and covered by the overlap of the vertical plates of the palatine, but posteriorly these are uncovered, and arch over outwardly and form the lower and posterior surface of the orbit. This free portion is pierced on each side near the centre by the large optic foramina, which pass backwards and downwards to the optic groove on the cerebral surface of the bone; the lower part of this surface is traversed by a strong ridge, below which the bone is roughened for the insertion of the powerful Fallopian muscle for raising the lower jaw. The roof of the ethmoidal sinuses is formed by a thick plate, narrow horizontally and anteriorly, but widening much posteriorly, at the same time curving downwards, so that it unites with the much thickened posterior end of the central mass at its junction with the presphenoid, forming at this pomt the homologue of the olivary process in man. Above, in front of this is the very deep transverse optic groove, the ends of which lead outwards, as before stated, to the optic foramina. The anterior portion of this, the cerebral surface of the bone, is the floor of the rhinencephalic fossa, the walls and roof of which are formed by the frontals. Anteriorly this surface rises centrally into a strong vertical ethmoidal spine, to which is anchylosed the cribriform plate, for the lower foramina of which the anterior edge of the orbito-sphenoid is deeply furrowed. The anterior edge of the vertical walls of the ethmoidal sinuses is articulated to the vertical walls of the palatine by firm sutures inclining forwards, to the frontals by nearly horizontal A2 PLEISTOCENE MAMMALIA. sutures, which pass above the optic foramina, and to the alisphenoid by very firm sutures of considerable depth, which pass round the posterior portion of the bone, leaving on each side a considerable free space to form the inner surface of the foramen sphenoidale, the central and largest of the five foramina near each other im this part of the skull, which, as before stated, is homologous with the sphenoidal fissure, or foramen lacerum anterius in man. The remaining muscles attached to this bone are the rectus externus of the eye; “orand abducteur”’ of Straus-Durckheim, and the ‘‘petit abducteur” of the same author, equivalent to a portion of the choanoid, which have their origin outside the optic foramen ; and the rectus inferior, “ grand abaisseur’”’ of Straus-Durckheim, and the “petit abaisseur” of the same author, equivalent to another portion of the choanoid, have their origin imme- diately under the same foramen. We have above indicated the only and very slight difference we have been able to distinguish on the guttural surface of the bone, between lion and Felis spelea on the one hand, and most, if not all other Feles on the other. § 7. Palatines (Plates VI, VII, X, XI, No. 20).—Of the palatines we have seen but a small portion in spelzan skulls from British localities but we are able to describe it fully from the skull, from the Sundwig cavern, preserved in the British Museum. It may be considered as composed of the horizontal naso-palatine portion, and the vertical plate that forms the lower surface of the optic fossa, its inner surface forming the floor and walls of the posterior nares. The naso-palatine portion presents a smooth horizontal surface joing its fellow by a thickened median symphysis, both forming a figure variably pentagonal in the bony roof of the mouth. Anteriorly it is firmly anchylosed to the maxillary by a serrated suture directed diagonally backward from the median lne; posteriorly it presents a free edge that sends back a process to articulate with the pterygoid. The free edge forms the infero-posterior border of the posterior nares. Externally it joins at a right angle the vertical naso-optic process, along a line passing from the sectorial fossa (4) of the maxillary diagonally backwards to the pterygoid and the palato-maxillary suture; and nearly equidistant from the interpalatal suture and the sectorial fossa (A), is the small posterior palatal foramen (2) for the transmission of .the palatal nerve. It is directed forwards, and opens upon a canal on the posterior surface of the maxillary. In some of the smaller Felide it is double, but in lion, tiger, panther, and all the larger species it is single, as in Melis spelea. In all the leonine and tigrine skulls which we have examined, the position of this foramen is constant. It is much nearer to the postero- exterior border of the palate in lion than in tiger, when skulls of equal size are compared. The only apparent exception to this rule is presented by the skull of a small lioness in the British Museum, in which it is roofed in by an abnormal growth of bone from the maxillary, so that instead of opening as it usually does on, or rather in rear of, the suture, it is carried forward and opens far on the maxillary. We also find that in all the lions” a ily FELIS SPELAVA. A3 skulls we have examined, a thin probe passed through this foramen passes directly into the orbit without showing itself on the nasal surface of the bone, while in all, except in one or two extremely small skulls of the tiger, it passes freely into the nasal cavity. In both these points Felis spelea agrees with lion. T'o these M. de Blainville would add a third point of difference between lion and tiger : that in the lion the posterior border of the horizontal plates terminates in sharp cusps, which form a somewhat deep notch at the interpalatal suture, while in the tiger it ends in a point without a notch (en pointe mediane sans échancrure).' We have carefully tested the value of this point of difference in a large series of recent skulls. In leonine skulls the notch is variable in size, and in some almost obsolete. In those of tiger on the other hand the median point often disappears, leaving the posterior border straight, and sometimes well defined cusps are present, and the notch more distinctly marked than in some leonine skulls. Although, therefore, M. de Blainville has rightly indicated the tendency of the two species in this respect, we cannot suppose that we have in this the means of absolute distinction. The only skull of Fehs spelea which gives us information on this point is that from Sundwig. It shows im its present state an affinity to the tiger. The bone, however, is abraded at the point where the cusp would be, had it ever existed, and on the other side the palatine is restored in plaster. We therefore do not consider the evidence afforded by it of any value as to the leonine or tigrine character of the animal. From the postero-external surface of the horizontal plate rises the vertical or optico- nasal, articulated by a slightly convex vertical suture to the ner side of the base of the malar process of the maxillary, and to the lower and posterior edge of the lachrymal, to the frontal above by a long horizontal suture, and to the lower edge of the orbito- sphenoid, and the anterior of the alisphenoid by a descending suture en échellon. The upper surface is slightly concave vertically and horizontally; the lower is convex ver- tically. It is pierced by two foramina, the larger of which is the spheno-palatine for the maxillary branch of the fifth pair of nerves; the smaller, situated more in front and at lower level, conducts the palatal nerve to the small posterior palatal foramen in the palato- maxillary suture. These two foramina are erroneously called by Straus-Durckheim “trous gustatifs.”* To the lower part of the orbital surface, and throughout the whole length of the horizontal process, is attached the Fallopian’ muscle, or external pterygoid of human anatomy, which is among those which elevate the lower jaw, and is inserted at the infero-exterior border of the horizontal ramus below the coronoid process. In man this muscle is inserted into the neck of the condyle and the meniscoid fibro-cartilage, and is a pretractor or rotator, while in Felis, being inserted much lower down, it serves merely for an elevator. 1 Ost. Felis,’ p. 28. 2 Op. cit., vel. i, p. 426. % *Straus-Durckheim,’ vol. ii, p. 217. AA PLEISTOCENE MAMMALIA. The only point of difference of specific value in this bone is in respect of the palatal foramen. In every case Felis spelea agrees with lion in a most decided manner, the space between the foramen and the postero-exterior border of the palatine being even smaller in it than in the lion. In all the specimens of the panther and jaguar that we have examined, it is proportionally greater than in the tiger. Fels spelea, then, is isolated from these two smaller species by this characteristic. 8. Mazillaries (Plates VI, VII, VIII, X, XI, No. 21)—The maxillaries of all the larger species of Felis resemble each other very closely, and yet it is in these very bones that we find minute differences which are specifically constant. Their surfaces may be described as the vertical or facial, the basal or palatine, the posterior or orbital, and the internal or ethmoidal. The first of these presents a roughly triangular outline, bounded behind by the malar, lachrymal, and frontal articulations ; in front by those of the frontal, nasal, and intermaxillary, and below by the alveolar border. At its upper angle it is slightly concave; at the infero-anterior convex, for the reception of the fang of the canine, and at the infero-posterior flat. The concavity immediately behind the canine is the canine fossa, the muscle of that name passing along it from the malar to the upper lip. The upper angle forms the “frontal process,” which is received into a deep notch in the frontal bone. It is truncated in the tiger, rounded in the jaguar, pointed or very rarely rounded in the lion and panther.' If a line be drawn joining the apices of the frontal processes, in the two former animals it falls below the extreme point of the frontal processes of the nasals, while in the two latter it falls above the nasals, and rests entirely on the frontals.? In two skulls of Felis spelea, that figured in Pl. VII, and that from Sundwig, the frontal process is pointed, and the line rests on the frontals; in the third (Pl. X) the processes of the maxillaries are unfortunately abraded. If, however, the outline were restored, it would be impossible to make it otherwise than pointed ; and if so, the line drawn from the frontal processses would also rest on the frontals without touching the nasal suture of these bones. Immediately opposite the superior portion of the malar suture is the great suborbital foramen, separated from the malar and the orbit by a stout bony arch, and giving passage to the suborbital nerve and artery. It is smaller, according to MM. Goldfuss, Cuvier, and de Blainville,* in tiger than in lion, and the arch is thicker; and those authors consider that in these respects Felis spelea is tigrine in character. The specific value of these points is by no means confirmed by the study of a large series of skulls of lion and tiger, in which we find great variations in the ' We have seen some skulls which are said to be those of the panther or leopard from Eastern India and the peninsula of Malacca, in which the formation of the frontals and maxillaries resembled that of a tiger. The Western panthers, as far as our experience goes, all resemble the lion in this respect. 2 This was first pointed out by Professor Owen, ‘Proc. Zool. Soc.,’ Jan. 1834, p. 1. 3 Goldfuss., ‘Nov. Act. Nat. Cur.,’ vol. x; Cuvier, ‘Oss. Foss.,’ vol. iv, p. 433, ed. 1825; de Blain- ville, ‘Ost. Felis,’ p. 108. FELIS SPELAA. AD proportions of their parts. We cannot, therefore, admit the tigrine affinity of Felis spelea to be shown in the slightest degree either of the suborbital arch or foramen. From the postero-inferior angle of the facial surface springs the stout malar process, firmly articu- lated to the malar bone by an oblique suture. It is vertically convex on the outside; the inner side, vertically convex, horizontally concave, joms the posterior or orbital surface, which is inclined downwards from the suborbital foramen to the alveolar border behind the molar series, and articulates with the lachrymal and the vertical plate of the palatine. The orbital surface presents many small foramina for the nerves and arteries which supply the teeth, and affords attachment to the “inferior oblique muscle’” for the rotation of the eye, immediately below the lachrymal suture. The inferior or palatine surface is horizontal and very slightly concave in both directions. Posteriorly it is articulated to the horizontal plate of the palatine, on the mner side to its fellow, by a straight symphysis, which rises on the nasal surface into a sharp crest for the reception of the vomer. From the posterior palatine foramen (¢),” which we have already described in the palato- maxillary suture, a broad shallow groove runs forwards the whole length of the bone, for the reception of the nerves and blood-vessels of the palate. In part it is joined to the premaxillary by a nearly straight suture, running obliquely forwards and outwards, passing into the alveolus of the canine, interrupted by a free oval space, which constitutes the posterior border of the naso-palatine canals. On its external edge is the alveolar border, for the reception of the canine and molar teeth, the alveoli of which will be described along with the dentition. At the postero-external angle is a round and deep cavity, which from its function of receiving the posterior blade of the lower sectorial molar may be called the ‘sectorial fossa (2). The internal or nasal surface of the bone follows the contour of the palatine and facial surface. The large fangs, however, of the teeth necessitate large alveoli, which leave very little space for the antrum of human anatomy. ‘To a ridge on the vertical portion of the surface the ethmoidal bone is attached. Muscles—Yo the facial surface of the bone the following muscles* are attached :— To the upper part of the frontal process the rhinzeus, a double muscle for the elevation of the nostrils and upper lip; to the anterior edge of the orbit, close to the lachrymal suture, one of the roots of the palpebral, a muscle for the closing of the eyelids. Between these two points arises the elevator of the upper lip. The buccinator is not attached to the alveolar border, as in man, but is reduced in size, and confounded with the labial. The smaller branch of the lesser zygomatic springs from the alveolar border in front of the sectorial tooth, and its function is to aid in raising the lip. In no respect do the attach- ments of these muscles m Hehs spelea indicate any difference between that animal and the lion. 1 Straus-Durckheim, op. cit., vol. ii, p. 207. 2 These are erroneously termed “‘ trous gustatifs ’? by Straus-Durckheim. 3 Straus-Durckheim, op. cit., vol. ii, pp. 203, 207, 210, 211. ~ A6 PLEISTOCENE MAMMALTA. § 9. Intermawillary (Pls. VI, VII, VII, X, XI, No. 22).—The inter- or pre- mazxillaries form the anterior end of the face, and consist, like the maxillaries, of an ascend- ing and an horizontal process. ‘The first of these, which may be called the nasal process, is wedged in between the maxillary bone below and the nasal above, forming part of the lower portion of the alveolus of the canine, and is to a great extent overlapped by those two bones. The horizontal, incisive, or palatal process is articulated behind to the palatine process of the maxillary, and in the median line to its fellow by a straight symphysis, strengthened by a ridge on the superior or nasal surface, which with its fellow ridge forms a trough, which is articulated to and forms a continuation of the vomer, ‘The palatine suture is interrupted by the two large oval naso-palatine foramina (4), which open into two grooves that pass forwards as far as the inner edge of the incisive border. The incisive border in front forms an arc of a large circle, and is separated from the alveolus of the canine by a shallow excavation for the reception of the canine of the lower jaw. The intermaxillary bones form the lower and lateral boundaries of the nostrils, and present us with a character of specific value’ by which we can separate lion from tiger. In viewing the lower half of the nasal aperture in front, its inner bounding line takes the form of an even curve, expanding regularly im the former animal, while in the latter it is distinctly a surface of double curvature. This character is more strongly impressed on the larger than the smaller skulls. In Felis spelea it is strongly marked, and its evidence as to the leonine character of that animal is beyond doubt. The Sundwig skull, however, is somewhat exceptional, showing a tendency towards the double curvature. of tiger, but the tendency is not greater than that presented by several small skulls of lion. The nasal, or ascending, is inclined backwards at an angle of from 50°’to 60° with the horizontal process in Leo, Tigris, and Felis spelea, the angle being greatest in the largest skulls. , Muscles.—The myrtiform’ muscle for the dilation of the nostril, and the moustache muscle for the protrusion of the lip, take their origin from this bone; the one from the sides of the nasal aperture, and the other from the median suture. § 10. Petrosal; Mastoid; Tympanic (Pls. VI, VIII, IX, Nos. 16, 8, 28).— In the description of these bones we adopt Professor Owen’s numbers and nomen- clature, without committing ourselves to his views of their homologies, rather than enter into a discussion which has no immediate bearing on our present work. We shall therefore describe the petrous bone, together with the posterior descending process, as the petro-mastoid, and the remainder of the acoustic organ as the tym- panic. 1 This difference is pointed out by M. de Blainville, ‘ Ost. Felis,’ p. 28. ? Straus-Durckheim, op. cit., vol. ii, pp. 208, 209. FELIS SPELAA. A7 The petrosal proper is so irregular in shape as almost to defy description. It may, however, be conceived as resembling a dried distorted pear, lymg diagonally across the axis of the skull, so that the pointed end or stem points forwards, inwards, and somewhat downwards. It has three well-marked sides or divisions, the mtero-posterior or cerebellar, the superior or tentorial, and the extero-anterior or tympanic. ‘There is also a smooth, rounded, triangular surface between the posterior edges of the tympanic and cerebellar surfaces, which is partly in contact with the paroccipital (paramastoid), and partly forms the inner surface of the “foramen lacerum posterius.” The names of these surfaces adequately show the position of the bone in the skull, for it is wedged in between the basi-occipital, the tentorium, and the tympanic, the inner surface alone being exposed on the side wall of the cerebellar cavity. The inferior edge between the cerebellar and tympanic surfaces is in contact with the exterior edge of the basi-occipital, the outer portion of the tentorial with the lower or inner edge of the squamosal, and the anterior. apex of the stem reaches as far forwards as the extero-posterior angle of the basi- sphenoid. The cerebellar surface of the petrosal proper is roughly elliptical in form, having the anterior end pointed. A low ridge, running diagonally upwards and forwards, divides it into two long, shallow depressions. Near the middle of the lower depression is the “meatus auditorius internus,” a foramen somewhat C-shaped externally, but divided internally into two canals, the one for carrying the facial-motor nerve, the other for the body of the acoustic nerve before its distribution in the cochlea and semi-circular canals. In the middle of the upper depression is the small foramen of the “aquzeductus vestibull,” as in man. ‘That of the aqueeductus cochlez is under the meatus auditorius, opening into the petrosal sinus, rather further back than in man. The antero-inferior edge of this surface is, as we have said, in contact with the basi-occipital. Between the two is the ’ which ends posteriorly in the “foramen lacerum posterius” (a), a large opening left between the petrosal, the tympanic bulla, and the basi- and par-occipitals. “infra-petrosal sinus,’ This gives exit to the jugular vein and the eighth pair of nerves. We are unwilling to pull to pieces the skulls of Fels spelea, and therefore cannot describe the other three surfaces of the petrosal that are hidden in the perfect crania. A glance at the disarticulated skull of lion or tiger will convey a more ade- quate notion of it in Felis spelea than we could convey by the most faithful word- painting. Mastoid—Firmly soldered to the posterior edge of the petrosal is a massive wedge- shaped-bone (No. 8), the thin end of the wedge passing upwards between the squamous bone and the paroccipitals, and the thick rounded head projecting freely downwards (/). Though cartilaginous in the young Feles, and incompletely ossified even at the age of five months after birth in the lion, it becomes very compact and hard in the adult animal, and perfectly coalesces with the squamosal and petrosal, the suture with the former being a continuation of the lambdoid suture, and rising into a sharp ridge continuous with the occipital crest. In the majority of leonine and tigrine skulls the narrow ascending process 48 PLEISTOCENE MAMMALIA. does not reach so high as the parietal. The lower internal surface of the thick rounded head of the wedge is concave and smooth, and is closely applied to the posterior surface of the tympanic bulla, with which it has no further connection. The free end bears the articular surface for the stylohyal. This point, therefore, represents the styloid process in man. We have ascertained that in this bone there are no mastoidal cells, similar to those in the mastoid of man, connected with the tympanic, their absence in the Felidae being compensated for by the large tympanic bulla. In spite of this significant fact, bearing on the “vertebral theory ” of the skull of Carnivora, we use Professor Owen’s name of “ mastoid” for the bone in question, without discussing the value of the many and conflicting theories of the homologies of the component parts of the cranium. Tympanic.—The tympanic bone consists of two portions—the tympanic proper, and the bulla or supplementary portion. The tympanic proper forms a somewhat compressed oval chamber, the outer, upper, and posterior walls of which are in part firmly articulated to, and partly formed by, the inferior edge of the squamosal between the glenoid cavity and the mastoid, and which is directed from the latter downwards, inwards, and slightly forwards, parallel to the tympanic bulla. Externally it presents an oval opening under the supra-mastoid ridge of the squamosal—the “meatus auditorius externus”’ (m). This is the original portion of the bone which is in the young animal simply a thin plate, resembling a horseshoe in form, attached by the two ends to the squamosal, which thus completes the ring. Across it is stretched the membrana tympani, like the parchment over the head of a drum, the centre receiving the handle of the malleus. The outer sur- face of the chamber curves slightly forwards, downwards, and inwards (x), in front of the bulla, and it is much roughened for the attachment of the posterior branches of the stylo-maxillary ligament. In the anterior part of the articulation with the squamosal is the glenoid fissure. The anterior end of the bone turns downwards, and forms one or two small, thin, hooked processes (z), which are irregular in shape and number, and overhang the foramen lacerum medium, a large irregular cavity formed by the junction of the bulla, the posterior processes of the basi- and ali-sphenoids, and the petrosal. It transmits the canals for the Eustachian tube, the groove for the Vidian nerve and artery, and has within it the external orifice of the foramen caroticum. It is called, erroneously, in our opinion, “le trou déchiré antérieur” by Straus-Durckheim. Inside the “meatus auditorius” a long septum or curtain partially divides the chamber, and forms a deep groove, open downwards, which passes under and in front of the “meatus.” The inner wall of the chamber is formed by a thin plate of bone, that divides it from the bulla. It is said by Straus-Durckheim to be double in the cat. We have, however, examined it in the adult cat with a powerful microscope, and it appears to be homogeneous in structure throughout. The lower surface of the petrosal roofs in and thus completes the chamber, bemg soldered to the tympanic by exceedingly thin though firm connections at the posterior and upper borders of the cavity. The tympanic FELIS SPELASA. 49 bulla, always present in the skulls of Carnivora, and very largely developed in the genus Felis, is a large chamber of oval shape, wedged in between the basi- and par-occipitals, the mastoids, and the petrosal. Its external walls are very compact and hard, though thin, and appear under the microscope to be somewhat fibrous in structure. It is closed on all sides, excepting at the top, where there is a long fissure, in which the promontorium of the petrosal lies m such a position that the “fenestra vestibularis” opens directly on the interior of the bullar cavity. The latter passes into the chamber of the tympanic proper. ‘The bulla thus performs the same functions as the mastoid cells in man, and is consequently called the mastoid by Straus-Durckheim and some other anatomists. In front the bulla sends forward a long, solid, pointed process, that passes between, and articulates with the posterior processes of the basi- and the ali-sphenoid. On the inside it is in contact with the external and lower border of the basi-occipital, and with the whole length of the petrosal. Its relations to the mastoid and paroccipital, to the foramen lacerum medium, and the ‘‘stylo-mastoid” foramen, have been already described. In comparing these bones with those of the large Feles we labour under the disadvantage of being able to see but a small portion of them im the perfect skulls. So far, however, as we have been able to institute a comparison, the difference between the spelzean and leonine bones is so sinall that it is scarcely worthy of note. There is absolutely no difference between the cere- beliar surfaces of the petrosal. As compared with those of tigers, the inter-tympanic width is greater in the Taunton skulls than in several skulls of tigers of similar size. The foramen lacerum posterius is shorter and rounder in lions and Fels spelea than in the majority of tigers’ skulls. These points, however, are of no great value, and are certainly not characteristic. ‘They show merely the leonine character of these bones of Felis spelea, which in the course of this Monograph we shall be able to trace throughout the rest of the skeleton. Malleus (Pl. X, figs. 2, 3)—_We found that in the tympanic of the larger skull at Taunton the malleus still existed in its original position, and nearly perfect. We extracted it, and are consequently able to give figures of it and describe it. The long process (the “‘manubrium mallei’’), which in the living animal rested on the drum of the tympanic, sloped downwards and forwards, while the condyle-like head was articulated very slightly to the upper part of the tympanic cavity, formed by the external edge of the petrosal and the inferior and inner edge of the squamosal. The neck of the bone is bent inwards and upwards, and posteriorly there is a small facet for the incal articulation ; on the opposite side of the neck from that to which the manubrium is attached a small sharp process rises, which affords insertion to the Eustachian muscle’ (“internus mallei”). Tn the living lion a thin plate of bone fills up the acute angle formed by the neck; this is broken in the fossil. ‘Two other small processes rise from the opposite side of the neck to-the Eustachian process, which appear to be homologous with the processi longus 1 Straus-Durckheim, op. cit., vol. i, p. 416. 50 PLEISTOCENE MAMMALIA. or gracilis and brevis of human anatomy. A comparison of the bone with those of lion, tiger, panther, and jaguar showed no essential point of difference. § 11. Sguamosal (Pls. VI, VII, VII, IX, X, No. 27)—The squamosal consists of a slightly convex scale-like process applied to the exterior of the sides of the cranium, and a stout articular process, which rises at right angles, and forms the pedicle supporting the lower jaw. ‘The former overlaps the ends of the following sutures, commencing ante- riorly, and passing upwards round the edge:—the alisphenoid, parietal, mastoid, petrosal, and tympanic. It is so firmly soldered to the mastoid posteriorly that all trace of the suture is obliterated in the adult. In the young skull, however, of Felis spelea (Pl. X, 0) it is articulated so loosely that the mastoids have been broken away along the line of weakness thus presented. In the fully grown animal, however, the squamosal, mastoid, petrosal, and tympanic, form one bony mass, which is the exact homologue of the temporal bone of human anatomy. With the parietal it is articulated superiorly by a long horizontal, and with the alisphenoid by a vertical, suture, that ends in the glenoid fissure below. A small part only of the centre of the bone appears in the mner wall of the cranial cavity. From the lower and anterior angle of the squamous portion springs the strong pyramidal articular process, triangular in section, with its antero-inferior border deeply excavated, so as to form a transverse horizontal groove, which is the glenoid cavity (p) for the reception of the condyle of the lower jaw. On the postero-inferior surface it is slightly convex, on the superior somewhat concave. At from two to three inches from its origin it suddenly turns forwards at right angles to its long axis, becomes much com- pressed vertically, and is articulated by a long splice or diagonal suture to the maiar, by which it is overlapped externally. A strong sigmoid ridge (7), equivalent to the “supra- mastoid ridge’ in man, passes forwards from the lower edge of the squamous portion at its juncture with the mastoid, is carried round the upper and posterior edge of the articular process, and forms the upper edge of the zygomatic portion of the bone. Underlying this ridge, at the point where the squamous and articular portions meet, is the meatus auditorius externus (m), or external orifice of the ear, of which the upper edge is formed by the free surface of the bone. At the origin of the ridge, which from its position we may call the squamosal, and abutting against the inferior process of the mastoid, to which it is firmly soldered, is a strong process, in length equal to the latter, that supports the stylo-articular ligament. The depression at its end is marked 7 in PL. VIL. Muscles —The whole space between the zygomatic arch and the cranium is filled by the masses of the great tearing and rending muscle which gives such enormous power to the jaws of the Felidee and Hyanidee—the “crotaphite,” or temporal. Its first branch is in part attached to the inner side of the articular process above the malar articulation. The second springs from the squamous portion above the articular process, and the third is partially attached to the general surface of the squamous portion of the base. ‘The FELIS SPELAA. 5d] second branch of the masseter also springs in part from the inferior edge of the zygomatic arch, and from the edge of the glenoid cavity. MM. Goldfuss and Cuvier’ agree in stating that the height of the zygomatic arch in Felis spelea is greater than in either the lion or the tiger, while according to M. de Blainville’ it is wider i Fels spelea and tiger than in the lion. He adds also that the articular process rises from the temporal portion of the squamosal at more nearly a right angle in (Melis spelea than im lion. On testing the value of these points in a large series of leonine and tigrine skulls, we cannot admit them to be of specific value; and after com- paring the two skulls of Fehs spelea in the Taunton Museum with those of both those species, we cannot lay hold of any character by which we can separate one from the other so far as this bone is concerned. We figure in Pl. IX, fig. 2, a fragment of the articular portion of a squamosal from Bleadon Cavern, which is very much larger than any other we have met with, either recent or fossil. § 12. Malar or Jugal (Pls. VI, VII, VIII, X, No. 26).—The malar is a thin quad- rangular bone which forms the anterior portion of the zygomatic arch, and stands out from the skull so as to form an angle of from 40° to 45° with the median plane, the angle being smaller in the younger than the older animals. This difference is very evident if we compare the young skull figured m Pl. X with the old one in PI. VII. In front it is articulated to the maxillary, and a small process passes inwards, forming the inferior border of the orbit, and the upper half of the bridge over the infra-orbital foramen, to articulate with the lachrymal. Behind it joins the zygomatic portion of the squamosal by a very oblique suture, which passes diagonally upwards, inwards, and forwards, as far as the plane of the suborbital process (s). This latter is a strong, flattened, triangular mass of bone, produced into a sharp angle pointing upwards and backwards, connected in the living animal with the supra-orbital process of the frontal (¢) by the fronto-malar ligament (gonio-malar® of Straus-Durckheim) which completes, with the frontal, malar, lachrymal, and maxillary, the orbit. From this ligament, as well as to the posterior part of the sub-orbital process, rises a portion of the large temporal or crotaphite muscle, which fills nearly the whole of the temporal fossa. Along the outer surface a ridge of con- siderable prominence runs parallel to the lower free concave edge of the bone, which affords an origin to the first portion of the masseter muscle, the second springing from the inner surface, without leaving any impression on the bone to mark its position. The infero-exterior surface also affords attachments in front to the “lesser zygomatic muscle’ 1 “Oss. Foss.,’ vol. iv, p. 463, ed. 1825. » “Ost. Felis,’ p. 108. 3 Straus-Durckheim, op. cit., vol. ii, p. 14. 4 Tbid., p. 210. 52 PLEISTOCENE MAMMALIA. for the elevation of the lip; and within this to the wide thin “canine,” which takes part in the same office. The malar bone presents no point of specific difference in lion, tiger, and Felis spelea ; a comparison of upwards of one hundred skulls having convinced us that the cha- racter of greater depth in the latter than the two former animals, insisted upon by MM. Cuvier and Goldfuss, is not of specific value. ‘The. sub-orbital process, however, appears to be set rather further backwards in the majority of leonine skulls than in those of the tiger, so that the orbit is wider and rounder in the former than the latter animal. In this point Lelis spelea certainly agrees with the lion. In the recent animal the whole orbit is surrounded by a strong deep ligament, resting on its edge, which renders it deeper and more complete. § 13. Lachrymal (Pls. VI, VII, X, No. 73).—The lachrymal bone occupies the anterior border of the orbit, and is articulated in front to the maxillary, behind to the frontal, below to the palatine, maxillary, and malar bones. It is a flat plate, of irregular form, varying from triangular to quadrilateral. Its greater part is within the orbit in the larger Feles, but the small portion adjoining the frontal process of the maxillary is con- tinuous with the external surface of the skull. At its superior angle is the palpebral tuberosity («) for the insertion of the palpebral muscle, and below it is the lachrymal foramen (v), sometimes excavated in the lachrymal, at others lying in the lachrymo- maxillary suture that runs downwards into the nasal cavity. The internal surface of the bone is ridged for articulation with a branch of the ethmoid. The sutures vary in direction according to the shape of the bone. In old animals they are almost entirely obliterated ; a sinall flat bone, “os planum,” is sometimes, though rarely, intercalated in Felis at the infero-internal angle. ‘There is no appreciable difference in the shape of this bone in tiger, lion, and Melis spelea. § 14. Lthmoid—lt cannot be expected that any large part of so fragile a bone as the ethmoid can be preserved in the fossil state; but as an important part occurs in one of our specimens, we describe the bone in the lion and tiger. In the genus Felis the ethmoid fills the great facial cavity, and may be considered as consisting of a central plate flanked on either side by a highly convoluted mass of bone, and a transverse vertical plate. The former is vertical, and firmly articulated to the ethmoidal spine of the pre- sphenoid, as well as to the vertical plate which divides the ethmoidal sinuses; it rests on the vomer;. it is also articulated to the median nasal crest of the symphysis of the frontal bones, and passes more than half way towards the anterior end of the nostrils. It is considered by Professor Huxiey’ as the continuation of the basis cranii 1 «Proc. Royal Society,’ No. 33, “ Croonian Lecture,’ 1858, p. 433. FELIS SPELAA. 53 formed by the basi-occipital and the basi- and pre-sphenoids, in opposition to the more strict vertebral theory of Professor Owen,’ who considers the vomer as the centrum of the nasal vertebra. The posterior edge of the plate expands laterally into a transverse, nearly vertical plate, concave posteriorly, which closes the anterior end of the cranial cavity ; from its being full of small foramina this is called the cribriform plate. From the lower and outer edges of these two plates spring others remarkable for their thinness and delicacy, which form a highly complicated tissue-like mass of bone, filling the greater part of the facial cavity, and sending prolongations upwards and backwards into the frontal sinuses, backwards and downwards into the anterior sinuses of the pre-sphenoid, and for- wards into the nostrils. This mass is attached to the maxillaries, the frontals, vomer, and presphenoid, by delicate and thin articulations ; and the whole is so arranged that the air breathed through the nostrils must pass over the greater part of its surface. Through the foramina in the cribriform plate pass the branches of the olfactory nerves, which are spread over the large surface afforded by the convolutions of the bone. ‘The whole mass thus described is called a sense-capsule by Professor Owen.” This bone is naturally highly com- plicated in animals endowed with a fine sense of smell, such as the Felidz. In the common cat it looks like a mass of lightly squeezed silver-paper. In the larger Feles it is of course somewhat thicker and coarser, and more slightly packed, but it is still of great delicacy and beauty. ‘The pattern of the foramina in the cribriform plate may, perhaps, vary in the different species, but the position of the bone in the skull renders a comparison difficult and uncertain. In the smaller skull of Hels spel@a the plate appears like a beautiful plate of Saracenic tracery filling the end of the cranial cavity when viewed through the foramen magnum. § 15. Wormian (Pls. VI, VIL, TX).—The Wormian or inter-parietal is a small triangular bone occupying the apex of the occipital crest, and firmly wedged in between the parietal and supra-occipital, the sutures being of considerable depth. ‘The develop- ment of its downward processes is very variable, but sometimes they extend down to the squamosal, and even the mastoid, thus separating the supra-occipitals from the parietals. Its inferior surface forms the highest part of the cranial cavity. Muscles —The Wormian bone gives origin to several small muscles that regulate the movement of the ear ; for their names we refer to the second volume of Straus-Durckheim’s great work on the cat, which we have so often quoted. § 16. Parietals (Pls. VI, VII, IX, No. 7).—The parietals form the roof of the greater part of the cerebral cavity, and appear on the exterior of the skull as two nearly rectan- 1 «Homol. Vert. Skel.,’ pls. 1 and 3. 2 Thid., pls. 2 and 3. 54 PLEISTOCENE MAMMALIA. gular plates, convex vertically and horizontally, and thin except at the point of junction, where they form the strong sagittal crest for the attachment of the third branch of the temporal muscle. This crest increases in size and height, as in the recent Felide, in pro- portion to the age, being almost obsolete in the young animal that has not yet had time to use his jaws for tearing and rending, and gradually increasing in size in proportion to the age, and consequently the increased use of the temporal muscle, until it reaches its maximum in the old tiger. From the lower anterior angle of the bone a process is sent downwards and forwards to meet the alisphenoid, on which frequently in old lions, and in the smaller speleean skull in Taunton, there is frequently a strong ridge defining the points of attach- ment of the second and third branches of the temporal muscle. ‘The size and situation of this ridge recall the vast parietal processes in the genus Otaria, in which also the ramal process of the lower jaw, described by us as characteristic of lion and Felis spelea, is enormously developed. Anteriorly they overlap the frontals by a straight suture; pos- teriorly they are overlapped by the Wormian; but in old age the latter suture is obliterated ; it is very variable in position in the larger Feles, generally running clear of, and in front of, the occipital crest, but sometimes occurring in the crest itself. In- feriorly they are to a great extent covered by the squamosals, which almost entirely conceal their alisphenoidal processes, and entirely their long posterior processes, which may be called “ petrosal,” from their close contact with those bones. Internal surface.—A plate, thin in the smaller but very thick and strong in the larger Feles, projects downwards from the cranial surface of the parietals, running diagonally forwards from the posterior angle of the symphysis to the petrosal process, where it is united to a small corresponding plate on the alisphenoid. This plate, with its fellow of the opposite side, forms the “tentorium ” (Pl. X), or ossified! curtain of the “dura mater,” that divides the cerebrum from the cerebellum. In the centre of the united processes is an arch rather more than half the height, and about one third of the width of the cranial cavity, which admits of the connection of the cerebrum and cerebellum. ‘The symphysis between them projects forwards into a ridge, which is high and sharp in the larger Felidae, and which sends down a sharp spine, which may be called the tentorial. This latter is present in tiger, but altogether wanting in lion and Fels spelea. In the former, also, the arch appears to be narrower than in the two latter. The whole of the cranial surface of the parietal and tentorium is ridged and furrowed for the convolutions of the brain. Nearly the whole of the temporal surface of the bone is occupied by the great third branch of the temporal muscle, which has its principal attachments along the posterior part of the sagittal crest. The other muscles associated with the sagittal crest are small, and spring rather from the Wormian, and the front of the occipital crest rather than from the parietal. They are all connected with the movement of the ear, and are termed by 1 Pl. X, “tentorium.” FELIS SPELAIA. 55 be) ere Straus-Durckheim “‘ sagitto-pavillien, occipito-pavillien,’ and occipito-scutien.! The first appears to have a retrorsal and the others a rotary action on that organ. There are also many other small muscles connected with the movement of the ear, for which we would refer to the pages of the author we have so often quoted on the myology of the cat. The proportions of the Wormian and parietals are not constant in the skulls of the larger species of Felidee. ‘There is, however, a tendency in those of the adult tiger to a greater development of the whole of the posterior and upper solid mass at the juncture of the sagittal and occipital crests than in those of the oldest lion. It also has a greater upward projection, which gives to the tigrine skull the “ serpentine” outline, which Cuvier considers characteristic.” The examination, however, of a very large number of skulls shows their variability in this respect, and compels us to look upon it as a tendency only. In the straightness, and even in one case the downward curvature, of this portion of the sagittal crest, Felis spelea agrees with Felis leo. § 17. Frontals (Pls. VI, VII, VIII, IX, X, No. 11).—The frontals in many, if not most, mammalia, present characteristics of more or less specific value, and especially in the leonine and tigrine skulls. They roof in the highest and most central region of the carnivorous skull, whether taken longitudinally or transversely. Hach presents the fol- lowing surfaces :—The superior or coronal, the antero-lateral or orbital, the postero-lateral or temporal, the postero-internal or cerebral, and the antero-internal, nasal, or ethmoidal. There are also two other surfaces, the median or symphysial, by which each is united to its fellow, and the parietal, a deep triangular suture, inclined diagonally backwards and downwards. Of this latter the great depth and deeply serrated structure is well seen in the spelzan skull, Pl. X, the parietals having been broken away. The superior or coronal surface of each half is somewhat triangular in form, the interfrontal portion of the sagittal suture being straight, up to the point of junction with the nasals, where the edge curves outwards and passes under the latter bones. ‘This edge, about one inch and a half more or less in length, ends in a sharp point, where it meets the maxillary. It then sweeps backwards, parallel to the median line, forming a deep notch for the reception of the frontal process (¢) of the maxillary, by which it is overlapped. The process of the frontal thus lying between the nasal and the maxillary may be called the naso-maxillary process (w). It is proved, by an examination of a large series of skulls, to be wider in lion and Felis spelea than in the tiger. From the top of the frontal process the fronto-maxillary suture passes forwards and downwards to meet the lachrymals. The edge of the bone, then passing backwards into the orbit, is articulated to the latter bone by a suture directed diagonally downwards and backwards, from the latter of which it is connected with the palatine and orbito-sphenoid by a long horizontal suture. It then on the I Op. cit., vol. ii, pp. 195 et seq. ? Oss. Foss.,’ vol. iv, p. 453. 56 PLEISTOCENE MAMMALIA. temporal surface is united to the temporal process of the alisphenoid by a small process passing into the anterior and upper angle of that bone. Thence it passes obliquely back- wards to join the straight, transverse, parieto-frontal suture, that has already been described. The boundary between the frontal and orbital surfaces is the “superciliary” ridge, that between the frontal and temporal the “temporal;’ and these two unite in the massive supra-orbital process (¢). ‘The frontal surface is generally wider and flatter in the fully adult lion than in the tiger; and in the latter animal it rises on either side of the inter-frontal suture into a well-defined rounded elevation, so that the entire mass of the supra-orbital process is directed more decidedly downwards than in the former animal. In lion the surface is more or less widely concave, and the mass of the supra-orbital process is nearly horizontal, the point only being directed downwards. Some aged skulls of lion, however, and particularly in the smaller varieties, approach those of the tiger in this respect; and the young of each species are, as one might expect, to be determined only with great difficulty as far as this character is concerned. But as we can discriminate adult leonine skulls from those of tiger in every case that has come before us by this character alone, we believe that it is tolerably constant. The smaller speleean skull (Pl. IX) agrees with the average and therefore typical lion in the possession of this characteristic, as also does the larger specimen (Pl. X), so far as its abraded condition allows us to judge. ‘The small skull in the British Museum, from Sundwig, on the other hand, is somewhat tigrine m this respect, but not more so than small skulls of Felis Jeo in the same collection. Across the inter-frontal suture, at a small distance from the nasal notch, a semilunar depression’ exists in many skulls of tiger, that is only slightly indicated in those of lon. We do not, however, find this constant. In all the three speleean skulls this depression is absent. The temporal ridges bounding the posterior part of the frontal surface pass diagonally backwards from the supra-orbital process on either side to unite in the sagittal crest. In the young of the larger and in the adult of the smaller Ve/es, they pass far over the parietal before they join, so that the sagittal crest does not reach as far forwards as the frontal bone. In the adult tiger it passes over the parieto-frontal suture, sometimes to the extent of an inch and a half before its point of junction with the ridges, while in the adult lion its extent on the frontal bone is very small, and in many cases the temporal ridges extend as far back as the parieto-frontal suture. In this respect the smaller skull of Helis spelea (Pl. TX), which is that of an old animal, is typically leonine, and the larger, though younger, figured in (Pl. X), shows the same tendency, as does also, though in a somewhat less degree, that from Sundwig. Also, if a line be drawn across the frontal surface joming the posterior edges of the supra-orbital processes, and from the point where it cuts the inter-frontal suture a measurement be taken to the fronto-parietal ' See Owen, ‘ Proceed. Zool. Soc.,’ Jan., 1834, p. 1. FELIS SPHLAIA. a7 suture, we get the temporal length of the frontal bone. If this be taken proportionally to the basal length of the skull, it is, so far as our experience goes, invariably greater in tiger than in lion; the nearest approach to an equality being afforded by a very small tigrine skull in the Oxford Museum, from the Himalayah, and a large Gambian lion (112 d) in the British Museum. | The orbital surface may be considered as a section of the interior of a hollow cone, of which the apex is directed obliquely backwards and downwards. The greater projection of the supra-orbital process causes the concavity to be deeper in lion and Felis spelea than in tiger. It is pierced near the palatine suture by the small “internal posterior orbital foramen,’ homologous with that in the human fronto-palatine suture. It serves for the passage of the ethmoidal nerve and artery from the orbit into the anterior end of the cranial cavity, close to the cribriform plate. The postero-external or temporal surface is highly convex vertically, and extends to the temporal ridges and the sagittal crest where the latter reaches the frontals. It roofs in the anterior portion of the cerebral cavity. In old animals of lion and tiger strong ridges are sometimes present in the lower part, directed obliquely upwards and forwards. ‘The mass of solid bone immediately over the anterior lobes of the cerebrum is of great thickness for the size of the animal, being 1°55 inches in the larger speleean skull (Pl. X), and 1:54 im a lion in our own possession. ‘The cerebral surface is deeply concave, and excavated for the convolutions of the brain, and for the arteries. A slight groove marks the inter-frontal suture. The vault formed by the cerebral surfaces of the frontals is interrupted anteriorly by an oval arch of about half its height. It leads anteriorly into a chamber, the rhinencephalic fossa, the end of which is closed by the beautiful tracery of the cribriform plate. The abutments of the arch are of great strength. The latter is the equivalent of the ‘‘ethmoidal notch” in man. It appears to be generally wider m lion and Felis spelea than in tiger. Its width in the skull figured in Pl. X is 0°55, in a lion’s skull of our own 0°54 inch. The ethmoidal surface follows for the most part the form of the thin optic plate. The great thickness, however, of the coronal plate makes the roof of the cavity underneath convex longitudinally. The inter-frontal suture is strengthened below by a long spine projecting far into the nasal cavity, and attached to the vertical plate of the ethmoid. It is called the “nasal spine of the frontal,’ and divides the posterior part of the cavity into two large hollows, which receive the upper. and posterior masses of the ethmoidal convolutions. The walls also of the optic plates are ridged longitudinally (see Pl. VIII). These cavities for the reception of the ethmoid are called the “ethmoidal sinuses of the frontal.” ‘The arrangement of the frontal, cranial, and optic, plates is such as to form opposite the supra-orbital process ‘two large pear-shaped chambers, connected by small orifices with the ethmoidal cavities described above. These chambers are the frontal sinuses, separated from each other by the nasal spine. This massive bone affords points of insertion to many important muscles, the largest of which is the great temporal or crotaphite, the antero-external branch of which rises 58 PLEISTOCENE MAMMALIA. from the temporal ridge and posterior border of the supra-orbital process. The anterior part of the third and internal branch springs from the lower portion of the temporal fossa, while the posterior takes its origin from the parietal. Several muscles of the eye, namely, those for the rotation and general movement of the eyeball and the eyelids, also have their point d’appui in this part of the skull. The rectus internus or “grand adducteur’ and “‘netit adducteur” of Straus-Durckheim, equivalent to a portion of the choanoid, rise near the orbito-sphenoidal suture. Above them are the roots of the rectus superior or “ grand. élévateur” and the “petit élévateur” of the same author, equivalent to another portion of the choanoid ; and above these is that of the elevator of the upper eyelid. ‘The palpebral, in addition to its other attachments to the maxillary, the lachrymal, and the fronto-malar ligament, springs also from the extremity of the supra-orbital process. Of the muscles of the ear, the “ sourcilio-scutien” of Straus-Durckheim does not appear to be represented in man. It rises within the. orbit, and performs the function of bringing forward the ear. Another muscle taking part in the same office, the fronto-auricularis, springs from the superciliary ridge. The scutiform cartilage is attached by means of the temporal aponeu- rosis to the anterior part of the temporal fossa. ‘To it are attached muscles which control the motions of the ear. In the temporal aponeurosis a considerable portion of the great temporal muscles takes its origin, besides other muscles of no particular im- portance. § 18. Nasals (Pls. VII, X, No. 15).—The nasals in Felis are two thin plates of bone forming the anterior part of the roof of the skull. They are scalene-triangular in form, deeply curved transversely, nearly straight longitudinally, and united somewhat loosely by a long, straight, deep suture. They form the upper covering of the nostrils, being articulated behind to the frontals by a deep overlap, as we have described in our description of that bone, laterally to the maxillaries by a suture, the inner edge of which projects into the nasal cavity, forming a sharp ridge for the attachment of the anterior branch of the ethmoid. They are also articulated for a short distance to the inner and upper edges of the nasal processes of the inter-maxillaries. In the living animal the anterior edges of the bones afford attachment to the epirhine cartilage anteriorly, and to the pararhine externally, which form the outer portions of the nostrils. The nasals appear generally to be stouter and more decidedly triangular and flatter posteriorly in the lion than the tiger, being depressed so as to form a median groove at the symphysis in the latter animal... M. de Blainville’ also notes as differences in the latter animal as compared with the lion—“ La déclivité des os du nez, qui sont aussi plus étroits, plus allongés, plus parallelogramiques, le lobe inférieur de leur bord libre étant plus pro- longé et plus detaché :” we have found, however, that the variations are so great in these 1 Owen, ‘ Proceed. Zool. Soc.,’ 1834. 2 «Ost. Felis, p. 28. FELIS SPELAA. 59 points in a large series of leonine and tigrine skulls that we cannot consider them of specific value. The frontal’ muscle is inserted into the lower extremity of these bones: it passes upwards to join the fronto-auricular muscle, another branch forming a part of the elevator of the nose and upper lip. These bones are not perfect in any skull of Fels spelea which we have seen, being almost entirely absent from the British specimens, and in that from Sundwig presenting only sufficient to show the flatness of the posterior portion which is characteristic of hon. 1 Straus-Durckheim, op. cit., vol. ii, p. 187. PLEISTOCENE MAMMALIA. 60 GL-T ¢8-1 82-1 69-1 ¢6-1 06-6 C1-G 08-6 £8-1 ie 8/-1 x OF-1 CE-1 OF-1 90-1 G&-T CPI 09-1 LL-1 OZ-1 ; GE-1 a Z9.¢ 0v-S CL-€ 0G-G CES 09-€ 96-¢ OL-T 0¢-€ = hd OI-€ |806-2 OF-€ 8V-§ 8L-€ CL-3 CZ-G || 0G-€ 08-€ 69-6 C9-¢ ao CE-E pe-€ 06-¢ C6-Z C0-€ 88-6 0€-G 80-€ C8-6 00-€ OF-G £8-6 ‘ 0Z:¢ aa 81-1 06-1 01-1 90-1 61-1 C3-1 06-1 II-T C6-0 ; 06-1 66-1 0€-1 CLE GL-€ LG-€ G1-€ 19-€ GL-€ €1-V CVs 6-0 69-€ 99-€ GL-€ O1-S 91-G 0¢-7 CO-V 68-P €€-G 8E-¢ €0-7 DL-G a 88-7 66-4 | . Eg | £ Bee sss | 5 Bo ; | Sa s | 5 Aputt DenTIrion. =e | Be | 28 |Saa s24| 8% 4 | S85] 62 | 23 zl Upper. et nose me |BEZ| Poa g < 2) Se ee OK = Ss ~fo|saco|] =k : reno|] S54 | Beze bo a datas Ss ales | =o = pat| He Tors ae I S26) 5 Fe pe AS) 5 fer ae} A) = c= J n R om 9] nw 3 er 4 a = (ea) fo) a) So ln Molar series. Length of eases 3°40 | 2°74 | 2°70 | 3:10 | 3:00} 2°80! 2°80 | 2°75 | 2:50 | 2:20 80 PLEISTOCENE MAMMALIA. Felis spelea. Felis leo. Felis tigris. 5 g , tof 23 3 i | Aputt DENTITION. E (ah 2 5 3 5 = 2 5 Lower. 3 2s ah: e es De ¥ S, aes Big «AN Use None tal wun a ce S&S |&8 PSE S| ZED Tey sles Wa | 7e5 5 Ss a = oe 3 & SP ia et ik ne E 6 Incisor 1. ag pen gth es essary te rene ees 1 Ofer Ee he His ie ane Ph, Valerled aXe Of GRO bo 546 Joc 2b- 029\5F 0:26 | 0:25 | 0°26 | 0°28 | 0°28 3, Width of ditto .................. 0:20 | 0°20 0:15 | 0°17 | 0:15 | 0°20 | 0°28 4. Circumference ................-. 0°60 | 0:60 0°45 | 0°50 I =e 26 33 = a & Incisor 2. ci Hs Teme th hie, aecha td enti aes Tess | Wests jp Wee) OOS | sa bee 1:0 ee ie 2. Height of crown.................. 0°35 | 0°38 | 0°32 | 0°25 | 0°32 | 0°30 | 0°30 | 0:34 | 0:34 See Widthyot ditto ee ere eter: 0:26 | 0:29 | 0°22 | 0°18 | 0:22 | 0°20 | 0°24 | 0:20 | 0°24 4, Circumference .................. 0°80 | 0°85 | 0°75 | 0°65 0°70 | 0°75 Incisor 3. jl Ufet V4] een ae Henan aa ta esc 1°65 | 1:51 1°20 nee B65 1°30 Ay eae 2. Height of crown.................. 0°45 | 0°47 0:40 | 0°68] 0:40 | 0°40 | 0-40 | 0:44 3. Width of ditto 0:42 | 0°33 0:29 | 0°28 | 0:27 | 030°} 0°30 | 0°35 4, Circumference .................. 1:05 | 1:00 0:98 0°87 | 0:90 Felis spelea. Felis leo. Felis tigris. Williams? Collection at Taun- | £2 : 3 Elie] e illiams’ Collection at Taun-| == | 22] | 3 i o| So g eel (ees ton, from Bleadon. aa ce alee (A) Nl cee gS 38 a a3 S226 |22| 2 |2e|" alee a | OS Se Ss) oO) | Sigs | Ba [OS wo las SRISS|SE)E EES SIF jae 3 3) o iB a 1) i} i] 9] 5 | sa q 5 PaaS hh ck B Canines. 1. Length of outer curve...... 5°40) 5:00) 5:50) 5°50\ 4°20) 4°78) ... 13°99 | ... |... |... |... | eee | eee 2. Length of crown ............ 2°10) 1:60] 1:70] 2:10] 1°50] 1-90} 1-30) 1-5 | 1°50) 1-60) 1-58) 1-95] 1°70) 1°65 3. Antero-posterior diameter | 1°10} 1°20] 1-20] 1-10] 0°98] 1-10) 0-90) 0°85) 0-90) 1°15) 0-87) 1-07 1:00) 0°82 4. Circumference ............... ean 3°20) 3°10) 2°60} 2°80) 2-10 2°10} 2°90} 2°50) 2:75} 2°45) 2-10 FELIS SPELAA. 81 Felis spelea. Felis leo. Felis tigris. Williams’ EGNOS Y wile Gateeien Beard Collection. W. B. a|.6 (3 ApuLt DENTITION. oTeanion Dawkins. | Silane. Sica Lower. from ; Be oF Woe ay ie SEES cf & 3 6 e 2. : | Oo |Hyzeena-den. eS |2S/ a |tni~ s& Bleadon. | =o || fenae |e er alle pat Vat 3 a i : : mite Z ae | ise e |AEISE| = (Salas Tm A) heel Sh 1 ey ee eis See ES Sian w|ORw O/;s8 1 | B® Alas 1 2 ro) o) wn ey PEs Rape VFS tle i NL AC a Premolar 3. 1. Antero-posterior measurement - | 0°80) 0°80} 0°78] 0°84] 0:62) 0-66) 0°60] 0 57] 0°80) 0-70) 0:58} 0°58) 0°78 2. Maximum transverse ditto ...... 0°50| 0°45} 0°42! 0-47] 0:34) 0°35} 0°34] 0-29] 0-40) 0:44}! 0°34) 0°34) 0°32 3. Height of crown .................. 0°54] 0-53) 0°46] 0°47) 0-39) 0°38) ... | 0°30} ... | ... |0°34) 0°34) ... 4. Circumference ...................-- 2°10) 2°10) 2°10) 2-20) 1-60} 1°72) 1°62) 1°52) 2-10) 1°80) 1-52) 1°70) 1-72 5. Transverse measurement, anterior Lobe a 5. eee ert 0°36) 0°34 0°37 0:30) 0-28 0°28 Felis spelea. Felis leo. Felis tigris. Beard Collection. 2 2 |e Williams’ Collection, from Bleadon |—_—_—___— 5 ae : =, | 6.3 Cave. 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Zz = 5 S S| Oo o o o iS) = (S) Ey S f aS < = tira) s S| s S| s = I Oe o 3 n RS = D4 iS 6) ) 6) S 2 eo |sk Es : > j ESI g 3g 3 3 3 a] = 2 |o 9 as _ isa] i ‘a Seo mieome ice We | gel 8 laa So ee ey Si n ee |e | | ee |e a | © Ie SF = ~ = o Extreme length ...... Soo rm ern ctw le vous Hes | bee [ld i[yoey Poeeel | ne ef LdtOh | L323) | M250 Minimum circumfer- ener ail sti inl eee Pe OPOOI ete rea | ee. [esse || swe fl ree | eee fy ALOR 4:4 SOR 3:00 Transverse measure- ment of proximal articulation ......... con N4LG nea dee all cee | uses lense A ieeeclienen Sex acre alt ere SHC7/ se typiL I Geen) Vertical ditto ......... rem A SO () emer ee te dy fel se le AO le Groin ie or Asie 323 () Transverse measure- ment of distal arti- culation ............ Pn eee ee ASSO 2-55 2-oos00 2°70) 2°40} 2°40} 2:4} 1:9 | 1:90 Vertical ditto ......... .. |... |... 14°00) 3°50) 3°50) 4°30 2°70 2°30) 2 2°0 | 2°20 Depth at distal end of deltoid ridge......... ee ere ener ee Hse OOD S2SGl oc leis oc OO Me Sole eGo oled, Transverse measure- MEME EAL ESA POU ty ieee meena ee lh el LVS) Le40p oh 20 TTA 0-95), 0292 16 H8 PLEISTOCENE MAMMALIA. CHAPTER XI. Femor, Pl. XVIII, figs. 4, 9. CONTENTS. § 1. Introduction. § 3. Measurements. § 2. Description. § 4. Definition. § 1. Introduction—The femur of Felis spelea very closely resembles that of the Lion and ‘Tiger, and, so far as we can judge from the fragments, it bore the same proportion to the pelvis and tibia that it does in those two animals, being much longer than the tibia, a proportion that is reversed in the smaller felines. As we have met with no perfect spelzean femur in Britain, we have adopted the-same artifice as in the humerus. We have used as the groundwork of our figure the cast of a perfect spelaan femur from Gailen- reuth Cave, the original of which is in the collection of Sir Philip Egerton, F.R.S. It is drawn in alight tmt. In a somewhat darker tint we have represented a large portion of the shaft of a British specimen, and in full tint a considerable portion of the head, which, equally with the above, exactly corresponds in size with the cast, and a distal end which is rather smaller, and in this way we have built up the bone from fragments found in Britam. A small portion of a still larger distal end and the entire distal end of a smaller specimen are with others in the Taunton Museum. ‘They are all from Bleadon Cavern. Since the figure was drawn we have found a nearly perfect shaft in the Jermyn Street Museum, obtained from the brickearth of Hartlip, in Kent, and slightly smaller than the specimen from Gailenreuth. A very fine specimen also of the shaft, slightly smaller than the femur from Gailenreuth and that from Bleadon, which we figure, has been found in the gravels of Barnwell, a suburb of Cambridge, and is preserved in the British Museum. § 2. Description.—The head (fig. 4, a a’) of the bone is hemispherical, and larger than the neck, which it overhangs distally (fig. 4, @). On its postero-internal surface is a very shallow depression, much less strongly marked than in most animals, for the ligamentum teres. ‘The neck connecting the head with the shaft is short and massive, and resembles FELIS SPELAIA. T19 in form the frustrum of a compressed conoid, the proximal surface being horizontal, while , the distal pomts diagonally upwards at an angle of forty-five degrees. The shaft is nearly straight, cylindrical, and slightly enlarged at the ends, and when compared with that of the Bear is far more massive. It gives the idea of immense strength combined with great lightness. _ A stout ridge runs from the posterior edge of the head, parallel to the distal surface of the neck, to the smaller trochanter, or “ trochantine” as it is termed by Straus-Durckheim, which is an oval process with its long axis parallel to the neck. To its smooth summit is attached the psoas muscle, the iliacus internus in the Cat being merely a second head of the latter. From the outer and lower base of the trochantine a sharp edge turns diago-. nally upwards, and forms the external edge of the great trochanter, and affords attachment to the quadratus. The trochanter is formed on the same plan as in Man, but is pro- portionally larger and higher ; it is separated from the neck behind by the great trochan- terlan cavity in which the tendons of the obturator muscles are inserted. Its external summit is chamfered and hollowed for the attachment of the pyriformis ; to its rounded outer summit (fig. 4, 6) is attached the gluteus medius, and anteriorly to the massive tuberosity (fig. 4, c) the gluteus maximus. The adductores longus and magnus are in the Feles extensor muscles of the thigh, and therefore require an attachment posterior to the lateral position they occupy in Man. Consequently the sharp ridge forming the outer lip of the linea aspera in Man is, as it were, removed to the extreme outer edge of the posterior surface of the bone, and the two above-named muscles, termed im their new position curvatus and arquatus by Straus- Durckheim, occupy the greater part of the posterior surface. The inner lip of the linea aspera is represented by a slightly roughened surface passing diagonally across the bone, and affords attachment in its upper part to the adductor brevis and the pectineeus. The outer lip of the linea aspera is roughened and enlarged immediately below the lateral tuberosity of the great trochanter, so as to form a rudimentary third, which affords attachment to the gluteus maximus. Inferiorly it may be traced to the external angle of the outer tuberosity above the outer condyle. This external position of the linea aspera causes the origin of the vastus externus to be entirely on the anterior and external surfaces, and it consequently occupies the whole of the upper part of this surface of the bone and the anterior edge of the linea, while the part corresponding internally is occupied by the vastus internus; between them are the origins of the crural proximally and the subcrural distally. The second head of the triceps cruris attached in Man to the outer lip of the linea aspera has no analogue in Felis. Hie The distal extremity of the femur differs remarkably from that of Man. ‘The two con- dyles (fig. 5, 27’) are subequal, the internal (figs. 4, 5, 7) being somewhat the larger, and reaching slightly further down. The intercondylian anterior articulation (figs. 4 5 7) for the patella is square in outline, and is defined by a high and well-marked ridge; it occupies 120 PLEISTOCENE MAMMALIA. almost the median line of the shaft. The lateral surfaces of the process on which it stands are very nearly symmetrical (fig. 5, g g’), the external (fig. 5, g) forming aslightly more acute angle with the patellar articulation ; the condyles extend further behind than in Man, and their lateral surfaces (figs. 4, 5, 77’) are roughened for the attachment of the lateral ligaments, and externally (figs. 4, 5, 7) for the attachment of the extensor digitorum or cnemodactylus of Straus-Durckheim. The condyles are separated by a very deep depression (fig. 5, /) for the crucial and other ligaments, passing slightly inwards, and rendering the inner condyle slightly smaller than the outer. i Immediately above the condyles the posterior surface of the shaft is flattened, and on its lateral edges are the internal and external tuberosities (fig. 4, mm), affording origin to the gastrocnemii muscles. In the angles between the condyles and the shaft are two small depressions (figs. 4, 5, 2) for the lodgment of the sesamoid bones, called the external and internal crithoids, which, with a third below the external, are usually termed fabelle. The two former are in the tendons of the gastrocnemii, and the latter in that of the popliteus. The position of the nutritive artery varies, sometimes piercing the shaft in the middle of the lesser linea aspera, sometimes in the greater. A fine specimen of the spelaan femur was discovered by Dr. Schmerling’ in the caverns of Liége, and is figured in his great work. It agrees in every respect with the German and English specimens. ‘The figure is produced by M. de Blainville’ in his ‘ Ostéographie.’ § 3. Measurements—The following table shows the variations in the size presented by the femora of J”. spelea, F.leo, and F. tagris. TapLte oF MrasureMENTS OF Femur IN Friis sprima, F. Leo, anp F. TIerRIs. Felis spelea. F. leo F. tigris. a Taunton Museum. » | a . a eB MEASUREMENTS OF | % & = BE j S23 | 25 ss Femur. 65 Ee |oe| 4 | 82 | 823 aS 3a 3 3 2 8 Die jms | = ee | oe | =e D ai 3 3S 3 3 ae | 2 = Ras | Re Eun 2 2 2 2 a a 5 ga 3 a + =? aa) a ea) i) an | ties = 1 16°65 ue a 16°60 14°40.| 14°5 | 12°4-| 14-2 2 5:00 ... | 4°80 ASQ S29 Ona oleae 4am nes 3 3°00 | 3°10 Ash Re ua 2250)\|\ 320i 2am eee 4 3°00 | 3-10 Bae PRIX | Be) || Bes} |) 83 5 3°20 3°20 | 3°00 3°40 3°00 | 2:3 | 261] 2°5 6 6°30 5°80 | 6:00 460 | 48) 4:5} 4:3 1 Oss. Foss. de Liége, tom. ii., Pl. xvi, fig. 2. ? Felis, Pl. xviii, fig. 0. FELIS SPELAA. 121 § 4. Definition—On comparmg the femora of the Cave Lion with those of other animals associated with it in Pleistocene deposits, the following points of difference may be enumerated. The straightness and cylindrical form of the shaft, and the symmetrical form of the distal end, distinguish it from that of the Cave Bear. In the Hyena it is also sym- metrical, but the great difference im size prevents the two being confounded together. In the latter animal, moreover, it is rather more bent and the patellar articulation is not so sharply defined. The large development of the third trochanter in the Horse is a point by which the most slender bone may be distinguished at a glance. 122 PLEISTOCENE MAMMALIA. CHAPTER XII. Trpia, Pl. XIX, figs.4,1’,1”,2, 2’. Fisua, Pl. XIX, figs. 3, 4. Parunxa, Pl. XTX, figs. 5, 5”. CONTENTS. § 1. Tika. § 2. Fibula. a. Description. a. Description. B. Measurements. B. Measurements. § 3. Patella. y. Definition from other Pleistocene a. Description. tibie. (@. Measurements. § 1. Tibia (PI. XIX, figs. 1, 1, 1,” 2, 2’).—The tibize of all the digitigrade Carnivora are remarkably alike in general form, and offer constant differences characteristic only of genera or widely dissociated species. They present variations of proportion in animals of the same, as great in degree as in the closely allied but distinct species, such as Lion and Tiger, and are therefore of comparatively small value in classifi- cation. We have met with several specimens of this bone in the caverns of Somerset ; the one (figs. 1, 1’) is perfect, with the exception of the proximal epiphysis, and belongs to a young animal; the other (figs. 2, 2’) consists of the proximal articulation of an adult; — the former was obtained from Sandford Hill, the latter from Bleadon. In the larger Feles generally the tibia is shorter, both proportionally to the femur and in relation to its own minimum circumference, than in the smaller forms. In Lion, Tiger, and Felis spelea, it is considerably shorter than the femur ; in the domestic and wild Cats, considerably longer. The articulations, as we have elsewhere remarked, are frequently larger in Tiger than in Lion, but the variations in this respect are so great that we agree with M. de Blainville in considering them of no specific value. § la. Description.—The tibia of the larger Feles is a strong bone of slightly double curvature, bent gently forwards distally, and backwards proximally, cylindrical distally, but expanding into a prismatic form proximally; so that the proximal vertical diameter of the shaft is more than double the minimum near the distal end. ‘The prism is so disposed that the narrowest side (fig. 1’ 6) forms the posterior face of the bone, while the two broader meet in the strong anterior crest (figs. 1, 1’ a), which curves gently outwards and may be traced in the adult as far down as the distal end. The head (figs. 2, 2’ c, d) of the bone is partially occupied by the two slightly concave semilunar facets for articu- lation with the femur separated from each other by the small bifid eminence termed the en ee FELIS SPELASA. 123 spine (figs. 2, 3’, e). They are deepened in the living animal by the semilunar cartilages that form an elastic or variable socket, and, with the adipose igament in front, make up a broad articulation for the support of the hind quarters of the animal, which would other- wise rest on mere points. At the intersection of the nearly plain tibial facets and the highly convex femoral condyles, these cartilages are firmly fixed to both tibia and femur by strong ligaments. The facets are separated posteriorly by a deep notch (fig. 2’), in which is implanted the posterior crucial ligament, and are strengthened laterally by the external and internal tuberosities (fig. 2, g, 4), affording attachment to the external and internal lateral ligaments. On the under side of the external tuberosity is a small oval articulation (fig. 2, g), for the head of the fibula, and under the posterior edges of the same tuberosity is a groove for the semi-membranosus muscle. In front of the spine, and forming a small shallow indentation in the anterior surface of the external facet, is a roughened space of triangular form (figs. 2, 2’, 7) that extends inwards, so as to cut off the internal facet from the base of the crest. It affords attachment to the adipose cushion or ligament, filling the space between the tibia, the fore part of the distal articulation of the femur, and the patella. A strong and massive anterior tuberosity forms the antero- external boundary of the external facet (figs. 2, 2’, m). The proximal end of the crest is covered by a projecting and slightly roughened mass called the tubercle (figs. 2, 2’, x), which extends considerably downwards in the larger Feles, and in the two bones at Taunton of Felis spelea, is nearly parallel to the posterior surface of the shaft. It affords attachment to the patellar tendon, which, through the medium of the patella, is the principal tendon of the muscles which act at once as the extensors of the tibia and flexors of the femur. Many of these muscles are also partially attached to the head and sides of the tibia, such as the rectus internus and sartorius, forming one muscle, the fascialis, the rectus anticus, vastus internus, and semi-tendinosus while others, viz., the arquatus, of Straus-Durckhem, the equivalent of the adductor magnus in Man, the popliteus and semi-membranosus and triceps cruris, act as the extensores cruris and flexores tibie, and leave their marks in the ridges and grooves on the posterior face of the bone. The posterior angles of the prismatic portion of the shaft are termed the extero- and intero- posterior crests (fig. 1’, 0, ), the latter of which is traversed by a minute ridge, which is’ the line of attachment of the mter-osseous membrane that binds the tibia to the fibula. Near this and at a distance from the proximal end of about one third of the entire length, is the small foramen for the passage of the nutritive artery and nerve. The outward sweep of the crest (fig. 2, m) forms on the external surface a large concavity (fig. 2, 9), affording origin to the large tibialis anticus, which acts as a flexor of the foot and extensor of the toes. A considerable portion of the length of the shaft is occupied by the origin of the flexor longus digitorum on the inner side, the tibialis posticus running in a parallel direction on the external surface close to the inter-osseous membrane. The shaft imcreases in size as it approaches the distal articulation (figs. 1, 1”, 7, s), which is of somewhat trapezoidal form, wider than deep in a vertical direction, divided 124 PLEISTOCENE MAMMALIA. diagonally by a strong ridge into two concave depressions for the reception of the condyles of the astragalus. The inner of these is by far the deeper, because the internal malleolus (fig. 1, 1”, r) descends much further than the internal (figs. 1, 1”, s). The internal malleolus is rough and massive, and traversed posteriorly by a small groove (fig. 1”, 4), running downwards and forwards, which is converted into a canal by an investing ligament, and receives the tendons of the tibialis posticus, and the flexor longus digitorum. The external edge of the outer facet (figs. 1, 1”, s) is rounded off to receive the small internal and lateral articulation of the fibula. § 1 B. Measurements.—The only difference to be found between the tibie of the Felis spelea and the living Lion and Tiger is the massive proportions of the former. The specimen (fig. 1) is perfect, with the exception of the proximal epiphysis. Had the latter been present it would have been of nearly exactly the same length as that of the lion in our own possession, whereas the followimg table of measurements shows how much they differ in bulk. Other fragments in the Taunton Museum corroborate this evidence, and the most slender of the larger specimens is more massive than any leonine or tigrine bone which we have seen. Others, however, differ very little from the propor- tions of those in the two latter animals, and the whole form a graduated series without any break. In the large spelzean tibize the tuberosities are somewhat larger, and the tubercle for the patellar tendon passes further down on the crest, so as to form an attachment proportional to the massiveness of the limb. CoMPARATIVE MEASUREMENTS. Felis spelea. Felis leo. F.tigris. Taunton Museum. | ie E | = 2 B 2 s ¢ =| ° = 53 S ae | 33 $ S 3 3 3 n =s BS | ee le EMS ME iil cueiu| 4, (el nee ee eee os oO 1) eI} o (>) o = Pas) = ee} || BS 5 = 8 E E = a aS ca Eee aby ef E) peal re ¥ jae] 3 re ees ile Aye iliesp | ies) Se || Be | & n 1. Maximum length ane ahi ee Ee eu as ace 12°50 | 11°80 | 11°6 | 11°4 | 2. Minimum circum- ference............ 4°80 ane me! 4:25} 4:50 | 3°90 | 4:50 3:28 3°00 3:2 3°3 3. Transverse measure- ment of proximal articulation ...... ie Se) | SAD | ae Le Sas si 3°00 2°70 29 29 4. Vertical ditto ...... bak 2°80 | 2°60 a see a a 2:20 1:80 16 16 5. Transverse measure- ment of distal articulation ...... 3°00 Bi a fie ae 3a Eo 2°30 2°08 1:9 1:9 6. Vertical ditto ...... 1:90 Lee Da ee 538 es ig 1:30 1:21 1:2 1:2 =e ee THE PALHONTOGRAPHICAL SOCIETY. INSTITUTED MDCCCXLVIL. VOLUME FOR 1868. LONDON: MDCCCLEIX. iam) el BRITISH PLEISTOCENE MAMMALIA. BY W. BOYD DAWKINS, M.A., F-.R.S., G.S., AND W. AYSHFORD SANFORD, F.G.S. PART III. BRITISH PLEISTOCENE FELID&. FELIS SPELAA, Goupruss. PALIS LYNX, Linnavs. (Pagans 125—176; Pruarrs XX—XXII, XXIa, XXII, XXIII.) LONDON: PRINTED FOR THE PALMONTOGRAPHICAL SOCIETY. 1869. PRINTED BY J. KE. ADLARD, BARTHOLOMEW CLOSE. FELIS SPELAA. 125 § ly. Definition from other Pleistocene tibie—The tibia may be distinguished from all other bones associated with it in Pleistocene deposits, by the following points :— From any of the Deer or Ox tribe, by the oblique direction of the astragaline facets ; from the Horse, by the slenderness of the shaft. It approaches that of the Hyzena very closely, but in the latter animal the crest is longer, both proximally and distally, so that the head is flatter and the whole shaft more decidedly prismatic. Its size, also, would at once stamp its character. In the Bear the shaft is decidedly prismatic throughout, the distal articulation is much wider, and the astragaline depressions shallower. The internal malleolus does not descend so far, while the whole epiphysis extends much further externally. The crest also is much smaller, as well as the patellar tubercle; the extero- anterior tuberosity is much lower, and the depression for the adipose ligament does not cut off the internal facet from the patellar tubercle, but passes down straight over the latter; the semilunar facets are flatter, and the posterior attachments for the muscles are far more distinctly marked. We know of no figures of any fossil tibiee of Felis spelea. § 2. Mbula (Pl. XIX, figs. 3, 4)—We have met with fragments only of the fibula of Fels spelea, one of which consists of the shaft, and the other of a perfect distal end; both were obtained from Bleadon Cave, and are figured in Pl. XIX, figs. 3, 4. § 2a. Description—As we have never met with the proximal end of the bone that” afforded origin to the soleus muscle, we omit all notice of it; analogy would show that it was identical in form with that of the living Lion. The shaft of the bone (fig. 3) at the proximal end is triangular in section, and its posterior surface is shown in the figure (a). The roughened surface affords point of origin to the peroneus longus. The shorter of the remaining two sides or the anterior is the fibular origin of tibialis posticus, which is attached to this bone in Felis. Below, the shaft becomes cylindrical, having a sharp high ridge (4) on the internal surface, which is the line of attachment for the fibulo-tibial inter- osseous membrane. On the posterior side of this ridge, about the middle of the bone, is the origin of the flexor longus pollicis, and lower down that of the flexor longus digitorum. It curves forward near the distal end and forms a sharp wedge-like tubercle, on the outer side of which (fig. 3, e) is the origin of the peronzeus tertius. At the inner angle of the distal end there is a roughened surface, which is developed in the adult into a second sharp ridge, that also runs spirally backwards half round the bone, so as to form the posterior distal edge. On the outer side of this, a little below the middle, is the origin of the peronzus brevis (fig. 3, d). The two ridges above described are opposite to each other at the distal end, and form a flat blade-like expansion, on the inner side of which is a slight polished elevation, which is the distal articulation with the tibia. The extensor communis digitorum or cnemodactylus of Riolan is not attached to the 17 126 PLEISTOCENE MAMMALIA. proximal end of the tibia in Felis, but passes upwards to the external face of the condyle of the femur. In fig. 4 we have represented the distal end of a left fibula which resembles in every respect, save that of size, the corresponding portion of the leonine or tigrine bone. It presents externally a deep groove between two tuberosities, which, in the living animal, is converted by a ligament into a canal for the tendon of the peronzeus longus (fig. 4, a). The posterior tuberosity forms the external malleolus, behind which is a second groove (fig. 4, c), deep and narrow, for the tendons of the peronzi brevis and tertius. Internally we find a small concave articulation close to the epiphysial division, by which the bone is attached laterally to the tibia, and below this is a flat surface which articulates with the outer side of the astragalus, and behind this, again, is a depression for one of the ligaments which bind the fibula to the tarsus. From the immature state of the shaft we have been unable to institute a rigid com- parison between it and those of the recent Feles. Dr. Schmerling' gives a rough and slight figure of a portion of a fibula, and states in the text that the upper portion is broken away; his figure, therefore, must be that of the external and posterior aspect of the distal end, together with about two thirds of the shaft ; with this exception, we have met with no figure of the bone. § 2 B. Measurements.—In the following table the superior massiveness of the spelzean fibula is shown over those of the Lion and Tiger. CoMPARATIVE MEASUREMENTS. Felis spelea. Felis leo. F. tigris. Taunton Museum. : SI gs 3 5 3 4 Oss! ada ; Ss | seh vae |S = S as (eiee| a Fa Beh is ae a =) 60 4 ar En aA 5S SY S 5 7 a he Mp aheng ion WOME oo acnoanoceonnscsonus 1 oadese cage se Aue 11:15 | 11°55 | 10:0 2. Minimum circumference ...............0-.-2--202-: 125 La 0°85 1:12 1:10 3. Transverse measurement of proximal articulation BS ree 0°35 0:75 0°45 4... Vertical ditto-s ce) | We eee dane RaneeM mea ae, Eve ba 1:30 1:30 1:05 5. Transverse measurement of proximal articulation ie 0°65 0°60 0°57 1°50 Gs Distallidittosy hy. fuG dah eel pe ae nie Eel) oe 1°23 1:10 1°26 0°45 1 «Oss. Foss. de Liége,’ vol. ii, pl. xvi, fig. 4, p. 80. FELIS SPELAA. 127 § 3 a. Patella (Pl. XIX, figs. 5, 5’).—The patella of Felis spelea, of which many specimens are preserved in the Taunton Museum from the caves of Sandford Hill and Bleadon, exactly resembles that of the Lion and Tiger in form, but surpasses them in size. As in the recent Feles, it varies considerably in size and proportion. In shape it resembles a flattened pear, the small end being that to which the great ligament is attached which unites the bone to the anterior crest of the tibia. Many anatomists consider that in this bone we have the analogue, in the hind hmb, of the olecranon of the ulna, a point that will be found fully discussed in the ‘ Cyclopedia of Anatomy and Physiology.’ The outer or anterior surface (Pl. XIX, fig. 5’) is roughened for the attachment of the tendons of the muscles, which we have elsewhere pointed out as the extensors of the leg and the flexors of the thigh, including the “ paracural” of Straus-Durckheim, which has no analogue in man. ‘lhe upper part of the proximal or posterior surface (fig. 5) is entirely occupied by the slightly convex articulation which fits the intercondylian or anterior facet of the femur ; its edges slightly extend beyond the body of the bone. The only patella lable to be confounded with that of Fels spelea is that of the Bear ; it may, however, be easily distinguished by its more oval form, and by the greater com- parative extent of its femoral articulation. § 3 B. Measurements——The variation in size is seen in the following table of measurements of the patella in Felis spelea, F. lev, and F. tigris :— Felis spelea. Felis leo. F. tigris. Taunton Museum. 8p 80 80 = 5 5 5 3 3 3 3 we 56 ne v2 Res EI 5 & < a 3 3 Oo oO iS) iS) oO oO ) = B sg g a = & s e i} ic) ° is} i=) oO onl eee) = oh ce a | & | 2 FS Ep fa fa a 3) Maximum length ...........0...:...--20.-00--- 2:90 | 3:00 | 2:80 | 2°50 | 2:40 | 2:42 | 1-85 |) 1:94 Maximum circumference ...................+. 473 | 5:00 | 458 | 437 | 400 | 4:1 3°7 3°4 Transverse measurement of proximal arti- OU) ENS TO Raa eemar nade seenaeocboHomsas ace haunt 185) | 2700 | 1:80) 1:70), I55))) 1:58 |) 1:4 1:3 Werticalidittor, nor ccesmascmeace etiemecoat 1:90 | 1°95 |} 1:95 | 1°85 |) 1:50 | 1:25) 1:1 1°33 1 Article “Skeleton,” p. 664 et seq. 128 PLEISTOCENE MAMMALIA. CHAPTER XIII. Carpus, Pl. XX, figs.1 1", 2;°3,°4, 5,5. CONTENTS. § 1. Seaphoido-lunare. § 2. Pisiform. a. Description. a. Description. B. Measurements. » § 3. Uneiform. a. Description. 3. Measurements. y. Definition from that of Ursus. 8. Measurements. Tn describing the bones of the carpus we shall consider the anterior or dorsal surface as that which is naturally so in all quadrupeds; the palmar and inferior or posterior as synonyms denoting the same portion of the bone. The carpus in the genus Felis is composed of seven bones, besides the small one which has no independent existence in human anatomy, called by Straus-Durckheim the ‘ phacoid.’ ‘The scaphoido-lunare, the cuneiform, and pisiform, compose the upper or proximal row ; the trapezoid, the trapezium, the magnum, and unciform, the lower or distal. Of these we have only met with the scaphoido-lunare, the unciform, and pisiform, in Felis spelea. 1. Scaphoido-lunare (Pl. XX, figs. 1, 1’, 2).—1 a. Description —The scaphoido-lunare is by far the largest and most important bone in the carpus; it extends throughout the whole width of the joint, and forms almost the sole means of attachment between the fore paw and the forearm. It is very massive and is roughly quadrangular in plan, with a large and strong tubercle projecting from the postero-imternal angle (fig. 1’, e) ; it is broader than long, and much thicker externally than internally. ‘The proximal or radial articulation (figs. 1, 2, a, 6), which covers the whole of that surface of the bone with the exception of that portion which is opposite the tubercle 1s convex, traversed intero-posteriorly by a depres- sion (figs. 1, 2, 6) running from behind forwards and inwards. On its external edge is a very small articulation for the upper edge of the pisiform, which just touches it at that point (figs. 1, 2, c). The tubercle (fig. 1’ e) is a somewhat pyramidal or conical mass, projecting diagonally FELIS SPELAVA. 129 inwards from the intero-posterior angle of the bone. It presents on its antero-internal surface a round flat articulation (fig. 1’, e) for the phacoid bone, to which is attached the adductor pollicis muscle instead of to the tubercle itself, as in man. It affords attach- ment to a large number of ligaments, the principal of which is the annular or armillary, which is attached by some of its lower fibres. It would serve but little purpose were we to enumerate them all, for they are extremely difficult to separate; they have been reckoned and described to the number of twenty-six by Straus-Durckheim.* They bind the radius to the carpus, and the bones of the carpus the one to the other. The distal surface of the bone is entirely articular, and is divided by well-marked ridges into three well-defined articulations, the internal being a slightly concave parallelo- gram (figs. 1, 1’, 2, 7) set diagonally outwards and downwards for the head of the unci- form ; the second (figs. 1, 1’, 2, g), beimg more concave than the preceding and wider posteriorly than anteriorly for the reception of the head of the magnum ; and the third, (figs. 1, 1’, 2, 2), beimg triangular and divided by a broad diagonal elevation into two slightly concave surfaces for the reception of the heads of the trapezium and trapezoid. We have met with no scaphoido-lunare of Lion or Tiger which equals in size several of those in the Taunton Museum belonging to Felis spelea, but we have figured one from Bleadon Cave (Pl. XX, fig. 2), which in no respect differs from those of either of the above animals. ‘The larger spelzean specimens are somewhat thicker proportionally than the smaller, as well as the leonine and tigrine. ‘That figured from Sandford Hill Cave, (Pl. XX, fig. 1), apparently belongs to the individual that has furnished us with a great many of the originals of our plates. § 1 6. Measurements—The following table shows the variation in size between the leonine, tigrine, and spelzean scaphoido-lunaria. CompPaRATIve MEASUREMENTS. Felis spelea. F. leo. |F. tigris. Taunton Museum. < - a e414 5 Bs | BE [kes = eS Weel [ace ie 4 BS |e Lem Wicxamamaylemotni. caer as Peete eee aie. wu sec ects. « lon |) 133] Wel | Uek@ | dene 2. Minimum circumference .........................-- Otel) | 7D | Hv | Sexo ra _3. Transverse measurement of proximal articulation | 2°30 | 2°10 | 1°87 | 1°80 | 1°8 Also NIGUAIIOR! GUMIG)o caamcmeargmentencs cst socben kA kaeee a anene ea We) jy shoves eats) is 5. Transverse measurement of distal articulation... | 2°08 | 1:96 | 1°65 | 1°64 | 1:42 GAMVeTUICH Mditvomen. sn ctgaa: Pose ee eee sees ene TOS) Al | SON SOs Orgs 1 Op. cit., vol. ii, p. 84 et seq. 130 PLEISTOCENE MAMMALIA. § ly. Definition from that of Ursus.—The scaphoido-lunare of the Bear may be distinguished at a glance from that of Felis spelea, the tubercle being much larger, cylindrical and projecting directly backwards, and the articulation for the magnum being parallel to the side of the bone instead of running diagonally across it, as in the latter animal. § 2. Pisiform (figs. 3, 4), a. Description.—The pisiform of Felis spelea strongly resembles in plan that of all the other Feles we have examined, those of the different species being only distinguishable by their size. Hven in the same species it presents small variations of shape and proportion. It may be considered, generally speaking, as a long three-sided pyramid, bevelled off proximally and distally for articulation, with the cunei- form and the styloid process of the ulna, the apex swelling into a tuberosity (figs. 3, 4, 6), for the attachments of the tendon of the ulnaris muscle, of those for the bending of the fifth digit, of the common flexor of all the digits, and of the transverse adductor of the first. It is firmly attached to its fellow carpals, to the metacarpals, and forearm, by a large number of ligaments, and thus it forms a powerful port d’appwi for the motions of the fore foot, analogous to the shaft of the calcaneum in the hind foot. The articulations may be distinguished as the proximal or ulnar (fig. 4, a), which is known by its semi- lunar form, and the distal or cuneiform, which presents the plan of a rectangular parallelo- gram with rounded angles (fig. 3, ¢). Both are nearly flat, and unite in a sharp and neatly straight edge. It may be known from the corresponding bone of the Bear by the possession of the following points :—By its greater length, by the flatness of the ulnar articulation as con- trasted with the concavity of that of the Bear, and by the straightness of the edge separating the articulations, which is concave in the latter. We have met with several specimens from the caves of Bleadon and Sandford Hill, some of which are but little larger than the living Lion and Tiger, while others (see fig. 4) are much larger than those of any of the living Feles. The original of fig. 4 probably belonged to the same skeleton as the large scaphoido-lunare figured in the same plate. § 2B. Measurements.—The variations in the size of the pisiform in Felis spelea, F. leo, and F. tigris are shown in the following table :— FELIS SPELAA. 131 Comparative MBasuREMENTS. Felis spelea. Felis leo. F, tigris. Taunton Museum. y I g o f=) =) se ee Mee ee 8 > > A See esr line iw! | Slee a5 m5 ag Ss 2 5 a ep | obo 3 3 7 a Go rs I § a ea ia) = ea is n Ils: leeetan iin NAM eA00 <6 -cacdocb6 concarinod sen Seeot MBSeeee DS 5e lO 2a CSO G77 ele 20m ele eel 2. Minimum circumference ..................-..--.2+. 210) 2250) |) 22007) 190 son | 1ESON I k35 3. Transversemeasurementof proximalarticulation | 1:05 | 0°95 | 0°96 | 0:93 : 0°95 ae A Vertical dittow acest ree eek cs ere seebnee aes 0°50 | 0°53 | 0-50 | 0°43 0°46 Bt 5. Transverse measurement of distal articulation | 1:00 | 1:00 | 0:82 | 0:74 | 0°75 | 0°80 | 0°75 Ome Vertical ditions mee eeepc cren ete ke pect crea 0:75 | 0°75 | 0:76 | 0°50 | 0°35 | 0°43 | 0:44 § 3. Uneiform (figs. 5, 5’) a. Description—The unciform of Felis spelea is a small, somewhat wedge-shaped bone, not distinguishable from that of the recent Lion and Tiger except by its massiveness. he head of the wedge occupies the anterior or dorsal aspect of the bone, and a very slight tuberosity is the only trace of the claw-like process which gives the name to the bone in man. The proximal or scaphoidal articulation (fig. 5, a) is much narrower than the distal, the sides being nearly parallel. It is much longer from front to back than broad ; convex anteriorly and slightly concave posteriorly, it ends on the palmar surface in a semicircular boundary. ‘he distal or metacarpal articulation (figs. 5, 5’, 4) partakes of the wedge- shape of the bone. It is deeply concave, and receives the heads of the fourth and fifth metacarpals. ‘The inner articulation (figs. 5, 5’, c) for the magnum is shaped something like the letter L, placed so that the stem coincides with the anterior edge of the bone. It is nearly flat. The remaining articulation, or the cuneiform (figs. 5, 5’, d), is for the most part slightly convex, and occupies the anterior part of the external surface. The anterior or dorsal surface assumes the form of an irregular pentagon (fig. 5), roughened and indented between the lateral articulations for the attachment of several interosseous ligaments. The thin end of the wedge, occupying the palmar surface, is a tubercle (fig. 5’, e) for the attachment of the adductor of the fifth digit; the opponens of the same digit not being attached, as in man, to this bone, but to the magnum. This bone strongly resembles that of the Bear, but is easily distinguished by the greater squareriess of the latter, as well as by the flatter distal articulation. We know of no other bone with which it can be compared. 132 PLEISTOCENE MAMMALIA. § 3. B. Measurements ——The variations in size of the unciform in Felis spelea, F leo, and F. tigris are shown in the following table : ComMPARATIVE MEASUREMENTS. Felis spelea. _ | F. leo. |F. tigris. Taunton Museum. o o =o is} <5 ES er ws Ney eh came a |e a) a "S a Lo Maximum: Tenet hi eee eee ees ee ee ree rere 1-40, | 1:82) “114 092 eer 2S ViinimumycireumierencCen ee etek seen nee 3:03 | 3°04 | 3°47 | 2:18 | 2°60 3. Transverse measurement of proximal articulation | 0°64 | 0°62 | 050 | 0:41 | 0:30 4.. Vertical Qittor5 ese eee ee eee ee nee NEO) eS Ue 7G || Ora! 5. Transverse measurement of distal articulation... 1:03 | 0°90 | 0°88 | 062 | 0°64 6." Vertical: dittonsstcen ee oe cee eee eee eee 1:20 | 1:10 | 1:20 | 0:80 | 0:98 FELIS SPELAA. 133 CHAPTER XIV. Mzracarpars. Pls. XIX, fig. 6; XX, 6,7; XXI, 1—5. Puauanezs. Pl. XXI, figs. 6—14. CONTENTS. § 1. Metacarpals. § 2. Phalanges. a. Introduction. a. First phalanges. 6. First metacarpal. B. Second ,, y. Second, Third, Fourth, Fifth, y. Measurements. metacarpals. 6. Measurements. § 1. Metacarpals. q. Introduction —As metacarpal bones of the genus Felis have to perform very much the same functions in all the species, they present but few characters of value in classification, the variation in the proportions observable in different individuals of the same being frequently as great as in the closely allied species. For the most part, however, the metacarpals of Felis spelea, from the bone-caverns, are stouter and stronger than those of the living Lion and Tiger; but, on the other hand, some of them are even smaller in every dimension than the average-sized bones of the two latter animals ; the large series also from the caves of Somerset proves that there is a gradual passage from the most massive to the most slender spelzean form ; size cannot, therefore, be considered of specific value. We are fortunate in being able to give figures of a complete set from the cavern of Sandford Hill (Pl. XXI, figs. 1, 2, 3, 4, 5), which belonged to an individual in the prime of life. They are remarkable for their straightness and stoutness, and the fourth presents a variation which we shall describe in its due place. We have also figured the proximal end of a fourth metacarpal of the small form from Bleadon Cave (Pl. XX, fig. 6), and a remarkably small fifth from the hyzena- den of Wookey Hole (Pl. XX, fig. 7). A gigantic second metacarpal from the lower brickearths of Crayford is also figured (Pl. XIX, fig. 6). The*metacarpals when united form a compact transverse arch, the convexity being dorsal, and the concavity palmar; the latter transmits the tendons of the flexor and adductor muscles to their points of insertion in the phalanges. 18 134 PLEISTOCENE MAMMALIA. § 1. B. First Metacarpal (P\. XXI, fig. 1).—The first metacarpal for the hallux or thumb, in its shortness and the obliquity of its articulations, differs from all the rest of its fellows. The proximal articulation with the trapezium occupies the end of the bone, and extends diagonally outwards and downwards on the dorsal surface, forming a shallow, pulley-shaped joint by which the thumb is freely moved in a transverse direction. On the internal edge there is a very small facet (4), which is in apposition with the phacoid bone in the recent Felidae. Hxternally also there is a small facet, which rests on the internal edge of the second metacarpal. The united action of these articulations and of the ligaments which bind this bone to the carpus, prevent any rotatory action ; and thus the thumb cannot be opposed, as in man, to the other digits. The shaft is short, stout, and slightly bent towards the outside, and traversed on its dorsal aspect by a groove running from the extero-inferior edge of the proximal articulation diagonally outwards and forwards, and receiving the tendons proper to the bone for the common extensor digitorum. The palmar surface is concave, and much roughened for the attachment of a large number of ligaments. The distal articulation (d) is very oblique, and faces inwards and downwards, so that the action of the phalange is still more transverse than it otherwise would be, and its tearing and grasping power is much increased. Immediately above it in the middle of the palmar surface is a small well- marked tubercle which affords attachment to the ligament, uniting the bone to the first phalange of the second digit, and to the adductor brevis pollicis muscle. ‘The majority of the tendons of the muscles that regulate the motion of this joint are attached, not to the bone itself, but to a large internal sesamoid, occupying the intero-inferior angle (e) of the bone. § 1. y. Second, Third, Fourth, Fifth Metacarpals (Pls. XIX, fig. 6; XX, figs. 6, 7; XXI, figs. 2, 3, 4, 5). The proximal end of the second metacarpal forms a wedge-like mass slightly shifted, as it were, outwards for a quarter of its width. In front it is rec- tangular ; behind, on the palmar surface, it throws out a small tubercle which is the thin end of the wedge. ‘The proximal or trapezoidal articulation (Pls. XIX, fig. 6; XXI, fig. 2, a) is triangular in outline; concave transversely, and nearly flat vertically. On the upper and outer edge of the palmar tubercle is a small flat surface (PI. XXI, fig. 2, 4), which articulates with the postero-internal facet of the os magnum, and immediately above is the somewhat triangular surface set at right angles to the trapezoidal articulation which is in contact with the antero-imternal facet of the same bone. At the antero-internal angle of the trapezoidal articulation is a small concave surface (fig. 2, c), which receives the convex external facet of the trapezium, and, extending from it backwards as far as the palmar tubercle, is a broad slightly concave surface for contact with the first metacarpal. Imme- diately under the articulations for the os magnum are two concave surfaces, the anterior being the larger and deeper (fig. 2, d), which overlap corresponding surfaces on the inner side of the third metacarpal. Between themis a large ligamentary cavity. On the dorsal FELIS SPELAVA. 135 or anterior surface of the proximal end runs diagonally a shallow groove (Pl. XXI, fig. 2, e), to the upper part of which is attached the trapezoidal ligament. The whole of the head of the bone is roughened for the reception of the ligaments binding the bone to the carpus and its fellow metacarpals. ‘The shaft presents a triangular section proximally, and is nearly circular in the middle and distally. At the poimt where it joins the distal articu- lation it is flattened in front, and develops a slight palmar ridge behind. The distal articulation of the four outer metacarpals (Pls. XIX, fig. 6 ; XX, fig. 7; XXI, figs. 2, 3, 4, 5, 7) bear a strong resemblance to each other; like those of the metatarsals they are bulb-shaped, and divided from the epiphysial line of the shaft by deep dorsal and lateral depressions (g); on the palmar or inferior surface they develop a short ridge in the median line (Pl. XX, fig. 7), which fits into the palmar notch of the first phalanges. On either side of it lies a sesamoid bone to which are fixed nearly all the tendons of the adductor and flexor muscles, for the movement of the metacarpals. At the point where the shaft joins the distal end is a tuberosity (Pl. XXI, figs. 2, 3, 4, 5,2) which catches the sides of the phalangeal articulations, and prevents these bones from bending backwards beyond an angle of about 60°, and thus forms a firm fulcrum for the support of the weight of the body. The distal terminations of the metacarpals may easily be distinguished from each other by the positions of the tuberosities, and by the form of the distal articulations. The distal articulation of the fifth is, as it were, cut off on the outer side (Pl. XXJ, fig. 5,/), while that of the second (fig. 2) is cut off on the inner side: the third and fourth are symmetrical, the former having the inner, and the latter the outer, tuberosity larger, and set lower on the bone. ‘These articulations are epiphysial. The proximal articulation of the third metacarpal (fig. 3) is set nearly at right angles to the end of the shaft, the dorsal face expanding considerably more than the posterior ; it is vertically convex, transversely concavo-convex. On its inner side is a broad oval surface (fig. 3, a) set on a tuberosity for articulation with the overhanging portion of the second metacarpal (fig. 2, 7); these two articular surfaces are divided from each other by a well-marked ridge (¢). On the external side are two concave surfaces (fig. 3, e), which overhang and articulate with the fourth metacarpal: they are not so deeply concave, and are more confluent than those of the second. ‘The proximal dorsal surface of the fourth (fig. 4) strongly resembles that of the third, but it is rounder and less excavated ; the _ articulation also is altogether different, the facet for the unciform (fig. 4d) bemg simply convex vertically and flat transversely, while that for the third metacarpal and the os magnum forms a continuous surface (fig. 4 a), partially interrupted on the inner side by a ligamentary notch (Pl. XX, fig. 6). It is slightly convex both vertically and transversely, and usually forms one curvilinear surface with the unciform articulation, bemg divided from it by a slight ridge. This form from Bleadon is shown in Pl. XX, fig. 6. In that figured in Pl. XXI, fig. 4, these surfaces (d, a) are set at a considerable angle to each other ; these two bones represent the extreme variation in the form of the proximal 136 PLEISTOCENE MAMMALIA. articulation. In the recent Felid also a similar amount of variation may be observed. The articulation for the fifth metacarpal (fig. 4 e) is much less concave than those of the second and third; the proximal edge is much thickened and flattened, the palmar is small. The form of the third and fourth metacarpals is much stouter than that of the others, and the arch-like curvature is less and the dorsal surface is more flattened. The fifth metacarpals of the Carnivora resemble each other so closely that it requires some attention to distinguish between forms of nearly the same size. In the Feles gene- rally, they have a tendency to be triangular in section, to be more taperig, and to arch more decidedly in a palmar and outward direction than any of the others. The proximal articulation for the unciform (Pl. XX, fig. 7a; XXJI, fig. 5 a) forms a continuous surface with that of the fourth metacarpal, and like it is convex only in a vertical direction; it covers the whole of the end of the bone. The inter-metacarpal articulation (Pls. X XI, fig. 56; XX, fig. 7 4) is a flattened surface, segmental in form, set at right angles to that for the unciform, and interrupted inferiorly by a large ligamentary notch (Pl. XX, fig. 7), in front of which rises an articular eminence (Pls. XX, fig. 7d; XXI. fig. 5d), which fits into a corresponding hollow in the fourth metacarpal. Externally the head presents a large tuberosity (Pl. XXI, fig. 5 e), which affords attachment to the strong ligaments that bind the bone to the unciform, cuneiform, and pisiform. On the palmar surface also there is a large tuberosity (Pl. XX, fig. 7/). Nearly all the ridges on the metacarpals are for the attachment of ligaments; the only direct muscular attachments being those for the very small muscles connected with the flexion of the digits. We have not attempted a more detailed description of these bones because of the great variation in the form of their articular surfaces, which renders it almost impossible to lay hold of characteristics common to a large series. The few points of difference that we have given are constant in all those of Fels spelea and Lion which we have examined. § 1. 8. AMeasurements.——In the following table of measurements we have given the extreme variations of size and proportion in Felis spelea as compared with corresponding bones of lion and tiger. The gigantic size of the second metacarpal from Crayford (Pl. XIX, fig. 6) is visible also in the metatarsal, fig. 7 of the same plate, and in an upper canine which was discovered after our plates were engraved. All these probably belonged to the same individual, and are far larger than any other remains of the animal that have yet been found. FELIS SPELAMA. 137 MEASUREMENTS OF METACARPALS. Felis spelea. Felis leo. F, tigris. 3 5 $ $ s s : 5 First METACARPAL. 2 & & g 3 2 - a w& I Ee | g = = rr) 3 3 Srconp METACARPAL. a5 & 3 & & 3 Ss 3 a g Ze 5 8 5 g = ea) = < = eae So} Bot Sb oe ears 5 = = — = = C= = Se 5 2 Q AQ pq a) = Ez a eS 1. Maximum length ...... 4:22 | 458) ... ae a 4:20 | 5:9 | 3°55 | 4:00 | 3°84 2. Minimum _ circumfer- EMCO eset eat omnis 2°20 | 2:25 | 2°24 | 2°33 ae 1:90 AGA Ni MR) 1 OE |) LORS} 3. Transverse measure- ment of proximal arti- GWETHOI, | Goeneesssoncnce 0:98 | 0°84 ee 1:08 | 1°15 | 0°81 0°8 | 0°72 | 0°78 | 0:80 BNerticall 25... careeeu seen lle 0) ae 1°31 | 1°45 ae 1:12 1-1 | 0°95 | 1:00 | 1:09 . Transverse measure- ment of distal articu- FACTO. wit taste 1:10 BCE AB AM ae 0°78 oO 1 GaaVierticalye no nae 1:95 chs ee ee ee 1°50 Oye 138 PLEISTOCENE MAMMALTA. Measurements or MetracarpaLts—continued. Felis spelea. Felis leo. F. tigris. o é $ E 5 Tuirp MpracaRPAL. = & g os 2 +=] = a g = < = ee al aa ie E ra Ea Es mM 1 Maximundilene thie teeseee eee eee eee eee ree 4:90 | 5:03 | 4:00 | 4°45 | 4:35 2. Minimum circumference ................2+00--.::0s 220 N50 e285) esl eeleoll 3. Transverse measurement of proximal articulation | 1°50 | 1°29 | 0-70 | 1:30 | 0°80 Al. Vierti@al) sci: hiseoiet bitoni oaths Sok Bees esau OR 1:20 | 1:44 | 0°80] 1:20) 0:91 5. Transverse measurement of distal articulation... | 1:25 | 1:11 | 0°85 | 0:90 | 0°80 Gi. Werticalld a0 Peek I oe VARee oe es ONO Se ene eo 2:10 | 2°30 | 1°30 | 1°60 | 0°39 Felis spelea. F. leo. ro) 2 2 2 2 FourrH MrtacaRPaL. = Ss 3 3 3 “a as 5 B § 8 eevee: ie sigh 4°40 2. Minimum circumference ..................02+00+2+ 2:08 | 2°12} 2°28 | 2:00 aoe 1°55 3. Transverse measurement of proximal articulation | 1:20 | 0°96 | 1:08 | 0°93 Le 0°85 AS Vertical (3.0.5 5 cn eon udescae eee oer eRe ace 1:40 | 1:96 | 2°02 ae if: 1:40 5. Transverse measurement of distal articulation... | 1:20 | 1:02 We An 1:05 | 0°80 Gis Vertieall. 3... FS Be eee 1°95 | 1:95 ie an 1:95 | 1°65 Felis spelea. Felis leo. F. tigris. Se eee ee : e i) . rs) Firra MEraAcaARPAL. S s & & a 2 wa 2 Bw gs 18 iS lt ey a 3 oO ea) (fa) fo a BS a (22) Q [2a) fa Fa fo He MER gente MeN 5oanancasen099 005209003 3°98 | 4°12 | 4°00 Be 3°21 | 3°70 | 3°24 2. Minimum circumference................-- TLetV} |) OKO) | Ne7/7/ || therday |) Se alfe} |) bess) |) Ihe) |] they) 3. Transverse measurement of proximal anticulacione ee eeeeee eee eeene 0:60 | 0°63 | 0°60 | 0°63 | 0°60 | 0:49 | 0°43 | 0°49 AveViertical? lca nuns tee eee oer re REE ee 1:80 | 0°90 | 1°77 | 1:75 | 2:13 | 1:30 | 1:30 | 1:49 5. Transverse measurement of distal arti- é r CUlAtHON a. bec nac cae EEL ne eee 1:12 | 0°85 | 0°74 | 0°87 ie 0°65 | 0°81 | 0°75 GamViertacal 7 200 ehhee SSeS At Coe bene 1°90 | 1°68 | 1:62 ahs 1°31 | 1:60 | 1°46 FELIS SPELAA. 139 § 2. Phalanges. a. First Phalanges.—The first phalange, or phalangiole, of the first digit, or thumb, is very short and of greater width than depth. The proximal articulation (Pl. XXI, fig. 6, a) is set obliquely to the dorsal face of the bone, so that the internal descends lower than the external edge. By this arrangement the first can be opposed to the other digits as in the Quadrumana, though the first metacarpal has no movement whatever of revolution. The palmar edge of the articulation is notched (fig. 6, 4) to receive the elevation on the same surface of the metacarpal, so that it gives but little hindrance to the action of the flexor muscles. On each side of this are two small eminences for the adductor and abductor medi, the flexor brevis pollicis not being differentiated in the genus Felis. The extensor longus is attached by an intermediate cartilage to a similar eminence on the dorsal surface, the tendon also passing on to the claw phalange. The distal arti- culation is terminal, and sits evenly on the bone; it is convex vertically, and slightly so transversely, so that a slight movement of rotation is allowed to the claw phalange ; it is much wider on the palmar than on the dorsal surface: immediately above the palmar edge is a deep depression for the reception of the flexor tuberosity of the claw phalange when the flexor muscles are in action. The small lateral expansion of this articulation prevents the retraction of the claw so completely in this digit as in the others, but as it is raised from the ground it is in no danger of being blunted by accidental contact; its position also on the inner side would prevent its being in the way of the animal. We have met with four specimens of this bone from Sandford Hill and Bleadon Caves ; they _vary in size from that of the ordmary Lion to that of the same proportions as the other large bones we have described. The first phalanges of the remaining digits (figs. 7, 8, 9, 10) are very similar in their general character; the shafts being nearly cylindrical and slightly arched, and much smaller than the wide proximal articulations (a); the latter are deeply notched on the palmar edge (4) to receive the palmar ridge of the metacarpal. The palmar surface also is furnished with small eminences for the insertion of abductor and adductor muscles and bear on either side two well-marked tuberosities to which are attached the ends of a strong ligament, the infra-phalangiole annular, through which, as through a pulley, pass the tendons of the flexor profundus and the flexor sublimis on their way to the claw phalange : the distal articulation is shaped like a pulley, and extends higher up on the palmar than on the dorsal surface, and bears a depression on the dorsal edge for the tuberosity of the second phalange. The first phalange of the third and fourth digits (figs. 8 and 9) resemble each other so closely that it would be impossible to determine to which of these two digits an isolated bone belonged; that of the second digit is very much the stouter, and slightly the shorter, and is curved outwards, while that of the fifth is very much more slender, and is bent considerably inwards. All are easily distinguished from those of the hind paw by their greater slenderness and by their cylindrical section. The large size of the proximal articulation and the tapering form of the bone m the Bear, and the small size of those in the 140 PLEISTOCENE MAMMALIA. Hyena, are points by which the first phalanges of those animals may be separated from those of Felis spelea. We know of no others that can be confounded with them. § 2 B. Second Phalanges.—The second phalanges of all the Feles are characterised by their triangular section, and by the outward projection of their distal articulation ; the latter allows the claw phalange when retracted to fall back outside the axis of the bone, so as to raise the point of the claw over the articulation, and thus protect it from injury. Those of Felis spelea (Pl. XXI, 11, 12, 18, 14) closely resemble their homologues of the hind limb, but are longer and more bent externally. The proximal articulation (a), which, following the section of the shaft, is triangular, is composed of two slight concavities divided by a median ridge; it is deeply excavated on the palmar ridge by a notch and pit, in which is inserted the tendon of the flexor sublimis muscle. On its dorsal edge is a small flattened tuberosity (4), forming the apex of the triangle for the attachment of the tendon of the extensor communis. The bone tapers gradually down to the distal end of the shaft. The distal end is a somewhat rectangular mass, of nearly double the width of the shaft and projecting outwards (d) from the axis of the shaft, and giving the whole bone a curved outline. ‘The articulation closely resembles that on the first phalange of the thumb, which performs similar functions. It affords a means of differentiating the digits of the same paw, that of the second phalange of the second digit forming an obtuse angle with the inner edge of the bone, that belonging to the third a nearly right angle, that belonging to the fourth being slightly acute, while that belonging to the fifth is more acute and very much shorter in its transverse diameter. Like the first phalanges these bones are much longer and more slender than those of the hind paw. We have been unable to detect any difference of form between the phalanges of Lion, Tiger, and Felis spelea. We have met with no British specimens of the third phalanges sufficiently perfect to describe, with the exception of Pl. V, fig. 14: they only differ from each other in size, those of the fore being in the main larger than those of the hind paw. § 2 y. Measurements——The following table of measurements shows that the supe- riority of size observable in the other bones of Felis spelea as compared with those of Tiger and Lion is carried out in every bone of the fore paw. FELIS SPELAA. MEASUREMENTS OF PHALANGES. 141 First PHALANGES. SEcoND PHALANGES. — Ne) Eee F. leo. \F. tigris. aoe F. leo. |F. tigris. mM a fo} Mm n B | = s 3 4 s E = 2 Fa = 2 FIRST DIGIT. i) a} a Ara Vcixcinna vanced). in’ ni eameen ee Pen sae. chinalccoc sem cdaciel oA |) ews |) lene: 2) Manian eimeunaterenceb eye. sseses secede -c+ s+ -seecesss sce 2°60 | 2:00 | 1°6 3. Transverse measurement of proximal articnlation ...... 0°92 | 0°80 | 0°7 AB BIVIET ECON 9s. ete aM ET eee tes fh ciciosaisloeiziae sie 0°70 | 0:55 | 0:4 od. Transverse measurement of distal articulation ......... 0:90 | 0:65 | 0°48 OR Ver tical), : oS eee eee ROR ee sie 1:00 | 0:90 | 0°8 SECOND DIGIT. | Ue AVE WOITA CRT PNET Yo. 2c tanccoscb6bp Cou SCORE COOL A SEO EE Eee 2°04 | 1:78 | 1:9 TOR2) || MOR ||. 11955 2 Minimum’ cmeumierence meeeeeneseeeeeeeceseeeeeee ccc. cee ZOOM mele OM ee lco 1°60 |} 1°50 | 1:2 3. Transverse measurement of proximal articulation ...... O:Gom ie Oi2n Ord 0°72 | 0°60 | 0°6 Al ViertiCal ethos MO eee EMEP elo oe Saiaececw ewe nds 0°60 | 0:50 | 0°5 0°50 | 0°50 | 0:4 5. Transverse measurement of distal articulation ......... 0:70 | 0°60 | 0:5 0°80 | 0°63 | 0°6 Gh Vertical... lose: cesee meee eee so ond peoO HE aeR EE pace aeee 0:72 | 0:80 | 0°8 0°63 | 0°60 | 0:7 THIRD DIGIT. | | tee Maximum: lengths. seperate cette acres se ace tacest sas 2°42) | 2:00) | 2-2 2°00 | 1:30 | 1°6 2. Minimum circumference ...............eeccccseececceceecees 2:00 | 1:80 an: 1°80 | 1°35 is 3. Transverse measurement of proximal articulation ...... 1:00 |} 0°85 | 0°88 | 0°82 | 0°64 | 0°65 Bilis NGA (CG WRB ee rele tele oe Sn 0:70 | 0°55 | 0°45 | 0°70 | 0°45 | 0:4 5. Transverse measurement of distal articulation ......... 0:70 | 0°65 | 0°55 | 0°80 | 0°54 | 0°55 Gs. NGA ICE Ra Meio senbind nde cuérco neo coe heeecr eee tee eee 0:80 | 0:90 | 0°8 0:70 | 0:50) 0°8 = © & Lo} 8 FourtH DiGi. Me iy diidimiantaten LEE oaaeenceee sbocedebcceavebectocoo 7 One eeReEe 9°20 | 1°85 | 2°05 | 1:75 | 1°30) 1°5 Oe Minwnumbcircumberencel ses -24. 2 -eeeeeeeteceererie ace enece: 2:00 | 1:74 | 1°6 7D) || eR |} Toe 3. Transverse measurement of proximal articulation ...... 0°98 | 0°85 | 0°75 | 0°84 | 0°60 | 0°75 Zh, AU GEION CE laced 2B ao beets oir ek SRR NES Se Fh te 0°69 | 0:45 | 0-4 0°61 0°45 | 0°35 5. Transverse measurement of distal articulation ......... 0:70 | 0°60 | 0°43 | 0:70 | 0°50 | 0:5 (i, “WEIGEL 3 none seen coton cp ooResroe cee Sone REPRE Cee ne Seen EEer ee 0:70 | 0°65 | 0°6 0°55 | 0°50 | 0:5 142 PLEISTOCENE MAMMALIA. MEASUREMENTS OF PHALANGES—continued. First PHALANGEs. SEconD PHALANGES. Felis ...| Felis ae spelza. F, leo. \F. tigris. spelen. F. leo. \F. tigris. = 5 z 5 4 S = < = FIrTH DIGIT. fF e a ca = 2 a Fay Fa} 1.)¢Maximumilengethitsyb 2.4.5. seee ae eeeeere sean ene eer ee U97 90375) 1565) 1570) tO R aes 2 SMinimumecircumterence -s-eeee see ee eee ee te eeeceeeeceeee: 160 | 1°50] 1:4 1:70) |) Vs255\eale2 3. Transverse measurement of proximal articulation ...... 0°75 | 0°75 | 0°6 0:70 | 0°60 | 06 A. Nertical:: ph Qi enu hts Meck ee ee eee 0°55 | 0°50 | 0°42 | 0°55 | 0:40 | 0°5 5. Transverse measurement of distal articulation ......... | 0°60 | 0°60 | 0°5 0°65 | 0°50} 05 6. -Vertical 5.20 5nt ee Cadac Se ae Seiad Mies Rec nee eae 0:70 | 0°65 | 0:5 0°55 | 0:50 | 05 FELIS SPELAA. 143 CHAPTER XV. Lims Bonrs oF tHe Wuetp. Pl. XXII. CONTENTS. § 1. Description. | § 2. Measurements. § 1. Description—M. Gervais, in his ‘ Zoologie et Paléontologie Frangaise,” states that M. de Serres and his coadjutors in the work’ on the fossil mammals of Lunel Viel have figured and described bones as leonine, which he considers to belong to young . Felis spelea, “ before it had lost its milk teeth,’ and that Dr. Schmerling had indicated ~ the presence of lion in the Belgian caves “without giving more certain proof of its existence.” ‘The bones from Lunel Viel (exclusive of the head) consist of the sacrum, the proximal half of a femur, both of which are figured, and an ulna that: is only described ; while those from Belgium consist of a nearly entire pelvis, and a perfect radius and ulna. All these, however, seem to be of by no means so young an age as M. Gervais supposed. In the figures given by M. de Serres’ and Dr. Schmerling,® there is no trace of lines of separation between the sacral vertebree ; while in a third sacral that passed through our hands, and is now in the Taunton Museum, and which had belonged to an animal very much larger than the average-sized Lion, the anchylosis was not yet completed. It is clear, therefore, that the former must have belonged to older animals than the latter. Tn our figures also of the radius and tibia (Pls. I, fig. 1, XIX, 1, 1’), the anchylosis is imperfect, so that the proximal epiphysis is lost in each case, although the full size and proportions have been reached. The limb bones from Belgium and Lunel Viel present epiphyses firmly anchylosed to the shaft, and strongly marked muscular ridges, and they therefore belonged to not merely full grown but to tolerably aged animals. In Britain we have met with several bones of the spelazcan whelp from the caves of 1 Edition 1859, p. 227, ’ 2 «Oss. Foss. de Lunel Viel,’ pl. viii, figs. 15, 16, p. 107. 3 «Oss. Foss. de Lidge,’ tom. ii, pl. xix, figs. 1, 2, p. 90. 144, PLEISTOCENE MAMMALIA. Bleadon, Hutton, and Sandford Hill. An ulna from the latter cave must from its extreme shortness have belonged to a very young animal. In Mr. Beard’s collection, now in the Taunton Museum, are two right fibulz, the larger of which we figure (Pl. XXII, fig. 9), with some doubt as to its correct determination. Its ridges, which are slightly developed, and the distal lateral articulations for the tibia, are of the same form as in the older Lion. So far as we can tell, from the limited means at our disposal, it is of the size of a Lion’s whelp six weeks old. In the same collection there are both ends of a left radius which correspond exactly with those of a young Lion of three months. They most probably were obtained in Hutton Cave, which has furnished two young humeri, one of which is figured (Pl. XXII, fig. 1), a large portion of a right scapula, both ulne (one figured, figs. 2, 3), both femora (figs. 7, 8), a left tibia, caleaneum (fig. 10), a fifth metacarpal (fig. 4), and some first and second phalanges (figs. 5, 6), including a part of the first digit of the fore paw. All these bones in size and proportions agree with those of a Lion’s whelp of three months. From Hutton also were obtained the maxillary and lower jaw, which have already been figured (Pl. XIII, figs. 1, 3), and described. The whole of them most probably belonged to one individual. The calcaneum and some phalanges of an older animal from Bleadon are also preserved in the Taunton Museum. All these bones coincide with those of the adult which we have figured and described in the course of this Mono- graph. They are, however, much more cylindrical, and present fainter nuscular im- pressions, and are considerably shorter. § 2. Measurements.—The followmg measurements are all that the condition of the bones would admit of being taken. Humerus: minimum circumference, 3°05; do., 3°04. Ulna: minimum circumference, 2°20; length of humeral articulation, 2:00; length of radial articulation, 1:15; depth of radial articulation, 0'40. A second specimen agrees with the preceding while in a third the minimum circumference is 2°20, and the length of the humeral articulation, 1:70. Radius: minimum circumference, 1°40. Femur: minimum . circumference, 2°78; do., 277. Tibia: minimum circumference, 2°60. Fibula: minimum circumference, 0°50; do., 0°60. FELIS SPELAA. 145 Table showing the variations in the size of calcanea of Felis spelea, from youth to old age. Adult Pl. IV, fig. 2. Tem Lotalelens ie emma ete ee aac/c che cece) cerinceas adeinedeseceencs owe 5°60 2. Minimum circumference.......... g coodbn Bao hed das CEHRC ODER on Rocce ana ee 4°73 oy Maximumbpverticalemeasumementy ee ec nese. ccececesecseese se «ceccsseace= 2°36 4, a transverse ERT oa see. Sod Da AGED CAE RG URCSO EA 2°38 5. From inner articulation to outer end of bone, articulation included 3°95 6. Transverse measurement of sigmoidal articulation ..................+6 0°92 7. Transverse measurement of cuboidal articulation.......................- 1°32 Gh, WGA GIHI@ “soc nbndcoocc ote dos odo ONCE DOC DE ORE CO EERE SEC REE EEE ea ee anee 117 9. Transverse measurement of inner astragaline articulation ............ 0°81 LD) AV GEG ION GWG) eee atic ode coon poe br abode SOUR DEDE CEE et oe eR CE OEE EEE eee EEE REE 0-74 Young. Pl. XXII, fig. 10. 3°00 1-40 1°50 0:50 1:00 1-00 0°60 0:60 Young. 1-08 068 0:36 0°50 0:50 The minimum circumference of the phalange of the first digit is 1:85; of the rest, 1:10, 1:42, 1°30, 1-46; and of the second phalange, 1:29. 146 PLEISTOCENE MAMMALIA. CHAPTER XVI. FELis sPELZA, GoLDFUSS, SPECIFICALLY IDENTICAL WITH Fruis Luo, Linnavs. CONTENTS. § 1. Introduction. | § 2. Various opinions held by naturalists. § 3. Conclusion. § 1. Introduction —In the preceding chapters we have analysed the differences observable in skeletons of Lion, Tiger, and Felis spelea, not founding our comparison on one skeleton merely of either of the former animals, but comparing and noting the variations in the form and proportion of the bones of all the individuals preserved in the museums of London and Oxford. One fertile source of error in the work of previous observers has been avoided—the use of the bones of animals kept in menageries, which are invariably affected in direct proportion to the length of the confinement of their possessors, and to the extent to which the natural habits have been restrained and curbed by domestication. They are so deformed, and, if the cub has been born in captivity, so small and puny, that they are absolutely useless as a means of comparison. Before we proceed to sum up the bearing of the evidence on the recent affinities of Fels spelea, we intend to quote the opinions of the naturalists in chronological order, following to a certain extent the method of M. de Blamville and Baron Cuvier. § 2. Various opinions held by naturalists—YVhe first evidence of the discovery of Felis spelea is afforded by a figure* of an unequal phalange, appended to a paper on the Dragons of the Carpathians, written by Dr. John Hain in 1672. It is most important, because it brings the range of the fossil animal into the Hungarian Basin of the Danube. Leibnitz, n 1749,’ figured a fragment of skull obtained from the cavern of Schartzfeldt. The plate contains four rudely executed figures, of which the upper may represent the * Recognised by Cuvier (‘ Oss. Foss.,’ t. iv, p. 449, 2nd edit., 1822), and ascribed by him to Dr. Vollgnad (‘Ephém. Nat. Cur.,’ an. iv, dec. 1, obs. clxx, p. 227). The latter, however, merely gives an outline of a paper published in the preceding year by Dr. Hain (‘ Miscell. Nat. Cur. Medico-Physic. Germ.,’ An. III, Obs. CXXXIX. “De Draconibus Carpathicis”’). 2 2 © Protogeea,’ pl. xi, fig, 1, p. 62. FELIS SPELAVA. 147 parietals and part of the occipitals of a Lion, while those underneath may be the fore part of the upper and lower jaws, of the same animal; but it is very possible that the originals may have belonged to the Bear. The whole are referred to in the text as “vera elephan- tium ossa,” the upper part of the head being taken for the “ tibia” of the Elephant. The fragment of skull is compared by Soemmerring* with the skulls of Lion and Ursus speleus. He considers that it differs in no respect from the former animal; but he adds that in most of the points relied upon it resembles other species of the genus Felis. He gives a more exact figure than that of Leibnitz. In 1774 Esper’ published an account of the mammals found in the Margraviat of Bareith, in which he figures an upper jaw from Gailenreuth. He obtained also detached teeth and bones. He believes them to belong to an unknown animal, more closely allied to the Lion than any other species. Rosenmiiller,? in 1804, states that he is about to publish a work on an unknown fossil animal of the genus Felis, and he adds that its bones differ in some respects from the Lion. Dr. Goldfuss* published, in 1810, a small work on the environs of Muggendorf, in which a nearly perfect skull from the cave of Gailen- reuth was figured and described under the name of Felis spelea, which was adopted by Cuvier, and became the recognised specific name of the animal. In 1821° he republished his determination of the species, and gave a full-sized figure of the skull, which he considered to belong to an extinct species, more closely allied to the Panther than to Lion or Tiger. Drs. Pander and D’Alton® state, in 1822, that Felis spelea differs specifically from i ‘ Felis leo, and refer to their figures in support of this conclusion. The figures, which are | those of a skull and lower jaw, exhibit no sutures. The second premolar of the upper { jaw is bifanged, as in the skull from Sandford Hill Cave (PI. XI, fig. 1). There are no . measurements of the skull given in the text, nor is any information afforded as to the 4 museum in which it is preserved. s Baron Cuvier, in the second edition of the ‘Ossemens Fossiles,” published in 1823, does not pronounce a decided opinion on the relation of Felis spelea to the large existing ' members of the genus, because he was unable to make a personal inspection of the type 2 specimens described by Dr. Goldfuss; but he states his belief that the real affinities of the animal are neither with the Lion nor the Tiger, but with the Jaguar (Felis onca), giving as his principal reasons the gentle curve of the profile and the form of the lower jaw.° * “Magasin pour I ‘Histoire Nat. de ’Homme’ de M. C. Grosse,’ t. iii, cah. 1, No. 3, p. 60; Cuvier, op. cit. We cannot verify this reference. 2 «Description des Zoolithes, &c., dans la Margraviat de Bareith,’ folio, Nuremburg,’ 1774, tab. TXSUXIITS p- 53. ° «Abbildungen und Beschreibung der fossilen Knochen des Héhlenbiren,’ folio; fig. 1, pp. 11, 19; Weimar, 1804. * “Die Umgebungen von Muggendorf, Erlangen, 1810. 5 * Nova Acta Physico-Medica Acad. Ces.-Leop. Cur.,’ tom. x, p. 489, tab. 45, 1821. 6 ‘Die Skelete der Raubthiere,’ tab. viii, a, b, c, d, 1822. 7 Tom. iv, pp. 451—455. * See ‘ Felis spelea,’ cap. i; cap. vi, § 20. 148 PLEISTOCENE MAMMALIA. Our great cave-explorer, Dr. Buckland,’ in 1823, was the first to ascribe the spelean remains to the fossil Tiger, without, however, giving any reasons for his conclusion. His rival, Dr. Schmerling, in 1833, in his réswmeé of the species of Felis in the caverns of Liége,’ considers that Felis syelea, was allied to the Lion, but of a distinct species. He figures, however, bones from the same locality as belonging to the existing Lion; but confuses them with those of the Melis antiqua of Cuvier, which was not a Lion, but a Panther (F. Pardus). MM. Marcel de Serres, Dubreuil, and Jeanjean,* writing in 1839, insist on the specific distinctness of Felis spelea from the recent Lion, assigning as the principal difference the shortness of the muzzle. Like Dr. Schmerling, they identify a second species with the latter animal. M. de Blainville, in 1841,* rejects the view advanced by Marcel de Serres and Dr. Schmerling, that the smaller bones ascribed to the Lion belong to a species differing from Felis spelea, on the ground that they were probably not those of an adult. He, however, offers no opinion on the exact affinities of Felis spelea. M. Pictet,’ in 1844, uses nearly the same words as Marcel de Serres and his fellow- workers in his notice on Felis spelea. We does not recognise the smaller remains as those of Melis Leo. M. Gervais,® in the first edition of his ‘ Paléontologie, published in 1848, regards the animal as a Lion (Felis leo major), without assigning any reasons for his conclusion. Professor Owen,’ 1842, adopted Dr. Buckland’s opinion, and terms the animal a “spelzean Tiger,” although he recognises the want of evidence sufficient to put the question of its species beyond dispute. He reproduced his views in 1846, in the ‘ British Fossil Mammals.’ In 1859, however, he published, in the ‘ Philosophical Transactions,” a figure of a skull with the nasal processes restored as in the Lion. It is clear, therefore, that he recognises the leonine nature of the animal, for his figure shows that characteristic which is of specific value in determining Lion from Tiger. Dr. Falconer is quoted by the eminent French paleontologist M. Lartet,” in 1864, as holding the view that Felis spelea was identical in species with the Tiger inhabiting the north of China and the region of the Altai, and that it was driven out of Hurope “par le développement progressif des sociétes humaines.” In 1858" he enumerated “Cave Lion” among the remains from Kent’s Hole. ‘Reliquize Diluviane,’ p. 261, 1823. 2 «Oss. Foss. de Liége,’ tom. ii, p. 93, 1833. “Oss. Foss. de Lunel-Viel,’ p. 101, 1839. 4 All these remains are preserved in the York Museum. 4 In the collection of R. Fitch, Esq., F.G.S., of Norwich. > “Brit. Foss. Mam.,’ p. 152. We have been unable to find out where these remains are preserved. 6 Oxford Museum. 7 Now in the British Museum. ET Bene ata Se ye ee ke ee eee oe 7 é ; —— ‘ FELIS SPELAA. 153 the Post-glacial brickearth of Otterham, near Sittingbourne, has furnished upper pre- molar 3 and a large upper sectorial to Mr. Hughes, F.G.S.; and a similar deposit near Hartlip, in the same neighbourhood, a femur to Mr. Bland.’ A very careful search throughout South Kent and the whole of Sussex has not revealed a trace of the former existence of the Lion in the heart of the dense Wealden Forest, that from the nature of the ground must have overshadowed those districts during the Post-glacial epoch. In going westward we meet with the animal again in the low-level river-deposits of Fisherton, m a lower jaw, found by Dr. Blackmore, and now in the Salisbury Museum. The low- level gravels also of Loxbrook, in the valley of the Avon, near Bath, have furnished a remarkably fine humerus (Pl. XVIII, fig. 2) to the energy of the Rev. H. H. Winweod, F.G.S. In the collection of the Right Hon. Earl of Enniskillen, at Florence Court, is a canine from the cave on Durdham Down, near Bristol, explored by Mr. Stutchbury. Remains of the leptorhine Rhinoceros of Owen and Lippopotamus major from the same cave are preserved in the Bristol Museum. But the district that, of all others, has furnished the most enormous quantity of the remains of the Cave Lion, and is entitled, therefore, to rank as its metropolis in Britain, is the western half of the Mendip range of hills in Somerset. Throughout the area extending from the ancient city of Wells westward to the new watering-place of Weston- super-Mare the Mountain-limestone is traversed by numerous caves that have afforded most valuable evidence as to the character of the ancient Post-glacial Fauna in the west of England to the Rev. D. Wilhams, Mr. Beard, and ourselves. Among the animals the Cave Lion stands out the most prominently. We found in Wookeyhole Hyzena-den twelve teeth (PI. XL fig. 9; Pl. XII, figs. 9, 18, 15), an ulna (PI. II, fig. 9), astragalus, and metacarpal (Pl. XX, fig. 7). Dr. Boyd also obtained a magnificent upper canine (Pl, XI, fig. 6).” All the remains were more or less gnawed, and bore indisputable traces of the animals to which they belonged haying fallen a prey to the Hyeenas. The Feline remains from Bleadon, Sandford . Hill, and Hutton Caves, explored by Messrs. Williams and Beard, are preserved in the Taunton Museum, where they con- stitute, perhaps, the most magnificent series in the world. They are as follows, in various conditions, some being perfect, others fragmentary. 1 Both these are in the Museum of the Geological Society of London. 2 In the Oxford and Taunton Museums, and in the collections of Mr. James Parker, Mr. Sanford, and Mr. Willett. PLEISTOCENE MAMMALIA. Bleadon Cave. Lo4 ADULT. Numbers. Miaxillaries > (ecu. jcassene act cement ata rpee 10 Squamosalli (Pls DXeitigss) 2593) eeer eee ete 1 Toower jaws? Wh. SiC ve. Pek c De eerie Beene 12 Vertebree (Pl. XIV, fig. 3; and Pl. XVI, figs. A 25, D105 75185 29 UO) lee meas cecen ets 50 Sterne bers (b)- OxCVilepicsn 0) eee ee see eeeeeee 5 Neapulae.. chev.cesepat cers seoteeeac cacti eeneecmes 1 Humenn (2 xavilh shea) peesecceeee rere eee 8 Ulnge GRIT, feti5)) pees. Bae ee ne ee ee , 14 RadiiCPl UE ig. A) ich eee act saeenes eee 8 Carpals (Cal XOX s fies 32573) meeeeners cee ee eer 8 Metacarpals (Pl. XX, fig. 6; and Pl. XXI, HSL Vo cos Seat PA alee eee Rene 61 Phalanges! ir gues u.tscces scree cece ee cere ema 38 1 AULT Cine Dat sAC HBB aA SEMA BE sos orc tin sos860 256 1 Remora, (BlSeOVill ly ie 4) eee ener eee 6 Tibides 228 veel cueinaceeits eae eee eee eer iene 9 ADULT. Numbers. Eibula (BIS XTX, fies), Caen cee ae aae ene 1 Marsalsy(QP 1, TV) |. i aie. e oe ee Aer een 39 Mefatarsals\(BI. iV, ital) hen eee eee 18 Phalanges,., .s,.2 4. ..jsa-\saodtenne sees Geka e eee eee 59 esamoids: . ¥..\ on2hi. sade -n ie. Renee eRe 2 Permanent teeth (Pl. XI, figs. 4, 8, 10, 11, 12, 13, 14; and Pl. XII, figs. 4, 6, 7, 8, LO. TO 14) a ee 95 WHELP. WY ersilllsmenetsy, (eal QUES area A eae ee 2 ower AWS Gee ror oc aan ne nee. Pome eee ee ee 8 IMetacanpal Ty) ie vieteenedreae te sate ee eee es 1 Phalanges) (Bl, XX fies 6) (ae ees penees 2 Calcanewme i) 502hite ks. Be I Bibulee (BEX TT, fig. 39). ee eee 2 Malk-teethy (Be) Xun siccrS)) ieeeae eee eee 12 Sandford Hill Cave. The following bones probably belong to one adult individual : Skull (Pl. X, fig. 1). Lower jaws (Pl. I, figs. 1, 2). Atlas (Pl. XIV, fig. 1). Sixth cervical vertebra (Pl. XIV, fig. 2). Second dorsal vertebra (Pl. XV). Second lumbar vertebra (Pl. XVI, fig. 3). Third sacral vertebra. Both scapulz (Pl. XVII). Left humerus. Both ulnze (Pl. II, fig. 8). Both radii (Pl. II, fig. 1). Right scaphoido-lunare (Pl. XX, fig. 1). Left pisiform (PI. XX, fig. 4). Metacarpals 2, 3, 4, 5, of right paw (Pl. XXI, figs. 2, 3, 4, 5). Metacarpals 2, 3, 4, of left paw. First phalanges of right paw (PI. XXI, figs. 6—10). Ossa innominata. Patella (Pl. XIX, fig. 5). Both tibize (Pl. XIX, figs. 1, 2). Right fibula (Pl. XTX, fig. 3). Both astragali. Scaphoid. Both calcanea. Left cuneiform. Metatarsals 2, 4, 5, of right paw (PI. V, figs. 2, 4,5). Metatarsals 3 (Pl. V, fig. 3), 4, of left paw. First phalange of the fifth digit of left paw. Oruzr ADULT Bonts. Numbers. Maxillaries and intermaxillaries (PI. XI, fig. 1). Lower jaws corresponding with the above. Wertebras,).2 3c. siz. igna6- ee eeeeee oeeeeeee 6 PPM CT oe Scat oa cine « leele cee eRe eee 2 Ulnz corresponding with humeri ............ 2 Carpal. ...4 325i. eee: one ee 1 Os pubis 4.4.25 62. eee ec eee 1 Marsal’ 00.052, Clause) eee nee nes ce ean ee ane 1 Permanent teeth (Pl xl) ican) eee eens RuEMaIns of WHELP. Maxillary bones .es.-0-nese eee eee eee 3 Wowersjaws (Pil. SUL iiiote4)) eee ene neerterens 2 Malk-tceth (Pl. SOUL sties15.sO507))earenpeeeres 4 FELIS SPELAA. 155 Hutton Cave. Bones oF WRHELP. Numbers. | Numbers. Maxillany (PI SX fest etna ctecectes tos nce 1 Silermeberets seco sa scaacessriaeayadeen west sere Lower jaws, a pair of (PJ. XIII, fig. 3) ...... 2 A pair of femora (Pl. XXII, figs. 7, 8) ...... 2 CAP UA see eM te Rem eaeee eee eielaeielec tise sien 1 AEUDAYS ESSE ccesgehe stole cnc he eee Mer ae aes ete 1 A pair of humeri (PI. XXII, fig. 1)............ 2 Calcaneum (PI. XXII, fig. 10). A pair of ulne (Pl. XXII, fig. 2)............... 2 ‘The skull and lower jaws, Pls. VI, VII, VIII, IX, were obtained either from Sandford Hill or Hutton Caves. | The accumulation of so enormous a quantity of the remains of the Lion in the caves of so small an area may be accounted for by the peculiar position of the Mendip Hills, that command fertile valleys on the north, and look out towards the south and west over a plain which, in Post-glacial times, occupied a large portion of the Bristol Channel. Around them were the feeding-grounds of incalculable numbers of the Reindeer, Bison, and Horse, of the Mammoth and tichorhine Rhinoceros, and, therefore, we might expect to find the carnivora present in very great abundance. ‘There is evidence, indeed, that a larger number, not only of Lions, but also of Bears and Hyzenas, existed in the district than have yet been proved to have lived in a similar area at any time in the past history of the earth. To the south of this district no leonine remains have been discovered as far as the outcrop of the Devonian Limestones on the shores of Torquay and Plymouth. Inthe Brixham Cave two phalanges’ were found along with flint flakes and the remains of the Hyzena, Bear, and other animals; in that of Kent’s Hole, explored by the Rev. J. McHnery, an upper jaw, four teeth, and an ulna;’ and in that of Oreston, near Plymouth, explored by Mr. Whidby, three canines, one humerus (Pl. X VIII, fig. 1),one metacarpal, and two metatarsals.* Nor were they less rare on the opposite side of the Bristol Channel in South Wales. The researches of Col. Wood and Dr. Falconer have resulted only in the discovery of an upper jaw and five teeth (Pl. XJ, figs. 1, 2, 5) mm the Cave of Ravenscliff, three canines and a fragment of skull in that of North Hill Tor,* and a few fragmentary remains from those of Spritsail Tor and Long Hole;> from a cave on Caldy Island also a carnassial has been obtained by the Rev. F. Smith. In North Wales, a cave at Cefn, in Denbighshire, is quoted by Dr. Falconer as containing the remains of Felis spelea, but we have been unable to submit them to a personal examination. These are all the cases of the occurrence of the animal in Great Britain revealed by a careful search in every public and private museum and collection of note in the kingdom. The absence, therefore, of the animal from certain districts cannot be accounted for on 1 Tn the possession of the Royal Society. 2 In the British Museum, that of the College of Surgeons, and of the Geological Society, and in the possession of the Earl of Enniskillen. ° In the British Museum, that of Leeds, and that of the Geological Survey of England. * « Paleontological Memoirs of the late Dr. Falconer,’ vol. ii, p. 458. Bl Opa eit., pl a2. 156 PLEISTOCENE MAMMALIA. the supposition that the animal remains have not been examined; and consequently the range of the animal through Britain, so far as extant evidence goes, is fairly represented, although, of course, it may be modified from time to time by future discoveries. Its metropolis was West Somerset, where it was incredibly numerous; thence it ranged throughout England as far as the North Riding of Yorkshire, being very rare in propor- tion to the other animals living at the time. Its absence from Scotland, Cumberland, and Westmoreland, and its extreme rarity in North Wales, may be accounted for by the fact that the mountains in those districts were crowned by glaciers during the Post-glacial epoch, which would necessarily involve a climate unfitted for the great development of the Herbivora in regions much broken up into hill and valley, and the consequent absence of the Carnivores. In Scotland, at least, there is no other hypothesis that will account for the absence of every animal that can be ascribed to the Post-glacial group, excepting the Mammoth, which has been found in a few places, and which has been proved by the Siberian discoveries to have been capable of existing in the zone of vegetation represented by the Scotch Fir. If it be objected to this view that the Reindeer flourished in count- less herds m a Siberian and North-American climate at least as severe as that of the Post-glacial winter in Britain, it may be answered that in Siberia and North America, where animal life is so abundant, the country consists of plains elevated but little above the sea-level, and capable of affording good pasturage in the short arctic summer, while in Scotland, Wales, Westmoreland, and Cumberland, the broken nature of the ground could not ever have admitted of the growth of feed for a large body of Herbivores. The animal also has not been found in Ireland, most probably because only one of the numerous caves of that country has been properly explored, the energies of collectors being directed towards the acquisition of prehistoric remains from the turbaries and alluvia. § 2. Mammals associated with the Cave Lion in Britain.—In the following table we have given a list of all the fossil animals associated with the Cave Lion im the bone-caves and river-deposits of Great Britain. ‘The varieties Cervus Bucklandi of Professor Owen and Cervus Guettardi of Baron Cuvier are included under the general specific name of Cervus tarandus ; and Strongyloceros speleus of the former under Cervus elaphus. Hquus fossilis is also intended to include Hguus asinus, which, up to the present time, has not been proved to have lived in North-Western Europe during the Post-glacial epoch. Hlephas antiquus also is intended to include Hephas priscus, a name which the author of the species, Dr. Falconer, gave up during the last years of his life. The Rhinoceros lepto- rhinus of Owen is used as the exact equivalent of the Rhinoceros hemitechus of Dr. Falconer, and of what M. Lartet’ takes to be represented by the R. Merchii of Dr. Kaup. With the exception of the lists of animals from Long Hole, Northhill Tor, Spritsail Tor, and Cefn Caves, for which we are indebted to Dr. Falconer, all the species have been determined by a personal examination of the remains. 1 «Paleontological Memoirs,’ 8vo, vol. ii, pp. 251, 592, 1868. 2 «Ann. des Se. Nat.,’ 5° sér. Zool. et Paléont., tom. viii, p. 157, et seq. FELIS SPELAA. 157 SPECIES ASSOCIATED WITH Felis leo (var. spelea) IN BRITAIN. Perret NG EAT Ghaas Post-GLAcIAL |Rrv.Dz- rs River Deposits. | POSsITs. cB 2 il = Ss : é 3 } B E S k é z A ce al |s SSI ol eB £ =|./s = Z| |g Blea ial (ole tel) eisic 5 ye O/P ial |s| |Sl=l.ls Soil eal Pea ae 16 ; s/8\= alo/= Sel slElelmplslslsialelelelels S|/s|o ale} a] =| 8 SIS/SI/SPeElSiLlSlel2lelolsl—- Sl(Slelelol/Sial els] nels lsis SslolsleclSiSlelel|s SI\KIAZISIS/SISIZIZ/SISISIEISIElSlelslEleleleiels SPelalSl|SIS/ofElelslS/SlielllalSleis|/2lolSps izle : AIAIO/OIAIS IMIS IS alolalal= [a lala S/S alels lola JE OVE) A Wee aaatcec ste decosc’ scnapbekce seen sai OS |e gel Beales Slat dee Pen ete ee 5c! ESA deel teleediecellaaelle.| Bscllodellone Rhinolophus ferrum-equinum, Leach... |...|...|...]... 5o|lse4| 36 beellane Bee Get ens ioe al tac aeliscel leslie. Seiad onl eral lab Vespertilio noctula, Schreb............. |--.|...|---|..« “pel bel eoel ee) ee Bal Be (SGe geal eed Geto ee\ baal BoB) lope Pelleod bepllosallaad (ORES GREGOR, MN, wenconteondsensasponseee lol lledl nes Se loncll SS llooall SS tlawall SO) / SX SKUSAI brcallecclloccllecellonallscolleo-| bos x|x Ubisnelceus; Golds Ve see ea ease XE 113< lovel] SMcall S< SM Hesell SS SXWSMWS< [etal &-¢| eel an fea eee baal Be eee BG Beeld Monies veal ersal bel bse x|x Gulovluseus: Habeas ee eee eae: Saleeelieebilece See yam ean ee eee (ee Reale pent Reel laee| bce aeal eal eclli sd Lcalleotlean IWGIGS AERIS LURE Seabed} oleic onttvendionet td eel lari [ee oll aes ea ee XM lecallowe Sle pallecell 94 (D4! Keallosollone tal Sasa ae ne a | WW OSAAID GHIDIW EL Wh arrandeabacdacaoonen |Peallve- 2 Al ecall sat Oecd S< PaaS Me ees Fea eR aC) feed ee oral ea) Lae TN a i all OY Fy ALLO VON sabia icoe ie Beene cee || BEA ae oe cllédalleeel Mel ete eee (eae ST cc ol OS (hire [ete elle a ltgeea( ase | es | MMR mantess Did ss Na ee ees ae S|: cal aed locel easels aal easel eal eee BPs ALL eed ee] NEI RUN eal laa) eval cy ba) Lally Ah IVA UGE, WP oe soa cene sh see ene X€|lsccflecofleon|l2% leoul/S< eee] lace bi eal el Pe ena! Bisel ecsell awe cecal elle elloe ul asad ial 2 Canis:oulpes; Ws ihc 5tn ee eee ee XWSSHEXS osell call |X CRUPUSH LAA hea oes ae SE td SSX eo Since ec lane ess Hyena spelen, Gold, areca: x| x x x Melisicatus, Ws | a eeeaes- cee eee ae 54 feel anal ee =2olacellead|/sac||obo Sa|[ece||ecallacd bac|looa|looa}face|[coo||aoe||a- F. antiqua, Cuv. (=F. pardus) ...... SGI alps 2 ee esdl DA le aoe ee CRA Nal ed [eal ee Boel ed ae end (eel lal 2 3 f ir spetcea: Gold... See shes ase acn ee X|X|X]X|X1x| x] x] x Lx] xT xT x] KY KY] | TX TX | KEKE Xx | XX Machairodus latidens, Owen ......... |...|... Soll tel bec eel le toe eee Pel oel Goalite allgeet Heallsaellgeell-teclias PAPA deat lo alee Megaceros Hibernicus, Owen ......... KX local] 34 lke of] S|] 4 loool OMI SS llecellonall <1 <| [DN |lecclloellanc|! <1] 5< 158] 124] 8 [5 Cervus Brownti, Dawkins ............ |-..J...]---[... 22) eel Atel Eee mh orl lee Pela i eee ee ee BP Sliced oa OStajandus! Wen. ao eae ee XP 9K |] SSK doll SCMNSK leoall SMHDS |< llecell SS 1/25) (2 Tecoll SS iloaallon: Salles allel a Cheapreolusyr lis 3. seep eee eee SGIEX |e eee ecallenal see See lat al Peel adel ee Oe Pedte-slocatloed lent onl| @Melaphus,) Ws iiscaqreeeeea eee 3€ <1 26] llecellcaa|| SX lee MME To ealloce SSCS SM loc cllocellaccllancl| Ovibos moschatus, Desm. ............... }...|...|-..}... sel ee ae is |e |e lh | ge SASS bel (oct Peralepe teal 2 Bos primigentus, Boj.......-....-00c..++ IS eee Aloo |e Bison priscus, Owen .......2.-06cees..42- 5 vel hal eee Re S21) lee Hippopotamus major, Desm. ......... |...|... zocl|S< lLoellodel |esaltaael eu eel eee] eel eee See ase ete eos eel ia Xia Ee Lepus cuniculus, Pall. .................+ DAIS] cee sodlenalh herd oct bari Cea Pea ear (ope (Se Ra ae) Peel eee ee Zh, COGS, UNSW ongeicadecacceacoacopnndo: pe ee ae Se eN ealtoslEN Sal eal ee aeoaltcmeal eine Wet Vbetligrend al ee I oat dl |e Lib CSA CACO: ADI) ie PG ee ee rales Ibi bel) 2S)lou || 2M leb a ceel[2Sliancl Meeligeldled\onetoacliballend sae x|X Wasimuseuluss, Weep. sss scss ss 2se+ 5s: pal sees |B eral Albae Fae |sere| le Ae) Belles ree) leesel less fessor eee laced rel (Mea teem Cision flein Wns aires aecee stereos ac [osevoad ee. [e Sl eaelaae Shee sale lk lee) Inala eal cel cel See kaneis! See ae x x x x xX x x x x3 x x x x x x x x X XK XK x x x DK >< SKE SK SSK Mx x xX x x 158 PLEISTOCENE MAMMATIA. § 3. Range in Time in Britain.—We have now to discuss the paleontological value of the remains of the animal in determining the age of the deposits in which they are found. On a reference to the foregoing Table it will be seen that the animal occurs more or less abundantly in Bone-caves and River-deposits that are beyond all doubt of Post- glacial age, that is to say, which contain the remains of the arctic group of animals that invaded Western Hurope during the great refrigeration of temperature at the close of the Pliocene epoch, such as the Mammoth, Musk-sheep, and Reindeer, and that spread over the area that had been occupied by the Glacial sea as the land gradually rose again above the waves. There are, however, in Britain certain deposits which contain the remains of Post-glacial mammals associated with those which have been considered characteristic of the Pliocenes of France, Germany, and Italy, and which, therefore, stand intermediate in the geological scale between Pre- and Post-glacial deposits properly so called. In two of these the Cave Lion has been found, in the ancient river-bed at Clacton, and in the Lower Brick- earths of the Thames Valley, at Ilford in Essex, and Crayford in Kent. We will, therefore, sum up the whole of the paleontological evidence as to their place in the geological scale. The occurrence at Clacton of Rhinoceros leptorhinus of Owen (R. hemitechus of Fal- coner), of Hlephas antiquus, Hippopotamus major, Irish Elk, Horse, and of Urus, may be accounted for equally well by the assumption of its Pre- or Post-glacial age, for these Pliocene animals dwelt side by side in the same area with the arctic group of mammalia during the Post-glacial epoch. A new species of Deer, Cervus Brownit, is closely allied to the Fallow Deer, that is now found wild only in the districts adjacent to the shores of the Mediterranean. The Bison is the only animal that points in the Post-glacial direction, and this even will very probably be proved by future investigations on the Continent to have lived in France, Germany, and Italy during the Pliocene period. With its exception, therefore, there is nothing to forbid the supposition of the Pliocene age of the deposit; but, nevertheless, as the characteristic mammals of the Pliocene, so abundant in the Forest-bed of Norfolk and Suffolk, are absent, it would be hazardous to ascribe it to that age. And in the same way, since the Reindeer, Mammoth, tichorhine Rhinoceros, and other equally common and characteristic Post-glacial mammals, are also absent, it cannot be said to belong to the class of deposits containing their remains. We are, therefore, justified im assuming that it represents, in point of geological time, an epoch during which some of the more hardy Pliocene species lived under a temperature too severe for the more delicate of their congeners, and not cold enough for the invasion of the Reindeer and the allied arctic mammals. The Lower Brickearths of the Thames Valley at Ilford and Crayford contain the remains of Rhinoceros meyarhinus, which has not yet been found in France, Germany, or Italy in any strata later than the Pliocene age, and are therefore brought into more intimate relation with that epoch than any other of the deposits undoubtedly Post-glacial. Nevertheless, strange to say, since the Essay on the “ Lower Brickearths’? was written, 1 «Quart. Geol. Journ.,’ May, 1867, “On the Age of the Lower Brickearths of the Thames Valley,” by W. Boyd Dawkins, M.A., F.G.S. Ee ee eS i t ; +t ? FELIS SPELAIA. 159 its author discovered the skull of a Musk-sheep at Crayford, which of all the arctic mam- mals now alive rejoices the most in a severe climate. How the remains of the megarhie or southern species of Rhinoceros, could lie side by side with those of the most northern in habit of the herbivora, in the same river-bed, is a problem very hard to solve. Could the two animals have coexisted under the same climate in the same area? So far as we know of the former range of the one and of the habits of its living analogue,’ and of the habits of the other, it would have been impossible. There is, however, one view which has the merit of explaining this conflict of evidence, and which therefore is probably true. During the depression of North Germany and the greater portion of Britain, those portions of the Phocene Continent now represented by France and the south of England were not submerged, for in that case they would present some traces of the deposit of the icebergs that were so numerous in the North Sea of the period; it is hardly within reason to suppose that all proof of submergence beneath the Glacial sea should have been removed by subaérial denudation from so large an area, while to the north of the Thames and in North Germany it is so abundant and so ample: it is, therefore, probable that the Thames Valley roughly marks the ancient coast-line of the Glacial sea in Britain, and that to the south the Pliocene land extended through France into Italy, while to the north the look-out was over a dreary expanse of sea, burdened with icebergs, like that off the coast of Newfoundland. The temperate Pliocene climate must of course have been lowered by the presence of so much melting ice as is implied by the presence of the boulder- clay, and especially in the neighbourhood of its coast-line, independently of any great change flowing from some other unknown and cosmical cause. This climatal change must have banished to a certain extent the Pliocene mammalia from the area over which it was felt ; but, nevertheless, it is highly consistent with what we know of the migration of the herbivores to suppose that now and then some of the Pliocene mammals, such as Rh. megarhinus, may have ventured northwards as far as the shores of the great glacial sea. Again, M. Lartet? has proved that the Quaternary mammals invaded Europe from their ancient home in Siberia, where they dwelt durmg the Pliocene epoch, at the commencement of the Huropean Quaternary period; the change in the Pliocene temperature coupled very possibly with the depression of land in North Siberia causing the animals inhabiting that area to advance westwards and to occupy the feeding grounds till then belonging to the Pliocene Fauna. ‘This immigration very probably began at the time that North-Eastern Europe was dezng depressed beneath the waves during the Boulder-clay epoch. If this be admitted there is nothing improbable in the hypothesis that the arctic immigrants would gradually creep round the shores of the glacial sea, and here and there occupy in the winter the same pastures that afforded food to Phocene mammals in the summer. ‘Thus, the remains of mammals indisputably Pliocene may have becn commingled in the deposits of the same stream. In this way the 1 See ‘Nat. Hist. Rev.,’ No. xix, p. 339, 1865. 2 «Comptes Rendus,’ p. 409, et seq., 1858. 160 PLEISTOCENE MAMMALIA. presence of the Musk-sheep, the only arctic mammal found, the Mammoth, and tichorhine Rhinoceros may be accounted for in the midst of the hardiest portion of the Pliocene mammalia, the Red Deer, Horse, Urus, and others, and even with 2A. megarhinus, in the brickpits of Crayford. On the whole, therefore, there is a high probability that the fresh-water deposit at Clacton and the Brickearths of the Thames Valley form the first terms of the Post-glacial series, that is to say, of a series characterised by the invasion of Western and Central Europe by the arctic group of mammals; that they are of a higher antiquity than the majority of British fluviatile deposits; and that they bridge over that interval between the Pliocene and Post-glacial or Quaternary epochs, which is sharply marked in Britain by Glacial phenomena, but which, in France and Italy, is not sharply defined. Such is the nature of the evidence on which we have founded our belief that these two deposits are more ancient than the ordinary Post-glacial brickearths and gravels, and that they con- sequently present the most ancient traces of the Cave Lion in Britain. The Cave Lion has also been found in association with the Pliocene Machairodus in Kent’s Hole, but the occurrence of that animal does not stamp the Pliocene age of the cave, because of the enormous number of Reindeer, Cave Hyenas, Mammoths, ticho- rhine Rhinoceroses, and other characteristic post-glacial mammals that were also found. Its presence can only be accounted for on the supposition that it strayed up northwards from its southern habitat very much in the same way as its congener the Tiger does now im Northern’ Asia. There is, mdeed, nothing more improbable in the idea that the — Machairodus of Kent’s Hole preyed upon the Reindeer of the neighbourhood than that a Tiger specifically the same with that of India should at times prey upon the same animal in Siberia at the present day. It proves, however, one important fact, that while the Post-glacial fauna were in full possession of the British area, the Pliocene fauna, of which it is a member, occupied a zoological province further to the south. What, then, is the range of the Cave Lion in time in Great Britain? It is found neither in the Forest-bed nor in the ancient land-surface underlying the marine Crag of Norfolk and Suffolk, whence the water-worn remains of terrestrial Mammalia were ultimately derived. It first occurs at Clacton, Ilford, and Crayford, and it subsequently lived in incredible numbers in the South of England during the occupation of the country by the arctic group of Mammals. At the close of the Post-glacial or Quaternary period it disappeared utterly, no trace of it having yet been found in any prehistoric deposit. $4. Continental range-—Nor on the mainland of Hurope has the Cave Lion been proved to have existed during the Pliocene epoch. In France it has been found in the caverns of Echenoz and Fovent (in Haut-Sadne), of Gondenaus (Doubs), of Lunelviel (Herault), of Pondres and St. Julien d’Ecosse (Garde) ;! and in that of Aurignac described 1 Gervais, ‘ Paléontologie Francaise,’ p. 123. SSeS Er ® en ee Ee ae FELIS SPELAA. 161 by M. Lartet.’ It has also been discovered in the caves of Bruniquel and Les Hyzies and in the Rock-shelter of the Madelaine under circumstances which prove that it inhabited France while the stone-using primeval hunters lived in the country, and engraved the objects of their chase on fragments of Reindeer antler, and tusks of Mammoth. In the extreme south it is quoted by Baron Cuvier from the bone-breccia of Nice. It occurs also in the river-deposits of Tour de Boulade (Puy de Déme), of Abbeville (Somme), of Paris (Seine), of Soute by Pons (Charénte Inférieure), and other localities. Throughout Belgtum and Germany it occurs more or less abundantly, and espe- cially in the caves, such as those of Liége, Goffontaine, Gailenreuth, Schartzfeldt, Altenstem, and Sundwig. The first case on record of its discovery is that by Dr. John Hain in the Carpathians im 1672, which is also very valuable because it is the most southern point in central Hurope in which its remains have been found. Up to the present time the animal has not been found in Spain, most probably because so few bone-caves have been explored im that country. In Italy it is proved by the dis- coveries of M. Ceselli’ to have been living in the neighbourhood of Rome, while the volcanos of that district were active. In Sicily, the labours of Dr. Falconer*® in the Grotto of Maccagnone have resulted in the proof that it inhabited the island along with Man, the Hyzena, Hippopotamus, and Mephas antiquus. Thus there is proof that the animal ranged throughout France and Germany, as far south as the basin of the Upper Danube, and throughout Italy as far as the extreme point of Sicily. It has not, up to the present time, been discovered in Scandinavia, Denmark, or Prussia. There is no reason to believe that any of the deposits in which it occurs through- out this great area are of other than Post-glacial or Quaternary date. Nevertheless, it would be rash in the present state of our knowledge of the Pliocene Felidz. of those countries to affirm that the Cave Lion was not an inhabitant of Europe during the Pliocene epoch. ‘ § 5. Identity with Felis atrox (Leidy), of North America—In 1852,* Dr. Leidy figured and described a left mandible from the neighbourhood of Natchez, in Mississippi, without angle or coronoid process, and enveloped in a coating of peroxide of iron which could not be removed. Sufficient of it, however, was shown to enable Dr. Leidy to recognise its leonine affinities, and to convince him that it belonged to an animal specifically distinct from Melis leo, F. tigris, or F. leo, var. a spelea. The two points that seemed to us mm our examination of the figure to separate it from that of the Cave ' «Ann. des Sc. Nat.,’ 1861, p. 177. 2 “Correspondance de Rome du 4 Mai,’ 1867. 3 Falconer, ‘ Paleontological Memoirs,’ vol. ii, p. 550, 1868. “ «Description of an Extinct Species of American Lion,” ‘ Trans. American Philos. Soc.,’ Philadelphia, n. s., vol. x, pp. 319—321, pl. 34. 162 PLEISTOCENE MAMMALIA. Lion were the enormous depth of the ramus, and the forward position of the ramal process underneath Premolar 4, as compared with our type specimens of the man- dibles of the latter animal.) _ In all other respects it was identical in form, those minor differ- ences brought forward by Dr. Leidy vanishing away at the comparison of the large series of leonine jaws in the Taunton Museum. The subsequent discovery, however, of a lower jaw’ of the Cave Lion in Mr. Beard’s Collection from Bleadon Cave has caused us to reconsider our conclusion, since it presents exactly those characters by which we believed Helis atrox to differ from the Cave Lion, its ramal process occupying pre- cisely the same abnormal forward position, and the depth of its ramus measuring 2°77 inches beneath Premolar 4, as compared with a corresponding measurement of 2°5 inches of Dr. Leidy’s figure. In the latter, moreover, the thickness of the coat of peroxide of iron is not taken into account. We are, therefore, compelled to admit that specific difference has not yet been proved to exist between the American and the Cave Lion, and to believe, on the evidence before us, that the jaw in question really belongs to the latter animal. Contrary to what might have been expected, it differs more from that of the great South-American Felis, the Jaguar, in the enormous development of the ramal process, than does that of the existing Lion of the Old World. The associated remains found at Natchez belong to Ursus, Bison, Equus, and Mas- todon, as well as to representatives (now extinct) of the South American Fauna of the time, Megalonyx and Mylodon. There is nothing @ priort unreasonable in the idea that a geographical variety of the Cave Lion should have lived im North America during the Post-glacial or Quaternary period of that area, when we recollect that the Mammoth, Bison, and Horse, which have not yet been proved to differ specifically from those found in the Europzo-Asiatic Post- glacial series, have a similar range. There is no doubt of the specific identity of the American with the European Mammoth. ison Americanus has been found in the fossil state at Big-bone Lick, Kentucky. The Bison associated with the American Lion at Natchez is considered by Dr. Leidy (‘Smithsonian Contrib. to Knowledge,’ 1852, vol. v, art. il) to belong to a new species, Bison latifrons, Leidy; but since we cannot lay hold of even one point of difference between it and the enormous Bisons of Post-glacial Hurope, we cannot think with him that Baron Cuvier was wrong in ascribing the remains to the Aurochs (‘ Oss. Foss.,’ 4to, t. iv, p. 50, pl. mi, fig. 2). We cannot detect a specific difference in the comparison of Hyuus Americanus with the many forms of Hguus fossilis in Europe. So far, then, as we have any evidence at all, the animal, is a link in the chain that binds the Post-glacial Fauna of North America with that of Europe and Northern Asia, and we may fairly argue that the American Lion bore the same relation to that of the Kuropean Caves as the Waipiti to the Red Deer, the American to the European ' See ‘Brit. Pleist. Mam.,’ article ‘“ Felide,”’ pl. i, figs. 1, 2 a. 2 «Cat. Taunt. Mus.,’ No. 1. Felis, No. 16 and p. 7. ee FELIS SPELAA. 163 Bison, or the Canadian Elk to that of the Old World. Its occurrence in America is not more startling than that of the Musk-sheep in the South of France. But it extends the ancient range of the Cave Lion eastwards through Russia and the vast steppes of Northern Asia, across Bhering’s Straits into the great treeless barren grounds of North America, and thence southwards into the zone of the woods, and over the great prairies of the Bison, down to the almost tropical region of the Gulf of Mexico. Subsequent investigation will, doubtless, prove its former existence in the intermediate area just as in the parallel case of the Mammoth. What we know of the living Carnivores, such as the Wolf, Fox, and Tiger, would naturally lead us to expect those found in a fossil state to have a far wider range than any of the Herbivora.’ 1 See ‘ Introduction,’ p. xlix. 164 PLEISTOCENE MAMMALIA. CHAPTER XVIII. Tue Retreat or tHE Lion From Huropr. § 1. Introduction. § 3. Evidence derived from Myths. § 2. Evidence derived from History. § 4. Cause of Disappearance. § 5. Conclusion. : : § 1. Introduction. In the last two chapters the specific identity of the Cave Lion with that living at the present day has been summed up, and its distribution over Post- glacial Europe and America unfolded so far as the materials at our commands would allow. In conclusion, we propose to discuss its retreat from Europe. It vanished away, as we have seen, from Britain, France, Germany, and Italy before the dawn of the Prehistoric epoch, or the epoch characterised by the introduction of the Dog, Goat, Bos longifrons, and Sheep into Europe, as well as by the appearance of the Neolithic and bronze-using races of men.’ It is, however, extremely probable that while those animals and peoples were spreading through Europe northwards and westwards, the Lion was retreating to the south and to the east; at all events, there is ample proof that it was living in Thrace at the commencement of the Historical period in Greece, and it is not unreasonable to suppose that its retreat from North-eastern and Central Europe was gradually brought about. The documentary evidence on which the former existence of the Lion im Hurope is based is of two distinct kinds ; first, that which is indisputably true, since it presents the same grounds for being accepted as any other fact recorded in history ; and secondly, that which is afforded by myths which we may expect @ priorz to have been based upon some foundation of truth that we are able to arrive at by using history on the one hand and paleontology on the other, as our analytical tests. * The Prehistoric Period is defined in the Introduction to the British Pleistocene Mammalia (§§ 1, 2, 3, 4, 5), and in an essay on The Prehistoric Mammals of Great Britain (‘ International Congress of Prehistoric Archeology,’ Norwich, 1868). FELIS SPELAA. 165 § 2. Evidence derived from history. We have seen that the first mention of the Cave Lion is that recorded by Dr. Hain from the Hungarian basin of the Upper Danube. Strange to say, the very first historical notice that we possess of the animal is that incidentally recorded of its attacks on the baggage Camels of Xerxes, in an area but a short distance to the south of this, in the mountainous district of Thrace, between Acanthus and the city of Thessalonica. The following exact account is given by Herodotus of an incident in Xerxes’ march through Southern Thrace and Macedonia before the battle of Thermopyle :—“ And Xerxes’ and the army marched from Acanthus, striking inland, wishing to come to Therma (Thessalonica), and he marched through the Pzonian and Crestonian districts to the River Echeidorus, which rises in the Crestonian district, and flows through the Mygdonian country, and opens near by the marsh that is close to the River Axius. And while he was on the march in this direction Lions fell upon the baggage Camels. For the Lions, coming down by night and leaving their usual haunts, touched nothing else, neither beast of burden nor man, but killed the Camels only. And I wonder what: on earth could have been the cause that made the Lions abstain from the other animals and attack the Camels only, beasts that they had never seen before nor tasted. Now, there are in these districts many Lions, and wild Oxen with very large horns that are objects of barter to the Greeks. Now, the boun- dary of the district inhabited by the Lions is the River Nestus, that flows through Abdera, and the Achelotis, that flows through Acharnania. For neither to the east of the Nestus is there a Lion anywhere in the whole of Europe, nor to the west of the Acheloiis in the rest of the continent, but its habitat is the district between these rivers.” We undoubtedly owe the knowledge that Lions dwelt in this district in the year 480 B.c. to the wonder at their strange choice of prey. The story was still fresh in the memory of the hunters of Chalcidice when it was chosen by Herodotus, in his travels some twenty-five years after- wards, to light up his wonderful narrative. The animal at that time ranged through the country south of the Balkans, through Roumania to the west of the River Carasu, and 1 Herodotus ; book vii, cap. 124-6. FépEnce o& kal 6 weCoc otparoc éwopévero ix tHc “AkavOov, THY pEcdyatav Tauvwv Tice ddov, BovAduevoc é¢ THY OZpunv amtkéoQar éropévero O& Ova Tie TlavoviKne Kal Konorwuixne emt worapov ’Exeldwpov, O¢ é« Konorwvawy ap&ayevoc, pee dia Muydoving xwpne, kai eict Tapa 7d EAoc 76 im “AEww Toraua. Tlopevopévy S& révty A~ovrec of émPixavTo Thor siToPdpoict KaphAotot’ KatapoiréovTEec yap ot Agovtec Tag viKTac, Kal AEiTOVTEC Ta oérepa HOEa, GAdoV piv dvdEVOE ATTOVTO OvTE Jro- Zuytov, ovte avOpwrov, oi St tac Kaundove zkepaiZov povvac. Owvpalw oé 7d altiov O,rb Kore jv, Tov GAdwy TO avayKalov ameyouevoue Tove Afovtag That Kayhroror éemitMecBar 7d pnre mporepov Oruwmecay Onprov ptr émeTEpéaTo avTov. "Evol of kata TavTa TA xXwola kal AEdvTEC ToXXol, kal dec Gyplol, TOV Ta Képsa Drepueyabea zor, Ta 2¢ TOG “ENAnvac poiréovTa. Ovpoe 88 roicr Noval zor O,re Of ABSipwv péwy Térapyog Néotog, Kat 0 du ’Axapvaving péwv "AyeAwoc’ ode yap TO Tpo¢ THY 7H TOV Néorov ovdauohk Taone ric tumpocBev Evewmne dot ric av Aéovra, ore Tpoc Eorépne Tov Axehwov év 7H UroASiTwjTElpw, GAN ev TH petakd TouTédy THY ToTénwv yivovral. 22 166 PLEISTOCENE MAMMALIA. through Thessaly as far south as the Gulf of Lepanto and the Isthmus of Corinth, having as its western boundary the River Potamo and the Pindus Mountains. The next mention of the Kuropean Lion we find in Xenophon’s ‘ Treatise on Hunting,’ which he composed on his banishment from Athens, after he had exchanged the court and the camp for the pleasures of gardening and hunting, in his splendid retreat in Lace- deemon, about the year 380 B.c.'—“ Now, Lions, Pardaleis, Lynxes, Panthers, Bears, and the like beasts, are caught in foreign countries in the neighbourhood of Mount Pangeeum and Mount Cissus, which is beyond Macedonia, and in the Mysian Olympus, and in Pindus, and in Nyse that is above Syria, and in other mountains that are able to support such animals.” Mount Pangeeum is situated near the sources of the Nestus, not far from the range of Rhodope (of the Balkans), Cissus is close to Thessalonica, and therefore this passage corroborates strongly the statement given by Herodotus as to the range of the animal, the only difference being that Xenophon states that it inhabited the Despoto Dagh Mountains of Roumelia, the eastern watershed of the Nestus, instead of its being restricted to the western bank of that river. Baron Cuvier,’ indeed, and the late Right-Honorable Sir George Cornewall Lewis* agree in refusing historical value to this passage, because other localities in Asia are mentioned, believing that all these animals were not found m any one of these localities. But the fact that the Lion lived in that area, both before and after Xenophon’s time, coupled with the fact that the Panther, Lynx, and Bear ranged through Europe in company in Post-glacial times, renders it very probable that he was scien- tifically accurate when he advised their capture in that district, by placing poisoned food near their drinking places. The Lynx and Bear still live in the same neighbourhood, and the Panther still remains in Asia Minor, bereft of his congener the Lion. The historical value of the account of the range of the Lion in Europe given by Herodotus and Xenophon is corroborated by the testimony of the great father of natural history, Aristotle, who flourished some fifty years after the time of the latter writer, and who, being a native of Stagira, lived in the very district said to have been inhabited by the Lion. He describes its European range very nearly in the same words as those used by Herodotus; but in the hundred and fifty years that elapsed between their dates the hunter and the husbandman had made great inroads on the last foothold of the Lion in Europe. The “ wodXot déovrec,” “the many Lions,” spoken of by the one had dwindled } Zevopwvroc Kuvnyeride. Cap. xi, 1. Agovrec 62, mapodAsc, AbyKec, TavOnpEc, apKror Kat radXa boca tort rovavta Ofpia aAlokera ev Esvatc ywpaie Tept 7d Ilayyaiov dpoe kai Tov Kirrov 7 Umep TG Makedoviac, Tad tv TH "ONOuTH 7H Mvoty kai év Livdw, ra © ev 1H Nboy 7H UTE Ti¢ Luplac kal Tpde tole adAote OpEct, Goa ola T oT TOEPELW TOLAUTA. * *Oss. Foss.,’ 3e €dit., 4to., t. iv, p. 425. 3 “Notes and Queries,’ second series, viii, 1895, “‘ Lions in Greece.”” We are indebted for several of the references to classical works to the learning of this eminent critic. FELIS SPELAA. 167 down into the “oanov yéoc,’ “the rare animal” of the other ; he adds also that the wild beasts of Europe are more courageous than those of Asia or Africa.’ We have no mention of the animal in Europe from this time recorded by any writer down to the year a.p. 80 or 100, when it is mentioned by Dio Chrysostom Rhetor’ in his « Hssay on Beauty.’ “The honorable,” he writes, “‘ have vanished away in time, as they say the Lions have done which formerly dwelt in Hurope, for there are no longer any more; but formerly they dwelt in the district of Macedonia and in other places.’ Philostratus, also, writing in his ‘ Lives of the Sophists,’ about the year 220 a.p., relates that Agathion, the athlete, who lived in the time of Herodes Atticus, 104—180, a.p., complained that he could not rival Hercules because there were no Lions in Acarnania. It is therefore clear that the Lion had deserted Europe before the end of the first century after Christ ; or, in other words, that the “rare animal” of Aristotle had become extinct during the four hundred years that followed his time. It is, of course, impossible to fix the exact date, just as in the parallel case of the Brown Bear in Scotland or the Beaver in South Wales. In the literature of Rome there is nothing that would lead to the supposition that the Lion lived in Italy during the Historic period. * At rabide tigres absunt et seeva leonum Semina.’’—Georg. i, 151. According to the high authority of Sir Cornewall Lewis, it is not even alluded to in Italian mythology.’ § 3. Lvidence afforded by Myths. In brmging Mythology to bear upon the question of the former existence of the Lion in countries where it was extinct before the Historical Period, we are justified only by the high probability of its truth, afforded by the fossil remains on the one hand and by history on the other. The evidence, indeed, afforded by the myths is so strong that Sir G. C. Lewis has admitted its value without knowing of the corroborative witness of the fossil remains. ‘That eminent critic sagaciously inferred 1 Aristotle, ‘Nat. Hist.,’ edit. Schneider, lib. vi, 28, 1: Lmrdviov yap 70 yévoc 76 THV AcdvTwY zor, Kal OK év TOAAY ylyveral TOTW, AXA Tie Eipwrne amaone év TH meTad Tov “Ayedwov kal rov Nécoov rorapov. Lib. vii, 27, 6.—'Ere 62 Novrec piv 2v 79 Evpway paddov, kal rie Evpwrne év TH METACD TOTW rou Axedvou kat rou Nécoov. TapddAac 8 gv 74 Acta tv & ty EKvpwiry ov yivovra. “Odwe 82 Ta piv dyola aypwwrepa tv TH Acta, dvdpedrepa 62 tavta Ta ev 7H Evpwirn, moAvpoppdrara Ot ra 2v AMiBby" Kat Aéyerat rie wapompia, bru det Pepe re AiBbn Karvdv. 2 «Dr. Falconer stated this to me in one of the many conversations on fossil bones which will ever remain in my memory.”—W, Boyd Dawkins. * “An. des Se. Naturelles,’ 5° série, tom. viii; ‘Deux Tétes de Carnassiers Fossiles.’ * “Animaux Vertébrés Vivaux et Fossiles,’ 4to, 1867-9, p. 68, pl. xv. 178 PLEISTOCENE MAMMALIA. ception of the canine belonging to Lord Enniskillen are preserved m the Taunton Museum. § 2. Comparison of fossil remains with most closely allied living forms.—Under the name Felis (Leopardus) pardus we intend to embrace the Panther properly so called, and the Leopard, the former consisting of the larger and stouter, and the latter of the more slender and smaller individuals ; smce an examination of a large number of skulls, skeletons, skins, and living animals, has convinced us that the differences are only varietal, and not of specific value. At the same time there exists m Northern China, and, we have reason to believe, also in other parts of Hastern Asia, a species which is undoubtedly distinct, and which differs from the Panther, in the comparative length of the nasals and frontal processes of the maxillaries, exactly as the Tiger differs from the Lion. This species has been named by Dr. Gray (‘Proc. Zool. Soc.,’ 1867, p. 264) Leopardus — Chinensis ; Dr. Gray describes a third species, under the name of Leopardus Japonensis (‘Proc. Zool. Soc.,’ 1862, p. 262); and the fourth is the Jaguar of America, Leopardus onca, Linn. ‘These are the only living species with which it is necessary to compare the fossils in question.’ On comparing the fossil teeth with those of the living Leopard and the Jaguar, we find that they agree with the former in their slender, delicate, and compressed form. In the Jaguar (Melis onca) the teeth are much stouter and the cusps of the molar series more obtusely conical. The canines (Pl. XXIV, figs. 1,3, 4) present a character which at once differentiates them from those of the latter animal in the two longitudmal grooves or sillons which traverse the outer and the inner sides of the crown. In the Jaguar the second sillon on the outer side is a mere rudiment, or in some cases is altogether absent. In all these pomts in which the teeth correspond with those of the Leopard, Melis pardus, they also agree with those of the allied species Zeopardus Chinensis of Dr. Gray. They may, however, be referred with considerable certainty to the former animal, since it has been found in several caves in France, while the latter has not been known to live out of China. A glance at the following table will give the relative size of the fossil lower true molar as compared with that of the Leopard and Jaguar (Pl. XXIV, fig. 2). Zegrdee. | seman. [Rea lager) Cat. 115. Africa. 117 C. Brit. Museum. | Brit. Museum. | Brit. Museum. Antero-posterior length ..................... 0°81 07, 07 0-74 Antero-transverse diameter ............... 0:3 0:3 0:3 0°42 Postero-transverse diameter ............... 0°28 0:28 0°28 0°38 LIENS Of CROWD cag casnnuscovesvovasyooasdaaos 0°44 0°38 0°39 0°39 * A fifth species of Leopardus, of large size, has been recently described by M. Milne-Edwards, from Northern China. ‘Ann. Sci. Nat.,’ ser. 5, tom. vill, pp. 374-76. FELIS PARDUS. . 179 _ This variation in the size of the fossil, as compared with the recent Leopard, is not greater than that presented by living individuals of the same species. The Femur.—The left femur figured (Pl. XXIV, fig. 5) corresponds with that of a large Panther from West Africa, in the Museum of the College of Surgeons. Its proportions may be gathered from the figure, and there is no necessity for a detailed description. The rest of the bones are too fragmentary to be figured or described. The remains from the Pleiocene beds of Mont Perrier in Auvergne ascribed by MM. Croizet and Jobert to the Melis antiqua are far too large’ to have belonged to the largest Leopard. Nor can any of the other fossil felines be identified with it with any certainty. M. de Blainville,” however, believes that the Fes pardinensis, and the F. Arvernensis of MM. Croizet and Jobert, are identical with the Panther. The Fels par- doides® of Professor Owen, from the Red Crag, differs from the Panther in the lowness of the crown of the last true molar. § 3. Associated Animals—The species with which the remains of the Panther have been found in Britain, France, Germany, and Gibraltar, are represented in the following table; Marcel de Serres, Dr. Falconer, Mr. Busk, and Dr. Goldfuss, being the authorities for those of Lunel-viel, Gibraltar, and Gailenreuth. The Ursus ferox from the last cave is based on the examination of a series of teeth obtained by Sir Philip Egerton and Lord Enniskillen. Caves. List or SPECIES. Sandford Banwell. | Bleadon. Hin, | Lnel-Viel.) Gibraltar. Gailen- reuth, Osu sta metos wun me epan Vance deceit Aa x x Werespelceust GOldey .taseycehe a. teh se he x x wie (OS ORO SUS Jue conopuctaos Qncoemeca deca tae ea ae Ba a aBh x Gralla (PISG0IS, WENT, codesoone sn ébo500ndaacd ube x WIGS GUIS, 1s So uodean soup eunedakerensepa neo WMastelaypuconcusslien sae. ecaceseene cee AY fo OCT Wis, WOR eis iieatp ein vin A cee a IDLE CTA TAD W183 16. Becbco dav oecodecos Habeoe COLTS SUNOS ioc nenonde poee roan eee Ola WIG ISS IU Mh eee eek ea Scene M aan Hyena (spelea, Gold.) crocuta, Zim. ... JIGHIS GIS WIR Sc cepdadae een eos ceaeaee ear JP, (CREED) POTTS, Ws soassooneobodon see x F. leo (war. spele@a), Li.............. D So NERS ee PEMD ORAULOR ag ce scttueloate se waite Saoaeecee ae sity ss aoe x IP's, GGFR, SONTRAD,, coo sndetesoaneancebaneoseee ya ee aia bee x a SK xX De xXx 2 XX MO 8 8 os XXxXxXXXXX: ~~: xX XX XX: x x QD VE Shw Sos ON Re 8 1 “Oss. Foss. de Puy de Dome,’ p. 214, 4to, 1828. ‘Ostéographie,’ article “ Felis,” p. 143, pl. xvi. ~ 2 «Ostéographie,’ article “Felis.”’ 3 «Brit. Fossil Mammals,’ p. 169. 180 PLEISTOCENE MAMMALITA. CavES. List oF SPECIES. Banwell. | Bleadon. sto Lunel-Viel.| Gibraltar. a : Cervus megaceros, Hart ..............+-...5 Ae x Ositarandus ites eco ee eee x x x 5 GOPRGOUTS Nis coaepaccsbaoncsodsscdesdsbe00 x iv sdk a Os GUTVOID, MWesonea0con cc con0boe padres ce x x x x Cxidama Mints. 33 on eee ee ie: Xx ISOS FOV DOUD) IB}o — areaccupdoncaonopons ee x x x 4 Se Bison priscus) OWn--ea)--o.0-4- de eheeeee tee x x x q x Capra abea, Mics un se oeen Peper oe x Hippopotamus major, Desm................ as i uh x 300 Susiscrofa, Mi pe eee pee eRe eee a < lah x x x J BUTE GOONS, Wisc soostagase codes0 one asec ba x x x x x Rhinoceros hemiteechus, Fale. ............ br to a x Xx ; 18 COOOHOTMIS, CRs soos00ccananancconsaccce ee ey x Ne. a. x Hlephasvantiquus hale. meena nee x LE prumigenvis,s sna ere ere ee eee x x x LCMMUSH SDs secede Eee oe x eae obo Jhernms GCN, VM, 35425555900 206 s00ese a ae Le oa x bio CHOTBUTOIG, SEP, soccodsndososesosuds0sa0 x . Tagomiysispel eis 10 were ee eee eee eee ae Spermophilus erythrogenoides, Fale. ...... 4 ae SS ae ears sry Goicrioridaso stan ons Arvicola pratensis, Bell cee eee eeseieere Lily CUPFORGIS) NEWT. 550008 002008500000 000 500000 Lily HHPHOUDOOIS, DOV 55 ..00c0ca0000 eooccose Mo 300 se Castorsyiber ties: a... Seer ee REEE nos aa ago x YES TROIXCOWUI, Wis scocswccosodones0nenooa09es xX XK XK XK X x § 4. Range of F. pardus—The present home of the Panther or Leopard is to be found in the warm regions of Africa and Asia, and not in Europe, or the colder districts of Asia. The occurrence of the animal in the caves of Gibraltar proves that in the Pleistocene period it passed northwards into Spain, while the discovery of its remains in France indi- cates that it ranged over that region, and those in Gailenreuth Cave that it lived in Central Germany. The remains in the Mendip Caves show that it passed northwards over what is now the Channel, to prey upon the Reindeer, Bisons, and Horses of Somer- setshire. Throughout this area it was very rare as compared with the contemporary Lions, Bears, and Hyzenas. The presence of this beautiful feline in Britain, now only living in a warm climate, may easily be explained by the hypothesis that it migrated northwards from time to time from the warmer regions around the Mediterranean; and it is very probable that the Arctic severity of the Pleistocene winter was the cause of its rarity in our latitude. SS PY ea FELIS CAFFER. 181 CHAPTER XXI. Tue Smatter FELID. PLATE XXIV. § 1. Introduction. | § 2. Felis caffer. § 3. F. catus. § 1. Introduction —The remains of the smaller Felidze have long been known to exist in the Pleistocene Caves and river deposits in Great Britain, France, Belgium, and Germany, and have for the most part been referred without any minute analysis of their characters to the common Wild Cat of Europe (77. catus, L.) M. Marcel de Serres! and his colleagues figure one lower jaw from the Cave of Lunel-Viel, and Dr. Schmerling’ another from the Caves of Liége, which are larger than any well-authenticated lower jaw of Wild Cat which has passed through our hands. The former ascribe their specimen to the Felis Jera, by which they probably mean the common Wild Cat, while the latter assigns his to the ‘Helis catus magna,’ regretting at the same time his lack of recent specimens with which to compare it. The measurements of the depth of jaw given in the ‘Ossemens fossiles de Lunel-Viel,’ p. 120, show that the jaw figured in Pl. IX belongs to the same animal as that figured by Schmerling, while those given by the latter do not agree with his own Plate, the confusion beimg probably caused by the substitution of height for thickness.’ The smaller feline remains which we have examined from the Caves of Great Britain * Marcel de Serres, ‘ Oss. Foss. de Lunel-Viel,’ p. 120, pl. ix, figs. 12, 13, and 17. * Schmerling, ‘Oss. Foss. de Liége,’ vol. ii, p. 88, pl. xviii, figs. 13, 14, 23, 24. 3 In the text these measurements are— . M. INCH—ENG. Hauteur de la machoire devant la premiére molaire : (by OWO7y = Oran la derniére : : (2) 0008 = 0:031 In the figuresthey are— ; : : é (1) 0012 = 0° 55 (2) 0-013 = 0-51 The other measurements exactly agree with the figure. 182 PLEISTOCENE MAMMALIA. seem to us to imply the existence of two species, one the Felis caffer of Africa, and the common Wild Cat, which is now rapidly being exterminated in our island. But on the very fragmentary evidence before us we do not attempt to define with absolute certainty the former existence of the Helis caffer in Britain. Nevertheless, the exact agreement in every particular of the lower jaw figured in Pl. XXIV, fig. 6, 6’, 6”, with that of the latter animal, and its disagreement in the same points with those of other animals, renders the specific identity almost certain; and with regard to the classificatory value of the points themselves in the recent lower jaws, we have found that they are present in all those of the former, and absent from all those of the latter which we have examined in the British Museum, the College of Surgeons, and elsewhere. § 2. Helis caffer—The right lower jaw from Bleadon Cave in the Mendip Hills, figured Pl. XXIV, fig. 6, 6’, 6”, differs from the jaws of all the smaller Leopardine Cats of both the Old and New Continents, as well as from the Lynxes, by the smaller size of the molar series as compared with the depth of the jaw, while the jaws in these animals are thicker in proportion to their depth. These smaller feline jaws being thus proved to be unlike that in question, there remain for comparison two groups of Cats, the larger, represented by the Fels chaus of India, which appears to have had a large share in the production of the Domestic Cat of that country; and the smaller, or that which is represented by the Wild Cat of Hurope, and the /. maniculata, the latter probably having a share in the breed of the Domestic Cat of Europe. Intermediate between these groups are two species, the F. caffer, distributed at the present day throughout Africa, and the /. torguata of the Himalayas, which are closely related together, and are probably representative forms in their respective districts. In the series represented by Felis chaus the lower true molar, m 1 1s larger in proportion to the premolars than in our fossil. ‘There remain, therefore, for comparison the Wild Cat of Europe, the /. maniculata, and the #. caffer. In all the well-authenticated specimens of Wild Cat which we have examined, as, well as in those of the large domestic cats with brownish-grey fur, which have run wild, the posterior inner alveolar border is much thickened (PI. XXIV, fig. 8, c), and rises higher than the outer border, so that the last true molar, and to a certain extent premolar 4, are thrown to the outside of the jaw. These characters are not to be found in our fossil, the molar series being set on the middle of the alveolar edge of the mandible. The ramus also in the fossil is deeper and transversely narrower than in the Wild Cat. In the Fels maniculata, and the smaller specimens of Domestic Cat, the Jaw is much less deep in proportion to its thickness than in the fossil. The jaw of Melis caffer, on the other hand, agrees with our fossil in the minutest detail, as well as with that figured by Dr. Schmerling, and measured by M. de Serres; and it is therefore impossible to resist the conclusion, that a species of Wild Cat most closely allied to the #. caffer lived in Britain, Belgium, and France, in the Pleistocene Period. There is indeed nothing unrea- a FELIS CATUS. 183 sonable in the suggestion of a Cat now found only in Africa having once ranged over Europe, since the Spotted Hyzna, the Hippopotamus, and the Panther were members of our Pleistocene Fauna, as well as being now associated with /. caffer in Africa. In the following table we have represented the relation of the fossil lower jaws to those of the Wild Cat and Felis caffer. ‘The measurements are taken in inches: Tosa Salmers one cae & Cues. a Catus. . ) oll. Surg., rit. uS., cotiand. ermany. Be ope NES: Bleadon. | ling’s fig. | “°4696.°”| 57 A. |Coll. Surg, | Brit, Mus. Wenethrot molar series| $..2...-4..-.22-2--+- 0-91 0:92 0-91 0°92 0°82 0°74 Wigibpeioniot ies: hasce ceca cke voutesscs. ans 0°35 0°36 0°34 0°33 0°32 0:29 Postero-transverse diameter ......... 0°16 me 0:16 0°14 0°13 O11 Antero-transverse diameter ......... 0°15 er 0-15 0°13 0:14 0:10 IRIGWELNG _ oie accseh Ron cooeneete oer teee nee 0-11 0:13 0-11 0°10 0:09 0:08 pre Wen ethy 225. y.escse-n: Heed oe Ue 0°34? ac 0°35 0°30 0°28 0°26 ON ECM OGM 2 ce oe neta. < sanaee tessa ect 0:26 0°25 0°26 0:26 0:20 0:20 MMrans verse ccoe ns cceein ec 0-14 sae 0°15 0°13 0°10 0:10 PCL DT busy teen ee Men et tee 0:20 0°20 0°19 0:17 0-18 0°15 Depthvok jaw: at Mil, aces. 022e-cesorcenen- 0°53 0°54 0°56 0°52 0°40 0°39 Fhhiclsnes sian. MS euppeme oe wet rat. 0°22 ae 0°23 0:21 0:25 0°22 Wen the ateP Vice. se ney SR as, 5: 0°45 0°45 er sie Bas ce The fragments of ulna (fig. 8) and of femur (fig. 7) of a large Cat from Bleadon Cave may most probably be referred to the 7. caffer, since they were obtained from the same place as the jaw which we have just described. They may, however, belong to the Wild Cat. § 3. F. catus—The lower jaws and bones of the Wild Cat of Europe, which is now so rapidly becoming extinct in Great Britam, have been discovered in the Cave of Kirkdale by Dr. Buckland, and in that of Kent’s Hole by the Rev. J. MacHnery, and differ m no respect from those of the living representative. If that figured Pl. XXIV, fig. 8, be examined, it will be seen that in its slender form it contrasts with that figured 6, 6, 6°. The lower jaw (Pl. XXIV, 9, 9’) is from the Brickearth of Grays Thurroch, and is in the Collection of Mr. Wickham Flower: its external aspect has been figured in the ‘ British Fossil Mammals,’ p. 172. ‘There are no points presented by these fragments which are worthy of a detailed notice. The remains of the Wild Cat have been obtained from several of the localities in Great Brita, and it lived in our country from the age represented by the Brickearths of the Thames Valley to the present day. 184 PLEISTOCENE MAMMALIA, CHAPTER XXII. Famity—FELID &. Genus—Macumrovus, Kaup. Species—MACH#RODUS LATIDENS, Owen. PLATE XXV. § 1. Nomenclature. § 6. Evidence that Macherodus latidens came § 2. Range of Genus. from Kent's Hole. § 3. History of British Remains. § 7. Continental Range of Species. § 4. Relation to Macherodus cultridens. § 8. Antiquity of Macherodus in Kent's Hole. § 5. Description. § 9. Macherodus from Norfolk. § 1. Womenclature.—The history of the discovery of the great sabre-toothed Feline Macherodus shows at once the difficulty with which naturalists have to contend in assigning fragmentary remains to their rightful possessor, and of the gradual steps by which that difficulty was removed. ‘The tooth of the animal now known as Macherodus cultridens, from having been associated with the remains of the Bear in the deposits of the Val d’Arno, was assigned to an abnormal species of that genus by Cuvier in 1824, or the Ursus cultridens,* and by Nesti in 1826 to the U. drepanodon ;? and similar teeth found in Auvergne have been described by MM. Croizet and Jobert® as U. cultridens Issidorensis. M. Bravard,* however, having discovered a feline skull in the same district, which was possessed of sockets such as would fit the large compressed canines somewhat resembling those im question, and of a mandible in which there was a deep outer depression for the reception of the upper canine when the jaws were shut, inferred that the animal was a * “Oss. Foss.,’ 4to, 3rd edit., 1824, vol. v, pt. ii, p. 517. 9° * “Lettera Terza dei alcune Ossa fossili, al S. Paolo Savi,’ 8vo, Pisa, 1826. ° «Oss. Foss. de Puy de Dome,’ p. 200. 4 ‘Monographie de deux Felis d’Auvergne,’ p. 143. MACHASRODUS LATIDENS. 185 Felis, and described it as F. megantereon. And that this conclusion of M. Bravard was true is proved, as Professor Owen remarks,’ by the fragment of skull of an allied form in the British Museum, from the Sevalik Hills. Dr. Kaup,’ on the other hand, who had met with the remains of the animal at Eppelsheim pointed out that the compressed and serrated canines, in which the two longitudinal grooves so characteristic of the larger Felines were absent, separated the animal to which they belonged frem the genus Felis, and he therefore proposed for them the name Macherodus, or the Sabre-toothed Carnivore. On the whole, the evidence which we possess as to the affinities of the animal prove that it belongs to the great family of the Felidee, although those pomts which Dr. Kaup has brought forward forbid its classification with the genus Felis, from which it differs in the enormous development of the serrated upper canines, as well as the presence of a third lobe on the sectorial edge of the upper Premolar 4. And that it is an aberrant member of the great family of Cats is the opinion of M. de Blainville, Professor Owen, and M. Albert Gaudry. Its dental formula is that of the true Felines. § 2. Range of Genus.—The genus Machzerodus is of very wide range both in space and time. It has been found alike in the Meiocene deposits of India by Falconer, the plains of Marathon by Gaudry, and the river-deposit of Eppelsheim by Kaup.? It has been known to occur in the Pleiocene strata of the Val d’Arno since 1812, and of Auvergne since the year 1828. It has also been found in the Pleiocene Caves of Brazil by M. Lund, along with the great Sloth, the Megatherium, and the peculiar Horses at that time living in South America. The best known and most widely spread European species is that which ranged over France, Germany, Italy, and Greece, during the late Meiocene and the early Pleiocene periods, under the name of Macherodus cultridens of Kaup, and which has been fortunate in being described by M. Gaudry in his classical work on the ‘Animaux Fossiles de lAttique. And the proof of the presence of a closely allied form in England we owe to the labours of the Rev. J. McEnery in Kent’s Hole Cave, near Torquay. § 3. History of British Remains.—The seven teeth which afford the proof of the ancient sojourn of the Machzrodus in Great Britain were discovered so long ago as 1826, and their history has been very remarkable. The Rev. J. MacKnery unfortunately did not publish the results of his explorations in Kent’s Hole, which he carried on from time to time after 1825 up to his death in 1840, and the manuscripts were lost, until they fortunately fell into the hands of Mr. J. Vivian, of Torquay, who published an abstract in the year 1859. ‘en years later they were published in full by Mr. Pengelly, F.R.S., and afford an authentic and circumstantial account of the discoveries, which had been lost to 1 «Brit. Foss. Mam.’ (1846), p. 176. 2 “Oss. Foss. de Museum de Darmstadt,’ 2nd part, 1833. 186 PLEISTOCENE MAMMALIA. science for nearly thirty years. The lithographs also which had been originally made to illustrate a work on ‘ Cavern Researches,’ which was never carried out, were published in part in 1859 by the permission of Mr. F. Buckland, into whose hands the stones of seven- teen out of the thirty plates to which Mr. MacEnery refers in his prospectus had passed at his father’s death, and now even these cannot be traced. Two more plates were fortu- nately added to these in the year 1869 ; and a third plate, which had not been known to be extant, was added subsequently, through which additional evidence as to the sojourn of the Macherodus in the Cave was obtained. Nor were the remains of the animals which Mr. MacEnery discovered, more fortunate. At his death they were sold by auction and divided up among private collectors, and for the most part irretrievably lost. Out of the seven teeth of Macherodus we are able only to trace five. ‘The canine in the Oxford Museum was purchased by Dr. Lovel Phillips at the sale, and given to Dr. Buckland; that in the College of Surgeons, and figured by Professor Owen, was presented by Lord Enniskillen to Professor Owen; a third, the original of pl. ¥’, figs. 4, 5, found its way into the Museum of the Geological Society ; the fourth, figured pl. rv’, figs. 1, 2, 3, is in the British Museum; and the fifth (pl. ¥, fig. 7) is in the Collection of Sir Walter Trevelyan, Bart., to whom it was given shortly after its discovery by Mrs. Cazalet." We are unable ‘to trace the two incisors, one of which is figured in the fossil mammals (fig. 70).” § 4. Relation to Macherodus cultridens—All the canines which had the above eventful history belong to the upper jaw, and are remarkable for their width as compared with the length of the crowns. ‘That in the College of Surgeons, which Professor Owen takes as the type-specimen of I. datidens measures 6°5 inches along the outer curve, and 1-2 inches across the base of the crown, while in the Italian Macherodus cultridens the corresponding measurements are 8°5 inches and 1-5 inches. If the proportions of the Italian specimen be constant, it is obvious that the British specimen in question must belong to another species, since the basal measurement of the crown is so much greater. Professor Owen attaches a specific value to this greater width, while Dr. Falconer, after carefully weighing the evidence, believes that the difference has merely a varietal importance. ‘The size of the canines is, to a certain extent, a sexual character, and therefore liable to variation in different individuals of the same species. And to what an extent this variation may take place within the limit of a species may be gathered from the comparative measurements of canines of the Cave-Lion in our Monograph on the animal. But although the character in question be not of specific value, the strongly marked serration in the incisors (woodcuts 1, 2, 8) differentiates, as M. Gervais remarks, the British from the French species of Letters to W. Boyd Dawkins, dated 11th and 26th May, 1869. 2 Mr. Pengelly has given a full account of the teeth in question, and of their singular history, in his series of ‘ Essays on the Literature of Kent’s Hole,’ published by the Devonshire Association. MACHARODUS LATIDENS. 187 Macherodus, since it is not presented by the teeth in the cranium found in the Pleiocene strata of Auvergne by M. Bravard, and admitted by Blainville, Owen, Kaup, and Gaudry, to belong to Macherodus cultridens, Kaup. For this reason, therefore, we consider the British MWacherodus latidens, Owen, to be distinct from the JZ. cultridens of the Continent. The recent discovery of a lower true molar, in the cave of Baume, in the Jura, renders it probable that all the teeth were serrated. § 5. Description.—The upper canines of Macheerodus are characterised by their com- pression parallel to the median line, and the strongly marked serration of the ridges which traverse the teeth in front and behind, and give it a sharp cutting edge, which Professor Owen describes as uniting the power of a saw with that of a knife." The regular curva- ture of the crown and fang causes the tooth to present an outline strongly resembling, according to Nesti, the crescent-shaped new moon when first appearing above the horizon. The crown is thicker in front than behind (PI. XXYV, figs. 3, 6), and thus possesses great strength without the penetrating power of the posterior edge being impaired. Altogether the tooth is the most perfect instrument for piercing and dividing flesh which is presented by any of the Carnivora, and doubtless belonged to an animal which lived solely on flesh. The general shape of the upper canines from Kent’s Hole may be gathered from Pl. XXV, which is a copy of the plate drawn by Mrs. Buckland, and lithographed by Mr. Scharf, for the work of the Rev. J. MacHnery, and kindly lent to us for the purpose by Mr. Pengelly, F.R.S. In figs. 1, 2, 3, the perfect crown is represented, which is now preserved in the British Museum, while figs. 4 and 5 represent the perfect fang. ‘The lower canines are proved by the lower jaw of Macherodus cultridens disco- vered by M. Bravard, along with the perfect cranium in the Pleiocene strata of Mont Perrier in Auvergne, to be very much smaller in every dimension.’ The incisors of Macherodus latidens are now only known to have been found in Kent’s Hole by three figures of the natural size in a lithograph which is deposited in the Museum of the Natural History Society of Torquay. The accompanying woodcuts have been drawn on wood from a photograph of the original, which has been placed at our disposal through the kindness of the Society, and fig. 1 representing the inner aspect of the left upper incisor, 3 is that which has been copied by Professor Owen. ‘The anterior and posterior ridges traversing the crown qare serrated, as in the canines, and at the base of each there is a well-defined cusp, 4 and c, both of which pomts are unknown in the incisors of any of the living Carnivores. ‘The incisors of the left lower mandible repro- duced the peculiar characters of the corresponding upper tooth, the serration being well 1 «Foss. Mam..,’ p. 180. * M. Bravard’s ‘Monographie,’ pl. iii. Blainville, ‘ Ostéographie,’ Article Felis, pl. 17. 188 PLEISTOCENE MAMMALTA. marked, and the cusps 4 and ¢ clearly defined. ‘he crown of both these teeth are consi- derably larger than those of any living or fossil species of the Felidz, and are of an eminently sectorial character. There is not the slightest evidence that the Machzrodus was more closely allied to the ‘Tiger than to any of the other larger Felines, and therefore the very tempting name of “Sabre-toothed Tiger” must be given up, as implying a relationship which does not exist. The size of the teeth may be gathered from the following measurements in inches: Basal width of Length of crown. crown. Length of fang. Pl) OXs fie Brit. Musas ener 2°4 Ne, bs 35 so, 4b. (GeoltiNoch ae eee eee te 1:18 3°2 5 pt Ze) OSID We Dreveliyanyeeeeiaee eee cee a 1°3 3°5 Coll iSurgeonst.5 fa.:)) Adore eee ee eee UT 22 3°3 Oxford Miase jojo sid ccs cee gee oe eee ee Soe a e § 6. Evidence that Macherodus latidens was derived from Kent’s Hole.—There can be no reasonable doubt as to these remarkable remains having been derived from Kent's Hole, and not from the Continent, since the animal to which they belong differs specifically from the Continental Pleiocene species. The MSS. also of the Rev. J. MacHnery point out the precise circumstances under which they were found :” ! Gervais, ‘Zool. et Paléont. Francaise,’ 1859, p. 231. 2 See Mr. Pengelly’s admirable series of “Essays on the Literature of Kent’s Cavern.” McEnery’s MSS., ‘Trans. Devonshire Ass.,’ 1869, pp. 55-6. MACHASRODUS LATIDENS. 189 “We now returned,” he writes, “to the excavation (in the ‘ Wolf’s Passage’), which produced the Wolf’s head. The stalagmite was about a foot and a half thick, and of ‘ excessive hardness, in which were imbedded rocky fragments rolled down the slope; but as we advanced inwards, the stalagmite became altogether free from foreign admixture and moulded itself upon the mass of bones. Of the quantity and condition of the remains here it is scarcely possible to give a just idea without appearing to exaggerate. ‘They were so thickly packed together that to avoid injuring them we were obliged to lay aside the picks and to grub them out with our fingers. They had suffered considerably from pressure after having first undergone violence from the force which impelled and congre- gated them in this narrow neck. ‘They were found driven into the interstices of the opposite wall, or piled in the greatest confusion against its side, with but a scanty covering of soil, and that of the finest and softest sand intermixed with greasy earth. To enumerate the amount of fossils collected from this spot would be to give the inventory of half my collection, comprising all the genera and their species including the cu/tridens, there were hoards; but I must specify the jaws and tusks of the Elephant, with the teeth m the sockets, and the bone of which was so bruised, that it fell to powder in our endeavour to extract it, a rare instance of the teeth occurring in their jaws or gums. The same may be observed of the jaws of the Rhinoceros, one portion alone of which was saved, but the teeth of both were numerous and entire. ‘The teeth of the Elk, Horse, Hyzena, were taken out whole; the teeth of the two last were gathered in thousands, and in the midst of all were myriads of Rodentia. ‘The earth, as may be expected, was saturated with animal matter; it was, to use the expressive words of my fellow-labourer Walsh, fat with the marrow and sinews of more wild beasts than would have peopled all the menageries in the world.” § 7. Continental Range of Species—Such as this is the evidence of the sojourn of the formidable Macherodus latidens in the Cave of Kent’s Hole. The proof that the species lived also on the Continent of Europe is due to the discovery of an upper incisor in every respect identical with figs. 1, 2, im a deposit near Puy, in Auvergne, by M. Aymard, which is doubtfully considered “ diluvium” by M. Gervais,’ and most probably belongs to the Upper Pleiocene, or the passage beds between the Pleiocene and Pleistocene formations. M. Gervais” has also recently determined the existence of the same species in the Cavern of Baume, which M. Lartet considers to be of preglacial age,* in the Jura, associated with horse, ox, wild boar, elephant, and a non-tichorhine species of rhinoceros, the cave bear, and the cave hyena.’ The two teeth of Macherodus are a lower canine (? upper incisor), and a portion of the lower sectorial, both of which have serrated edges. The serration in _1 ‘Zool. et Paléont. Francaises,’ 1859, p. 231. 2 Gervais, ‘ Animaux vertébrés vivants et fossiles,’ Ato, 1867-9, p. 78, pl. xviii, 3, 3a, 30. * “Congrés Internationale d’ Anthropologie et d’Archeologie Prehistoriques,’ Paris volume, p. 269, 4 M. Lartet considers the rhinoceros to be non-tichorhine. 190 PLEISTOCENE MAMMALIA. the latter, as well as in the canine (incisor ?), renders it probable that all the teeth of this species were serrated. § 8. Antiquity in Kent’s Hole——We have now to consider the very difficult question as to the relative antiquity of the Macherodus latidens in Great Britain. Is it Pleiocene or is it Pleistocene? Was it a contemporary of the woolly Mammoth and Reindeer, or had it disappeared before the lowering of the temperature in the Glacial period? Untfor- tunately the peculiar physical conditions under which Kent’s Hole has been filled with its present contents forbids an answer which is absolutely decisive. In the present cave- earth, and underneath the stalagmite which now constitutes the floor, are large masses of breccia and of stalagmite, which evidently had formed a floor that had been broken up before the introduction of the cave-earth. They are remarkable for their hard crystal- line structure, and in one or two cases they have yielded fragments of very dense mineralized bones. In a portion of the cave, called the gallery, there is evidence of the undisturbed portion of the crust in a “ceiling” or uppermost floor, that extended from wall to wall, “without further support than that afforded by its own cohesion. Above it there is, in the limestone rock, a considerable alcove. ‘This branch of the cavern, therefore, is divided into three stories or flats; that below the floor occupied with cave- earth, that between the floor and ceiling entirely unoccupied, and that above the ceiling also without a deposit of any kind.” For a ceiling of this kind to have been formed it is absolutely necessary for the cave to have been filled up to its level with materials of some kind. It would, indeed, be as impossible for a solid calcareous sheet to be formed in mid-air as it would be for a sheet of ice to be formed without resting on water. From some cause or other this ancient stalagmite has been in part broken up, and the materials by which it has been supported have disappeared; and that it was deposited on cave-earth, like that now occupying the lowest story, is shown by its red colour. Prior, however, to its formation, animals dwelt in the cave, smce bones are imbedded in the large fallen masses of stalagmitic breccia. Moreover, there is reason to believe that certain fragments of bone and splinters of teeth, remarkable for their mineralization, that have been found in the earth now occupying the cavern, were derived from this more ancient deposit, for they differ essentially from the remains with which they are now associated, being heavier and of a more crystalline structure. Some splinters have assumed the fracture of greensand-chert. So hard, indeed, was one of the canines of Bear that it has been splintered by the hand of man into the form of a flint-flake, and has evidently been used for a cutting purpose. Its fracture proves that it was mineralized before it was splintered ; and as it was found in the present cave- earth, it must have been fashioned while the cave was being inhabited by palaolithic man, prior to the accumulation of the earth. For these reasons, the evidence in favour of these denser remains having belonged to the deposit which once supported the ancient floor seems to us incontrovertible. MACHAIRODUS LATIDENS. 191 To which, then, of these two periods of accumulation of cave-earth in Kent’s Hole can the Macheerodus be referred? Was it living at the time of the older deposit, and did it become extinct before the newer had been formed? It is impossible to give a distinct answer to these questions; but a careful examination of all the circumstances tends to the belief that the older period was that to which the Macherodus belongs. Since it is a species which differs but slightly from the I/. cultridens, and belonging to a genus which inhabited Europe in the Meiocene and Pleiocene ages, its affinities are undoubtedly Pleiocene, and it belongs to a group of animals that inhabited Europe before the lowering of the temperature brought about the mvasion of the Arctic Mammalia from the north and the east. On the other hand, in the teeth-marks on the incisors figured, as well as on the canines, we recognise the unmistakable traces that the animal to which they belonged fell a prey to the Hyena; and since the Pleistocene Hyena crocuta (var. syelea) is abundant in the cave, to its teeth the marks in question may pro- bably be referred. It seems, therefore, to us, to be almost certain that the animal inhabited Devonshire during an early stage of the Pleistocene, and most probably before the Arctic invaders had taken full possession of the valley of the English Channel, and of the low grounds which now lie within the hundred-fathom line along the Atlantic shore of western France. Along a great, fertile, low-lying region, which then was offered by what is now the bed of the sea, there must necessarily have been a swinging to and fro of animal life; and before the temperature of France had been sufficiently lowered to exterminate or drive out the southern forms, it is most natural to suppose that im warm seasons some of the southern Mammalia would find their way northwards, and especially a formidable Carnivore such as the Macherodus. ‘The extreme rarity of its remains forbids the hypothesis that it was a regular inhabitant of Britain during the Pleistocene age. It seems, therefore, to us that it belongs to the earliest stage in the complicated history of the deposits in Kent’s Hole, and that it probably became extinct before the great majority of Pleistocene caves in Great Britain had been filled with their present contents. This view of the extreme antiquity of Machzrodus in Kent’s Hole is materially strengthened by an animal which has been determined by Professor Busk among the Mammalia from the fissures of Oreston, near Plymouth. The Rhinoceros from that cave, considered by Professor Owen to belong to the tichorhine species, so common in the Pleistocene period, turns out to belong to the megarhine, which is a well-known Pleio- cene species. In that case, also, it is evident that the Southern forms of life still lingered on the British side of the valley of the English Channel, while the Pleistocene Mammalia were the normal dwellers in the British caves. Both these animals, therefore, may be taken to indicate an early stage in the Pleisto- 1 The Rey. J. MacEnery made the same remarks on the gnawed condition of the canines. MSS,, op. cit. 192 PLEISTOCENE MAMMALIA. cene period, and both, most probably, may be referred back to a time before the maximum point of cold was reached, in the Glacial period. § 9. Macherodus from Norfolk.—A small fragment of a right upper canine, in the collection of Mr. Jervis, probably from the Forest bed of Cromer, has been iden- tified, by Mr. E. R. Lankester’ with the Machzrodus. It consists of a portion of the crown, with the serrated edges very well marked. It is too small a fragment to enable any conclusion to be drawn as to the species. It shows, however, that the genus was living in the Eastern Counties at a still earlier period than that indicated by the remains in Kent’s Hole. 1 «Geol, Mag.,’ 1869, vol. vi, p. 440. CONCLUSION, 193 CHAPTER XXIII. CONCLUSION. We have now described all the members of the family of the Felidee which have been proved to have lived in Britain. Out of the six species we have been able to add two to the catalogue of British animals—the Panther or Leopard, and the Melis Caffer—and the latter of these has been hitherto unknown in Europe. Our investigations into the osteology of the living Lion and the Felis spelea have resulted in the identification of the fossil with the living animal, and the probable ex- tension of its range into North America. We have also brought forward the very curious historical evidence as to its retreat from Europe some two hundred years before the Christian era. The Lynx, and the Leopard or Panther, and the Felis catus and Fels Caffer, living during the Pleistocene age, have also been shown to be specifically identical with the living forms. The Macherodus latidens, an aberrant member of the Felide, is the only Pleistocene member that has become extinct. And since it is specifically distinct from the Pleisto- cene and Meiocene J. cultridens by the serration of its incisors, it is very probably a form that characterises an early phase of the Pleistocene period, a modification of the Pleiocene type that lived on into the succeeding geological age. In the following table we have given the range and distribution of the British Fossil Felidee : BRiIvTain. Evroprgan ConvmInEnNT. id List oF SPECIEs. 79 6 a Pleiocene. ie Be Historic. | Pleiocene. Res ice Historic. | © 5 Felis leo (spelea), Li. ...... x ae aah x x x x JEG Ii ard ON erew ion op ecoc karen aS x awe 36 x x x IP, POORONISY We, cooecoacasan nos 2 x Bee ben x JE, (Chih WORM: co codca sae x oe x x Scotus uit, te ere x x x A *« x ~< Macherodus latidens, Ow. x bee zh x< The presence of the Lion in Europe in Prehistoric times is rendered necessary from the fact that it is both Pleistocene and Historic, although it has not been discovered in any Prehistoric deposit. 26 194 PLEISTOCENE MAMMALIA. It may be that in the long interval which elapsed between the Pleistocene and the succeeding age, it had retreated from Northern and Central Europe, partly from the competition with man, and partly from the operation of the same obscure causes which banished the Spotted Hyena and the Hippopotamus to Africa. It will be seen from the above table that the Wild Cat is the only British feline which still lives, and the time is not far distant when it will become extinct in our island. TABLE OF CONTENTS. INTRODUCTION Felis spelea.—Lower Jaw — Ulna and Radius — Os innominatum — Tarsus —— Hind Paw Felis spelea.—Skull — Teeth -— Vertebree, Sternum — Scapula — Humerus . —— Femur — Tibia, Fibula, Patella Felis spelea.—Carpus . : 2 = Metacarpals, Phalanges — Limb-bones of the Whelp . — Identical with F. leo — Range of Dae p. I, chap. I, pls. I, VI. p. 6, chap. II, pls. 11, XXI1a. p. 10, chap. IIT, pls. III, XXiIs. p. 13, chap. IV, pl. IV. p. 21, chap. V, pl. V. PART II. PART — Retreat of the Lion from Europe Felis lynz Felis pardus . o Car - PE CALUS) : Macherodus latiden ConcLUSION PART p. 30, chap. VI, pls. VI, VII, VIII, IX, X. p. 65, chap. VII, pls. I, VI, VIII, XI, XII, XIII p. 85, chap. VIII, pls. XIV, XV, XVI. p. 109, chap. IX, pl. XVII. p. 113, chap. X, pl. XVIII. p. 118, chap. XI, pl. XVIII. p. 122, chap. XII, pl. XIX. Il. p. 128, chap. XIII, pl. XX. p. 133, chap. XIV, pls. XIX, XX, XXI. p. 143, chap. XV, pl. XXII, XXTTa, XXIIz. p- 146, chap. XVI. p. 151, chap. XVII. p. 164, chap. XVIII.” p. 173, chap. XIX, pl. XXIII. IV. p. 177, chap. XX, pl. XXIV. p-. 181, chap. XXI, pl. XXIV. p. 183, 5 55 4 p. 184, chap. XXII, pl. XXV. p. 193, chap. XXIII. PHATE at: Felis spelea, Goldfuss. (Natural size.) Brent : ; ay ‘i oy: 1. Lower jaw, external aspect. | These are from Bleadon Cavern, and form part of the — 2. Lower jaw, internal aspect. Collection of Mr. Beard in the Taunton Museum. Beiac _ 3. Lower jaw, posterior aspect, showing condyle, coronoid process, and angle. From — the Brickearth of Crayford. In the possession of Dr. Spurrell. W.A.S. del. J. Dinkel ith. IPILAVICN) Il hh An A W.West imp. PELIS SpgLEA. Lower Jaw, ee PLATE 1. SERRE Sy ieaeeseeeeate eae ‘W.West imp. / Fic. i Or PLATE II. Felis spelea. FOREARM. (Half natural size.) Radius. Sandford Hill Cavern. Mr. Beard’s Collection in Taunton Museum. RS.'S. 4. Bel. . Radius, proximal articulation of fists . Radius, distal articulation of ney: - Radius, small form. Bleadon Cavern. Mr. Beard’s Collection at Taunton. Bl. B. FS(L). 1. . Ulna, largest specimen known to authors (proximal half). Bleadon Cavern. Mr. | Beard’s Collection at Taunton. BI. B.U.FS.1. . Ulna, external view of fig. 5. . Ulna, internal view of fig. 5. . Ulna, proximal end. Sandford Hill. Belongs to same individual as the radius, fig. 1. FS.U.1.S.H.B. Taunton. Mr. Beard’s Collection. - Ulna, small form, external view. From Wokey Hyzna-Den. Mr. Boyd Dawkins’s Collection. FELIS SPELABA. | PLATE IL. hah. STH: WBidgood dal. JDinkel lith, Sc Prats vid PLATE III. Felis spelea. OS INNOMINATUM. (I'wo thirds of natural size.) Fic. 1. Os innominatum of Felis spelea. Brick-earth, Slades Green, Crayford. Museum. 2. Os innominatum of Fels figris. India. In British Museum. British meee crea Cheeta beng ce. PLATE IV. Felis spelea. LEFT TARSUS. (Natural size.) Fic. 1. Astragalus. 2. Calcaneum. Naviculare, reversed from right paw. Cuboid. 4’, Cuboid, internal aspect. 4”. Cuboid, distal or metatarsal aspect. 5. Hcto-cuneiform, reversed from right paw. 5’. Kcto-cuneiform, internal aspect, right paw. . Ecto-cuneiform, internal aspect. Lion. 5”. Keto-cuneiform, internal aspect. ‘Tiger. 6. Meso-cuneiform. i Endo-cuneiform. Figs. 1, 2, 3, 4, 5, 6, 7, show the front or superior aspect of these bones ; all those of Felis spelea ave from Bleadon Cavern, and formed part of the Collection of Mr. Beard, now in the Taunton Museum. FELIS SPELAA. dap, West. Var 7 WAS del. J_Dinkel th, Ae ¥ oo ll 0 . ? thle : ‘ , * iy A . ' ri \ S : = 4 2 ' A y d ie. c > \. ri . y . ~ et neat nn EE Ree are ee ogi PLATE V. Felis spelea. RIGHT HIND PAW. (Natural size.) . Metatarsal 1. Bleadon Cavern. . Metatarsal 2, : 5 BN iets ro From Sandford Hill Cavern. ‘These have the appearance of having . Metatarsal 4. . Metatarsal 5. from a left paw. all belonged to the same individual. Metatarsal 3 is reversed | First phalanges. From Bleadon Cavern. Second phalanges. From Bleadon Cavern. . Third phalange, probably of the fifth toe. From Bleadon Cavern. i Sesamoids. From Bleadon Cavern. V. PLATE : eal fa dp) dp) =! eI cx] Ay a5 del, J. Dine oO e PLATE VI. Fehs spelea, Goldfuss. (Skull: small form: lateral aspect.) The skull belonged to Mr. Williams; the jaws to Mr. Beard. . From either Sandford Hall or Hutton Caves in the Mendip. Now in the ‘Taunton Museum. { ‘ $ i ~ WwW Bidgood del. et. lith PELA A. PLATE VI. W.West imp £41 0) igi FELIS SPELAA PLATE VI. W West imp W Bidgood del ot ith SKULL Pont he re oe i PLATE VII. Felis spelea, Goldfuss. Skull: small form: natural size.) Skull: the same as in Plate VI; inferior or palatal aspect, showing the dentition. | W Bid good, del et lith LIS SPELAA PLATE VIL. W. West imp W Bidgood,del et lith FELIS SPELEA PLATE VIL. W. West imp. nM z PLATE VILL. Felis spelea, Goldtuss. (Skull : small form: natural size.) Skull: the same as in Plate VI; upper or frontal aspect. N.B.—The reference to Plate VIII, with regard to Dr. Spurrel’s metatarsals, in page 22, must be transferred to Plate XXIII. * fey ~ - ne ' 4 \ ‘ ; f “X e : R x , i 3 iG . . fy i byt a A ; 7 , a4 as = : ee Y 1 { i i} [ i ai) oe te f i }, 7, ns f ~ = Fie. ely: PLATE XII. Felis spelea, Goldfuss. LOWER PERMANENT DENTITION. (Natural size.) First left incisor, anterior or outer and posterior or inner aspects. 2, 2', 2”. Second left incisor, anterior or outer, posterior or inner, and inner lateral aspects. 3, 3’, 3. Third left incisor, anterior or outer, posterior or inner, and outer lateral aspects. These three 4. “SI teeth are from Sandford Hill. No. 2 and 3 belong to mandible figured in PI. I. Right canine, outer lateral aspect: the largest known tothe authors. Probably from Bleadon Cave. Mr. Williams’s Collection, Taunton Museum. Right canine, inner lateral aspect ; rather small size of large form. Sandford Hill Cave. Mr. Beard’s Collection, Taunton Museum. Left canine, outer lateral aspect, of very oldanimal. Probably from Bleadon Cavern. Williams’s Collection, Taunton Museum. Third right premolar; largest known to the authors; outer lateral aspect. Bleadon Cave. Mr. Beard’s Collection, Taunton Museum. Third left premolar, inner lateral and coronal aspects, of the average size of large form. Bleadon Cave. Mr. Beard’s Collection, Taunton Museum. Third left premolar, inner lateral aspect: the smallest known to the authors. Wookey Hyzna- den. In possession of Mr. Boyd Dawkins. Fourth right premolar, outer lateral aspect : the largest known to the authors. Bleadon Cave. Mr. Beard’s Collection, Taunton Museum. Fourth left premolar, inner lateral and coronal aspects: of the average size of large form. Probably from Bleadon Cave. Mr. Williams’s Collection, Taunton Museum. Fourth left premolar, inner lateral aspect: smallest known to the authors. Probably from Bleadon. Mr. Williams’s Collection, Taunton Museum. Right true molar much worn, outer lateral aspect: the largest known to the authors. Wookey Hyzna-den. In possession of Mr. Boyd Dawkins. Left molar, of the average size of large form, inner lateral and coronal aspects. Probably from Bleadon. Mr. Williams’s Collection, Taunton Museum. Left molar, inner lateral aspect: smallest known to the authors. Wookey Hyzena-den. In possession of Mr. Sanford. PLATE, Xi J fe ILS (Sue ve TG AeA, = EN Se Moy West. imp 5; plug > 4 i} =i IN eae LOWER PERMANI} W.A. Santora del W™ Bidgoodlith 1, 8 i ) i ri . f Rial i i { ne * ‘ fh ” i \ . k Cana te U i i ; : ‘ ‘ + 5 . ‘ 1 a i { t = i F j Hi ! : i i i i ind i Y Wy! FIG, Lal ell Oy 3, 3. Or 6. PLATE XIII. Felis snliea, Goldfuss. MILK DENTITION. (Natural size.) . Outer, inner, and inferior aspects of left maxillary. The small premolar 2 inserted in fig. 1” is from that figured in fig. 2. Hutton Cave. Outer and inner aspects of anterior portion of left maxillary. Bleadon Cave. Outer and posterior aspects of right lower jaw, the crown of the canine and the incisor copied from other specimens, the lines of restoration bemg shown in the canine, and by the junction of the incisor with the jaw. Hutton Cave. Inner aspect of part of right ramus of young individual. From Sandford Hill Cave. Upper canine, inner aspect. Probably from Sandford Hill Cave. Upper milk molar 3, inner aspect. Sandford Hill Cave. Lower canine, inner aspect. Probably from Sandford Hill. This tooth furnished the restoration for that in fig. 3. Lower milk molar 3, inner aspect, of large size. Bleadon Cave. All the above are in the Taunton Museum, and were found by Messrs. Beard and Williams in the caverns of the Mendip Hills. iat 18h JIL SPELAA. PLATE XII. dy 3 WA.Sanford del, W. Bi dgooa Lia WWest. timp MILK DENTITION ; ' i ; ~ , ‘ t - 7 i Os +4 7 . re ‘ Ad S) ap A a ae i Vib bevptabore eri PLATE XIV. felis spelea, Goldfuss. VERTEBRA. (Natural size.) Fic. ee 1A Proximal, dorsal, and distal aspects of atlas. Sandford Hill Cave. The restoration of the transverse process is slightly enlarged from the cast of a specimen from Gailenreuth, in the possession of Sir Philip Egerton. It is to be seen in the British Museum and the Museum of the College of the Surgeons. 2, 2’, 2”, 2”. Dorsal, lateral, proximal, and distal aspects of sixth cervical vertebra. Sandford Hill Cave. 3, 3, 3°, 3. Proximal, dorsal, lateral, and distal aspects of seventh caudal vertebra. Bleadon Cave. All these specimens were found by Mr. Beard, and are now in the Taunton Museum. The following letters are used for the different parts of the vertebrae in Pls. XIV, XV, XVI: ce, centrum. d, diapophysis. * ae, anterior epiphysis. a, anapophysis. pe, posterior epiphysis. m, metapophysis. m, neurapophysis. az, pre-Zzygapophysis. ms, neural spine. pz, post-zygapophysis. pa, parapophysis. ne, neural canal. pl, pleurapophysis. v, canal for vertebral artery. hy, hypapophysis. W.A.Sanfourd del, W bids pod hth PLATE XIV. W.Weat imp An) ldel, W B eee in At PLATE XV. Felis spelea, Goldfuss. VERTEBRA. (Natural size.) Fic. 1, 1’, 1”. Proximal, lateral, and distal aspects of a perfect second dorsal vertebra. ford Hill Cave. Mr. Beard’s Collection, Taunton Museum. Sand- ee VUdaludaA IvySsuod aNooags BH PAU LP Tp PLIOFWEG YM AX SRE! Vai lads sitma Rt att i end! Viale , AL ee PLATE XVI. Felis spelea, Goldfuss. VERTEBR& ; STERNUM. (Natural size.) Fic. eal Seventh dorsal vertebra, distal and lateral aspects. 2, 2’, 2”, 2”. Eleventh dorsal ; lateral, distal, proximal, and dorsal aspects. 3, 3’, 3”, 3” Second lumbar vertebra, proximal, lateral, dorsal, and distal aspects. A, 4’. Fourth caudal vertebra, ventral and distal aspects. 5. Ninth caudal vertebra, dorsal aspect. 6. Tenth caudal vertebra, ventral aspect. ed Eleventh caudal vertebra, dorsal, proximal, and distal aspects; very large specimen. 8. Twelfth caudal vertebra, lateral aspect. 979", 9" Fourteenth caudal vertebra, dorsal, proximal, and distal aspects. 10, 10°. Third sterneber, lateral and ventral or superior aspect. The above are in the Taunton Museum, and were in Mr. Beard’s Collection. All were derived from Bleadon Cave, except No. 3, which was from Sandford Hill. Nos. 5, 6, 8, 9, and perhaps 4, have the appearance of having belonged to one animal. W™ Bidgood del et. Jith. May,18 67, XW JE MG AMAL 18, linp. West W ZS LEENA S). eS ~— PLATE XvVt Wo Went snp W® Bidgood dol at.lith May 1867 TERR TERY ATER R an VEBE-*SRE, STERNALS. shone PLATE XVIL Helis spelea, Goldfuss. _ SCAPULA. (Natural size.) Fie. | EDs 1. Glenoid cavity and distal surfaces of the right scapula. From Sandford Hi Ca _ Found by Mr. Beard; now in the Taunton Museum. ie 2. Outer or superior surface of the same bone. W* Bidg ood del. et Jith JP yA IS) JOVI, W. West imp. ) FELIS spELEBA. PLATE XVII. W Westimp W™ Bidgood del.et, hith PLATE XVIII. Felis spelea, Goldfuss. HUMERUS. FEMUR. (Natural size.) FIG. 1. Composite figure of the posterior or palmar aspect of the left humerus. The ground- work in pale tint is taken from Dr. Schmerling’s great work on the caverns of Liege (‘Oss. Foss. de Liege,’ vol. u, pl. xv, fig. 2). The distal end is from a humerus found in Bleadon. The small compressed shaft was found in Oreston, and is now preserved in the Bristol Museum. The imperfect proximal and distal articular portions are from Bleadon Cave. All the specimens from this latter cave are in the Taunton Museum. 2. Anterior aspect of distal end of left humerus, obtained by the Rev. H. H. Winwood, F.G.S., from the gravel of Larkhall, near Bath. 3. Distal articulation of very large humerus, left side. Sandford Hill Cave. Taunton Museum. 4. Composite figure of left femur. A cast of a specimen obtained by Sir Philip Egerton, from Gailenreuth Cave, supplied the groundwork in light tmt. The proximal and distal ends and the fragment of shaft were obtained from Bleadon Cave by Mr. Beard, and are now preserved in the Taunton Museum. 5. Distal articulation of femur. Bleadon Cave. ‘Taunton Museum. ~~~ Ne, I FELIS PLATE XVIII. ue eh Ibe UTS aoe f ow ‘) = fz] B. so) te) ag ied = = =) FO els PLATE XVIII. HUMERU cae ite: oe ve wa th oh a Or Or PLATE XIX. felis spelea, Goldfuss. TIBIA. FIBULA. PATELLA. METACARPAL. METATARSAL. (Half natural size.) Left tibia of young adult, anterior aspect ; perfect, with the exception of the proximal articulation. Sandford Hill Cave. Taunton Museum. Proximal end of shaft. _ Distal articulation. Anterior aspect of proximal end of left tibia. Sandford Hill Cave. Taunton Museum. _ Proximal articulation of the same bone. Posterior aspect of right fibula, which probably belonged to the same animal as fig. 2. Sandford Hill Cave. Taunton Museum. External aspect of distal end of left fibula. Bleadon Cave. Taunton Museum. Anterior aspect of patella. Sandford Hill Cave. ‘Taunton Museum. . Posterior aspect of ditto. (Natural size.) Second left metacarpal of gigantic size. Lower Brickearths, Crayford. Dr. Spurrell’s Collection. Second left metatarsal of gigantic size. Lower Brickearths, Crayford. Dr. Spurrell’s Collection. These are referred to in p. 22 as figured in Pl. VIII. JE IWATE 1S, F.0D. SIP a IL, Aa, AN PEGS W West imp. J Dinkel del etith, BONES OF LIMBS Fic. ce PLATE XX. Felis spelea, Goldfuss. CARPUS. METACARPALS. (Natural size.) Scaphoido-lunare, anterior or dorsal aspect. Sandford Hill Cave. ‘Taunton Museum. Distal aspect of same. Scaphoido-lunare of the small form, anterior or dorsal aspect. Bleadon Cave. Taunton Museum. Distal aspect of pisiform. Bleadon Cave. ‘Taunton Museum. Proximal or ulnar aspect of pisiform. Sandford Hill Cave. ‘Taunton Museum. Anterior or dorsal aspect of unciform, right side. Sandford Hill Cave. ‘Taunton Museum. Distal or metacarpal aspect of same bone. Proximal end of fourth metacarpal of the ordinary form and size. Bleadon Cave. Taunton Museum. Fifth metacarpal internal aspect. Wookey Hyana-den. Mr. Boyd Dawkins’ Collection. PREIS SPHL AA. + PLATE XO W A.Sanford del. W™ Bidg oad lith. W West imp FIG. py WD 2 oR wo we PLATE XXI. Felis spelea, Goldfuss. RIGHT FORE PAW. (Natural size.) First metacarpal. Bleadon Cave. Remaining metacarpals from the same individual. Sandford Hill Cave. The first phalanges, apparently belonging to the individual as the last. Second phalange of second digit. Bleadon Cave. Second phalange of third digit. Bleadon Cave. Second phalange of fourth digit, apparently belonging to the same individual as the set of metacarpals and first phalanges. Second phalange of fifth digit (reversed), from left paw. Bleadon Cave. All these were found by Mr. Beard, and are now preserved in the Taunton Museum, W.Weat amp. Tey AVI I), HOS SPBILAGA. FORE—PAW ls JS) W.A Sanford del W" Bidgood hth. bs \ t 5 ~ , ° t “ ¢ i 4 1 i ‘ * d] . Bary Cyt OU . hae ’ aye Lid Coe ont DB Ot FS WwW WHO = pd S PLATE XXIL Felis spelea, Goldfuss. Bonzs or WHELP. (Natural size.) . Left humerus, posterior aspect. . Right ulna, external aspect. . The same, internal aspect. . Left fifth metacarpal, dorsal aspect. . Hirst phalange of one (second or fifth) of the outer digits of the hind paw. . Second phalange, dorsal aspect. . Right femur, posterior aspect. . Left femur, front aspect. . Shaft of left fibula, tibial aspect. . Left caleaneum, external aspect. These bones, with the exception of 6 and 9, were found in Hutton Cave by Messrs. Beard and Williams, and probably belonged to one animal. The two exceptions were found in Bleadon Cave. Museum. All are now in the collection of cave-mammals in the 'l'aunton Bs } Oe ) y fe aap . W.West IPIL VAMP, 2O-QUE ae) r ; ca. Rs a ‘i y i a { * ee AMANO PG HAG my, ts tt HAL itt F Bey } Or 6. PLATE XXII a. Felis spelea. Goldfass. ULNA. RADIUS. TIBIA. FIBULA. PATELLA. (Natural size.) Internal or ulnar aspect of right radius, represented in PI. I, figs. 1, 2, 3, described at pp. 8, 9. Sandford Hill Cave. Internal aspect of proximal half of right ulna represented in PI. II, fig. 8, described at pp. 6, 7. Sandford Hill Cave. Composite figure of external aspect of right tibia. The proximal portion is repre- sented in P]. XIX, figs. 2, 2’, the shaft and distal end in Pl. XIX, fig.1@. Both are described at pp. 122—4. Sandford Hill Cave. External or tibial aspect of shaft of right fibula, represented in Pl. XIX, fig. 3, described at pp. 125—6. Sandford Hill Cave. Internal aspect of distal end of right fibula, represented in Pl. XIX, fig. 4, described at pp. 125—6. Bleadon Cave. Lateral aspect of patella, described at p. 127. Bleadon Cave. We have added this and the following supplementary Plate X XII B in consequence of the desire, which has been expressed, that all the plates should be of life size. In no case has the same aspect of the same bone been repeated. PLATE Kxe A RELI S SPILL Stee sen ~ W.West imp. , nat.size LIMBS BONES OF THE W* Bidgood del.ct lth. 1) Ri, A €) i} PLATE XXILA. FELIS SPELAA WWest imp BONES OF THE LIMBS, nat.size ° dSood del.et th. q WwW? B PLATE XXII s. Felis spelea, Goldfuss. OS INNOMINATUM. VERTEBRA. (Natura! size.) Fig. 1. External aspect of right os innominatum. Sandford Hill Cave. 2. Internal aspect of left os inmominatum. ‘This pair probably belonged to a Lioness, on account of the slenderness and curvature of the os pubis. The texture of the bone closely resembles that of the sacral vertebra, fig. 4, and several other young bones from the same locality. The animal, therefore, to which it belonged, though of full size, was probably young at the time of its death. Sandford Hill Cave. 3. Os pubis of very large animal, much stouter and less curved than that of fig. 2. Sandford Hill Cave. 4. Third sacral vertebra, with portion of the second (see p. 96). Sandford Hill Cave. i W™ Bidgood del et lith. v7 +a” é s "7 t y | es ad FHLIS SPELAA. PLATE XXIIB. W'Bidgood del eb lith PELVIC BONES, nat.size —E——————— Fic. PEATE SOX: Lclis Lyne, Linneus. SKULL AND LOWER JAW. (Natural size.) Occipital aspect of skull. Yew l'ree Cave. Dr. Ransom’s Collection. Basal aspect of fig. 1. External aspect of right lower jaw. Yew Tree Cave. Dr. Ransom’s Collection. Internal aspect of fig. 3. Superior ditto. Posterior ditto. Exterior aspect of right lower canine, taken from the lower jaw. "LXX XX LLIVId XNATT SL TBC ae WL? BP Top ¢ Fic. —~ DP oe PLATE XXIV. Felis pardus, Linneus. (Natural size.) Internal aspect of upper canine. Bleadon Cave. Taunton Museum. Lower true molar. a 5; Internal aspect of lower canine. Re 9 Kixternal aspect of lower canine. Banwell Cave. In the Collection of the Earl of Enniskillen. Posterior aspect of left femur. Taunton Museum. Felis Caffer, Desmarest. Left mandible. Bleadon Cave. Taunton Museum. Posterior aspect of upper portion of femur, which probably belongs to this species. Bleadon Cave. Taunton Musenm. Radial aspect of ulna, which may belong to Melis catus. Bleadon Cave. ‘Taunton Museum. Felis catus, Linneeus. Left mandible, from the Lower Brickearths of Grays Thurrock, Essex. In the possession of J. Wickham Flower, Esq., F.G.S. The letters attached to the figures of this plate indicate the same parts as those which have been used in the plates of Felis Ico, var. spelea. In this work we propose to use one system of letters for the teeth and another for the bones of the Carnivora, since in that way the homologies can be shown with the greatest precision. SOL. 4 PATE IP CAIUS CAFFER, PUAIRUD US 5 AE ILLS) Sey aS W. West imp delet lith dfgooa Bid WwW FELIS PARDUS= ANTIQUA (CAI IMFIRE|(S) 3G CAIUS) Tee Ce eee yo oe Pa ae ee ee ak ae te ee ee PLATE XXV. Macherodus latidens, Owen. (Natural size.) This plate is copied from that which was intended to have formed a portion of the ‘Cavern Researches’ of the Rev. J. MacKnery. Fig. yy Left upper canine of young adult, inner aspect. British Museum. Ch) outer aspect. > FA posterior aspect. 3 4, Right upper canine of adult, outer aspect. Museum of the Geological Society of London. inner aspect. 2) 39 6. Right upper canine of adult, inner aspect. In possession of Sir Walter T're- vellyan, Bart. ~ 2» 5: posterior aspect. Plate XXV, roe ae, Litecea een steams ad Mary Buckiina de! « GScharf athog Nat, size. “Teeth of Ursus (Cuiltridens Found in the Cave of Kents Hole near Torquay, Devon, by Rev &M! Me Knery vont! 1646. ium diluvial Mud mixd with Teeth and gnowd Bones of Rhinoceros, Elephant, Horse, Ox, Elk 2 Deer, with Teetn &.Bones of Hycenas, Bears, Wolves, Foxes &c. MACHARODUS LATIDENS. (OWEN). reper rey er Ae an a iia | + eee | ||. = SSeS ees pra. Lm Ps, tel aie FPA, PAP AP Ah ea \ ae aie! 2 Ma! 114 Vana yA Poe EOE PT pet tat Fe Ina \\a*Na VA PRAT | i rhe AAAs apnan, aH poea hataiitaiiy, AA’ map fia’ Aap | AMMA Muh WM vena, ha An A. pPaAARP EA? Aw kg ot Paeasgel ie a AAna AA, pAAAAAAP AA, nA t | on aN ee ae eo Be - - = ae al = (a Va ys Nae A Raa PhaPePar”p, LOPS @ « TY, a ashe Par aT Beata os ee ee ETAT OOAANT AE anne Aal as, : A f a Caton fr PRPs) . 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