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Table of Contents of a Vol. IT =" 1911 February, 1! Cave Hyena, pp. 1—25 ; -—y 1902 December, Bears, pp. 1—35 LWiaee 1906 | December, Canide, pp. 1—28 _I—VI 1909 December, 1£ 09 . Mustelide, pp. 1—28 I—VI 1911 =| February, 1912 ae ae =F ia =p ae gut gs 6 >: A MONOGRAPH OF THE BRITISH PLEISTOCENE MAMMALIA VOR LT BRITISH PLEISTOCENE HYAINIDAL, URSIDA, CANIDAZ, AND MUSTELIDAL. SDN NOSES ON ODS Ma HGS. PROFESSOR OF GEOLOGY IN THE UNIVERSITY OF BRISTOL. LONDON: PRINTED FOR THE PALHONTOGRAPHICAL SOCIETY. 1902—1912. PRINTED BY ADLARD AND SON, LONDON AND DORKING. TABLE OF CONTENTS PART I Hyzxna crocuta.—Historical Introduction an Best Sopa le — Distribution ae 8g a - — Comparison of Cave, Brown, and Gerzrly Bears 56 son JO PD — Bibliography ... wi Au 600 se soa Os BPA, | i | PART III. Canis.—Historical Introduction pale —_— Distribution of Canis lupus : ao: a won 100 De — — C. familiaris me on a tog [D0 Bb — — C. vulpes Dp tho — — C. lagopus... As aa 1 Ch — Skeletal Differences between Common and Arctic Foxes p. 8. Lycaon anglicus .. oo ry bp foe eS Canis. Sy mibation | in British River Deporte 3 — Distribution in British Caves , one as ace (D4 10) — Skull p. ll, pls. I—IV. Canis.—Dentition ; — Vertebral Column —_— Limb Girdles _ Limbs ... as, m HB i Mutual Relations of Pleistocene and Post-Pleistocene Canide ... Conclusions Bibliography Au alive Mustelidz. —Introduction Mustela martes Mustela robusta Mustela putorius Mustela erminea Mustela vulgaris Gulo luscus ... Meles tazus ... Lutra vulgaris af Me a aya 2 Distribution in British Pleistocene ae? ee! naa peal ee Skull _ shel ogee e; | p13, pk, =e Dentition . . Pae bo ae. 33. a. (p. 15, pls: 1 Vile Vertebral Column... er me ae Ta Deeps aval Limb Girdles nae oe oe “ 2) ieali7s Limbs Be a aa ne ae ey soo ob PAD), Conclusions ... Pera dy a ae ea deo 0b PDs Bibliography ae oe dat wie re peo: THE % - PALBONTOGRAPHICAL SOCIETY INSTITUTED MDCCCLXVIL. VOLUME FOR 1902. LONDON: MDCCCCII. 6) a =e ek eer on 2 a a i 2 Seo fe A MONOGRAPH OF THE BRITISH PLEDSTOCENE MAMMALIA VOL. IL, PART TI. THE CAVE HYAINA. BY SIDNEY H. REYNOLDS, M.A., F.G.S., PROFESSOR OF GEOLOGY AND ZOOLOGY, UNIVERSITY COLLEGE, BRISTOL. Paces 1—25; Puates I—XlLyV. ILO IW IDOINS PRINTED FOR THE PALHONTOGRAPHICAL SOCIETY. 1902. PRINTED BY ADLARD AND SON, BARTHOLOMEW CLOSE, E.C.; 20, HANOVER SQUARE, W.; AND DORKING. ~ a MONOGRAPH ON THE BRITISH MAMMALIA PLEISTOCENE PERIOD. THE CAVE HYAINA. Order—CARNIVORA. Faminy—HY ANID. Genus—HYaANa. Species— Hyena crocula, Krxleben. I. HISTORICAL INTRODUCTION. Tun early history of the recognition of remains of the hyzena in Europe is dealt with by Cuvier, and much use has been made of his account in the following pages. The first evidence for their occurrence is afforded by a figure of part of the right mandibular ramus given by Kundmann in his ‘ Rariora Nature et Artis,’ published in Breslau in 1737. He regarded this as similar to that of a calf, but its hyeenine nature was recognised by Cuvier. Thirty-seven years later (1774), Hsper figured bones from Gailenreuth—an atlas which he regarded as hyzuine, but which Cuvier says is that of a bear,—and some teeth, which he regarded as belonging to a lion, but which Cuvier says are hyzenine. Again, in 1784, Collini gave an excellent figure of a hyzena skull found near Mannheim. Unfortunately, however, he was disposed to regard it as perhaps that of a seal, l 2 PLEISTOCENE MAMMATIA. The first full account of the cave hysena was that given by G. Cuvier in 1812." He mentioned a number of Continental localities in which bones of hyzenas had been found, and considered that the fossil hyzena was distinct from any living species, basing his opinion at that time mainly on the great size of many of the fossil bones. The occurrence of the cave hyzena in England was first clearly established by Dean Buckland in his account of the Kirkdale Cave.’ ‘The full title of this important paper, which was published in 1822, is “ Account of an Assemblage of Fossil Teeth and Bones of Elephant, Rhinoceros, Hippopotamus, Bear, Tiger, Hyzena, and sixteen other Animals discovered in a Cave at Kirkdale, Yorks, in the year 1821, with a Comparative View of five similar Caverns in various parts of England, and others on the Continent.” In this paper, and in his ‘ Reliquize Diluvianz’ (1824), he clearly showed that the caves in which the hyena bones were found were the actual dens of the animals. Buckland’s discovery of hyana remains at Kirkdale was closely followed by Clift and Whidbey’s discovery of them at Oreston, near Plymouth. Goldfuss,* writing in 1823, was the first to apply the distinctive name Hyena spelea to the cave hyana. He gave a detailed comparison with figures and measurements of the bones of the cave species and of the spotted hyzena, In the second edition of the ‘ Ossemens Fossiles’ (1823), Cuvier, in giving a further account of the cave hyzena, referred specially to what he held to be the differences between it and the spotted hyena, and mentioned, with regard to the metacarpals and metatarsals, that all the bones measured were, without exception, shorter and thicker in the cave hygena than in the spotted hyzena. With regard to the teeth, however, the general tendency of lis remarks implies that it is impossible to distinguish those of the one from those of the other. Meanwhile the discovery and study of hyzena remains were actively pursued on the Continent, and a number of new species of hyena, some allied to the living ZH. crocuta and some to the living /Z. striata, were described by Croizet and Jobert* (1828), and by Marcel de Serres, Dubrueil, and Jeanjean® (1839). ‘Throughout the first half of the nineteenth century little doubt apparently was felt by paleeontologists- that the cave hyena was distinct from the spotted hyena. ‘hus de Blainville’ (1844), Pictet ® (1844), and Owen ® (1846) all accepted this view. De Blainville discusses the question in detail (vide postea), and bases his opinion mainly on the form of the upper molar. The first paleontologist to express strong doubts as regards the specific distinction of the cave and the spotted hygenas was Gaudry ” (1863), but Boyd Dawkins,” writing in 1865, was the first definitely to conclude that no distinction could be drawn between 1 “Oss. Foss.,’ ed. 1, iv. 2 © Phil. Trans.,’ exit (Gl) apsevale Seo hil ransom Cxiiemp OS: 4 «Saiig. Vorw.,’ vi. 5 «Oss. Foss. Puy de Dome.’ 6 «Oss. Lunel Viel.’ f 7 © Osteographie,’ livr. 14. 8 «Traité Paléont.,’ i, p. 180. 9 «Brit. Foss. Mamm.,’ pp. 138—160. 10 « Bull. Soc. Géol, France’ (2), xx, p. 404, ‘Nat. Hist. Rev.,’ n.s., v, p. 80, i | HYAINA CROCUTA. 3 Hf, spelea and FL. crocuta. He laid stress on the variable character of the tubercular portion of the lower carnassial, and considered that several of the supposed species that. had been founded by Croizet and Jobert, and by de Serres, Dubrueil, and Jeanjean, mainly on variations in this tooth, were not valid, but were varieties of the cave hyena (vide postea). In his paper on the mammal fauna of the Creswell Crags,’ published in 1877, the same author says that, after comparison of the skulls of 7/7. crocuta and H. spelea, he has been unable to detect points of difference of specific value, and definitely states that he believes the two to be identical. Busk, however, writing in the same year,’ while recognising the close relationship between the two forms, said that he did not consider it proved that 7. spelea was a mere variety of H. crocuta. Since the publication of Boyd Dawkins’ paper in 1863, almost all authors have accepted the view of the identity of the two forms. This has been done, for example, by Newton® (1883), Lydekier* (1884-5), Forsyth Major’ (1885), Woodward and Sherborn ® (1890), Gaudry’ (1892), and Zittel® (1893); so that the fact of their identity may be considered to be clearly established. Schlosser,’ however, expresses doubt as to their identity, mainly on account of the geographical distribution of “ZZ. erocuta at the present day. A later phase in the study of hyaenas has been the discussion of the mutual relation- ships of the fossil forms, and the probable ancestry of the living ones. ‘This subject has been most fully dealt with by Lydekker,” Schlosser,’ and Gaudry.’ Lydekker, basing his opinion largely on its occurrence in the Pleistocene Caves of Karnul, in the Madras Presidency, considers that Hyena crocuta originated in India, being derived from the Siwalik (Lower Pliocene) Hyena Colvini, Lyd. The lower carnassials of the two forms agree closely, especially as regards the development of the cingulum, differing chiefly in the relatively large development of the hind talon in HZ. Colvint. Schlosser derives the cave hyena, and eventually HZ. crocuta, from the Upper Pliocene 7. Perrieri of Croizet and Jobert. He derives H. Perrzert from an unknown form whose nearest ally was /7. sivalensis, and he regards H. Colvini as altogether off the line of descent in question. Gaudry also derives H. crocuéa (including the cave hyena) from ZH. Perrieri, but expressly states that he has not taken account of the Indian species, not being personally acquainted with their fossil remains. The subject of the mutual relationship of the different species of hyzena lies, however, too much beyond the scope of the present monograph to be fully dealt with. 1«Q. J. Geol. Soc.,’ xxxiui, p. 596. 2 «Trans. Zool. Sve.,’ x (2), p. 53. 5 * Geol. Mag.,’ 1883, p. 433. 4 «Pal. Indica,’ ser. 10, ii, p. 275; ‘Catal. Foss. Mamm. Brit. Mus.,’ i, p. 69. 5 ‘Zool. Record,’ 1895, p. 28. 6 Hyzena antiqua, Laukester, ‘Ann. Mag. Nat. Hist.,’ ser. 3, vol. xiii, 1864, p. 56, pl. viii, figs. 5, 6, from Red Crag, Suffolk. ’ “Geol. Mag.,’ 1883, p. 433. 8 , oe PLEISTOCENE MAMMALIA. COMPARATIVE MEASUREMENTS OF HymNA VERTEBRE (continued). Gth cervical. 7th cervical. crocuta, Wookey (Taunton Surgeons). Museum’, A. Museum), B. HI, crocuta, No. 522 (College of Surgeons). Museum), A. eum), B. EL. crocuta, No 522 (College of Mus Wookey (Taunton Wookey (Taunton Wookey (Taunton H, spelea = crocuta, HH. spelea = crocuta, EZ. spelea = crocuta, EZ. spelea Maximum length of centrum | from dorso-anterior to ventro- posterior edge Length from dorso-anterior to dorso-posterior edge of cen- Width across transverse pro- CBHI. cooaacoonssooavqneseds20000 Width across postzygapophyses Height from roof of neural canal to top of neural spine ... Length of inferior lamella of transverse process ........... Sno 1st thoracic. 2nd thoracic. 3rd thoracic. 4th thoracic. crocuta, (College of Surgeons). Surgeons). Surgeons). HI, crocuta, 2, 52: Museum), a. Surgeons). Museum), a. Museum), B. TZ. crocuta, No. 522 (College of Museum), A. HT. crocuta, No. 522 (College of Museum), a. Museum), zs. Wookey (Taunton HI, spelea = crocuta, Wookey (Taunton Wookey (Taunton HT, spelea = crocuta, Wookey (Taunton HZ, spelen = erocuta, Wookey (Taunton H. spelea = crocuta, Wookey (‘Taunton Museum), B. H., crocuta, No, 522 (College of HI. spelea = crocuta, H, spelea = crocuta, Wookey (Taunton HT, spelea No. Length from dorso-anterior to dorso- posterior edge of centrum ......... bo ivy) ou a Ww) a Width across prezygapophyses ...... LS) ror) Width across postzygapophyses...... SP Si Width across transverse processes... Length of neural spine from notch between prezygapophyses eR ee ee ee ee ee ee eee 5. ae (— ae —— “a “(TUNESH & ‘nynv0.00 = AOYOO'M « 8 Ned : D H ; & = vajads * FT a Ge) | | s = e Nel = | = i = 6 vy ‘(ainesnyy o S OQ Nn : - 4 _noqunuy,) Lax0 [38 3 Nu : ra = Dynd0L0 = 7 310.0 A BG 19 a ; 1 (CALOS AHL = a =vajads "FT | x N lo BI a ToyuNR,) L300 Ay ve) ==== = 6 = i - —= pyN< Pi = — wa S|, es ne: Bl oes ; Lome Reniear | EBS : $ 962T190) GGG ‘(ON 2 10 a a | % + . = *(mmmesnyy be) o2 N oaks ene LE ae ilies ee s | gemma sea | : | as | ie = = 45 “nyna a4 3100 MA —j : a ‘(\uanes O | = O10 = DD cars | = . uoyunt SL ee: | 5 jeds “7 a rat a : ney) & : i ae e C a8 & “pyna0.00 D) ASIN st 19 ee Sou ges rma) £230 ee = | A iS = vajads “7 2 5 = im | ojumey,) Aex00 | low | & Te N 19 : = 10 | pna000 = 300M | @) ) a) | 8 vy ‘(umesnyy 7 is ; o nN 19 d=vajads-E | N 2 z = ie eci mojumey) « | $2) : a La) | | 3 2 Ge Ge : = | es) Pre ee ‘eos iS) jx SH a J(stoasamg = Bet ic : g = 0) s Fe a najads “ET ae col B a Bie a8 at109) ze ‘ON | ; = = e “(stoasang © E= nS | DINIOD “HE tad ae S JO a59T10D) Zz S | | = N a Len! =e | =e = 9 S Sy ey GEG ON | ee | - ~ H a : Vv ‘(umesnyy i ee ee ee a Z| xommg) & : g = > 1 : oe |g % AOYOO MN eA iS 3 ae a ‘(UIMeSsnyT : : ; | | Hg Ow = pajads “ET Be S es = 5 a oa DINIOM) = Ne) |) a OLN pa — E S) = dajads * : i=) 2 | & JO 95aT[0D) Zz 10 == 4 g = { = TT N co - 1 | ~~ G TL99) GZg “ON fon) 19 cy ei B on nes ; 8 ~ + | MvOGD “TEL a =I 8 q C ae 5 3 > o = > | Gin tae AOHOO AK ay ee 3 a oS * | ‘a ‘(caMeEsnyL “a = ¢ a | 0 = papas Fr BS eS 1 E a jee) Laxoo AA 10 . ~ D Q Hq ) A S (stoasmg ~ a ce) ~ f s 8: 7nI0.09 = DHjads “FT 9 ee) 2 3 © . | JO 9891109) BEG “ON 10 a nN + : EI “y ‘(tmmesnyy pee Geese me ee Oe ee eee Z| cmon joe a 2 ee 5 a} ° D () = @ a ‘(mMESnTT Ss jna0wo = nayads ori ; a : q is ; moyuney,) & | ° : Go) STARS Oe Se . : oor | S x “nani 300M. —_ ° “(stoas.m he DANS | 3 ynI0U) = najads * | Jo asa S j = Ooo = a z v ele 19 | gale ene lowest ces = | : 6) j wines’ No) Ne) NI060 * gi = és a = Z, zi a eerie SSA HN a x § x 2 a . = IL ol Se = | S| ss “pyn90.W0 iL) Aox100 | eres 10 a Aes +H = V¥ ‘(tanesuyy ie | R a 0) = pojads | : 00 oS z b 2 uoquney,) & = 7 f | 2 = = AVZOO | 19 . Q oS We N 219 B 19 g PGA 3100 MM 5 man | “(suoesaMg = = = 2S : 5 woos = varjas “FL fee ES Ho esa. & 5 JO a59]109) Zag * a = N 28 : is TeTOss co HH mae: S 2 Cael nD | “ngno0 GGG “ON F G | a || ©) ro i OacIng - i) 8 qt a sel Bes ore Ne Be ‘ i = | gossip) eon | © Bak | 5 | "a ‘(UU Ne) = 12 bY eae ON: 48-2) AS @ =e) op Ae) Osny Xe) : Eile: n = : : , Boas eS f A a rowuna) £0300M | iG ves : a ny “(uans = ees reese A a= Dyna0u = 1 i | Jo) 2 5 (Tans | 4 Hus aA a = 2 = BES TAL ek es | a _moqumuy,) Saxo 2 a a. 28255 Ie a (canes = o eo) || No) 10 10 q DINIOLI = DD aa > N = Sa 3 g i 5 2 | wo Nl 3 a fAB: ade3 ae 32 Bon 29 Sf Ele 8 geaqos N don | Dae 8 gag enog | Baas” 5p aR Eo © iy 6 BP * 8, 60.4 Pa | Ps oS) a2 ooo, lac, & e es ne) RY Serq oaq@aatt aH sl G3 <8 ~_ 3 ocr q SESS OR =, @ Sex © aa & te 2 : | Sp o GBs SbF q a8 28 A = — >] a oS q ator =) = a H R S| [es g gq oe Ap 2 | R= pera oO nS Bap Bo { S = 0 S iS a A i 18 PLEISTOCENE MAMMALITA. p. Tun Riss anp Srernum. Hyena possesses fifteen pairs of ribs, which are much arched, causing the cavity of the chest to be large as compared with that in Canzs, and very large as compared with that in /eis. ‘The sternum includes eight sternebrae. Neither ribs nor sternum present any features of special importance. bn. Tax SHountprer Girput (PI. IX). The scapula in ZZ, erocufa is straighter, and the postscapular fossa relatively smaller than in H/. striata. ‘The coracoid process is very little marked in Hyena, and the clavicle, which is minute and more or less oval in outline, is entirely suspended in the muscles. | TABLE OF MEASUREMENTS OF THE SCAPULA. HL. crocuta, HH, spelea = crocuta, | H. spelex = crocuta, No. 522 College of Creswell (Owens Wookey (Taunton | Surgeons). College Museum). Museum), a. | ee a | Length along line of spine...............6+ 178 | 19:55 ' 20:95 Antero-posterior diameter of neck ...... | 42 5:25 46 Maximum length of glenoid cavity ...... 4°45 45 45 . . . . . | Diameter from highest point in spine | to point on inner surface of scapula | immediately below .........se0.esseesenees 4°95 4°25 4:0 5 | Length from end of coracoid process | to surface of bone behind glenoid | CANADY: aaactattenlaccommmuesnestastesms esata tes 6:0 6-0 1 Figured. F. Tue Anrertor Limp (Pls. X, XI). The humerus of Hyena is a well-marked bone. Its form is short and robust, with an exceptionally large great tuberosity. ‘The condyle is larger and more pronounced than in Canis, the radial part being specially large. ‘The deltoid crest extends further down the shaft in Z/. erocuta tian in HZ. striata. ‘The humerus differs from that of Canis, Ursus, and Mustela in nearly always having a supra-trochlear foramen. ‘There is no entepicondylar foramen such as occurs in Canis and Mustela, and neither an ectepicondylar foramen nor crest occurs. ‘The manus of “Hyena differs from that of all other Carnivora in having the pollex represented by only a rudimentary metacarpal, which resembles a sesamoid bone. ‘Uhe metacarpals are longer and less enlarged above the phalangeal articulation than in Fels. HYANA CROCUTA. 19 TasLes oF CoMPARATIVE MEASUREMENTS. | | | H. crocuta, H.. spelea = crocula, H. spelea = crocuta,| H. spelea = crocuta, . No. 522 (College of Wookey (Taunton | Wookey (Taunton Creswell (Owens : | Surgeons). Museum), a. | Museum), B. College Museum). > | | | od = a = = — z = = | | | i Humerus (right). | Extreme lencth .................. agbep-seu cae 0 21:6 223 23°8 | Diameter of proximal end passing | across centre of articulating surface | and greater tuberosity ..............-.....- 6:4 | sis 67 ! | | i : Vertical diameter of shaft at middle of Wedeltoidhrid @ es i. saci. csereasseescota thee Wars eee ccc. 24-15 25°65 | 25°3 Maximum vertical measurement ............ 4-7 46 | ay il | 43 Maximum transverse measurement of | | | olecranon........ Bostioctana rgaeen Dean teaisee car 2°55 2715 2°95 30 Transverse diameter at carpal articulation 115 | ileal 1-2 | aes Vertical diameter at carpal articulation .. 18 | 18 | 16 | 1 Figured. PLEISTOCENE MAMMALIA. TABLE Of ComPARATIVE MEASUREMENTS (continued). a ‘ I, crocuta, FH, speleaa = crocuta, | H. spelea = crocuta, sey No. 522 (College of | Tor Bryan, Torquay | Wookey (Taunton A Surgeons). (Brit. Mus.). Museum). mS Length of first metacarpal.................. 2:35 | 335 : +4 second metacarpal ............ 15) “y 6 U7 uy third metacarpal ............... : 9-2 8:15 8:95 Ss fourth metacarpal ............ | 9:0 86 8°75 | PRA aRRBAETAET coo scesn ee | 76 TAS | 1 Se Figured. The above measurements tend to confirm Cuvier’s observation that the metacarpals of the cave hyana are relatively shorter than those of the living spotted hyena. c. THE Purvie Girpiy (PI. XII). The pelvis of Hy@na is characterised by its shortness, its comparatively large size, and its obliquity with regard to the sacrum. ‘he ilium is decidedly larger in proportion to the size of the animal than in bears, and is prolonged into an anterior downwardly directed hook. TABLES OF MEASUREMENTS. HT, crocuta, HH, spelea = crocuta, a No. 522 (College of | Wookey (Taunton Surgeons). Museum), a. Ricut InNoMINATE Bone. Wibprotaniyien Were), 5550090c0000000 coaonaadonoaconoeaaee 19°55 22°5 | Length from acetabulum to border of ilium 111 12°6 Vertical measurement of ilium ............... 10:55 10:9 Thickness of ilium at middle of surface...... 0:95 0:95 S Antero-posterior diameter of acetabulum ... 3°25 3°55 Length from acetabulum to posterior border (0) ae f=(6) ct) Vdd Wee aeeinernGn Soonerachasenecaccogdeer 4:85 65 Lert INNOMINATE Bone. Maximum diameter of obturator foramen ... 41 Gils)! Measurement along ischium from Speyer to end of ischial FOUND oocccancoses 75 1 Fioured. H. ‘lau Posrertor Limp (Pls. XIII, XIV). ‘The intercondylar notch of the femur is less deep than in Canis. The cnemial crest of the tibia gradually dies away instead of being strongly truncated as in Canis. ‘The hallux in Zye@na, as in Canis and Felis, is represented only by a vestigial metatarsal. HYAINA CROCUTA. TABLES oF MEASUREMENTS. IT. crocuta, Hi. spelea = crocuta, | Hl. spelea= crocuta, No. 522 (College of | Wookey (Taunton | Wookey (Taunton Surgeons). Museum), a. Museum), B. Ricut Femur. | Wteyabonmura WeyaeRH0 5.4 5nn0c9s00800595R0000D000 600 23°7 25°71 26:0 | Transverse diameter at condyles ......... 51 55 53 | Antero-posterior diameter of head ...... 3°25 315 3:15 | Vertical or antero-posterior diameter of | SLAY BYE TACKNE —.sabcccancoecooac Hacena500 18 19 19 Transverse or right to left diameter of | Slane Bye AMC json000c0nsds00con00080..000 25 2°35 2°35 | Transverse diameter at proximal end | measured across head and great LHEDOINENOINEID Gon andolicnadeaesobocdaes AOA Sea eBE | 6°35 69 6°55 Ricur Trista. | Wikyabaainian ISBVEROAN 550560000000 296ae6b0n 008660008 18°9 2071 NG)" | Transverse or rieht to left diameter at | jortor-aaMe| GIMNCl! Aoh.G5 coadesscodbouee Gesodee: 5°3 53 iss) | Vertical or antero-posterior diameter at | | proximal end measured from notch | between articulating surface for | femur and! top of crest -0......+.+..-... | 525 54 525 Transverse diameter at distal end ..... | 4:05 39 42 Vertical diameter at distal end............ 2°45 2°6 2°85 Transverse diameter at narrowest part | ObyShiahity Uae pastas asco shi ies ea. sores ae | 175 18 2:15 1 Wieured. HT, crocuta, No. 522 (College of H, spelea = crocuta, Wookey (Taunton H. spelea = crocuta, Tor Bryan, Torquay Surgeons). Museum), B. (Brit. Mus.). i} RicuHr FrBuna. Marxamamm! Venethy -.. 0.00.22. ..cseeeneceecee se. 178 17°85 | Transverse diameter at distal end ...... 1:25 1-25 | | Vertical diameter at distal end ........... | 2710 2715 Transverse diameter at proximal end ... | 10 0-9 Vertical diameter at proximal end ...... | 14. 1-4 CALCANEUM. IU@RIGR EE. Seeuns codebesba cosmos rearee ae ce ea eres 59 67 Maximum transverse diameter............ 2°8 | 2°8 | ASTRAGALUS. Right to left diameter...................0005 315 37 Merararsats. Length of first metatarsal.................. 175) 63 et second metatarsal ............ 7-35 78 es third metatarsal ..... ......... 84 8:0 | aA fourth metatarsal ............ 82 7-95 ma fifth metatarsal ............... 72 6°75 22 PLEISTOCENE MAMMALIA. IV. CONCLUSIONS. The evidence for the view that the yena from the caves is a species distinct from the modern H/. crocuta may now be more fully considered. Cuvier,’ writing in 1825, mentions the following features as characteristic of H. spelea: 1. The upper surface of the skull is less arched than in H. crocuta, and the temporal ridges do not unite so quickly to form a sagittal crest. 2. All the bones of the metacarpals and metatarsals measured are without exception shorter and thicker than in /7. crocuta. Reference is also made to the relatively thick character of the bones in the cave hyena by Gaudry,’ who concludes that the animal was more thick-set than its living repre- sentative, and suggests that it may have had a more crouching gait. Cuvier remarks, however, with regard to the teeth, that it 1s impossible to distinguish — . — < e those of H. syel@a from those of H. crocuta. De Blainville® gives the following features as characteristic of H. spelea: ]. The form of the upper carnassial with the large size of the third lobe (talon).* 2. The absence of the inner cusp on the tubercular portion of the lower carnassial. 3. The size, which is # larger than in H/. crocuta. A, The greater extension and compression of the occipital crest. ee a ee ee 5. The increased thickness and shortness of the muzzle. 6. ‘The increased thickness and shortness of the limb bones. He remarks that of these characters the least important is the increased relative size, and the most important the increased thickness of the limbs and elevation of the occipital crest. Relative size depends on conditions of life, and comparisons as regards size have too often been made between fossil individuals at the maximum of their development, owing to savage life, and individuals raised in menageries. ‘The character of the occipital crest also differs much, according to the age of tlie animal. Owen * states that the upper true molar in /. sye/ea is monoradicular, and quotes this as a character distinguishing ZH. spelea from ZH. crecuta. Dawkins,’ on the other hand, shows that the tooth in question is sometimes mono-, sometimes bi-radicular, and that the method of implantation cannot be quoted as a character of specific value. The modern view that there is no specific distinction between HH. spele@a and H. crocuta was first clearly stated by Boyd Dawkins in 1865,° and is now almost universally accepted. 1 «Oss. Foss.,’ ed. 3, iv, p. 396. 2 “Mater. Hist. Temps Quat.’ (4), 1892, p. 118. 3 ‘Ostéographie, Hyénes,’ p. 39. 4 TI cannot trace any difference as regards the form of the upper carnassial or the size of the third lobe between H. spelea and LH. crocuta. » * Brit. Foss. Mamw.,’ p. 150, 6 * Nat. Hist. Rev.,’ n.s., v. ——. = = SS —— : _._—"_" 7 HY AINA CROCUTA. 23 The measurements quoted above, however, show that some of the skulls of the cave hyeena, especially those from the German caves, are considerably larger than those of any modern hyena measured. ‘They also show that it is true that the metacarpals of the cave hyena tend to be shorter than those of the modern form, hence it seems reasonable to follow Gaudry* in regarding the cave hyena as a distinct race of fT, crocuta. 1737. J. C. Kundmann, ‘ Raviora Natures et Artis.” Breslau. 1774. J. F. sper, ‘ Ausfiheliche Nachricht—Zoolithen, Bayreuth.’ Nirnberg. 1833. 1839. 1844. 1844. 1845. 1846. J. de 1s . C. Collini, ‘ Acta Ac. Theod. Palat.,’ v. J. . G. Cuvier, “ Sur les Ossemens fossiles des Hyénes,’ ‘Ann. Mus. Hist. Nat.,’ . G. A. Goldfuss, ‘Die Umgebung von Muggendorf.’ Erlangen. . G. Cuvier, ‘ Recherches sur les Ossemens fossiles,’ iv, p. 396. 22. W. Buckland, “ Account of an Assemblage of Fossils—Kirkdale Cave,” ¢ Phil. 3. W. Clift and J. Whidbey, *‘ On some Fossil Bones—Oreston,” § Phil. ‘I'rans.,’- . G. A. Goldfuss, ‘Saugethiere der Vorwelt,’ vi. . W. Buckland, ‘ Reliquize Diluviane.’ . G. Cuvier, ‘ Recherches sur les Ossemens fossiles,’ ed. 3, iv, p. 381. . J. B. Croizet and A. C. G. Jobert, ‘ Recherches sur les Ossemens fossiles du . S. ‘I. von S6mmering, ‘Ueber die geheilte Verletzung eines fossilen Hyanen . J. de Christol and A. Bravard, “Mémoire sur les nouvelles Espéces d’Hyénes V. BIBLIOGRAPHY. Mannheim. C. Rosenmiller, ‘ Beitr. zur Geschichte—fossile Knochen.’ | Weimar. Wily [Os Lens ansaliS 225 pel 7. exiii, p. 88. London. Département du Puy de Dome.’ Schadels.’ fossiles,” ‘ Mém. Soc. Hist. Nat., Paris,’ iv, p. 368. P. C. Schmerling, ‘Recherches sur les Ossemens fossiles—Cavernes de Liege,’ Mm, chi. iv. Marcel de Serres, J. M. Dubrueil, and B. Jeanjean, ‘Recherches sur les Ossemens humatiles des Cavernes de Lunel Viel.’ H. M. D. de Blainville, * Ostéographie,’ livraison 14. F. J. Pictet, ‘ Traité de Paléontologie,’ i, p. 180. C. G. Giebel, “ Die fossile Hyane,” ‘Isis’ (Oken), 1845, i, p. 483. R. Owen, ‘A History of British Fossil Mammals, etc.,’ pp. 138—160. 1 *Matér. Hist. Temps Quat.’ (4), p. 122. I AI AE NO 24. 1859. 1860. 1863. 1865. 1866. 1867. 1868. 1868. 1869. 1870. eee 1874. 1876. USAT: Sate PLEISTOCENE MAMMALTA. J. MacEnery, ‘ Cavern Researches.’ H. Falconer, ‘‘ On the Ossiferous Caves—Gower,” ‘ Quart. Journ. Geol. Soc.,’ xvi, p. 487. A. Gaudry, “Sur les Liens que les Hyénes fossiles établissent entre les Hyénes vivantes,” ‘ Bull. Soc. Géol. France’ (2), xx, p. 404. Summary in ‘ Quart. Journ Geol Sccsaxa@) pact W. Boyd Dawkins, ‘On the Dentition of Hyena spelea and its Varieties, with Notes on the Recent Species,” ‘ Nat. Hist. Rev.,’ n.s., v, p. 80. W. Boyd Dawkins and W. A. Sanford, ‘The British Pleistocene Mammalia,’ Part 1, Mon. Paleont. Soc. H. Wicks, ‘ Discovery of a Hyena Den near Laugharne, Carmarthenshire,” ‘Geol. Mag.,’ iv, p. 307. H. Falconer (Notes on Hyena), ‘ Paleont. Memoirs,’ 11, p. 464. G. Busk, ‘‘ On the Cranial and Dental Characters of the Hyzenidee,” ‘ Journ. Linn, | Soc.,’ 1x, p. 59. 3 W. Boyd Dawkins, “On the Distribution of British Post-glacial Mammals,” ‘Quart. Journ. Geol. Soc.,’ xxv, p. 192. W. Pengelly, “The Literature of the Caves near Yealmpton,” ‘'Trans. Devon. Assoc.,’ iv, p. 81. G. Busk, “ Report on the Exploration of Brixham Cave—Animal Remains,” ADail, We. Chau, jo, 47/11. W. Pengelly, ‘‘ Report on the Windmill Hill Cavern, Brixham,” ‘Trans. Devon. Assoc., vi, p. 779. . W. Boyd Dawkins, “ On the Mammalia and Traces of Man found in the Robin - Hood Cave,” ‘Quart. Journ. Geol. Soc.,’ xxxii, p. 245. W.-Boyd Dawkins, “On the Mammal Fauna of the Caves of Creswell Crags,” ‘Quart. Journ. Geol. Soc.,’ xxxi, p. 596. G. Busk, “On the Ancient or Quaternary Fauna of Gibraltar,” ‘Trans. Zool. Soc.,’ x (2), p. 53. . P. N. Bose, “ Fossil Carnivora from the Siwalik Hills,” ‘Quart. Journ. Geol. Soc., xxxvil, p. 130. . E. D. Cope, “On the Extinct Dogs of North America,” ‘Amer. Nat., xvii, p. 243. . E. T. Newton, “ On the Occurrence of the Cave Hyena at Corton Cliff, Suffolk ,” ‘Geol. Mag.,’ 1883, p. 433. . R. Lydekker, ‘‘ Post-tertiary and Tertiary Vertebrata; Siwalik and Narbada Carnivora,” ‘ Palzeont. Indica,’ ser. 10, i, p. 275. . R. Lydekker, ‘Catalogue of the Fossil Mammalia in the British Museum,’ Mita ls \05 Wale | Sasa . : 1 Se — es Sl ———— =< 1885. 1886. 1886. 1886. 1886. 1889. 1890. TSSMOe Isle 1892. NSO. 1893. 1893. 1894. 1895. ESO HYAANA CROCUTA. 25 C. J. Forsyth Major, “On the Mammalian Fauna of the Val d’ Arno,” ‘ Quart. Journ Geolssoc., t ' pee ty, ‘ ’ i ns , ua » a ms f 3 Fie. BS CS B) = 7. Dorsal 9. Anterior 11. Dorsal 12. Anterior Sa S G: h. ag PLATE VIII. Cave Hyana. Vertebre. (Natural size.) Sixth cervical, posterior view. Thirteenth thoracic, seen from the left side. Thirteenth thoracic, front view. Fourth lumbar, posterior view. 5. Fifth lumbar, dorsal view. 6 Sacrum, dorsal view. 8. Ventral view of the first free caudal vertebra. 10. The same vertebra seen from the left side. } view of probably the nimth free caudal vertebra. 13. ‘The same vertebra seen from thie left side. 14. Anterior view of a late caudal vertebra, perhaps the twelfth. 15. ‘The same vertebra seen from the left side. Neural! spine. Neural canal. Vertebrarterial canal. Foramen in sacrum for exit of spinal nerve. Prezygapophysis. Postzygapophysis. Transverse process. Notch for exit of spinal nerve. ‘he specimens shown in figs. 1—6 are from Wookey Hole, and are preserved in the Taunton Museum. ‘The other specimens figured are also without doubt from Wookey Hole, but I was unable to find them in the Taunton Museum. Figs. 1 and 4 are from Skeleton A; figs. 2, 3, and 6 from Skeleton B. TAS et ‘206! ‘ALBIOOS IWODIHdVYSOLNOW 1Vd te th Py eue2kyy Ae) ‘spjouAayy PLATE IX. Cave Hya@na. Scapula. (Natural size.) Outer aspect } of left scapula. Posterior aspect a. Glenoid cavity. 6. Acromion. Gs Prescapular fossa. d. Postscapular fossa. e. Glenoid border. 5 ° 0 x a . : Fe oe en This specimen is from the Creswell caves, Derbyshire, and is now preserved in the Museum of Owens College, Manchester. Reynolds, Cave Hyzena. J.Green del, et lth. PALZONTOGRAPHICAL SOCIETY,1902. PLATE X. Cave Hyana. Flumerus, Ulna, and Radius. (Natural size.) Right humerus, Right ulna, { all anterior aspect. Right radius, Head of humerus. Great tuberosity. Lesser tuberosity. Deltoid ridge. Supra-trochlear foramen. Trochlea, Iixternal condyle. Internal condyle. Olecranon. Sigmoid notch. Surface for articulation with radius. — Distal end of ulna. Surface for articulation with humerus. Distal end of radius. The three bones are all from the Skeleton B found at Wookey Hole, and ne served in the Taunton Museum. oe -Reynelds, Cave Hyena. West Newman anrp W.Bidgood del, et lith. Everenneles Uliaa, aac Humerus, PLATE XI. Cave Hymna. Manus. (Natural size.) Fie. 1. Dorsal or anterior view of right manus. 2. Ventral or posterior view of right manus. a. Bone representing the fused scaphoid, lunar, and centrale. 6. Cuneiform. c. Pisiform. d, 'Trapezoid. e. Unciform. J. First metacarpal. g. ¥ifth metacarpal. h. Sesamoid bone at metacarpo-phalangeal articuiation of second digit. 2. Ungual phalanx of third digit. ‘The specimens from which these figures were drawn are from the Tor Bryan caves, near Torquay, and are now in the British Museum (Nat. Hist.). Reynolds, Cave Hyena. J.Green del. et lith. PALAZEONTOGRAPHICAL SOCIETY,1902. Manus. PAL. J Mintern Bros.imp. PEATE XO. Cave Hyana. Pelvis. (Natural size.) Fie. 1. Left innominate bone, seen from the outer side. 2. Right innominate bone, seen from the inner or sacral side. a. Acetabulum. | Obturator foramen. c. Supra-iliac border of ilium. d. Sacral surface. Pubic border. f. Ischial border. y. schium. 4. Ischial tuberosity. Qaeelaubise Both the above specimens belong to Skeleton A, in the Taunton Museum. “UNTL 4? 19P POBPIE MN. dua, weurmeahy "982. “euURATT oAe) ‘sppouAa ; oe = ee ee ae ee PLATE NITI. Cave Hymna. Femur, Tibia, and Fibula. (Natural size.) = Q SOWA TR ww Right femur, viewed from the left side. The same, proximal end, The same, distal end. Right tibia, anterior aspect. ‘The same, proximal end. The same, distal end. Right fibula, anterior aspect. The same, proximal end. No} The same, distal end. a. Head of femur. 4. Great trochanter. c. Lesser trochanter. | d. Outer condyle of femur. e. Intercondylar notch. f. Surface for articulation with fibula. yg. Cnemial crest. 4. Surface for articulation with astragalus. All the above specimens are from Wookey Hole, Somerset, and are now preserved in the ‘Taunton Museum. ‘The specimen from which figs. 1, 2, and 3 were drawn forms part of Skeleton A; the remainder are from Skeleton B. NS ————————— — W. Bidgood del et hth. q 2 a6) oO 5 oS GN a a Fie. PAGE xeeVE CavE Hyana. Pes. (Natural size.) 1. Dorsal or anterior view of right pes. 2. Ventral or posterior view of the same. a. b. 6, d. ~ The specimens from which these figures were drawn are from the Tor Bi near Torquay, and are now in the British Museum (Nat. Hist.). Astragalus. Calcaneum, Cuboid. Navicular. External cuneiform. Middle cuneiform. First metatarsal. Fifth metatarsal. Sesamoid at metacarpo-phalangeal articulation of second digit. Ungual phalanx of third digit. PAL AONTOGRAPHICAL SOCIETY,1902. Reynolds, Cave Hyena. 2. PAL 2a, | J.Green del,et lth ; Mintern Bros.imp-= | Pes. | >> 9 Ph ee i sstisesaemaamhalaa ieateaaaa Pepe 2 : Palsontographical Society, 1906. A MONOGRAPH OF THE BRITISH PLEESTOCENE MAMMALIA WO, JUL” eee JOE IE BEA RS. BY SIDNEY H. REYNOLDS, M.A., F.G.S., PROFESSOR OF GEOLOGY AND ZOOLOGY, UNIVERSITY COLLEGE, BRISTOL. Pacts 1—35; Puates I—VIII. LONDON: PRINTED FOR THE PALMHONTOGRAPHICAL SOCIETY. 1906. PRINTED BY ADLARD AND SON, LONDON AND DORKING. MONOGRAPH THE BRITISH MAMMALIA PLEISTOCENE PERIOD. THE BEARS. Order—CARNIVORA. Famity—U RSID Ah. Genus—Ursus. I. HISTORICAL INTRODUCTION. Tue fossil bears form a group of animals whose study is by no means easy, not from any scarcity of their remains, but from the difficulty of coming to a decision about the mutual relationship of the various living and fossil forms. Very divergent opinions have been expressed with regard to the number of species of bears, and the literature dealing with the subject is remarkably extensive. Cuvier’ and de Blainville* treat of the early discoveries of fossil bears very fully, and their accounts have been freely used in the following pages. Fossil bones, which eventually proved to be those of bears, were first mentioned by J. Paterson Hayn?® (1672), who considered them to be the bones of dragons. He obtained representatives of nearly all parts of the skeleton from a cave in Mount Krapacks, Hungary. H. Vollenad* (1673) referred to the same bones, again considering them to be the remains of dragons. 1 “Oss. Foss.,’ ed. 1, 1812, tom. iv, part iv. 2 * Ostéographie,’ tom. 11, K. BG Ephém. Curieux de la Nature,’ dec. i, an. ii, obs. exxxix, p. 220. 4 Thid., an. iv, obs. clxx, p. 226. 2 PLEISTOCENE MAMMALIA. F. E. Briickmann! (1732) was the first to compare these caye-bones with those of bears. J. F. Esper? (1774) gave figures of a large number of bear-bones found in a cave in Franconia, but in default of material for comparison was unable to decide definitely that they belonged to bears, though he noted the resemblance. Later on Hsper® (1784), having obtained the skull of a polar bear, adopted the view that the cave remains were to be attributed to the same species. In 1794 John Hunter* compared a fossil skull, which had been referred to the polar bear, with the skull of the last-mentioned species, and noted various differences, though cautiously observing that great changes in the shape of the skulls of Carnivora occur during their growth to maturity and old age. In 1795 J.C. Rosenmiiller® recognised differences between the brown, white, and cave bears, and gave a table of comparison between the skulls of these three forms printed in parallel columns. He was also the first to apply the name Ursus spelxus to the cave bear. In 1804 Rosenmiiller® published a folio volume in French and German dealing solely with the cave bears, and fully described their remains, concluding with a suggestive chapter on the conditions under which bones found in caves might have accumulated. He also emphasised the fact that differences in skulls depend not only on age (as noted by Hunter), but also on sex. Meanwhile, the study of fossil bears was undertaken by Blumenbach and Cuvier. The former’ arrived at the conclusion that the German caves contained not only Ursus speleus, which he regarded as distinct from all living species, but also another form which he named U. arctoideus, intending thus to indicate its relationship to the brown bear. Cuvier* (1806) confirmed Blumenbach’s statement that some of the larger bones from the German caves indicated specific differences from all living bears, and also agreed with the suggestion that they represented two extinct forms—U. speleus with the forehead arched, U. arctoideus with the forehead flat—the latter approaching living species more closely than the former. 1 «Breslauer Samml.,’ 1732, p. 628; and ‘ Hpist. Itin.,’ 32. 2 « Ausfithrliche Nachricht zoolith. Bayreuth.’ 3 «Herits Soc. nat. Berlin,’ v, p. 56. 4 «Phil. Trans.,’ lxxxiv, 1794, p. 407. 5 ¢ Beitr. Geschicht. fis. Knochen,’ p. 44 (German reprint of the same author's ‘ De Oss. foss.,’ Leipzig, 1794). 6 Abbild. u. Beschreib. der foss. Knochen des Héhlenbiren,’ Weimar. 7 Quoted by Cuvier, ‘Bull. Sci., Soc. Philomath.,’ no. 50. This reference is taken from de Blainville, ‘ Ostéographie,’ Carnassiers, p. 46. It is quoted apparently from him by Owen, ‘ Brit. Foss. Mamm.,’ p. 86, and by other subsequent writers. In the official catalogue of Cuvier’s papers the title appears without any reference as to where the paper can be found. The paper cannot be traced, and was probably suppressed. 5 «Annales du Muséum,’ vii, p- 324. URSUS. 3 In 1810 Goldfuss! published a memoir in which he attempted to distinguish a third species of fossil bears, named U. priscus. This was accepted by Cuvier.” Meanwhile the fossil bones of bears were being discovered in several localities in England, notably in the caves of Kirkdale, Yorkshire (whence Buckland * described rare fragments), and Oreston, near Plymouth (Clift and Whidbey).* In 1825 Cuvier in the third edition of ‘Ossemens Fossiles’ reverted’ to the conclusion that the forms called U. speleus and arctoileus were only varieties of the same species. De Blainville, however, remarked® that Cuvier’s unfortunate establishment of a new species on insufficient evidence gave an impulse to this practice, which was exaggerated in the hands of less skilful paleontologists. In proof of this he referred to Croizet and Jobert’ (1828) who believed they could recognise U. cultridens by a single canine, and sought to establish a new species U. arvernensis on a fragment of the anterior part of the skull, a humerus and other isolated bones. The work of M. de Serres* is an instance of the same method. P. C. Schmerling’? (1833), although he corrected certain mistakes of Cuvier, was led by his example to establish several new species on material more or less incomplete. He concluded that no less than five species of bears lived in the Liége district—viz., U. speleus and arctoideus, Blum., U. priscus, Goldf., and two new species, U. giganteus and leodiensis. In 1842 Owen" described a fine skull of the brown bear from Manea fen, Cambridge. With the increase of knowledge and facilities for comparison, the extreme difficulty of recognising specific distinctions between the various bears began to be apparent, with a tendency to group together several forms which had previously been regarded as distinct species. This tendency was first shown by de Blainville! who in 1844 gave a detailed and critical account of the different kinds of fossil bears with splendid illustrations. Further reference to his conclusions follows later, but it may be mentioned here that he considered all the bears, hymg and fossil, found in Hurope to belong to one species, but thought there were two races of fossil bears, a larger race the male of which was represented by U. giganteus and U. speleus major and the female by U. arctoideus and U. leodiensis, and a smaller race in which the male was represented by U. speleeus minor and the female by U. priscus. He considered that a second small species wes represented by Ursus = ‘Verhandl. kaiserl. Leopold.-Karolin. Akad. der Naturforscher,’ x, 2, p. 260. ‘Oss. Foss.,’ ed. 3, 1825, iv, p. 380. ~w 3 ‘ Reliquie diluviane,’ ed. 2, p. 17; and ‘ Phil. Trans.,’ 1822, p. 171. 4 «Phil. Trans.,’ cxii, 1823, p. 88. > Tom. iv, p. 358. 6 «Osteographie,’ Carnassiers, p. 50. 7 “Rech. Oss. foss. Puy de Dome,’ p. 628. 8 ‘Bull. univ. des Sci. Nat.,’ 1830, xii, no. 19, p. 161. 9 « Recherches Oss. foss. Cavernes de Lite.’ 10 «Rep. Brit. Assoc.,’ 1842, p. 69. 1 « Ostcéographie,’ Carnassiers, p. 38. 4. PLEISTOCENE MAMMALIA. arvernensis. He doubted the distinction of the bears of north-western America (t.e. the grizzly bear) from the Huropean species. This view was also accepted by Middendorff (1851),! who concluded that all the bears of the arctos group from both eastern and western hemispheres were varieties of one species. On the other hand, Owen in 1846 in his ‘ British Fossil Mammals and Birds’ dealt fully with the British fossil bears, recognising three species, U. spelexus, U. arctos, and U. priscus, Goldf., to which species he attributed a lower jaw from Kent’s Cavern. J. A. Wagner? also (1851) agreed with Owen in recognising more than one species among the bears of the arctos group and considered de Blainville’s views on the subject to be retrogressive. Gray ”* (1864) went farther in the process of subdivision than anyone else, separating the living bears not only into a number of species, but also into several genera. The descriptions of the bones of bears from a number of Irish localities now commenced—e.g. by R. Ball, A. Carte,° and H. Denny.® Some of the bones found were even attributed to the polar bear. Miller’ (1872), in a beautifully illustrated work on certain bears’ skulls from Russia, doubted the possibility of distinguishing between the different species of fossil bears even by their teeth. In 1867 appeared the first of a series of important communications from Busk dealing with the fossil bears. In this paper, of which, unfortunately, only an abstract was published,® he mentioned that the teeth on which reliance was to be placed in distinguishing the different species of fossil bears were pm. 4, pm. 4, m.2,m. 3. He expressed the opinion that U. priscus was identical with U. feroz. In 1873 appeared his very important paper? on the animal remains found in the Brixham cave, in which he fully discussed the mutual relationship of the various species of fossil bears. He established the fact that U. priscus, Cuv., was identical with U. fossilis, Goldf., and U. fervor, the modern grizzly, and considered that all the Irish specimens were referable to the latter species. He thought that U. ferox (priscus) was commoner even in England than U. speleus. He discussed the differences by which, according to Owen, the teeth of U. speleus, U. arctos, and U. feroe could be distinguished, but thought that these differences were not all constant and considered that it would be impossible to distinguish between the ‘Untersuch. Schideln des gemeinen Landbaren,’ etc., St. Petersburg. ‘Abhandl. k. bay. Akad. Wissensch.,’ vi, I Abth., 1851, p. 193. ‘Cat. Carniv. Pachyderm. and Edentate Mamm. in Brit. Mus.,’ p. 217. ‘Proc. R. Irish Acad.,’ iv, 1849, p. 146. ‘Journ. R. Dublin Soc.,’ ii, 1860, p. 344; and ‘Journ. R. Geol. Soc. Dublin,’ x, 1864, p. 114. ‘Proce. Yorks. Geol. and Polyt. Soc.,’ iv, 1864, p. 347. ‘Drei in der Provinz Preussen ausgegrabene Birenschiidel.’ ‘Q. J. Geol. Soc.,’ xxii, 1867, p. 342; and ‘ Phil. Mag.,’ xxxiv, 1867, p. 399. ‘Phil. Trans.,’ clxiii, p. 532. a nN for} ox ~ os ~w - Cc ae URSUS. 5 three species in respect of the teeth. He returned to the subject four years later in his ‘Report on the Ancient or Quaternary Fauna of Gibraltar,’! m which he discussed minutely the characters of the cave, brown, and grizzly bears as based upon their teeth, stating his belief that no character of specific importance could be drawn from any part of the bear’s skeleton except the teeth. The question of the relationship of the fossil bears was further considered by R. Hensel? (1876). Basing his view on a study of the teeth, he urged the distinc- tion of the cave bear, but did not express a clear opinion as to whether the other Pleistocene bears represented more than one species. In 1877 Boyd Dawkins,® who had already* commented on the extreme difficulty of distinguishing between the brown and grizzly bears by means of their hard parts, adopted the view of their identity. About this time A. L. Adams commenced a series of important papers in which he discussed the Irish specimens. In the earliest? of these (i878) he referred the specimen described by Carte as U. maritimus to U. ferow, and in the second® he gave a critical account of all the Irish bear-remains, referring them all to U. ferow, the grizzly bear, which he concluded was the only bear whose remains had been proved to occur in Ireland. In a later paper’ the same author, in describing further remains of Irish bears, was the first to suggest that those known as U. speleus might be only those of large individuals of U. ferow. He confirmed Busk’s and his own previously expressed opinion, that all the remains of Irish bears were referable to U. ferox. ‘The paper included a table of dimensions of bears’ crania. The following year (1881) Adams published another paper® further developing his suggestion that the differences between fossil bear-remains may be racial, sexual, or even individual, dependent on mode of life or character of food, and that the different British fossil bears may best be regarded as races of one species. Later writers have also discussed fully the mutual relationship of the bears, and very varying opinions have been reached. Lydekker,’ writing in 1854, gave dental characters by which the brown and grizzly bears might be distinguished, but a year later doubted!? whether a valid distinction of this kind was possible. Im 1897 he separated" U. spelxus as a Species, grouping all the bears of the arctos group (i.e. all those of the northern 1 «Trans. Zool. Soc.,’ x (2), 1877, p. 60. 2 «Sitzb. Naturf. Freunde, Berlin,’ p. 49. 3 «Q. J. Geol. Soc.,’ xxxiii, 1877, p. 598. * Thid., xxxii, 1876, p. 248. > ‘Proc. R. Irish Acad.,’ 2nd series, iti, 1878, p. 94. 6 «Journ. R. Geol. Soc. Ireland,’ iv, 1877, p. 247. 7 ‘Sci. Proc. R. Dublin Soe.,’ ii, 1880, p. 49. 8 «Trans. Roy. Dublin Soc.’ (2), i, 1881, p. 201. 9 *Paleont. Indica,’ ser. 10, u, 1884, p. 202. 0 «Cat. Foss. Mamm. Brit. Mus.,’ pt. i, p. 173. MU « Proc. Zool. Soc.,’ 1897, p. 412. 6 PLEISTOCENE MAMMALIA. hemisphere except the polar bear, the American black bear, and the blue bear of Thibet) as one species. A. EH. Brown! (1894) went farther than this, including even the American black bear as a subspecies of U. arctos. In this view he was in accordance with that previously reached by Allen,* who, however, afterwards changed his opinion * with regard to this point. In the paper just referred to® he gave a valuable table of measurements showing the great individual variability in bears’ skulls from the same locality, and considered that, though U. americanus might be distinct, there was a complete passage between the brown and grizzly bears. The remarkable individual variability was still more impressively shown by H. Schiff* (1889) in a paper on a collection of thirty-five skulls all obtained from a limited area in Russia. The variability of the European bears was shown to be more than paralleled by those of America in C. H. Merriam’s paper,’ which was based on a study of more than two hundred skulls, a series of as many as ninety-five haying been obtained from one locality. The conclusions which he drew were, however, widely different from those drawn by Brown and Allen; for he not only considered that the American “brown” bears (of which he made a number of new species) were specifically distinct from the Huropean, but separated the black bear subgenerically. The difficulty of distinguishing between the different species of fossil bears is further illustrated by the important papers by Gaudry and Boule,® and H. T. Newton.’ The former authors showed that even the loss of the three anterior upper premolars is not absolutely characteristic of the cave bear, as individuals of the smaller race from Gargas retain pm. 3. The close connection between the bears of the brown and grizzly types is illustrated by the fact that they considered U. priscus to come nearer to U. arctos, especially as regards the humerus, than to U. horvibilis, with which it is usually thought to be identical. Newton, in describing the Vertebrata from the Forest Bed, agreed with Owen in assigning the jaw figured in the ‘ British Fossil Mammals,’ p. 106, to U. spelzus, in spite of its small size, while he assigned another specimen to U. spelxus in spite of its retaining pm. 1. 1 «Proce. Acad. Nat. Sci. Philad.,’ xlvi, 1894, p. 119. 2 «Bull. Mus. Comp. Zool.,’ i, 1869, p. 184. 3 «Bull. U. S. Geol. Surv. of Territories,’ 11, no. 4, 1876, p. 334. 4 «Archiv fir Naturgeschichte,’ 1889, p. 244. 5 «Proc. Biol. Soc. Washington,’ x, 1896, pp. 65-83. 6 ‘Mat. pour histoire des temps quaternaires,’ fase. iv, p. 105 (1892). 7 “ Vertebrata of Forest Bed Series,” ‘Mem. Geol. Surv.,’ p. 5 (1882). “URSUS. 7 Il. DISTRIBUTION IN BRITAIN AND ELSEWHERE. The oldest British formation from which the fossil remains of bears have been described is the Suffolk Crag. Owen,! writing in 1846, says “the oldest fossil referable to the genus Ursus from British strata is the crown of a molar tooth, which was found at Newbourn, near Woodbridge, Suffolk. The bear’s tooth is the antepenultimate grinder of the right side, upper jaw; it is smaller than the corresponding tooth in U. speleus.” Newton” was, unfortunately, unable to verify this determination, and suggests that the tooth may be attributable to Sus. In 1864 Lankester® described and figured a slender canine tooth, also said to have come from the Red Crag of Newbourn, near Woodbridge. This specimen, which is in the Reed collection at York, he referred with little hesitation to U. arver- nensis. Boyd Dawkins, and Newton? have both doubted the correctness of this identification, the latter saying ‘‘it seems more probable it will prove to be an anterior tooth of Squalodon and, therefore, cannot be taken as evidence of the occurrence of Ursus in the Red Crag.” The Forest Bed is the oldest British formation in which undoubted bears’ bones have been found. ‘The best specimens were originally described by Owen,' and they have been re-examined and described by Newton.’ The specimens found were at first attributed to as many as four species of bears—U. speleus, arvernensis, etruscus, and priscus (= horribilis). The best specimen described by Owen is a small mandible,° which, nm spite of its small size, is referred to U. speleus for the following reasons: (1) There is a long diastema between the canine and the first tooth of the molar series, pm. 4; (2) pm. 4 has a complicated form; (3) m. 3 is broad as compared with the same tooth in the brown and grizzly bears. Owen’s identifi- cation is endorsed by Newton. Another specimen? has pm. | present, but in spite of this fact 1s referred by Newton to U. spelzus, while a third and larger specimen‘ agrees with the normal speleus in the complete absence of the anterior premolars. Of the sixteen specimens found in the Forest Bed, nine are referred by Newton without hesitation to U. speleus, and probably two more belong to this species. The supposed occurrence of U. arvernensis is based on a fragment of the right maxilla with two teeth, now in the Museum of Practical Geology. It has been regarded by Dawkins as probably referable to U. arvernensis. Newton considered that there was no evidence to show the correctness of this attribution, and regarded 1 «Brit. Foss. Mamm. and Birds,’ p. 105 (1846). * “Mem. Geol. Surv.,’ “ Vert. of Pliocene Deposits,” p. 15 (1891). 3 «Ann. Mag. Nat. Hist.’ (8), xiv, p. 358. 4 «Brit. Foss. Mamm. and Birds,’ p. 89 (1846). > “Mem. Geol. Surv.,’ “ Vert. of Forest Bed Series,” pp. 5—16 (1882). 6 Fig., ibid., pl. i (1). 7 Fig., ibid, pl. i (2). 8 Fic,, ibid., pl. i (3). 8 PLEISTOCENE MAMMALIA. He con- the size of the teeth as too great, U. arvernensis being a small species. sidered it far more probable that the fragment was to be referred to the grizzly bear. Newton summarised his views with regard to the occurrence of U. arver- nensts a8 follows!: “No description of a specimen of U. arvernensis from these deposits has ever been published, although the name has been admitted in the list of mammals. If such a specimen is in existence its resting place is not known and one is compelled, therefore, to omit the species until evidence of its existence is TABLE SHOWING THE DisrripuTiIon oF THE British PLEIstocENE Baars. i | . | ale River Deposits, ETC. E é s CAVES AND FIssuRES. B 6 S Oo} mA] & 1S) eon eo IA CUOMR saee Ne vee eee iye tee eres ... |... | * | Bacon Hole, Gower.................. 6 ll aes Ballymahon on borders of Long- tea co. Waterford Ty. | oe ford vandWiestmeathu sss seer lesan Banwell.. Pee ene moeiauucasaltees. ||) coc Ballinamore, co. Leitrim............ teal) and |) ae | Bleadonin on. ceececce ree eee eens * | * ISNRANTEHOM, jcocdoocsenospocdaenssosced x | x Bosco’s Hole, Gower ............... * chee Bedtord®. at eacrneet acckeet eee P Brixham Ss, -eecsaose- ce eeneoee een ee * * | * Bourne, Lincoln % Burrineton: 25. scessmeeceeae % iBurwelllticnve eee eee eee x |... | Cae'Gwyn, Clwyd Vale 7.57.2..2:| teen eee Clonburne, King’s co. ............... x | Cefn, near St. Asaph ............... * | eee Cophovdlen Whe. paint e tenon errr wos | Sct | a a RChudlleto hey, scereccsenaenteneccs reese || = Craiyhord: 85. Wace ceene ne metre Ba rss Clévedom: 45 hci seaketne eee eee oe), eee ID mIMATIES) 4) cscewman aden eee ic eee ... | * |... | Cresswell Crags, Derbyshire ...... Pees (Cait Wy /Siied BYSISEV'C ts Sey canboneodsneshoneebosll aaa || LSP 2 “Trans. Roy. Irish Acad.,’ xxxii, B, pt. 4, p. 201. 3 Ibid, xxxii, B, pt. 1, p. 48. 2 Te, i ee a a 10 PLEISTOCENE MAMMALIA. III. DESCRIPTION OF THE REMAINS. It may be well to begin with a statement of the distinctive osteological characters of bears. They agree with the other Arctoidea in the following respects ': (1) there are five well-developed digits; (2) the auditory bulla is simple with no trace of a dividing septum, and the inferior lip of the auditory meatus 1s considerably prolonged ; (5) the paroccipital process of the exoccipital is more or less triangular and is directed backwards, outwards, and downwards, standing quite apart from the bulla; (4) the mastoid process of the periotic is widely separated from the paroccipital and generally very prominent; (5) the carotid foramen is large and placed on the inner margin of the bulla, usually near the middle, but occasionally more posteriorly ; (6) the condyloid foramen is distinct and exposed and never sunk into a common opening with the foramen lacerum posterius; (7) the glenoid foramen is always present and usually conspicuous; (8) a large penial bone occurs. The family Urside is characterised by the followimg features:* In existing forms the true molars are = and have broad flat tuberculated crowns. The three anterior premolars of both jaws are rudimentary and often deciduous. The fourth upper premolar, the carnassial tooth, has no third or inner root. An alisphenoid canal is present. The auditory bulla is depressed and scarcely at all inflated. The feet are plantigrade. There is no entepicondylar foramen to the humerus. As noted by de Blainville, a bear’s skeleton presents certain resemblances to that of man, dependent partly on the animal’s habit of sitting on the ischia, partly on the plantigrade method of walking. A. Tur Sxuun (Plates I—YV),. (1) Distinctive Features of the Skull in the Genus Ursus.—The skull is more or less elongated. The orbits are small and the post-orbital bar is complete. The palate is prolonged considerably behind the last molar tooth. An alisphenoid canal is present. The pterygoid has a well-developed hamular process. The following are the features in the skulls and teeth of bears, in which the ereatest amount of variation takes place, and to which special attention should be paid in attempting to discriminate between the different species : (1) The presence or absence of the anterior premolars ; (2) The length of the mterspace between c. and pm. 4, and between c. and pm. 4; (3) The form of pm. 4 and m. 3; (4) The width of the posterior narial opening ; (5) The shape of the jugal arcade ; 1 Flower and Lydekker, ‘ Mammals Living and Extinct,’ p. 586. 2 Tbid., p. 556. 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"On iS Tes ot 5 a8 Bos Peel bee | Be | eet | of (Fee eee eee | seul see | POS | oa lee | es | ee | 8s | ees peo |rae| #2 | Pee| 2s (UESl Sf | SEP |SCE |SEL| Par leas | ee lez? ef | ee | fee rSe| gel oe [Psi] ex | e2e)288) Beloea| ge|eee|/Pei| £8/£S8| ge | FE | Ze ° ee ee ee eo eee eee | aoe ee % ne 1 = eal a 28 a aie 5 SFist lee one Be ie) Be ‘STIONG UVAG {O SINGWHUASVEP[ WAILVUVAWOH JO mTaYy, (T) 12 PLEISTOCENE MAMMALIA. (6) The shape of the forehead; and (7) The form and size of the mastoid process of the periotic. Although the conclusion arrived at is, that it is impossible to separate U. horribilis (ferox) from U. arctos by the study of the skeleton, it has been thought advisable in the tables of comparative measurements to quote without comment the name previously assigned to any specimen. B. Dentition (Plate VI). (1) Distinctive Features of the Teeth in the Genus Ursus.—The dental formula is i, 3, c. +, pm. 4, m. 2, as in the dog. In Hyena it isi. 2, ¢. + pm. & m. 4, andem Felis 1. 3, c. +, pm. 3, m. +. Although the upper incisors increase somewhat in size from the first to the third, Ussws agrees with Felis and differs from Hyena in presenting a marked contrast in size between c. and i 3. The canine is dis- tinguished from that of the lion by the more massive character of the root. The three anterior premolars above and below are very small, one-rooted, and often early deciduous, especially the second, which is rarely present in the adult animal. Pm. 1 is situated close to the canine, pm. 3 close to pm. 4, which is the upper carnassial. This tooth lacks the antero-internally placed inner tubercle supported by a distinct root, which is so characteristic of Felis, Hyena, and Cais. Pm. 4 possesses, however, a postero-internally placed inner cusp which, as in other Ursidee, is not supported by a distinct root. The sectorial characters of pm. 4 are very little marked, and it is much smaller than m. 1. The crowns of both the upper true molars are longer than broad, and have flattened tuberculated erinding surfaces; the second has a large backward prolongation or heel. The lower carnassial (m. 1) has a small and indistinct blade and a greatly developed tubercular heel. The second molar is of about the same length as the carnassial, but with a broader and more flattened tubercular crown. The third is smaller. The milk-teeth are comparatively simple and shed at an early age. The following descriptions are of teeth of Ursus speleus, but in each case the differences presented by the teeth of bears of the arctos type are noted. It has been thought best when describing the teeth not to use terms involving assumptions of homology and require long explanatory prefixes. The terms cusp and tubercle are regarded as synonyms for small elevations on the surface of a tooth. The terms cone or lobe are used as denoting a rather larger elevation, and the terms talon or heel for posteriorly placed segments of a tooth. (2) Permanent Dentition of the Upper Jaw (see Pl. VI).—I. 1 and 2 are very similar teeth showing a prominent anterior, pointed, and somewhat backwardly directed cone or cusp, and a depressed triangular posterior area not bearing any & j ct , 4 URSUS. 13 definite cusp. The marginal portion of the triangular area is shghtly raised. The root, which in each case is rather more than twice as long as the crown, is con- siderably laterally compressed. In bears of the arctos type the posterior area is more sharply marked off from the cone than in the cave bear. I. 3 is a larger and more caniniform tooth, with a prominent, sharply reflected cone. There is no depressed posterior area as in I. 1 and 2, but a slight cusp is developed on the inner side, and from it a cingulum extends along the inner and posterior margin of the crown. The root is not so much laterally compressed as ml. l-and.2. In bears of the arcfos type this tooth differs only in size and in the slight development of the cingulum. C. The canine has the form usually met with in Carnivora. Its crown constitutes about one third of its length, and is frequently, though not invariably, marked by a wide, shallow groove along its inner face, and by a slight ridge along its posterior face. The crown is slightly longer and more recurved and pointed than in Hyena, and the root tapers more than in that animal, and is more massive than in Felis. Size constitutes the only difference between the canine of the cave bear and that of bears of the arctos type. Pm. 1 and 2, which are almost mvariably absent in the cave bear and irregular in their occurrence in bears of the arctos type, are small and simple teeth, with a low crown and long cylindrical root. Pm. 3, which is absent as a rule in the cave bear, is in bears of the avctos type a small tooth with well-developed cone and slight mdications of anterior and posterior cusps. The cingulum is slightly developed on the inner side. Pm. 4, the upper carnassial, has the blade divided mto a prominent anterior cone (a)! and a less elevated posterior cone (b), behind which is commonly a slight additional cusp. Lying postero-internally to the blade is a large inner cone or lobe (Pl. VI, fig. 1, c), which shows considerable variation. The cingulum is often strongly marked, especially antero-internally. There are two roots, a smaller anterior one supporting the anterior cone of the blade, and a larger posterior one supporting the posterior cone of the blade and the inner lobe. In bears of the arctos type the cingulum is not so strongly marked, the inner cone or lobe tending to be relatively larger than in U. speleus, and often having a slight additional cusp cut off from its posterior edge, sometimes also from its anterior edge. This inner cone or lobe, which is posteriorly placed, must not be confused with the inner tubercle characteristic of the upper carnassial mn Felis, Canis, and Hyena, which is anteriorly placed. M. 1 has a large, somewhat quadrangular crown with the surface raised into 1 This and the following letters refer to Plate VI. \\ ij 14, PLEISTOCENE MAMMALIA. a double row of low cusps lying along the inner and outer edges of the tooth. Of the four cusps along the outer border, the anterior and posterior are very small, the second from the anterior end being the largest. The cingulum is well marked along the inner surface. The tooth is fixed in the jaw by three roots, one lying internally and the other two antero- and postero-externally. In bears of the arctos type the cingulum is less marked, and the two principal cusps along the outer border do not appreciably differ in size. M. 2 is the largest tooth belonging to the molar series. The grinding surface is completely tuberculated, the two most prominent cusps or. elevations (() occupy the anterior half of the outer border, and behind them a third and much smaller cusp is often found. The anterior cusp tends to be the largest. There are four roots, one placed anteriorly, one near the middle of the outer surface, one postero- internally, and one on the inner surface near the anterior end. In bears of the arctos type the two antero-external cusps tend to be equal in size, and the tooth to narrow posteriorly more than in the cave bear. These distinctions, however, do not always hold, and are of little practical value. (5) Permanent Dentition in the Lower Jaw (see PI. V1I).—I. 1 is a small tooth with both root and crown much laterally compressed. The crown forms a single very slightly recurved cone, with a small tubercle on the outer side. I can detect no valid difference in bears of the arctos type. T. 3 has the root similar to that of I. I, but the crown is not so much compressed and the tubercle on the outer side is larger and placed lower down the crown than in I. 1. There is also an indication of a tubercle on the inner side of the crown, while from each tubercle a slight ridge runs downwards and backwards to meet its fellow at the base of the crown. In bears of the arctos type the outer tubercle is relatively more prominent than in U. speleus, while the inner tubercle and pair of downwardly and backwardly directed ridges are not present. T. 3 is a slightly larger tooth with the root triangular in section, the apex of the triangle being directed backwards. The outer tubercle (PI. VI, fig. 2, ¢) is very prominent and sharply divergent from the crown. A slight ridge passes backwards and downwards from it to meet another bounding the inner side of the crown. In bears of the arctos type the tooth differs only in its smaller size, and in the shghter development of the ridges. CO. This tooth differs from the corresponding one in the upper jaw in haying sometimes at any rate, both crown and terminal part of the root shghtly inwardly inflected on the main part of the tooth. The smaller size is the only respect in which c. of bears of the aretos type differs from the corresponding tooth of the cave bear. Pm. 1, 2, 3, which are absent as a rule in the cave bear, are all small conical URSUS. 15 teeth with low crowns and rather long cylindrical roots, pm. 1 being the largest and the one which most commonly persists. Pm. 4 is a small tooth showing much variation. As a rule, in addition to the principal cone, one or more of three little cusps may be developed, two placed respectively at the antero-internal (Pl. VI, fig. 6, 1) and postero-internal (Pl. VI, fig. 6,3) edges, and the third (Pl. VI, fig. 6, 2) slightly behind and to the imner side of the principal cone. In bears of the wictos type the development of inner cusps Is not so great as in the cave bear, and in many cases the tooth is entirely without them. M I. This is an elongated tooth, divided by a constriction into a posterior square portion (y) whose length is about one third of that of the entire tooth, and an anterior more elongated portion. The posterior portion, which represents the greatly enlarged heel or talon of the corresponding tooth in Canis, has the surface raised into a series of low cusps, the two most marked of which he on the imner border and are nearly equal in size. The anterior portion generally shows two prominent cusps, one forming the anterior extremity of the crown, one farther back and on the outer side of the tooth. There are several smaller and generally ill-defined cusps along the inner border. Hach portion of the tooth 1s supported by a strong root. The angle of divergence between the two roots varies much. In bears of the arctos type the constriction between the two portions of the tooth is not so marked as in the cave bear, and the cusps are less prominent. The hinder of the two, lying on the inner border of the posterior square portion of the tooth, tends to be larger than the anterior. The cusp forming the anterior end of the tooth is less marked than in the cave bear, and often has a small accessory cusp on its inner side. M. 2. The sides of this tooth are parallel, and the length is nearly twice the breadth. A shght constriction divides the tooth into anterior and posterior halves. The surface is somewhat uniformly tuberculated, the greatest elevation lying antero-internally. The tooth is fixed in the jaw by two stout roots, the posterior being the larger. I cannot detect any difference from the above in teeth of bears of the arctos type. M.3. The crown, which has parallel sides, is shehtly rounded im front and more markedly rounded behind. The length as compared with the breadth is about 5—1. The posterior border is sometimes rounded, sometimes more or less obliquely truncated externally. The surface of the crown is very uniformly covered with low tubercles (PI. VI, fig. 5), the largest being placed at the antero internal angle. There are three roots, one placed anteriorly and two posteriorly, but all three roots sometimes coalesce. In bears of the actos type the tooth is rather longer in proportion to its width 16 PLEISTOCENE MAMMALIA. than in the cave bear, and is as a rule rather more contracted posteriorly. The surface of the crown tends to be ridged rather than tuberculated. It proved impossible to obtain anything approaching a complete series of milk- teeth, and it therefore seemed best not to attempt a description of them. (2) TasLe or MEASUREMENTS OF THE SERIES or PERMANENT Trento FROM TORBRYAN, NEAR ToRQUAY, FIGURED ON Puats VI. il || oy ; : AH ea a cel th ee aL OS. | ips ; Call esealiaies |) |S 1 a rel : | A| aI EVE Vie Ha ile |e 1. Antero-posterior extent at base of | | CLOW Huiee. gat idan crn teo nen pean | 10 | 1:1 | 1-1 | 2:1 |0°75) 20 | 2°5 | 3-95) 0-9 | 0'9 | 0-95) 21 | 1°55) 2°7 | 3:05 2°55 |2. Maximum transverse measure | | TLOIG! sectemeceeitenee eeaaceere sec ouaen ie | 09 | 0-9 | 1:05, 1°6 | 055} 1°5 | 1-8 | 1-9 | 0-5 | 0°75) 1:05) 1-4 | 0-95) 1:2 | 1°8 | 1°85 (3. Maximum length measured along | a straight line from root to | GLOW caisson cee ec cane eneennee see 29 | 3:0 |3°7 | 89 29 |33 34 |825 o. THe Verteprat Cotumn (Plates VII, VIII). The vertebral column of bears shows few characters distinguishing it from that of other Carnivora. Probably the most noteworthy feature is the tendency to ankylosis in the sacral region in old animals, which may have as many as five sacral and pseudo-sacral vertebree (see Text-fig. 1, p. 19). There are fourteen thoracic and six lumbar vertebree as compared with thirteen thoracic and seven lumbar in Felis, Canis and Viverra, and fifteen thoracic and five lumbar in Meles. (3) Tastes oF CoMPARATIVE MEASUREMENTS OF BraR VERTEBRA. U. speleus, U. horribilis, U. horribilis, U, arctos, | Banwell Sandford No. 854 (College No, 218d (Taunton Mus.). | (Taunton Mus.). | of Surgeons). (Brit. Mus.). ! | ATLAS. 1. Maximum width ...... Renee eens Ade Haze sob ahs | 17:3 19°6 2. Median dorso-ventral diameter............ ee 52} | 44, 5°35, 3. Extreme width of the condylar articular SUELACES hs actos sangsoo mbar serie tee 8:1 65 82 4. Maximum width of neural canal. ... . | a 3°55 31 4-0 AXIs. 1. Length from anterior end of odontoid process to postero-ventral extremity Of -cembrumavin agen co ceaeeeeone eee 89 8:15! 69 81 2. Height from roof of neural canal to top OL TAC UN) (SOE) o00 002 590000 009000003 000050 6°35 3°55 | 402 44. 3. Transverse diameter across prezygapo- | DW SCS ihicuenaa rece carer reseee ARatau anti &4 6:85 | 56 68 4, Transverse diameter across postzygapo- physes............ Lise afeaieetehnn sted eee anteater 7:65 55 5°35 59 5. Length from anterior end of neural | spine to notch between postzygapo- DUYSES rnc queton eh onnaa nee 6:3 67 61 il 1 Fieured. & URSUS. 17 ComPARATIVE MEASUREMENTS OF BEAR VERTEBR® (continued). 4 3rd cervical. 4th cervical. 5th cervical. ow fai wa. A oop Ai ee) S38 Red o3 Rea RES Soa 0 Se See | 2 || S38 | Se £00 5a a) 2a £02 parca 238] 8 S53 Ss | 235 | oe 19 DS hey b= Yo) = See | ae Pe | Seas eed ae | be 1. Maximum length of centrum... measured from dorso-anterior to ventro-posterior edge ...... 4°75 48 45 4°55 4°35 5:0 2. Width across transverse pro- CESSES ......5. Adeiama eee lasiaeiee 10°7 12°6 Bae 13°35 12°5 13:3 3. Width across postzygapophyses| 6:0 6°25 78 6:95 6°35 7-0 4, Height from roof of neural canal to top of neural spine 2°15 1°35 wae 44. 37 445 q ! Figured. 6th cervical. 7th cervical. ( > ¢ a aS Z 3 i E 2 hp a ace 2 bo = | Bae Ss ad se ss OS papel eSB 288 Bal sob se aga SS b sa ; | cs 2 og <8 os SBn Dx alia BA De it aa a ie es cS) 1. Maximum length of centrum from dorso-anterior to ventro- posterior edge ...............0+5 43 50 4-0 45 2. Length from dorso-anterior to dorso-posterior edge of cen- PEUIMMCreee tinea oases ele aicete 3:2 37, 3:15 2°95 3. Width across transverse pro- GEESE Goonapocosonoobsvedsanas donna. 116 12°95 a6 12:0 12°15 4. Width across postzygapophyses 615 6:45 70 6:3 67 : 5. Height from roof of neural| q canal to top of neural spine 36 485 11:4 46 66 1 Fieured. 18 ~~ — A a ee oe PLEISTOCENE MAMMALIA. Comparative MEASUREMENTS OF BEAR VERTEBRZ (continued), 1st thoracic. 2nd thoracic 3rd thoracic. 4th thoracic. 4 a4 4 a4 = = aS} a URW ee eal b 5 7 || Bea |r eee eoa eee BEd | £28 | SS | Sak Sag £ag 208 S48 Us Sa See e= 2 So@ | 882 BoB | 882 5 B8 882 S58 “ae | S55 5 = Fe Mone See (Pat | see | ase “an eI E 5 E z E n | n 8 8 | a i) a 2) 1. Length from dorso-anterior to dorso- posterior edge of centrum ............ 3:11 2°9 3715) 2°65 2°81 29 2. Width across postzygapophyses ...... 6°55 4°35 AT 3°45 31 3:0 3. Width across transverse processes ...| 12°8 10:0 11:0 8°85 84 8°25 4. Leneth of neural spine from notch between prezygapophyses ............ 60 6:9 8°35 8:2 8:35 5th thoracic. 6th thoracic. 13th thoracic. | 14th thoracic. 1. Maximum length of centrum ......... vias 301 30 3°95 4-1} 2. Width across transverse processes ... 775 79 3. Length of neural spine from notch between prezygapophyses ............ 8:45 8:55 4°45, 45 ! Figured. 2nd lumbar 3rd lumbar 4th lumbar. J 5th lumbar. | 6th lumbar. is iS iS e bb ae | oa ae eee eee sD 22S we D - so ae D ~~ oD x S28 Bas Sag 27 Sag Sq 2a5 2-5 245 Sa5 R45 248 S28 E24) See Wee mee eles | “eel | Soe as bo2 | ir) ns i) 3) iC) ©) E EB E z E E 5 a 5 5 5 5 ea} -Q a 2 a 1. Maximum leneth of centrum ......... 4°35 4045 } 435 AAS 445 4-0 2. Width across processes bearing pre- PAF SENSO VSS. cocosccocaandacssacenou08 008 55 8:3 58 5°45 59 6:35 3. Height of neural spine from notch between postzygapophyses............ 48 65 5:15 5°25 5:1 44. 4, Width across transverse processes ... 11:4 13:0 14:7 14°3 13°4 1 Fioured. URSUS. 19 Fre. 1.—A dorsal, B ventral view of a sacrum (No. 48827), from Brixham, preserved in the British Museum; a, neural spine; b, foramen for exit of spinal nerve; c, articular surface for ilium; d, anterior face of centrum of first sacral vertebra; e, posterior face of centrum of third caudal or pseudosacral vertebra; f, ventral surface of centrum of first sacral vertebra; g, ventral surface of centrum of first caudal or pseudosacral vertebra. D. THE SHOULDER GiRDLE. The scapula in bears (Text-fig. 2) does not present any features of special interest. The British fossil specimens are almost invariably in a very frag- mentary state. j (4) Tasie or MeasuREMENTS OF THE SCAPULA. U. horribilis, No. 854 (R. Coll. of Surgeons Mus.). U. arctos, No. 218d (Brit. Mus.). 1. Length along line of spine measured from ACKOMIVOMY qeimep west vitiere cok nevis sm cioabiasietiastibe cussion palleuats 26'6 31°85 | By WSR eOTTA WACKHM one opcnoosnnnabsonaosenoaaacandaoaenpcecan 20°1 23°8 3. Maximum length of glenoid cavity .................. 5°95 61 4. Height from top of acromion to inner edge of llemor dic anyatiyg cr pe nsta- acct. era mais daisies seciaeies cei ee 8°35 109 5. Length from end of coracoid process to surface of bone behind glenoid cavity............:.:1ecceeeeeee 7°35 E. Tur Anrerior Lime. The humerus (Text-fig. 3) has a strong deltoid ridge. A supra-condylar foramen is not present. PLEISTOCENE MAMMALIA. Fie. 2.—Articular view of a portion of the left scapula (N.H. 93), from the Newhall Caves, Edenvale, Co. Clare, now preserved in the National Museum, Dublin. (1) (2) OS Zz Gigs a, glenoid cavity ; 6, acromion. Fia@. 3.—(1) Back view of right humerus, Sandford (Taunton Mus.). (2) Front view of left humerus, Grays, Essex (Brit. Mus.). (8) Antero-internal view of right ulna, Sandford (Taunton Mus.). (4) Front view of right radius, Sandford (Taunton Mus.). Ursus horribilis, and are drawn 4 natural size. All the above specimens are attributed to ‘ f| , H | 4 4 cv URSUS. (5) Tasne or Comparative MnasureMENTS OF Bones oF ANTERIOR Limes. 21 U. ferox fossilis= U. spelaus, U. speleus, horribilis, U. aretos, U. horribilis, Banwell Sandford Grays, Essex, No. 218a (Brit. |No. 854 (R. Coll. of (Taunton Mus.). | (Taunton Mus.). | No. pee Mus.). Surgeons Mus.). us.). HumMeERvs. 1. Extreme length ..................... +373! 39°15! 34:0 2. Diameter of proximal end pass- | ing across centre of articulat- ing surface and greater tuber- OHI coudononn cpa55boonpoRDoDBeaRa0no0 9:0 8:15 8:3 3. Vertical diameter of shaft at middle of deltoid ridge ......... 6°35! 36°87 4rd 4:95 38 4. Transverse diameter at same [OGWIMIS scooncooqosca0se00n05 00008550000 55 5°85 39 4:05 37 5. Maximum transverse diameter at distal @nd ...c.ccccss2ces-seee es 168 16°75 11:3 108 a 6. Maximum width of trochlea one ists nee on 71 Ravtivs. 1. Extreme length ..................... 42°40 $378? 339! 30°37 2. Right and left or transverse measurement at humeral arti- GIN HIOED Soo coo snencbaeneenona noo annnoe 5°55 61 46 41 3. Antero-posterior or vertical measurement at humeral arti- CUNATIOMY coesceiger cosemetnnneateeeiesee 44, 48 3°65 3:05 4. Transverse diameter at carpal articulation ...........00csseeee es 81 9:2 6°85 5:9 5. Vertical diameter at carpal arti- culation ............4. oooonsetoocoanc 4°8 5°45 AA, 35 6. Transverse diameter at middle ONS ARGE arcepcriew see see aeateinece ats 51 53 2-9 4. Vertical diameter at middle of Seu bGe ga terse it ers i alstaneeioctaetores 2°83 2:5 21 1 Left 2 Right. 3 Figured. U. speleus, Banwell U. horribilis, Kew Bridge, No. 24361 (Brit. U. ferox fossilis= horribilis, Tiford, No. 38512 | U. horribilis, No. 854 (R. Coll. U. arctos, No. 218a (Brit. (Taunton Mus.). Mus.). (Brit. Mus.). ize Surgeons Mus.). Mus.). ULNA. | | | 1. Extreme length ..................... 46:5 AAO | 38:6! | 3415 38°45! 2. Antero-posterior or vertical | | measurement at distal end of | | Siomoid motchwee reer eet 103 9°35 PU | 64 TB 3. Maximum transverse measure- | ment of olecranon.................- 10:7 9°75 8:25 6S 78 4. Transverse diameter at carpal EVAGKOWIENHO}0 Gooonenpspsno0oKbosobe5o 3°4 3°25 30 2:2 21 5. Vertical diameter at carpal articulation yescersesecceeeeeeeee eer 48 48 39 39 Length of Ist metacarpal............ 8:1 Me 2nd pes aE oenae ae 8:8 3 3rd Bo Mine Seraces 9:0 Ms 4th Bre esas CCS: 9°3 ty 5th Be We er eee ccre 9:4 1 Left. 22 PLEISTOCENE MAMMALIA. ¥. Tne Penvic Grrpie. This is characterised by the shortness of the ila and their evertion above. The British fossil specimens are almost always in a very fragmentary state. | (6) Taste or ComPpaRATIVE MrasurEmMeEnts. U. horribilis, No. 854 (R. Coll. of U. arctos, No. | | Surgeons Mus.). 218d (Brit. Mus.). | INNOMINATE Bonz. idle Wile psriremeneA NOMEN, 5s cononoxcoonnsn0eco9s oon csnHsB000n00000 31°85 375 2. Length from edge of acetabulum to dorsal or | anterior border of Uium ..............:ecceeeseeeneee ens 15:1 18°65 3. Vertical measurement of ilium at widest point... 121 14-7 /4. Thickness of ilium at middle of surface............ ae ee | 5. Antero-posterior diameter of acetabulum ......... S05 53 6. Length from acetabulum to posterior border of ASCH | cise secs casey, vo cacisesioroadaeea creates cestaosneeeien 11°25 117 7. Maximum diameter of obturator foramen......... 8:2 we 8. Measurement along ischium from symphysis to | EMG OE WSOTEM! SOWA “sosccc0ndonsaa> acksc0000 cvosaasoa090 12:2 | 16°9 Fig. 4.—Ventral view of left innominate bone (N.H. 197), from the Newhall Caves, Edenvale, Co. Clare, preserved in the National Museum, Dublin (4 nat. size). a, ilium; b, ischium; c, pubis; d, acetabulum ; e, obturator foramen. G. Tue Posterior Lime. This, as noted by Gaudry and Boule, tends to be somewhat shorter in propor- tion to the size of the animal in U. spelxus than in the bears of the arctos type. URSUS. 23 : The tibia is specially short, and Gaudry and Boule have suggested that this is perhaps 3 a disposition favourable for descending into the caves in which the animal lived. Owen! attempts to discriminate between the femur of the different species of fossil (1) ae \i\ Cty hi a? Oger eal un : \ OAC ENN BERL Fie. 5.—(1) Front view and (2) right side view of left femur attributed to U. horribilis, from Sandford (Taunton Mus.) (4 nat. size). bears. He says that in the brown and grizzly bears the femur is broader in proportion to its length, and the tuberosity above the internal condyle is larger than in the cave bear. He also states that in the cave bear the lesser trochanter _ projects a little beyond the inner margin, while in the grizzly and brown bears it is thrown wholly on the posterior surface of the bone. 1 «Brit. Foss. Mamm.,’ p. 97. 24 Fia. 6.—(1) Front view of right tibia, from Sandford (‘Taunton Mus.). (2) Postero-external view of right tibia, from Ilford, Essex (Brit. Mus.). (3) Patella, from Banwell (Taunton Mus.). All are drawn 4 natural size. The patella is attributed to U. spelzus, the tibia to U. horribilis. (7) TaBLEe oF CompaRATIVE M#ASUREMENTS OF Bones oF Posterior Lims. | ieee, | Wh hora U~ horribilis, wanton vstaatord Sandfordid No eon xGol No. 218% (Taunton Mus.)./(Taunton Mus.). Mee (Brit. Mus.). | Femur. i, Mipgapenban WMA pooeqnocn00000000 v02 o60 000000 600 352°5 } ate 40:2? 46°01 2. ''ransverse diameter at condyles ............ 115 ible 73 8:3 3. Antero-posterior diameter of head ......... 56 Fe 45 4°85 4, Vertical or antero-posterior diameter of | JOON ANG WANCCNS Sraanasco0neaea godaern0u00 savage ano 4:0 43 2°8 2:95 5. Transverse or right to left diameter of | shatbiatimaid diel eres. -eeerreeeerereee heer ceren reer 5:0 5:2 34 4:85 6. Transverse diameter at proximal end measured across head and great trochanter 13°45 260 98 1 Left. 2 Right. 3 Figured. URSUS. 25 TABLE OF CoMPARATIVR MrasuREMENTS—continued. pana extra U. ferox fossilis | U. horribilis, U. horribilis, | U.horribilis, | 2+) 0710 U. aretos, Sandford’ | Sandford” |= horvibilis, U- |No. 854 (R. Col. yo.’ >i8¢ (Taunton Mus.).|(Taunton Mus.). (Brit, Mus.), Mus.). (Brit. Mus.). TIBIA. 1. Maximum length . 36°5 | 31:5? 34°25 } 28°92 33°2 2. Transverse or right to left “diameter at | [oRosaiieney| ys aye [ase eer sao ceavenonaccosnoserancaaceacoee 11°35 9°85 8-9 79 8°25 a 3. Vertical or antero-posterior diameter at q proximal end measured from notch between articulating surface for femur and top of erest ......:. rice aces 7:3 Et 83 6°55 6°95 e Transverse diameter at distal Bidens. 81 gall 5:95 6:45 67 . Vertical diameter at distal end measured across elevation between articular faces for a (0B ERWINIA Fa wcngdanonadouadanod esd see coead oor ACdaeE NC: 5:1 4:25 2°95 6. Transverse diameter at narrowest part of RSINENEN cagmcoo caegdsetn Gd scoe ca BAO SEA aSR Ee cae aCoonBes 4:0 35 3:0 2°45 2°65 1 Wieured. 2 Right. eye Een U. horribilis U. horribilis, | U. horribilis, : 2 U. aretos, Sandtc fale Banaiordl oe ae (R. Col. No. 2180 (Brit, | (Taunton Mus.) |(Taunton Mus.). o MS Mus.). FIBuna. 1. Maximum length ..... 27-2 26°25 30°45 2. Transverse diameter at ‘distal end . exci n ei 17 2852 ii tae | 8. Vertical diameter at distal end............... 34 38 1:45 26 4. Transverse diameter at proximal end ...... 175 17 sels 5. Vertical diameter at proximalend ......... 2°5 2°05 CALCANEUM. 1. Length.. eeistels Uae wenieceee 11-25 11°4 83 gyi 2. Maximum transverse diameter. eee eee 715 75 Onl, 5°55 ASTRAGALUS. Right to left diameter .................006 88 a 5:3 46 M@raTarsats. Length of 1st metatarsal ! 6:45 7:2 5 2nd » 7-25 8:65 3rd WN NARS HR. 73 9:05 ia Ath i afl Met ie a 8:3 10:25 i 5th Bs 8:8 10°5 | | 1 All measured along plantar surface. = A large bone lacking proximal end. } IV. COMPARISON OF THE CAVE, BROWN, AND GRIZZLY BEARS. The subject of the mutual relations of the Pleistocene bears is one of very great difficulty, and very varying opinions have been expressed as to the number of species. Most paleontologists have recognised three species, viz., U. speleus, U. 4 horribilis (= ferox, = ferox fossilis, = priscus), and U. arctos. Owen, Busk, Boyd j A, | 26 PLEISTOCENE MAMMALIA. Dawkins (previous to 1877), Lydekker (when writing in 1885), Woodward and Sherborn, adopt the above three-fold division of the Pleistocene bears. With very few exceptions paleontologists have considered U. speleus to be dis- tinct from the rest, but as early as 1844 de Blainville expressed doubts on this head, considering that the differences between the fossil bears were merely racial. Adams too, in 1880 and also in 1881, suggested that the bones attributed to U. speleus might be only those of larger individuals of the same species as U. ferow, and that all the British fossil bears might be regarded as races of one species. CoMPARTSON OF THE Cave Brar with Brars or tHe arctos Typr. The following characters have been quoted by various paleontologists, princi- pally Owen,’ Busk,” and Lydekker,?’ as distinguishing the cave bear from those of the arctos type : (a) Distinguishing Characters drawn from the Teeth. Cave Bear. Bears of arctos Type. (1) The three anterior premolars of both The three anterior premolars, espe- jaws are generally lost very early, all cially pm.1 and pm. 3 of both traces of their alveoli commonly dis- jaws are far more persistent. appearing. The complete loss of the three anterior premolars is undoubtedly almost or quite universal in the large cave bear skulls, but is not universal in the case of smaller individuals attributed to the cave bear. Thus Gaudry and Boule* have shown that in the small race from Gargas pm. 3 1s not always lost, and Owen? mentions a jaw from Torquay which retains pm. 1. Newton® too attributes to U. speleus a small jaw from the Forest Bed in spite of the retention of pm. 1. Cave Bear. Bears of the arctos Type. (2) M. 2. has a more or less oblong form, the M. 2 is more constricted behind, and sides being nearly parallel, and the hind ~ the erinding surface of the un- end not much narrower than the middle, worn tooth is more compressed and never or hardly ever pointed. The from side to side than in U. erinding surface when unworn is com- speleus. Of the three outer cusps paratively flat. On the outer border the two anterior are more nearly are three cusps of which the hindmost equal in size than in U. speleus is very low and soon worn off (Busk). (Busk). 1 «Brit. Foss. Mamm. and Birds,’ p. 86, et seq. * «Trans. Zool. Soc.,’ x, 1877, p. 60. 3 «Palaont. Indica,’ ser. 10, vol. ii, p. 210; and ‘Proc. Zool. Soc.,’ 1897, pp. 412 4 «Matériaux pour lhistoire des temps quaternaires,’ p. 109. » «Brit. Foss. Mamm. and Birds,’ p. 91. 6 «Vert. of Forest Bed” (‘Mem. Geol. Sury.’), p. 5. 426. ——_-— , SO ee ee en Ly Fy ae eye URSUS. 27 I cannot detect any constant differences between m. 2 in the two cases, though in some instances there is a tendency for the crown of the tooth to show less posterior contraction in the cave bear than in the others. Cave Bear. Bears of arctos Type. (3) In pm. 4 besides the principal cone Pm. 4 has either only the principal there are usually on the inner side two cusp or at most a very small and always one smaller cusp, of which internal tubercle corresponding one is anterior in position to the prin- to the hinder of those met with cipal cusp (Busk). Hensel’ and Owen in U. speleus (Busk). make similar statements, and Owen also mentions a ridge extending along the outer and back part of the base of the crown as characteristic. Lydekker’ says pm. 4 is relatively short, the mner tubercles are very large and the first is placed more on the inner side than in U. aretos. It is undoubtedly the fact that there is a greater development of accessory tubercles in the case of the tooth im the cave bear than in bears of the actos type, and this tooth probably affords better characters for the separation of the cave bear than any other part of the skeleton. An examination of a large series of skulls, recent and fossil, of bears of the arctos type, shows that although very often pm. 4 is without any internal cusps or possesses only one small one, and though they never show the development that occurs in U. spelxus, yet that in some cases two or even three may be present. Further information with regard to the develop- ment of these cusps is given in the table on p. 31; cf. also Pl. VI, fig. 6. Cave Bear. Bears of arctos Type. (4) M.3 is broader in proportion to its M. 3 is subtriangular and narrower length than in bears ef the arctos type. behind than in U. spelwus. In The outer surface is divided into two typical examples there is no sulcus distinct but low cusps by a deep sulcus. on the outer border. The grinding The grinding surface is minutely tuber- surface is coarsely ridged,not tuber- culated (Busk) (cf. Pl. VI, fig. 5a). culated (Busk) (cf. Pl. VI, fig. 5 0). Great stress is laid especially by Busk on the structure of this tooth. It is certainly somewhat broader in proportion to its length in the cave bear than in bears of the arctos type. While no example of m. 3 from a bear of the actos type 1 «Sitzb. Naturf. Freunde Berlin,’ 1876, p. 49. * “Proce. Zool. Soc.,’ 1897, pp. 412—426. 28 PLEISTOCENHK MAMMALIA. was met with showing the peculiar uniform tuberculation of some of the cave bear specimens, some of the small mandibles attributed to this species have the surface of m. 3 wholly or partially ridged as in bears of the arctos type. (5) Boyd Dawkins! states that the canine is on the whole more massive in the cave bear than in the grizzly, especially as regards the root. “It is also generally but not always absolutely larger in the crown, as Prof. Busk has remarked in his description of the teeth from the Brixham Cave.” (b) Distinguishing Characters drawn from Parts other than the Teeth. (1) The relatively enormous size of the cave bear.—The size of the cave bear’s skull, though as a rule much greater than that of the fossil representatives of the brown and grizzly bears, is not so much greater than that of the huge grizzlies of Alaska and brown bears of Kamtchatka. Also, as pomted out by Owen, a mandible from the Forest Bed which he figures,” and which on account of the complete _ absence of the anterior premolars he attributes to U. spelxus, is a good deal smaller than a second mandible from Manea fen, which, owing to the large alveoli for pm. 1, 3, he regards as belonging to U. arctos. (2) Lhe relatively great length of the interspace between c. and pm. 4 in the cave bear.—This certainly is subject to a very large amount of variation, as the table of measurements on p. 11 shows (cf. also Pl. V). Busk says it is a feature distin- guishing the cave from the brown bear but not from the grizzly. Owen quotes it as also distinguishing the cave bear from the grizzly. (3) The velatwe narrowness of the posterior narial opening in the cave bear.— This, again, is a feature showing much variability (see the measurements on p. 11). (4) The arched character of the frontal region im the cave bear's skwll—In many of the huge skulls from French, German, and Belgian caves this is very marked, the skull rising into two considerable bosses at the junction of the frontals and nasals owing to the enlargement of the frontal sinuses. In some of the smaller cave bear skulls, on the other hand, it is scarcely more noticeable than in those of the brown and grizzly bears. It is probably a character which increased with age and became specially marked in the old males. Owen refers to de Blainville’s suggestion, that the development of the frontal sinuses depended on the cave bear’s breathing a fresher, dryer, and more invigorating atmosphere than its present-day allies. (5) The rapid approach of the temporal crests so as to form an obtuse angle posteriorly. 1 «Quart. Journ. Geol. Soc.,’ xxxi, 1875, p. 251. 2 «Brit. Foss. Mamm. and Birds,’ p. 106. URSUS. 29 (6) Uhe convex character of the lower jaw (Owen and Falconer).—Neither of the characters 5 or 6 seems to be constant. (7) The relative shortness of the limb bones, especially of the tibia in the cave bear (Gaudry and Boule).—This appears to be a constant character. (8) The relative weakness of the hind limbs (Gaudry and Boule).—Gaudry and Boule’s suggestion in this connection has been referred to on p. 22. (9) The relative narrowness of the femur in proportion to its length, the small size of the tuberosity above the internal condyle, and the projection of the lesser tro- chanter « little beyond the inner margin in the cave bear (Owen). CoMPARISON OF THE VARIOUS Brars or THE arctos Tyer. With regard to the distinction between the brown and grizzly bears, there is by no means such a consensus of opinion as there is concerning the distinction of the cave bear. Not only de Blaimville (1844), but Middendorff (1851), Miller (1872), Busk (1873), Boyd Dawkins (1877), Lydekker (1884 and 1885), and Brown (1894), doubt whether the brown and grizzly bears can be separated from one another. The points of difference, whether valid or otherwise, have been noted as follows, and are mainly due to Owen and Busk.} U. arctos. U. horribilis. 1. m. 2. The unworn crown is much compressed; The unworn crown is less compressed, and there there are only two cusps on the outer border of the tooth, of which the anterior is con- siderably the larger, and the posterior has in most cases a small portion im front con- stricted off so as to form an accessory tubercle between the two cusps (Busk). . pm. 4 tends to be relatively long. . There is a relatively narrow space between c. and pm. 4. . The inner posterior cusp or tubercle of pm. 4 is very small or absent, and if present there is no bifid posterior talon projecting from it (Busk). : are occasionally three outer cusps ; the anterior two are more nearly equal in size than U. arctos, and the third is always small and often wanting. There is no accessory tubercle cut off from the anterior border of the posterior cusp (Busk). pm. 4 tends to be relatively shorter, and there is more of a shelf-like projection of the cingulum at the antero-internal corner (Brown).? There is a relatively wide space between ¢. and pm. 4. The inner posterior cusp of pm. 4 is better deve- loped than in U. arctos, and the posterior talon is commonly bitid or marked by two longi- tudinal ridges running back from it to the end of the tooth (Busk). 1 «Trans. Zool. Soc.,’ x, 1877, p. 60. * «Proc, Acad. Nat. Sci. Philad.,’ 1894, p. 119. 30 PLEISTOCENE 5. The crown of m. 3 is usually more angular behind than in U. horribilis. There is usually no sinus or constriction on the outer border. The grinding surface presents a few coarse folds, but is never tuberculated in the slightest degree (Busk). 6. The jugal arcade is more circular (Busk and Adams). 7. The posterior narial openings are wide (Busk and Adams). 8. The angular crotchet is less thick and incurved than in U. horribilis (Busk). The coronoid process is rather less broad and high (Owen). 9. The claws are less long and_ straight (Merriam). MAMMALIA. The crown of m. 3 is usually less angular behind than in U. arctos. In teeth of the typical triangular form there is no sulcus on the outer border. When the tooth is more elongated it presents a shallow sinus dividing the outer border. The grinding surface is coarsely ridged, rarely tuberculated (Busk). The jugal arcade is more elliptical (Busk and Adams). The posterior narial openings are of medium width (Busk and Adams). The angular crotchet is thicker and more incurved than in U. arctos. (Busk). The coronoid pro- cess is rather broader and higher (Owen). The claws are longer and straighter (Merriam). The constancy and importance of the above supposed distinctions may now be considered. (1) The differences to which Busk refers are very slight, and so far as my own observation goes, quite imconstant and unreliable. (2) The skulls of U. horribilis in the Zoological Department of the British Museum do not show any marked projection of the cingulum at the antero-internal corner of pm. 4, or that the tooth tends to be shorter than in U. arctos. (3) Busk considered that the relative length of the interspace between c. and pm. 4, on which Owen laid stress, was not constant. This is also shown by the measurements in the table on p. 11. (4) Nearly all paleontologists have laid stress on the structure of pm. 4, this being specially the case with Busk. Lydekker considered that Busk attached undue importance to the structure of the talon. Brown, too, remarks that two skulls of U. horribiis in the British Museum do not possess the longitudinal ridges con- sidered by Busk to be characteristic of pm. 4 in this animal, while on the other hand a skull of the Isabelline bear, a variety of U. arctos, possesses them. (6) The elliptical character of the jugal arcade is variable. In the case of two grizzly bear skulls in the College of Surgeons’ Museum, in No. 856, the jugal arcade is more elliptical than in the brown bear skull No. 836, while in the grizzly bear skull No. 854 it is not more elliptical. The skull from Ballymahon in the British Museum, attributed by Adams to the grizzly bear, has the jugal arcade not more elliptical than the brown bear skull No. 218e in the British Museum. (7) It is generally the fact that the posterior narial opening is wider in the , “I? @ Se ee a a ee Pe ee a a a re URSUS. 31 brown than in the grizzly bear, but this difference does not always hold, and skulls sometimes show a remarkable amount of variation in this respect. Dr. H. Woodward! quotes Dupont as stating that skulls of brown and grizzly bears may be distinguished by the fact that U. arctos has only the last small upper premolar (7. e. pm. 3), while the grizzly has also pm.1. Whether this were a true distinction could only be determined by reference to recent skulls, and in these it emphatically does not hold. The following table shows the distri- bution of the small premolar teeth in a number of bears’ skulls, recent and fossil, and it will be seen that all the recent skulls of the brown bear referred to, show pm. 1 in addition to pm. 3; in one case also pm. 2 is present. | (8) T'aBLE sHOWING DEVELOPMENT OF SMALL PREMOLARS AND OF CUSPS ON PM. 4 IN BEARS OF THE arctos TYPE. | Cusps 0 pm. 1. | ee 2. | pm. 3. pm. 1. pm. 2. pm. 3. = ik | CRANIUM AND MANDIBLE. U. arctos, Kamtchatka, No. 91.12.18.13 (Brit. + + + + ae de No cusps. Mus.) | U. arctos, Jesso, No. 96.4.27.1 (Brit. Mus.)...... + aa + + Oneside aa 1, 2,3. U. arctos var. piscator, N.W. Asia, No. 93.9.10.1 + Fae + + Hoe ae 2 (small). (Brit. Mus.) U. arctos, No. 61.4.1.3 (Brit. Mus.) ............... Alv. one 355 Aly. Aly. aa ae No cusps. side U. horribilis, No. 78.6.18.1 (Brit. Mus.) ......... + Es +, Fe ae as 2 U. horribilis, No. 58.6.18.10 (Brit. Mus.)......... + aes Oneside + ee oe No cusps. U. horribilis var. horriezus, No. 67.2.23.3 + zie + + + + No cusps. (Brit. Mus.) U. horribilis, Muggendorf (Brit. Mus.) ......... + a + + nee it No cusps. U. horribilis,! Ballynamore (Brit. Mus.) ......... Alv. Alv. Alv. U. horribilis, Clonburne (Brit. Mus.) ............ Aly. one Alv. U. horribilis, Ballymahon (Leeds, cast Brit. Aly. os Aly. Mus.) U. arctos,! Bourn, Lincoln (Mus. Pract. Geol.) + oe + Alv. a a ae U. arctos,! Burwell fen (Sedgwick Mus.)......... 208 re + + rt. Sea Aly. rt. | No cusps. side side U. arctos,! Manea fen (Sedgwick Mus.) ......... Alv. + + U. arctos' (labelled U. speleus), Crayford,| Alv. Aly. + No. M.5041 (Brit. Mus.) | MANDIBLE. | U. arctos, Manea fen (Fig. Owen, p. 106, fig. sis ee fe Alv. BAY) 1673. H. Vollgnad, ibid., an. iv, obs. clxx, p. 226. 1732. F. EH. Briickmann, ‘ Bresl. Saml.,’ p. 628, and ‘ Hpist. Itin.,’ p. 32. 1774. J. F. sper, ‘ Ausfiihrliche Nachricht—Zoolithen’ (Bayreuth). 1784. J. F. Esper, ‘ Herits par la Soc. Nat. de Berlin,’ v. 1794. J. Hunter, “ Observations on Fossil Bones presented to the Royal Society,” Slelomll, hens, Ibooany, 70. 4407. 1794. J. C. Rosenmiiller, ‘De Oss. Foss.,’ p. 18 (Leipzig). 1794. J. C. Rosenmiiller, ‘ Beitr. Gesch. Foss. Knochen,’ p. 44 (Weimar, a German reprint of the above). 1803. J. F. Blumenbach, ‘Spec. Archeeologie Telluris,’ 1, p. 12 (Gottimgen). 1804. J.C. Rosenmiiller, ‘ Abbild. u. Beschreib. der Foss. Knochen des Hohlen- biiren’ (Weimar). 1806. G. Cuvier, “Sur les ossemens du genre de l’ours,’ ‘Annales du Museum,’ vu, p. 301. 1810. G. A. Goldfuss, ‘ Verhandl. k. Leopold-Karol. Acad. der Naturforscher,’ x (2), p. 260. 1822. W. Buckland, “ Account of an Assemblage of Fossils—Kirkdale Cave,” lela, Meine, Caan, [Oo 7. 1 «Proce. Acad. Nat. Sci. Philad.,’ 1894, p. 119. 1823. 1823. 1825. 1828. 1830. 1833. 1842. 1844. 1846. 1849. 1851. 1851. 1851. 1860. 1860. 1862. 1864. 1864. 1864. 1867. 1869. 1871. 1872. 1873. URSUS. 33 W. Buckland, ‘ Reliquiz Diluviane,’ p. 35. W. Clift and J. Whidbey, “‘ On some Fossil Bones—Oreston,” ‘ Phil. Trans.,’ exiu, p. 88. G. Cuvier, ‘ Recherches sur les Ossemens fossiles,’ ed. 3, iv, p. 340. J. B. Croizet and A. C. G. Jobert, ‘ Recherches sur les Ossemens fossiles du Dépt. du Puy-de-Dodme,’ p. 828. M. de Serres, “‘ Recherches sur les Ossemens fossiles du midi de la France,” ‘Bull. Univ. des Sci. Nat.,’ no. 19, xu, p. 161. P. C. Schmerling, ‘ Recherches sur les Ossemens fossiles—Cavernes de Liége.’ R. Owen, “ Report on British Fossil Mammalia,” ‘ Rep. Brit. Assoc.,’ p. 62. H. M. D. de Blainville, ‘ Ostéographie,’ ii, K. R. Owen, ‘ A History of British Fossil Mammals and Birds,’ pp. 77-108. R. Ball, ‘ Proc. Roy. Irish Acad.,’ iv, p. 416. A. Massalongo, ‘ Osteologia degli Ossi Fossili del Veronese’ (Verona). J. A. Waener, ‘“‘Charact. der in Héhlen um Muggendorf . . . Siugthier- arten,’ ‘ Abhandl. k. bayr. Akad. Wiss.,’ vi, 1. abth., p. 198. A. 'T. v. Middendorff, “ Untersuchungen titber Schadeln des gemeinen Land- baren als krit. Beleuchtung der Streitfrage u. d. Arten fossiler Hohlen- baren,” ‘ Verhandl. Mineral. Gesell. St. Petersburg.’ A. Carte, “Description of Fossil Bones discovered . . . at Shandon,” ‘Journ. Roy. Dublin Soe.,’ 1, p. 352. H. Falconer, “On the Ossiferous Caves . . . of Gower, Geol. Soc.,’ xvi, p. 487. W. Boyd Dawkins, “On a Hyzna Den at Wookey Hole, near Wells,” ibid., xvi, p. Lilo. A. Carte, ‘‘ On the Former Existence of the Polar Bear in Ireland,” ‘ Journ. Geol. Soc. Dublin,’ x, p. 114. Fie Dennyeyenoc: Yorks, Geolvand Polytasec... tv, Pp. O38. J. H. Gray, ‘ Catal. Carniv. Pachyderm. and Edentate Mamm. in Brit. Mus.,’ [De Paletie G. Busk, ‘‘ On Certain Points in the Dentition of Fossil Bears, and on the Relation of U. priscus to U. ferow”’ (abstract), ‘Quart. Journ. Geol. Soc.,’ xxi, p. 342. J. A. Allen, “Catal. of Mammals of Massachusetts,’ ‘Bull. Mus. Comp. Zool.,’ i, p. 184. H. Woodward, “ Visit to Royal Museum of Natural History, Brussels,” “Grell, Wiayee.° sant, jos 17, A. Miller, ‘Drei in der Provinz Preussen ausgegrabene Birenschidel ’ (Konigsberg). G. Busk, “‘ Report on the Exploration of Brixham Cave—Animal Remains,” oe inleieanis:,; (c¢lxams ps 499: 2) 6 Quart. Journ. 5 34 PLEISTOCENE MAMMALIA. 1875. W. Boyd Dawkins, “On the Mammalia found at Windy Knoll,” ‘ Quart. Journ. Geol. Soc.,’ xxxi, p. 250. 1876. W. Boyd Dawkins, “ On the Mammalia and Traces of Man found in the Robin Hood Cave,” abid., xxxu, p. 248. 1876. R. Hensel, “‘ Ueber die Unterschiede zwischen Ursus speleus u. U. arctos,” ‘Sitzb. Naturf. Freunde, Berlin,’ p. 48. 1876. J. A. Allen, “ Geographical Variation among North American Mammals,” ‘Bull. U.S. Geol. Surv. of Territ.,’ 1, 4, p. 334. 1877. W. Boyd Dawkins, ‘On the Mammal-Fauna of the Caves of Cresswell Crags,” ‘ Quart. Journ. Geol. Soc.,’ xxxiui, p. 598. 1877. G. Busk, ‘On the Ancient or Quaternary Fauna of Gibraltar,” ‘ Trans. AXON, \XOCry 56 (C)), 10> (GO- 1877. R. Johnson, ‘‘ An Approximate List of the Extinct Mammalia of Norfolk,” “Trans: NorkNat. Soc.,c1. p00: 1877. A. L. Adams, “ Observations on Remains of Mammals found in a Fossil State in Ireland,” ‘Journ. Roy. Geol. Soc. Ireland,’ iv, p. 246. 1878. A. L. Adams, “ Report on History of Irish Fossil Mammals,” ‘ Proc. Roy. Irish Acad.,’ 2nd ser., i, p. 94. 1879. A. L. Adams, ‘“‘ Report on the Exploration of the Shandon Cave,” ‘ Trans. Roy. Irish Acad.,’ xxvi (Sci.). 1880. A. L. Adams, “On Recent and Extinct Irish Mammals,” ‘Sci. Proc. Roy. Dublin Soc.,’ ui, p. 49. 1880. H. T. Newton, “ Notes on Vertebrata of Pre-glacial Forest Bed of H. of England,” ‘Geol. Mag.,’ dec. ii, vol. vu, p. 1538. 1881. A. L. Adams, “ Report on Animal Remains of Bone Cave of Bal aan atv, co. Waterford,” ‘ Trans. Roy. Dublin Soe.’ (2) 1, p. 201. 1882. H. T. Newton, ‘‘ Vertebrata of Forest Bed Series of Norfolk,” ‘Mem. Geol. SWIPE JO> Oe 1884. R. Lydekker, “ Post-tertiary and Tertiary Vertebrata, Siwalik and Narbada Carnivora,” ‘ Palzeont. Indica,’ ser. 10, vol. u, p. 202. 4 1885. R. Lydekker, ‘Catal. of Fossil Mammalia in the British Museum,’ pt. i, p- 173. 1887. A. Gaudry, “Le petit Ursus speleus de Gargas,” ‘Comptes Rendus,’ civ, p- 740. 1889. H. Schaff, “‘ Ueber den Schiadel von Ursus arctos,” ‘ Arch. f. Naturgeschichte, p. 244. 1890. A. Smith Woodward and C. Davies Sherborn, ‘ A Catalogue of British Fossil Vertebrata,’ p. 388 (London). 1892. A. Gaudry and M. Boule, ‘ Matériaux pour Vhistoire des temps quater- naires, fasc. iv, p. 105. 1893. 1894. 1896. 1897. 1903. 1906. A. A. URSUS. 35 Smith Woodward, “The Har Dalam Cavern, Malta... . Report on the Organic Remains,” ‘ Proc. Roy. Soc.,’ vol. liv, p. 278. H. Brown, “On the True Characters and Relationships of Ursus cinna- momeus,’ ‘ Proc. Acad. Nat. Sci. Philad.,’ xlvi, p. 119. C. H. Merriam, “ Preliminary Synopsis of the American Bears,” ‘ Proc. Biol. R. R. Soc. Washington,’ x, p. 65. Lydekker, “The Blue Bear of Thibet, with Notes on the Members of the Ursus arctus Group,” ‘ Proc. Zool. Soe.,’ p. 412. F. Scharff, “ Exploration of the Caves of Kesh, co. Sligo (Mammals except Man),” ‘Trans. Roy. Irish Acad.,’ xxxii, B, pt. 4, p. 201. . F. Scharff, “ Exploration of the Caves of co. Clare (Animal Remains except Birds),” ibid., xxxiul, B, pt. 1, p. 48. PLATE I. 3 ; ~Puotsrocenn Buars (U. speleus). ie E a Z ae : . Cranium and Mandible. F — . Pi, a a 1. Lateral view of a skull from Banwell (one quarter natural siz ‘i . or } view of a cranium from yedtord (one third se iene # 3. Lateral ia : 3 Both specimens are in the Taunton Museum. ie 7 . a. Mastoid process of periotic. a : f . | b. Post-orbital process of frontal. se c. Jugal. & 2 3 d. Hxternal auditory meatus. % Nasal. ; PALZONTOGRAPHICAL SOCIETY. 1906. Reynolds, Pleistocene Bears. Ee d.Green del. lith.et imp. URSUS SPELAUS. Cranium & mandible. PLATE It. PLEISTOCENE Bears. Cranium and Mandible. (The cranium one third, the mandibles one half natural size.) | Fic. 1. Posterior view of cranium of U. speleus from Banwell. 2. Palatal view of mandible from Bancvelll. 3. Left mandibular ramus from Sandford, seen from the outer side. 7 All these specimens are preserved in the Taunton Museum. The are attributed to U. spelzus. | a. Occipital condyle. b. Mastoid process of periotic. d. Condyle of mandible. Angle of mandible. Coronoid process. ' PALZONTOGRAPHICAL SOCIETY. 1906. Reynolds Pleistocene Bears. deal ll. J.Green del.iith et imp URSUS SPELAUS. Cranium & mandible. a SS - PLATE III. Pretstocenge Bears (U. arctos). Cramum. (One third natural size.) Fic. 1. Dorsal 2. Lateral - view. 3. Ventral a. Post-orbital process of frontal. b. Occipital condyle. c. Post-orbital process of jugal. d. Mastoid process of periotic. e. Anterior palatine foramen. Ff. Post-glenoid process of squamosal. g. Post-glenoid foramen. h. Infra-orbital foramen. ?. Lachrymal foramen. This specimen, which is preserved in the British Museum (Nat. Hist.), was found near Ballinamore, co. Leitrim. It is referred to as U. ferow fossilis by Busk, ‘ Phil. Trans.,’ clxi, p. 543 (1873), and Leith Adams, ‘ Trans. Roy. Irish Acad.,’ xxvi, p. 225 (1879). PALZONTOGRAPHICAL SOCIETY.1906. Reynolds, Pleistocene Bears. d.Green del. hth.et imp. URSUS ARCHOS: 1UTM.. Cran Fia. PLATE IV. Prierstocene Buars (U. arctos). Cranium and Mandible. (Two fifths natural size.) 1. Dorsal view of the cranium. 2. ‘he skull seen from the left side. 3. Ventral view of the cranium. This specimen was found in the peat of Burwell fen, and is preserved in the Sedgwick Museum, Cambridge. Lettering as in Plate IIT, with the addition of— Condyloid foramen. End of sagittal crest. Foramen lacerum posterius. Carotid foramen. Hxternal auditory meatus. Hustachian canal. Posterior aperture of alisphenoid canal. Paroccipital process of exoccipital. Angele of mandible. PALZONTOGRAPHICAL SOCIETY. 1906. s, Pleistocene Bears. ; a l Reyno oul imp ae + vo 5 4 3 i" xo) (S) o o iI 9 bro) o ro inte “4 ame) 5 3 O a & a6 ni (9) ae > @ Cc (©) PLATE V. PuErsroceNne Brars (U. arctos). Mandible and Portions of Cranium. (Fig. 3 one half, the others two thirds natural size.) Hach figure shows the palatal aspect. | Fie. 1. Part of cranium from Crayford (No. M. 5041). Only the teeth of the right side and the neighbouring portion of the palate are figured. In the British Museum this specimen is labelled U. speleus, but as all the upper premolars are represented, it seems better to attribute it to U. arctos. 2. Part of the right maxilla and premaxilla from Manea Fen (No, 40405). 3. Mandible from Burwell Fen. 4, Anterior part of cranium from Manea Fen. 5. Anterior part of cranium from Bourn, Lincolnshire. The Manea Fen cranium (fig. 4) and the Burwell Fen mandible (fig. 3) are preserved in the Sedgwick Museum, Cambridge. The Bourn cranium (fig. 5) is in the Museum of Practical Geology, Jermyn Street. The other specimens are in the British Museum (Nat. Hist.), South Kensington. a. Mandibular condyle. b. Coronoid process. —— === ae = ——s SSE See SESS — ——= ——S = Se eee PALZONTOGRAPHICAL SOCIETY.1906G. 4d -iltth.e deli Cc @.Green crama. > WRSUS ARcTlaS: ortions of Mandible &p SAPS. € ene B 1Stoe Ple lees re ty. for pi pin p : TO ” Reynolds . = | Yl. 6 d. In fig. 6, 1 = anterior PLATE VI. PLEISTOCENE Brars. Permanent Dentition. (Natural size.) Left upper teeth seen from the inner side ; U. arctos, Torbryan Caves, Left lower teeth seen from the inner side Torquay (Mus. of Pract. Left upper teeth seen from the outer side Geol., Jermyn St.). Left lower teeth seen from the outer side Grinding surface of left m. 3 of U. speleus, Kent’s Cavern, Torquay (Brit. Mus.). Grinding surface of left m. 3 of U. arctos, Torbryan, Torquay (Brit. Mus.). Left pm. 4 U. speleus, mandible No. M. 5995, Cromer Forest Bed (Brit. Mus.). Left pm. 4 U. ferow (horribilis), mandible No. 22029, Grays, Essex (Brit. Mus.). Right pm. 4 U. spelexus, mandible from Bacton (Brit. Mus.). Right pm. 4 U. arctos, a recent skull, No. 96.4.27.1, from Jesso, Japan (Brit. Mus.). All the teeth in fig. 6 are seen from the inner side. a. Anterior cone. h. Posterior cone. c. Inner cusp or lobe. d. Anterior external cusp. e. Outer tubercle. | of the small cusps referred to in the last column of 2 = middle | the table on p. 31. 3 = posterior “OTA AWE ‘duit 72 491] 8p 4992 “SANSHN (Pe Fe zy WA Nel “Sueog 9us007}sta[q spjousey “9061 ALHFIO0S TYOIHdVYDOLNOFZ1Vd PATE Sane PLEISTOCENE Brars. Vertebre. (Two thirds natural size.) Fie. 1. Atlas, dorsal aspect. 2. Axis, seen from left side. 3. Fourth cervical, front view. 4. Fourth cervical, seen from the right side. 5. Seventh cervical, back view. 6. First thoracic, front view. le All the above specimens are from Sandford Hill, Somerset, and are preserved in the Taunton Museum. b. C. d. & Second thoracic, seen from the left side. Vertebrarterial canal. Neural spine. Neural canal. Odontoid process. Anterior articulating surface for atlas. Pre-zygapophysis. Post-zygapophysis. Transverse process. Foramen for exit of spinal nerve. PALZONTOGRAPHICAL SOCIETY. 1906. Reynoids, Pleistocene Bears. d.Green del. ith et imp IR SIGS Vertebre. PLATEH VIII. PLEISTOCENE Bears. (Vertebrex.) (Two thirds natural size.) Fie. 1. Third thoracic, front view. 2. Fifth thoracic, left side view. 3. ‘Twelfth thoracic, left side view. 4. Twelfth thoracic, front view. 5. Fourteenth thoracic, dorsal view. 6. Third lumbar, left side view. 7. ‘Third lumbar, front view. Figs. 1, 2, and 5 are drawn from vertebree found in the peat at Burwell Fen, and now preserved in the Sedgwick Museum, Cambridge; the others are from Sandford Hill, and are preserved in the Taunton Museum. a. Neural spine. b. Neural canal. c. Pre-zygapophysis. d. Post-zygapophysis. e. Notch for exit of spinal nerve. jf. Transverse process. g. Anapophysis. h. Metapophysis. 1. Facet for articulation with tubercle of rib. j. Facet for articulation with head of rib. dur 32 WAI] [ep uUsean: P Palexontographbical Society, 1909. A MONOGRAPH BRITISH PLEISTOCENE MAMMALIA NOL. ie Aukay ile THE CANIDA. BY SIDNEY H. REYNOLDS, M.A., F.G.S., PROFESSOR OF GEOLOGY IN THE UNIVERSITY OF BRISTOL. Paces 1—28; Pratres I—VI. LONDON: PRINTED FOR THE PALHONTOGRAPHICAL SOCIETY. 1909. a le} a a a <4 a eI A a MONOGRAPH ON re els Sa MA MA fh A OF THE PLEISTOCENE PERIOD. THK CANID/. Order—CARNIVORA. Famity—CANID 4h. Genus—Canis. I. HISTORICAL INTRODUCTION. Av the commencement of a previous memoir dealing with the Pleistocene * bears reference was made to the difficulty which the study of those animals presented owing to the practical impossibility of coming to a satisfactory conclusion with regard to the mutual relationship of the various species and varieties. That diffi- culty presents itself in perhaps an even more marked form in the case of the Canide. The earliest reference to the existence of fossil Canide is Hsper’s* account (1774) of the finding of bones in the cave at Gailenreuth which he recognised as those of wolf. Rosenmiller ® (1794), in a pamphlet written in Latin and dealing principally with the fossil bears, stated that bones of dogs and foxes, as well as of wolves, had been found in caves, but considered that the bones of foxes were intro- 1 The terms “ Pleistocene” and ‘“ Prehistoric” are used in the following pages in the sense as defined by Dawkins and Sanford, ‘Monograph of the British Pleistocene Mammalia,’ vol. i, p. 7. 2 « Ausfiihrliche Nachricht—Zoolithen Bayreuth.’ 3 “Quzedam de Ossibus fossilibus animalis,’ Leipzig, p. 27. 2 PLEISTOCENE MAMMALIA. duced by diluvial action, and were not contemporaneous with the associated bear and hyena bones. ‘T'he earlher writers were disposed to doubt the identity of the canine bones found in caves with those of living species. Thus Goldfuss, who in 1810 had figured! the skull of a wolf from Migegendorf, when describing thirteen years later ® (1823) a wolf’s skull from Gailenreuth, regarded it as specifically dis- tinct from Canis lupus. Cuvier,® too (1812 and 1825), was apparently disposed to regard the wolf remains in his possession as specifically distinct from the modern species. He made further comparisons of the skeletal characters of wolves and dogs, and agreed with Daubenton * (1758) in recognising the extreme difficulty in distinguishing between the skull of a wolf and that of a dog. The first author to express himself decisively as to the identity of the fossil remains of the wolf with those of the living species was Schmerling® (1833) in his description of bones from the caverns of Liége. M. de Serres, Dubrueil, and Jeanjean © (1839), though not expressing themselves very definitely, attributed the canine bones found in the caves of Lunel Viel to the living species. The question as to the specific identity of the recent and fossil species was, however, fully considered by de Blainville’ (1844), who, in his ‘ Ostéographie,’ discussed and summarised all the evidence, strongly supporting the view that no distinction could be drawn between the wolves, dogs, and foxes of the caves and those living at the present day. Owen, too, in his ‘ British Fossil Mammals and Birds’> (1846), in which a full account of the fossil Canidae was given, agreed that “the wolves which our ancestors extirpated were of the same species as those which . . . left their bones in the limestone caverns ee Since then almost all zoologists who have considered the subject have agreed as to the specific identity of the fossil remains of the wolf with those of the living species, but Pomel (185+) and Bourguignat as lately as 1875 maintained the con- trary view, the latter author retaining the name Canis spelzus of Goldfuss for the fossil wolves of the caverns. Meanwhile the bones of wolf and fox had been described from many British caves, such as Kirkdale (Buckland,’ 1822), where, however, they were very scanty, 1 «Die Umgebungen von Miggendorf’ (Erlangen). * «Satgethiere der Vorwelt,’ p. 451. 3 ‘Oss. Foss.,’ tom. iv, iv, pp. 5—9 (1812), and ibid., ed. 3, tom. iv, pp. 457—467 (1825). 4 In Buffon’s ‘ Histoire Naturelle,’ tom. vu, p. 53. 5 «Recherches sur les Ossemens fossiles des Cavernes de Liége,’ tom. 11, pp. 22—46. 6 «Recherches Oss. humatiles des Cavernes de Lunel Viel,’ pp. 72—74. 7 * Ostéographie—Carnassiers,’ pp. 101—104. 8 « British Fossil Mammals and Birds, p. 132. 2 eh iranss ac xls pamlo2r CANIS. 3 Paviland (Buckland,! 1824), Oreston (Clift and Whidbey,” 1823), Banwell (Rutter, 1829), Yealm Bridge, Devon (Bellamy,* 1839). Buckland, in the ‘ Reliquie Dilu- viane ° (1824), gives a table showing that these animals had been recognised in various other Pleistocene deposits, both in Britain and on the continent. The question as to the specific identity of the fossil Canide of the caves with those living at the present time being settled, the far more difficult one concerning the mutual relationship of the wolves, jackals, and dogs occupied attention. This question had, as has been already mentioned, been considered by Cuvier and Daubenton. It was fully discussed in 1844 by de Blainville,° who went beyond those anatomists in being unable to recognise any osteological distinction between dogs and wolves, and by Pictet’ (1853), who was the earliest author to suggest as the origin of the domestic dogs, not any known species of Caiis living or fossil, but an unknown species assumed to have existed in Pleistocene times. Between 1859 and 1885 appeared a long series of papers dealing with the Pleistocene and Prehistoric Canide of Ireland, which were described from the followine localities: Dunshaughlin, co. Meath (Wilde,® 1859, dog); Shandon, co. Clare (Adams,’ 1879, wolf and fox); Knockninny, co. Fermanagh (Haughton,!° 1876, wolf, dog, fox); Ballynamintra, co. Waterford (Adams," 1881, wolf, dog) ; Knockmore, co. Fermanagh (Ball,” 1885, wolf). More recent are the records from Kesh, co. Sheo (Scharff, 1903, wolf, dog, fox), and Edenvale, co. Clare (Scharff, 1906, wolf, dog, fox, Arctic fox). At Shandon and Kesh it is probable that deposits of both Pleistocene and Prehistoric date occur. At all the other Irish localities in the above list it is probable that the remains belong solely to the Prehistoric period. Other important records of Pleistocene canine remains are those of Kent’s Cavern, Torquay (MacHnery,” 1859, wolf, fox), Wookey Hole near Wells (Dawkins,” 1862 and 1863, wolf, fox), Creswell Crags, Derbyshire (Busk,” 1875, wolf, fox, Arctic fox). The occurrence of the Arctic fox in Britain had not been previously noted. The records from Norwich (Denny, 1859, dog) and from 1 « Reliquie Diluviane,’ p. 85. 2¢ Phil. Trans.,’ cxiu, p. 88. 3 *Delin. Co. Somerset,’ p. 156. 4 Nat. Hist. 8. Devon.’ Bellamy’s account is reproduced by Pengelly in his paper on “The Literature of the Caverns near Yealmpton, 8. Devon,” ‘Trans. Devon. Assoc.,’ iv, 1871, p. 92. > “Reliquie Diluviane,’ facing p. 1. 6 « Ostéographie—Carnassiers, pp. 101—104. 7 «Traité de Paléontologie,’ tom. i, p. 202. 8 «Proc. Roy. Irish Acad.,’ vu, p. 193. 9 «Trans. Roy. Irish Acad.,’ xxvi (Sci.), p. 227. 10 «Proc. Roy. Irish Acad.,’ (2), 11 (Sei.), p. 482. 1 «Trans. Roy. Dublin Soe.’ (2), i, p. 205. 2 Thid. (2), 11, p. 335. 13 « Cavern Researches.’ 14 «Quart. Journ. Geol. Soc.,’ xviii, p. 124; and xix, p. 267. 15 Thid., xxxi, pp. 684—687. 16 «Proc. Yorks. Geol. Polyt. Soc.,’ iii, p. 538. ————— SS Se ess ——— = SS 4. PLEISTOCENE MAMMALIA. Burrington Combe, Somerset (Dawkins,’ 1864, wolf, fox), are of Prehistoric, not Pleistocene remains. More recent are Newton’s’ account (1894) of the fauna of the Ightham fissure near Maidstone, in which the Arctic fox was again met with, and Newton and Arnold Bemrose’s* account (1905) of the Hoe Grange Cave, Derbyshire, where scanty remains of the wolf and common fox were found. More comprehensive records are those of Falconer* (1868), who showed that bones of both wolf and fox had been recognised in all eight of the Gower caves, and Dawkins ° (1869), who, in his well-known paper on the ‘ Distribution of the British Post-Glacial Mammals,’ gives a long list of localities for canine bones. Much additional information with regard to both the Pleistocene and Pre- historic Canidee is contained in Dawkins and Sanford’s introduction to their Memoirs on the British Pleistocene Mammalia (Paleontographical Society, 1866). Harting’s ‘ Extinct British Animals,’ published in 1880, though chiefly concerned with the wolf during the historic period, has some account of its occurrence in Britain in Pleistocene and Prehistoric times, and adds some further localities® to Dawkins’ list. Pennington’s ‘ Notes on the Barrows and Bone Caves of Derby- shire’ (1877), though treating the subject in a more or less popular fashion, contains some further information. During the middle and latter part of the last century, too, the question of the mutual relationship of the Canidze was not left unconsidered, being discussed by Riitimeyer “ (1862), Jeitteles* (1872 and 1877), and Bourguignat’ (1875), while more recently the subject has been taken up by Huxley 1° (1880), Woldrich * (1881 and 1886), Lydekker (1884), von Pelzeln * (1886), Wilckens* (1886), Nehrmg* (1888), Boule” (1889), Vieira” (1894), Gaudry and Boule’ (1892), Studer” (1902) 1 «Proc. Somerset. Arch. and Nat. Hist. Soc.,’ xii, pp. 161-176. * «Quart. Journ. Geol. Soe.,’ 1, pp. 201—208. 5 Thid., xi, pp. 49 and 50. 4 «Pal. Mem.,’ ui, p. 525. > “Quart. Journ. Geol. Soc.,’ xxv, p. 192. o Oo, Gt, yo. LUIS. 7 «Untersuchung der Tierreste aus Pfahlbauten der Schweiz’ (1862). 8 ‘Mittheil. d. anthropol. Gesell. in Wien,’ ii, p. 169 (1872), and ‘Die Stammviter unserer Hunderassen’ (1877). 9 «Ann. des Sciences Géol.,’ vi, p. 33. 10 « Proe. Zool. Soc.,’ 1880, pp. 238—288. 11 « Mittheil. der anthropol. Gesell. in Wien’ (1881), xi, and ‘Anz. Akad. Wien’ (1886), p. 12. 12 «Palezont. Indica,’ ser. 10, vol. ii, p. 240. 13 «Zool. Jahrbuch,’ i, pp. 225—240. 14 «Biol. Centralbl.,’ v, pp. 719 and 751. ls “Naturwissenschaft. Wochenschrift,’ ii. 16 «Comptes Rend.,’ eviii, p. 201. 17 « Ann. Sci. Nat. Porto,’ i, p. 109. 18 «Mat. pour l’Hist. des Temps quatern.,’ fasc. iv, pp. 123—129. 19 “Abh. schweiz. pal. Ges.,’ xxviii, art. 1 (1902). ee CANIS. 5 and Keller! (1903). A very brief summary of their several conclusions is given subsequently when dealing with the mutual relationship of the Pleistocene and Prehistoric Canidee. EE IDES TR INBSIUIMIUOUNT ONT esas MIWA. Although this memoir is, strictly speaking, only concerned with the Pleistocene Canidz, when the range of a species extends into other strata, whether pre- or post-Pleistocene, brief reference must be made to such remains as occur. Tus Wotr (Canis lupus”). The oldest British formation in which the remains of the wolf have been found is the Red Crag. Owen,’ in 1856, first noted their occurrence in British Pliocene deposits, describing certain teeth from Woodbridge, which he attributed, with some hesitation, to this species. Newton* (1891) described two canine teeth from the Red Crag of Boyton, which he believed to be those of the wolf. Owen also identi- fied a humerus from the Forest Bed, but Newton remarks that it is very doubtful whether there is evidence of the wolf being represented at this horizon. The bone in question is now preserved in the Museum of the Geological Survey at Jermyn Street, London. These scanty records comprise the whole evidence for the occurrence of the wolf in Britain in pre-glacial times. During Pleistocene times, however, wolves abounded throughout England, their remains having been found in nearly every bone-cave of this period (see list, p. 10) and im many river deposits,” etc. There have been comparatively few records of the wolf from Scotland, this probably being largely due to the lack of caves and deposits suitable for the preservation of the bones. ‘This explanation will not, however, account for the scarcity and generally fragmentary condition of the wolf-bones found in Ireland. Fragmentary bones were recorded by Adams° from the Shandon Cave, where they were associated with the Mammoth and were clearly of Pleistocene age, and in 1 «Vierteljahrschr. Ges. Ziirich,’ xlviii. * Tt has been thought desirable, following Flower and Lydekker (‘ An Introduction to the Study of Mammals, Living and Extinct,’ p. 546), to include wolves, jackals, dogs, and foxes in the old com- prehensive genus Canis. 3 «Quart. Journ. Geol. Soc.,’ xii, 1856, p. 227. 4 «Vertebrata of the Phocene Deposits of Britain,’ p. 8. > The lengthy account of the wolf in Harting’s ‘ Extinct British Animals’ is mainly concerned with its distribution in historic times. 6 «Trans. Roy. Irish Acad.,’ xxvi (Sci.) (1879), p. 221. SSE < = TI Eo Se 6 PLEISTOCENE MAMMALIA. recent years they have been recorded by Scharff’ from the prehistoric caves of Hdenvale, co. Clare, and from those of Kesh, co. Sligo, which yielded remains referable in all probability to both the Pleistocene and Prehistoric periods. Bones of the wolf have also been found in Prehistoric deposits at Knockninny ” and Knockmore,’ co. Fermanagh, and bones somewhat doubtfully referable to the wolf at Ballynamintra,* co. Waterford; but with these exceptions no wolf-bones have been recorded from Irish Prehistoric deposits, a somewhat remarkable fact in view of its great abundance in Ireland in historic times.” THe Doc (Canis famiiaris). Owing to the frequent references to the bones of dogs in various papers dealing with the Irish Mammalian remains some allusion must be made to them here, though it is at least doubtful whether any animal that could be called a dog existed in the British Isles in Pleistocene times. Owen °® admits the dog to the number of his British fossil mammals, but does not describe any British specimens. The dog is not included by Dawkins’ in his table showing the distribution of British post-glacial mammals, and is not mentioned by Lydekker in his ‘Catalogue of the Fossil Mammalia in the British Museum.’ Woodward and Sherborn® admit it among the British fossil vertebrates, but (?) Pre- historic deposits at Norwich and Walthamstow are the only occurrences in Great Britain to which they allude. Skulls attributed as a rule, owing to the length and slender character of the muzzle, to the large extinct Irish wolf-hound, have been repeatedly referred to by writers on Irish mammals. Wilde’ (1859) described examples from near Dunshaughlin, co. Meath ; Haughton '° (1876) referred to the occurrence of the dog in Knockninny cave near Lough Hrne ; Adams“ (1880) and Ball” (1885) referred to the skulls described by Wilde, and agreed with him in attributing them to dogs; and Adams! (1881) described slender mandibles from 1 «Trans. Roy. Irish Acad.,’ xxxii, B., pt. 4 (1903), p. 201, and xxxiu, B., pt. 1 (1906), p. 43. 2 «Proc. Roy. Irish Acad.’ (2), ii (Sci.), 1876, p. 482. 3 «Trans. Roy. Dublin Soc.’ (2), 11, 1885, p. 335. 4 Ibid. (2), 1, 1881, p. 205. 5 See Adams, ‘ Proc. Roy. lish Acad.’ (2), ii, 1878, p. 99; and Scouler, ‘Journ. Geol. Soe. Dublin,’ i, 1838, p. 225. 6 « Brit. Foss. Mammals and Birds,’ p. 1383. 7 ‘Quart. Journ. Geol. Soc.,’ xxv, 1869, p. 192. 8 «A Catalogue of British Fossil Vertebrata,’ 1890, p. 324. 9 «Proc. Roy. Irish Acad.,’ vii, 1859, p. 194. 10 Thid. (2), ii (Sci.), 1876, p. 482. 11 «Sci. Proc. Roy. Dublin Soc.,’ 11, 1880, p. 66. 2 «Trans. Roy. Dublin Soe.’ (2), 111, 1885, p. 340. 18 Tbid. (2), 1, 1881, p 205: So —— Se. oe. eT a a a a eee eS a a ee ‘F CANIS. @ Ballynamintra cave, co. Waterford, which he attributed to the Irish wolf-hound. There can be little doubt that all these are of post-Pleistocene date, and belong to the Prehistoric period. Numerous bones, clearly of dogs, have been found in peat and other Prehistoric deposits in many parts of Great Britain, especially in the alluvium of the lower part of the Thames valley. THe Fox (Canis vulpes). The occurrence of the fox in the Red Crag is well authenticated, a well-pre- served palate from Boyton in Suffolk having been figured and described by Lydekker.! He gives measurements showing that its size considerably exceeds that of a full-grown recent specimen, but in spite of this concludes that the Specimen is to be referred to the fox—an opinion in which he is supported by Newton. The evidence for the occurrence of the fox in the Forest Bed is not very good. It is based on part of a humerus which Newton? hesitated to refer to the fox. Liydekker, however, thought that the specimen was correctly referred to this species. From Pleistocene times onwards the distribution of the fox throughout the British Isles has been practically universal. In the cavern deposits its distribution shows a remarkable correspondence with that of the wolf (see Table, p. 10). Tue Arctic Fox (Canis lagopus). As yet the remains of the Arctic fox have been recognised at only a very few localities in Britain. The earliest record is that of Busk*® (1875), who found among the bones from the rock fissures of the Creswell Crags an axis vertebra which he carefully described and figured, referring it to the Arctic fox on account of (1) its small size; (2) the slenderness and abrupt divergence of the transverse processes; (3) the prominence of the median keel on the ventral surface of the centrum ; (4) a difference in the form of the anterior articular facets from those in the common fox. The second record is by Newton,* from the Ightham fissure near Maidstone. Newton figured and ascribed to the Arctic fox a femur, a tibia, a humerus, a mandibular ramus, and part of the upper jaw. Many other bones of the Arctic fox from the same locality are in the collections of Dr. F. Corner, of Poplar, and 1 “Geol. Mag.,’ dee. ili, i, 1884, p. 443. 2 Thid., dec. 11, vii, 1880, p. 152. 3 «Quart. Journ. Geol. Soc.,’ xxxi, 1875, pp. 685—687. 4 Thid., 1, 1894, p. 202, pl. xii, figs 5—9. = a ee ee a 8 PLEISTOCENE MAMMALIA. Mr. W. J. Lewis Abbott, of St. Leonard’s (see Pls. V and VI, and Text-figs. 1—7). In the same paper by Newton a reference is made to a skull belonging to Dr. H. P. Blackmore, of Salisbury, who obtained it in 1875 from the brick earth of Fisherton, near Salisbury, where it was associated with the following Arctic animals!: Lepus variabilis (the mountain hare), Microtus nivalis, Myodes torquatus (the lemming), Ovibos moschatus (the musk ox), and Rangifer tarandus (the reindeer). The reindeer was also found associated with the Arctic fox at Creswell Crags and Ightham, and Newton” is further of opinion that certain vertebree and other bones from a small cave at Walton near Clevedon are to be attributed to the Arctic fox; here the presence of another arctic animal, the lemming, is indicated. The only record of the occurrence of the Arctic fox in Ireland is contained in Scharff’s* account of the Newhall cave, Edenvale, co. Clare, where a jaw clearly to be attributed to this species was met with. Here again it was associated with the reindeer and lemming. SKELETAL DIrFERENCES BETWEEN THE Common and Arctic Foxss. The common fox is, as a rule, very considerably larger than the Arctic fox, but as small individuals may occur this difference is not always a safe criterion. There are, however, many differences in the skull. The skull of the common fox is the larger, and has the length of the jaws relatively greater in proportion to the size of the cranium, and hence the anterior premolars are more widely spaced than in the Arctic fox. On the other hand, the cranium of the common fox is somewhat narrower in proportion to its length than that of the Arctic fox, especially just behind the post-orbital processes. These tend to be longer in the common fox than in the Arctic fox. Scharff mentions that the length of m. 2 is somewhat ereater in the common fox than in the Arctic fox, but this difference seems scarcely appreciable in the British Museum specimens. ? Lycaon anglicus, Lyd. This name is applied by Lydekker * to a left mandibular ramus from the Sprit- sail Tor cave, Gower, which was originally described and figured by Falconer ° under the name of ‘ hyzenoid wolf.’ The specimen was subsequently fully des- 1 «The Geology of the Country around Salisbury,” ‘Mem. Geol. Surv. of England and Wales,’ 1903, p. 68. 2 «Proc. Bristol Nat. Soc.,’ 4th ser., i, pt. 8, p. 186, 1907 (ssued for 1906). 3 “Trans. Roy. Irish Acad.,’ xxxiii, B., pt. 1, p. 48. * «Geol. Mag.,’ dec. 111, 1, 1884, p. 443. 5) Pal. Mem,” 11, pl. xxxya, figs, 1) 2t a ae : | | | : CANIS. 9 eribed by Lydekker' in his ‘Catalogue of the Fossil Mammalia in the British Museum.’ The feature upon which he principally relies in the attribution of this specimen to the genus Lycaon is the presence of a ‘ distinct anterior talon” to the fourth premolar. There is no doubt that an anterior cusp is very distinctly present in this specimen (see PI. V, figs. 7, 8); and in a large series of wolf- skulls in the British Museum no specimen was found showing any comparable development, though in certain cases, e.g. the skull of a wolf from Kandahar (168a) and a North American example (165d), slight indications of an anterior cusp occur. But on the other hand pm. 2 and 3 of the Spritsail Tor specimen are identical with those of the wolf, being considerably longer in proportion to the height of the crown than are the corresponding teeth in any of the skulls of Lycaon examined in the British Museum; pm. 4, too, agrees precisely in the characters of its main lobe and posterior cusps with the corresponding tooth of the wolf, and differs considerably from the Lycaon type. In view of the known variability in the teeth of Canidee and of the difficulties of geographical distribution involved in the addition of a southern form like Lycaon to the British faunal list, it seems the most satisfactory course, on the whole, to regard the Spritsail Tor specimen as a somewhat abnormal wolf. TABLE SHOWING DistripuTion or BritisH PiEistoceENe Canipm 1N RIVER Deposits, ETC. Wolf. | Fox. mee ‘OX. ]Byerill omveltas), NOS). Seon i een ee Bracklesham, Sussex | Craytond peep tere ee eur Se aehene oto cata omen lide it wees Dartford . Bs) Pie ear cee ob es Sa, hey 7 eRe ee Bee | x Fisherton, Salisbury eee Ore eer ena hen eae eae (COHEN SC hts aCe eRe an cae x xK X Hy Sipvl Claw tyes recite ese okie Waco wan teats, dein dictum bras Saaaeh Murston, cee oe Newbury aiacectar Sete ERR iota rt ic STARE Gna vin Slade Green, near Erith spade ote de LoavOL GR AED RRA ESRC aM CN ERE Tewkesbury ... MORIN). 5 scone adegosdosgochioa ts Ae tee teen ieee eee ene | We Petece A\WGSOINAS, STANTS, colade ceded Ce Bae toe ORR eD ECR a lie bcc tu | Windsor x & KKK XK KX XK x | «Catalogue of the Fossil Mammalia in the British Museum,’ pt. i, p. 122, 1885. 10 PLEISTOCENE MAMMALIA. TaBLE SHowING Disrrigution or British PrLeistocenr Canina 1n CAVERNS. Arctic Wolf. Fox. Dox. | Anstey 's: Came, Tox quay ar1.1- acer e eee ee eee peer Does Bacon Hole; ‘Gower ~ ois. cestecn cece eee nee ee eRe eee eee Bamiwell” swsi.ds can Gee dalle cakeegne ee eR Ronn Rt ea ne ana Bench Cave, Brixham Bleadon : nen cocnhenanso tates uacamaruned gms Bosco’s Hole, Gower ccccaouileslethe (2 te eter tee ana | Cae Giwyn, Clwyd) Valen sist anes eee eeee rere nee epee Caswell Bay; (Gower. Sans. eset ee een ee eee Cefn, near St. Asaph .. PA Rr Rear neeok Anieiia ade quiet dae Creswell Crags, Derbyshire . BENET rad sods TEN Osa aN Sed acne atc Crow Hole, Gower ........... ......... Deborah Den, Gower ...... Ffynnon Beuno, Clwyd Vale. Shs) sete scaeee Ramee Durdham Dowans: a.cacku.csdewc th. eee eee Eee nee Hoe Grange, near Longcliff, Derbyshire Hutton... : SaaS ans Ey Cee Ichtham Fissure 1 near r Maidstone. Azan atten Moses eee ee Kent’s Cavern, Torquay Bi Giia (qe eo tne nan ear AON a ABA gn Na abae Cone LaciccS Langwith Bassett, near Mansfield Saeustharenee Reyne hong Holes Gowers ach meecceceisiecr ant Sonate eee REESE XK XKOOX XE KS XK Oe eX ex ae CO) eS) A050 amen ote nn Le er We RMSE. ety Pied in 8S cs ah ane Paivallian Gis Gower esheets Cee eee | Pleasley Viale) Derbyshire.cecs.c-ces scarce narra eee eee Ravens clitts (GOwer t.sceacatepicenee ae ee lon eee Gee Saiivdiordl 2e.. 9: Scns brasserie anaes e NA ee RE eT EE | Shandon Cave, Dungarvon een) ge ae a ey ee Spratsanl Wor; "Gower sss eaaasacdie a eee ceeranee cree reen tee | MMOGSC BG: Savlx sucis nome rowel acces bens tas ME ea ECE | ee ee | Walton, - near (Clevedon! Ree RT rs ie Crane ie. ol Wraterhousesm Sta tiord shire sess setter ete reece ere Ee res | ivauas iyp 1GaOll, CRSMETOMY © 355000200000000600. 000 0ac 0a 00004 and annus Wrookey Golo crcl so scons: cnccone eee eee eee ERECT Yeealim: Bride iy oc scecsnin uns cite Ce enen ae eee EEE a ee > Gs io Ko KK KK: x X x SE SEK SE NE OOK OK OK KKK RRR RRM ME x xXx: Il]. DESCRIPTION OF THE REMAINS. The Canide! comprise the only family of the section Cynoidea, the second of the three into which the Carnivora Vera are divisible. With regard to the denti- tion they show less specialisation than any other group of living Carnivora, and in other respects approach relatively near to the primitive type. The structure of the auditory bulla and adjacent parts of the skull is intermediate in character between that of the Alluroidea and that of the Arctoidea, as the Cynoidea agree with the Hluroidea in having the auditory bulla inflated and the paroccipital process of the 1 See Flower and Lydekker, ‘Mammals, Living and Extinct,’ p. 544, et seq. CANIS. idl exoccipital in contact with it, while they agree with the Arctoidea in the almost complete absence of a septum dividing the auditory bulla, in the large size of the glenoid foramen, and in the presence of an alisphenoid canal. In the living forms there is no entepicondylar foramen. The upper molars have a triangular crown, and the blade of the upper carnassial consists of two lobes. A. Toe Sxuwt (Plates I—IV). The cranium is moderately elongated, the jaws long, tapering, and somewhat compressed. ‘I'he zygomatic processes of the frontal and malar are short, so that the orbit communicates widely with the temporal fossa. The pterygoid has a well- developed hamular process. The auditory meatus forms a short but fairly promi- nent bony tube. The following are characters upon which most stress has been laid in attempt- ing to discriminate between the skulls of wolves, dogs and foxes: (1) The relative proportions of the jaws and cranium ; (2) The extent to which the temporal ridges, always widely separate in young animals, approach and coalesce into a sagittal crest in the adult; (3) The greater or less backward extension of the nasals; (4) The character of the post-orbital process of the frontal ; (5) The union of the nasal processes of the frontals with the ascending pro- cesses of the premaxille, or the separation of these processes from one another by the meeting of the maxille and nasals ; (6) The length of pm. 4 as compared with that of m. 1 and 2 taken together ; (7) The orbito-frontal angle or the obliquity of the opening of the orbit to the brow. B. Dentition (Plate V). (1) Distinctive Features of the Teeth in the Genus Canis.—The typical dental formula isi. 3, ¢. +, pm. 4, m. 2, as in Ursus, but in aberrant forms (Cyon) the molars are 3, and in a fossil form, Canis (Lycorus) nemesianus, the premolars are 4. Further, in Canis cancrivorus the missing last upper molar is occasionally present. The formula embracing these variations is i. 3, c. 4, pm. 34, m. 5-3. The contrast in size between the canine and incisor teeth is not so great as in either cats or bears. The upper carnassial tooth, pm. 4, differs from that in bears, and resembles that in cats and hyenas in possessing an antero-internally placed inner tubercle supported by a distinct root. In accordance with the method adopted in previous memoirs it has been SRR SS SR aS A = eS ST Se a ce ee ee ee “‘SOTJOUIIZUWID Ul WATS OLB SJUATOINSvEW [[ Y— 270N ‘BIUBIO OY} 0} SUOTEC SoTqIPULUE esa} TeYyZeYA UTez1eOUN st 4yT 4 ‘pomnsny » | E Exe rae Gre 0-F Bs aie Ap Gree | IQ |p ore ssa00id prowo109 Fo do} 0} apour woIF FYSIOH * 66.6 |b GT-R l4x8-6 2-11 200 @.31| 181 P27 Bod ve GNA ea eer eas Pe ea anes ene estes a, Apuoa Jo puta 07 PIMSVIUT SNTUVI IV[NGIPULUE Jo YASUE, WMUIXE PL LLP 197 46g Lgp LBP sae LOF RVakeuatsinialalvinisieliaelelviuldistedaheleiwisielalslslninhulstetrtai aie ad ev [VP WOIF-071qG.10, 9.¢ 2-9 8.9 eee 1-8 | 69.6 8.6 eae eee eee G8.3[ ZS wee eee eee eens Cee eee ee ee ee ee ee ene . aqvyjed jo Youle] Ol GT GT | ¢9.T 9T 8.[ OGG bell ee mS | CORE (oar ae WNUGLU WIMV.LOF FO LoJOUUIVIP VstOASTVA], = DRE ar PP 1-7 CBP 2.9 0-9 Pd Bol) 2.9 eae Dol E Ce SCC ai i Ci i CCS snzveul iS ALOJIPNY [VUIO}XO SSOLOV JUOUIOINSVIU S.LOASURL], < BG GZ 9.3 Le 1% 6-8 6-8 9.7 | BFP Gp | ¢3.p | Ge.p | sopAptod [eztd1O90 ssoaov YUOTMAINSvEUT OS.1eASTRA, = Gg G8 GQ. G6. Gr Gt || Geri G-1 Tell SP Nona eC fa ne ees ees SUILI}UV WINALL] WOULVIOF MOTO UIPTAA S| a ELS a oS 10,9 | so S IA48 aes | Ss Ss Sao aB§ |e | Aas aos HES | Ges | PBs | oes | Sas (see) SA s\eae| Pos | Pos le os | 22 He 2 Ee 7) He O~ = S$ OO | = we He Qe? wo S Bee lta | oe Tel tag lose lace | Ste | oa a] T2a|e as Ea: Sam aoe | Feel ear | So's | Fe iy| Peg | 4S )| Se | Bee | so a) ees SER EOS | SEa|FDE|Fok| tne | ses | S28 | 80S | eae) eae |S ALS | Vos | OLS loom |CSs ie B: BEl| ese |o 8 |-os (ees) ou Sen eee 9 we ool aed aol pet ells (ea eo) ae Sis | Sie ray | ESE SSeS | esos | se | eels eee “Re | SE ee | xs PSS esl a Se Se SIS” © ‘STTOMG ANINVO JO SINTWAYASVET, TAILVAVAWOD #0 ATAVY, (7) al ee CANIS. 13 thought best when describing the teeth not to use terms involving assumptions of homology and requirme long explanatory prefixes. The terms “cusp” and “tubercle” are regarded as synonyms for small elevations of the surface of a tooth. 5] The term “cone” is used as denoting a rather larger elevation, and the terms “talon” or “heel” for posteriorly-placed segments of a tooth. (2) Permanent Dentition of the Upper Jaw in Canis lupus (Pl. V).—In drawing up the following description, the skulls and teeth of a number of wolves, recent and fossil, now preserved in the British Museum have been examined, but the principal part of the description is based on a skull of a male wolf from Pekin, No. 90.7.8.2 in the British Museum collection. I. 1 and 2 are very similar teeth, differing only in the shghtly larger size of 1. 2. The principal cone is somewhat recurved and there are small laterally-placed accessory cusps. ‘The cingulum is rather strongly marked. In some cases the lateral accessory cusps are scarcely noticeable, but the cingulum is raised into slight cusps posteriorly. The root is about three times as long as the crown and is much laterally compressed. I. 3 is a rather larger and more caniniform tooth than i. 1 and 2 and shows some variability, the postero-internal face being sometimes marked by a cingulum, sometimes raised into a pair of laterally-placed cusps. The root 1s about twice as long as the crown and is triangular in cross-section, not laterally compressed as iekeand 2. C. The canine has the form usual in the Carnivora. Its crown constitutes about two thirds of its length. Pm. 1 is a small, single-rooted tooth with a fairly well-marked cingulum surrounding the principal cone. Very slight accessory cusps may be developed on the cingulum posterioriy and antero-internally. Pm. 2 is a larger and more elongated two-rooted tooth with a conical crown triangular in outline. A posteriorly-placed cusp may be present or absent. Pm. 3 closely resembles pm. 2, differimg only in its larger size and in the ereater prominence of the posterior cusp, which may be double. Pm. 4, the upper carnassial, is a large three-rooted tooth with a powerful trenchant blade divided into an anterior more conical portion and a posterior portion with a chisel-like edge. Placed anteriorly is a low inner tubercle supported by a distinct root. M.1. This is a large tooth, somewhat wider than long. The outer portion of the crown is formed by two prominent cones of which the anterior is the larger ; the inner portion of the tooth is much depressed, but the inner edge is raised into amore or less prominent ridge and two low cusps lie between this and the outer portion of the tooth. ‘Two roots support the outer portion, and a third and stouter root the inner portion. 14 PLEISTOCENE MAMMALIA. M. 2 is a considerably smaller tooth than m. 1 but is constituted on the same plan, having two relatively prominent cones or cusps on the outer part of the tooth, and a depressed inner area bearing several slight cusps. Two roots support the outer portion of the tooth, a third and larger root the inner portion. (3) Permanent Dentition of the Lower Jaw of Canis lupus (Pl. V).—I. I, 2 and 3 are all very similar teeth, differmg only by their progressive increase in size, and in the fact that while i. I and 2 have the root strongly laterally compressed 1. 3 has it more or less triangular in section. Hach tooth has a somewhat chisel- shaped edge with a small accessory cusp placed externally to the main cone. C. This tooth is of the usual type and differs from c. only in the fact that the inner border tends to be rather more sharply curved. i: Pm. 1, which is sometimes wanting, is a small single-rooted tooth with a simple conical blade, bearing, as a rule, a very shght cusp posteriorly placed. Pm. 2, 3, 4 are very similar two-rooted teeth each with a triangular blade, which in pm. 3 and 4 may be slightly recurved. There is a slight cusp posteriorly placed in pm. 2, and this becomes larger in pm. 3 and 4. The cingulum is well marked on the inner surface of the teeth, and in pm. 3, and more often in pm. 4, may give rise to a second posteriorly-placed cusp. Im some cases each of these teeth bears a slight cusp anteriorly placed. M. 1. This is a large tooth supported by two stout roots. The anterior two thirds is formed by the powerful bilobed blade, the posterior lobe being shghtly the larger and having a small cusp placed postero-internally. The last third of the tooth forms a depressed talon or heel and bears two cusps placed side by side, the outer one being somewhat the larger. M. 2 is a rather small, somewhat oblong tooth supported by two roots. The anterior part of the crown bears a pair of cusps placed side by side, while a third cusp lies postero-externally. In some cases there are indications of a fourth cusp placed postero-internally. M. 3 is a very small one-rooted tooth with a nearly oval crown bearing one or more slight cusps. (4) Milk or Deciduous Dentition (Pl. V).1—The formula for this is di. 3, d.c. 4, d.m. = = 28. The first permanent premolar has no milk predecessor. (a) Milk Dentition of the Upper Jaw.—D.i.1,2 and 3. These are all small and extremely simple teeth with short crowns and long, somewhat tapering roots. D.c. is a reduced representative of the permanent tooth and requires no special description. 1 The description is drawn up from a made-up set of milk-teeth of Canis ? lupus from Torbryan, Torquay, now in the British Museum. CANIS. 15 D.m.1 is a good deal like pm. 2, the blade having a principal cone and often a slight posteriorly-placed accessory cusp. The two teeth differ, however, in the fact that dm. 1 has the roots strongly divergent. D.m. 9. the milk earnassial, is the largest of the deciduous teeth. The major portion of the crown 1s formed by a powerful bilobed blade, of which the anterior lobe is conical, the posterior more depressed and chisel-hke. In front of the blade are two low cusps. There is further a prominent inner cusp anteriorly placed. This is supported by a special root, and two other strong roots support the outer part of the tooth. D.m. 3 is a very irregular tooth. The outer edge, which bears certain ill-defined cusps, is raised anteriorly and supported by a small root, while posteriorly it is depressed and supported by a strong triangular, divergent root. There is an mner cusp supported by a third root. (3) Milk Dentition of the Lower Jaw—D.i. 1, 2 and 3. These are even slighter and simpler teeth than those of the upper jaw, and have very long, tapering roots. .c. This is practically identical in character with that of the upper jaw. Dm. 1.—Except for its smaller size and the more divergent character of the roots, this is identical with pm. 2. D.m. 2 very closely resembles pm. 3, but the accessory cusp anterior to the principal cone is better developed than in that tooth. D.m. 3, the milk carnassial, is a large tooth supported by two strong divergent roots. Four fifths of the crown is formed by a bilobed trenchant blade, and behind this is a considerable cusp. o. THe VerrepraL Cotumn (Plate V1). Little or nothing can be mentioned as specially characteristic of the vertebral column in the Canidz in comparison with that in other Carnivora, but the variable number of the caudal vertebree may be alluded to, the number ranging from seventeen to twenty-two. ‘There are thirteen thoracic and seven lumbar vertebree. Only three vertebrz are fused together in the sacral region, while in bears the number may be as many as five. D. Tae Lime GIRDIES. The Shoulder Girdle—The scapula (Text-fig. 1) scarcely calls for special comment, though it may be mentioned that the coracoid process is very slightly developed. The British fossil specimens are almost always in a very fragmentary condition. The clavicle is im a much reduced state, though better developed than in bears. SS RE RS EES LEER OE SIE E SE: 16 PLEISTOCENE MAMMALIA. Trxt-ric. 1—a. Right scapula of a common fox (Canis vulpes) seen from the outer side. 3. Left scapula of an Arctic fox (Canis lagopus) seen from the outer side. Both from Ightham (Lewis Abbott Coll.). 3 natural size. 1, glenoid cavity; 2, spine; 3, acromion; 4, coracoid process. Text-ric. 2.—A. Pelvic girdle of an Arctic fox (Canis lagopus) seen from the left side. 8. Pelvic girdle of a common fox (Canis vulpes) seen from the left side. c. Pelvic girdle of a common fox (Canis vulpes) seen from below. pb. Pelvic girdle of an Arctic fox (Canis lagopus) seen from below. The Arctic fox is from the Pleistocene of Ightham near Maidstone (Lewis Abbott Coll.), the common fox from the Prehistoric alluvium of the Thames Valley, Walthamstow (Brit. Mus.). 4% natural size. 1, acetabulum; 2, obturator foramen; 3, ischium; 4, sacral surface of ilium; 5, gluteal surface of ilium ; 6, ischial tuberosity ; 7, pubis. CANIS. 17 (2) TasLes oF CoMPaARATIVE MEASUREMENTS OF THE LIMB GIRDLES. C. vulpes, Pleis-- OC. lagopus, tocene, Igh- | Pleistocene, Table of measurements of the scapula. Se eae pee a (Lewis Abbott | (Lewis Abbott Coll.) Coll.) ey Maxaman lenethii wen saeree-ebea css een acae 8:25* 6:45* Zee Viaxammuin willbe aay race e seer ac actin wn peices stele — 3°25 3. Minimum width at the neck .................. eg 13 4. Width at proximal end measured to end of COMO AONE! THROTITS ao, abo casono dup oeaneosencbo ead sne eg) 1-45 5. Height from top of acromion to inner edge om slenordicamb yer a cecsusenainnse esses == 18 Table of measurements of the pelvic girdle. IL, Mileamamtoram: Neaveailal, yb ooaancmae saeoop aps eadeauson eno 9:3 70 2. Length from acetabulum to supra-iliac WONG Cree oeallAUTIM eeritare ie ae eset vare, cc ccesrebscieree een ans) 4-0 3. Dorso-ventral measurement of ilium at widest point ...... ee 315 2-1 4, Antero-posterior diameter of acetabulum ... 1-45 ell 5. Length from acetabulum to posterior border CATT... ct en = 2°35 6. Maximum diameter of obturator foramen ... 2°15 16 7. Measurement along ischium from symphysis 18O) GING! GIE uSolaNAIL KYowNS) Goocaossassedocnesea roo 38 31 * Figured. (2) The Pelvic Girdle—This (Text-fig. 2) does not present any specially characteristic features. E. Tar Lines. These are of moderate length, and agree with those of the Felide and differ from those of the Urside in being sconghy geonerade. All the digits are termi- nated by non-retractile claws. vic: (1) The Anterior Limb.—The humerus (Text-fig. 3) is rather short. There is no entepicondylar foramen, but a supra-trochlear foramen is always present. The metacarpals are longer and more curved than in Felis. Five digits are present, but the pollex is much shorter than the others and does not reach the ground. ois) = 18 PLEISTOCENE MAMMALIA. TEXxT-FIG. 3. TEXx?T-FIG. 4. Texr-ric. 3.—a. Left humerus of a common fox (Canis vulpes) seen from the inner side. 3. Left humerus of an Arctic fox (Canis lagopus) seen from behind. Both from Ightham (Corner Coll.). 3? natural size. 1, head; 2, greater tuberosity ; 3, lesser tuberosity ; 4, supra-trochlear foramen; 5, internal condyle; 6, trochlea ; 7, deltoid ridge. Text-ric. 4.—a. Right radius of a common fox (Canis vulpes) seen from front. 8. Left radius of an Arctic fox (Canis lagopus) seen from behind. Both from Ightham (Brit. Mus., No. M.7232, and Corner Coll.). natural size. 1, surface for articulation with ulna; 2, surface for articulation with carpus. (3) Taste or Comparative MuAsureMENTS or Bones or ANTERIOR Limes. C. lupus (left). C. vulpes C. lagopus (left). (left). Pleistocene. Pleist Pleist Humerus. Banwell. CAS LOSERS: CISLOCeMe? ; Ightham, near | [ghtham, near Ns 4f029 “Maidstone Wile (Brit. Mus.). : , (Corner Coll.) | (Corner Coll.) 1. Maximum length “slabs «REE Ae ERE ae See 19°75 12°3* 9-4%* 2. Diameter of proximal end passing across centre of articulating surface and greater tuberosity ............ 366 2°6 19 3. Antero-posterior diameter of shaft at middle of deltoid 1g KG (24: ee ee eee MnChn nd ayacodgacdesghdte coma adesess 2°5 115 10 4, Transverse diameter at same point ..............000.e00 eee 1-75 0°8 0-7 5. Maximum transverse diameter at distal end ............ 4:4 21 1137/ * Figured od CO — CANIS, Text-Fric. 5.—a. Left ulna of a wolf (Canis lwpus), incomplete at distal end, seen from the left side. From Kent’s Cavern, Torquay (Brit. Mus., No. M.830). x. Right ulna of a common fox (Canis vulpes) seen from the left side. From I¢htham (Brit. Mus., No. M.7232). c. Right ulna of an Arctic fox (Canis lagopus) seen from the right side. From Ightham (Lewis Abbott Coll.). 32 natural size. 1, olecranon ; 2, surface for articulation with the trochlea; 3, surface for articulation with the radius. 19 20 “PLEISTOCENE MAMMALIA. (3) Taste or Comparative MrasvremMEntTS or Bonzs or Anterior Limp—continued. C. vulpes C. lupus (left). (right). o. He me Pleistocene. Pleistocene. Pl rs Pe Radius. Ightham, near | Ightham, near i eistocene. 5 5 ghtham, near Maidstone. Maidstone. Maidst (Corner Coll.) | No. M.7232 Me os (Brit. Mus.). (Corner Coll.) 1. Maximum lenoth® Stace see ee eee ee eee 18:15 11:05* 87 2. Shorter, 7. e. right to left, diameter at humeral articu- Jatiom: «.aheachacghtscentasaece nase eee eee oe CPE I: 1:4 0°75 0-6 3. Longer, 7. e. antero-posterior, diameter at humeral articulation: oi Seite sdecnneqenceeene caeee oe cece 2°4 115 0:95 4. Shorter, 7. e. right to left, diameter at carpal articu- TEHLONL.. a Secaagadonssmptrteeeino rarer ROR elm eee Ee 16 0°75 06 5. Longer, 7. e. antero-posterior, diameter at carpal ATtICULATIOM ,,.c2isacwareuemestecmanmae seal ae O OGRE eR ETRE 3°05 1:45 12 C. lupus (left). Giaht) : yes. eae Pleistocene. Pleistocene. Ulna. a BS ASE Ightham, near | Ightham, near Nounay. | “Maidstone. | “Maidstone. (Brit Mus). | No.M.7232 | (Lewis Abbott : |) (Brit. Mus.). Coll.) 1. Maximunt length 235). syssdses cece te aeeeoen eer eee eee 12:9* 10:05* 2. Antero-posterior or vertical measurement at sigmoid 10) 1(0) Cen Aaa era amoneeu ana auododean cb oooCse oan da Ap 19% 1-0 0:7 3. Maximum transverse measurement of olecranon ...... 1:55 0-7 05 (2) The Posterior Iimb.—The tibia (Text-fig. 7) has the crest somewhat sharply truncated. The second to fifth digits are well developed, but the hallux is absent, or vestigial and suspended in the skin without bony connection with the rest of the pes. (4) TaBsLEe or ComPaRATIVE MEASUREMENTS OF Bonezs oF PostuRion Lime. Femur. C. lupus (right). Pleistocene. Ightham, near Maidstone. (Corner Coll.) C. vulpes (left). Pleistocene. Ightham, near Maidstone. (Corner Coll.) C. lagopus (left ) Pleistocene. Ightham, near Maidstone. (Corner Coll.) 1. Maximum leneth 2. Transverse or right to left diameter at condyles COCR i i ri ee a ay 3. Antero-posterior diameter of head ...... | 4. Vertical or antero-posterior diameter of | shaft at middle 5. Transverse diameter at proximal end measured across head and great tro- chanter 21-4 4-0 24 eS | 5:0 * Ficured. CANIS. Trext-Fic. 7. TExtT-FIG. 6. Text-Fric. 6.—a. Left femur of a common fox (Canis vulpes) seen from behind. x. Left femur of an Arctic fox (Canis lagopus) seen from front.. Both from Ightham (Corner Coll.). 2 natural size. 1, head; 2, great trochanter ; 3, lesser trochanter ; 4, internal condyle; 5, external condyle. Trxt-rie. 7.—A. Left tibia of a wolf (Canis lwpus) seen from front. From Torbryan Cavern, Torquay (Brit. Mus.). 8. Left tibia of a common fox (Canis vulpes) seen from behind. From Ightham (Brit. Mus., No. M.7232). c. Right tibia of an Arctic fox (Canis lagopus) seen from the right side. From Ightham (Lewis Abbott Coll.). % natural size. 1, cnemial crest; 2, facet for articulation with fibula, 22 PLEISTOCENE MAMMALIA. (4) Taste or ComparaTivE MrasurEMENTS OF Bones or PostERion Limp—continued. Chupa Got) | upns Gat) | iuege et) agar as Torbryan Tor- lesions. Ightham, near Pleistocene H Tibia. N Ightham, near Wes igo, aio Tein : ; quay. No. Maidstane aidstone. No.| Ightham, near } M.4563 (Brit. | , Ce Coll.) M.7232 (Brit. Maidstone. ‘ Mus.). ‘ : Mus.). (Corner Coll.) I Wikrahinbren WENN 5.05 oona0as0s00sa05080002 21:45* 21-7 14:3* 10°75 2. Transverse or right to left diameter at . proximalendiy ere eaee eee eye AA, 44, 2°45 1:85 3. Vertical or antero-posterior diameter at proximal end measured to top of crest 34 43 17 ley 4. Transverse diameter at distal end ...... 3°05 3:0 16 1:25 5. Vertical or antero-posterior diameter at distal iémdy ie. he tvaees sasneren eee eater 2°3 2:05 Hell 0°85 i * Figured. IV. MUTUAL RELATIONS OF THE PLEISTOCENE AND POST- PLEISTOCENE CANIDA. This most difficult subject) has puzzled zoologists from the time of Buffon and Daubenton to the present day. It cannot be entirely overlooked in such a memoir as the present, but no attempt will be made to deal exhaustively with it. Two questions are involved, which, though distinct, have the most intimate bearing upon one another. The first of these is, whether any valid and reliable distinction can be found between the dogs, on the one hand, and the wolves and jackals on the other. The second is, whether the origin of the domestic dogs is to be sought wholly or partially in the existing wild Canidee—wolf, jackal, or certain kinds of wild dog, or whether it may be found in one or more fossil species known or as yet undis- covered. The former of these questions may be first considered. A large number of points have been referred to by zoologists in their attempts to find valid osteological distinctions between dog and wolf. To each point in the following list the names of certain authors who allude to it are appended; but it 1s not implied that in every case the points are accepted as valid distinguishing characters by the authors who allude to them. (1) Wolves have the triangular part of the cranium between the orbits a little narrower and flatter than in dogs (Cuvier, Denny). (2) The sagittal crest is longer and more elevated in wolves than in dogs (Cuvier, Denny, Vieira). (3) The teeth, especially the canines, are longer in wolves than in dogs (Cuvier, Denny). = CANIS. 23 (4) The length of the upper carnassial pm. 4 is superior or at least equal to that of the molars m. 1 and 2 in wolves, while in dogs the length of pm. 4 is less than, or at most equal to, that of the m. 1 and 2 (Gaudry and Boule). (5) The plane of the eye-socket is more obliquely inclined to the brow, 1. e. the orbito-frontal angle is less in wolves than in dogs (Studer) (see Text-fig. 8). (6) The brow is more swollen at the base of the muzzle in dogs (de Blainville). (7) The zygomatic arch is less curved and shorter in the wolf (Vieira). (8) The coronoid process reaches above the zygomatic arch in the dog but not in the wolf (Vieira). (9) The caudal vertebree are more variable in the dog (de Blainville). Denny' also refers to the following points: (a) The intermaxillaries (? nasal Text-Fi¢. 8.— Anterior view of the skull of a dog, and instrument for measuring the orbito-frontal angle. processes of the premaxillz) and nasals are longer and narrower in dogs than in wolves; (b) the nasal cavity is wider in wolves; (c) the orbital projections (probably the post-orbital processes of the frontals) are more acute in dogs; (i) the jaws are wider; and (e) the tympanics are larger in wolves. The above is a considerable list, but the great majority of the points amount to very little and are quite inconstant and unreliable. Probably the most important character is No. 5, for which we are indebted to Studer.” The angle to which he alludes can be better realised by the preceding diagram (Text-fig. 8) than by description. He regards as belonging to wolves, skulls in which the angle between the plane of the orbit and that of the brow measures 40—45°, and as belonging to dogs, skulls in which the angle is greater 1 «Proc. Yorks. Geol. Polyt. Soc.,’ iii, 1857 (1859), p. 538. Dy} * « Abh. schweiz. pal. Ges.,’ xxviii, p. 13. 24 PLEISTOCENE MAMMALIA. than 45°. The effect of this greater obliquity of the plane of the orbit in the wolf is to produce the “ oblique leering eye which gives the wolf a false expression as compared with the noble, trustful expression of a dog, whose eye, with rounder opening, is more directed forwards.” He remarks that if we look at a wolf’s or jackal’s skull from above, more of the orbit is visible than in a dog’s skull. The reliability of this distinction, which is accepted by Scharff! and used in discriminating the canine skulls from the Edenvale caves, has been to some extent tested by examination of skulls at the British Museum and at Bristol with the following results : | Number of | Set; Fr |Maximum | Minimum| Average | Specimens. ae a angle. | angle. angle. (a) 8 Wolf (recent) Brit. Mus. 45° 40° 42° (b) 5 | Dog (prehistoric) 25 » 53° 46° 493° (c) 27 Dog (recent) * *s 62° 45° 504° (d) 9 3 i Bristol Univ. | 54° | = 48° 512° Belonging to each of the first three groups there were, however, certain exceptional skulls which are not included in the above table. Thus two additional wolf skulls belonging to group (a) gave angles of 47° and 48° respectively, two dog skulls in group (0) had angles of 42°, three skulls in group (c¢) had angles below 45°. The angle is not very easy to measure even with a clinometer such as is shown in the figure, and it was found that when the same skull was measured on different occasions slightly varying results were sometimes obtained. The measurement was in each case taken over the ends of the post-orbital processes of the frontal and jugal. Though it can hardly be claimed that the results of the measurement of the fifty-six skulls referred to in the above table afford a complete test of the reliability of the orbito-frontal angle as a distinguishing character between dogs and wolves, they certainly confirm Studer’s contention that the angle tends to be decidedly less in the wolf than in the dog, and that it affords a useful distinction of practical value. The occurrence, however, of dog skulls with an angle of less than 45°, and of wolf skulls with an angle of over 45°, shows that the distinction is not absolute, and cannot be relied on in all cases. The second point, that of the origin of the domestic dogs, is the subject of a most voluminous literature. It is beyond the scope of the present memoir and no attempt can be made to discuss it. The different opinions which have been maintained are, however, briefly the following: Daubenton and Cuvier were 1 «Trans. Roy. Irish Acad.,’ xxxii, B., pt. I, p. 208. | | \ ee ee ee ae AO cite ee ee ——— Beene 3 ‘* CANIS. 25 disposed to derive the domestic dogs from the wolf; Nehring, partly from the wolf and partly from the jackal ; Gueldenstiidt, and G. St. Hilaire, from the jackal ; de Blainville, Pictet, Boule (1889), Gaudry and Boule (1892), Bourguignat and Woldrich, from one or more extinct types of dog, neither wolves nor jackals; Studer, partly from wolves, partly from extinct types of dog; finally, von Pelzeln has recourse to all the above-mentioned sources—wolves, jackals and extinct types of dogs, and in addition derives certain races from the existing wild dogs, Canis sinensis and Canis pallipes. Jeitteles also has recourse to several living Species, including the jackal and Caiis pallipes. V. CONCLUSIONS. These may be very briefly stated and contain no element of novelty. In Pliocene times the wolf (Canis lupus) and common fox (Canis vulpes) were already inhabitants of Britain. In Pleistocene times they abounded and the Arctic fox (Canis lagopus) was sparingly represented ; but no animals which can be distin- guished as dogs have been recognised in Britain in Pleistocene deposits. In Pre- historic times, however, true dogs abounded. Doubt is expressed with regard to the desirability of recognising the occurrence of the genus Lycaon in England. My sincere thanks are tendered to Mr. W. J. Lewis Abbott and Dr. F. Corner for kindly placing their large collections of canine bones from the Ightham fissure at my disposal for examination and figuring. I am much indebted to Mr. H. Bolton, Prof. T. McK. Hughes, Mr. A. Somervail and Dr. A. Smith Woodward for the loan of specimens preserved respectively in the Bristol, Sedgwick, Torquay and British Museums. I further wish to thank Dr. A. Smith Woodward, Dr. C. W. Andrews, Mr. R. Lydekker and Dr. R. F. Scharff for help and information, and Mr. J. Green for the care he has taken with the illustrations. VI. BIBLIOGRAPHY. 1758. Daubenton in Buffon’s ‘ Histoire Naturelle,’ tom. vii, p. 53. 1774. J. F. Esper, ‘ Ausftihrliche Nachricht—Zoolithen, Bayreuth.’ 1776. J. A. Gueldenstadt, “‘ Schacale Historia,’ ‘Nov. Comment. Acad. Scient. Imp. Petropol., xx, 1775 (St. Petersburg). 1794. J. C. Rosenmiller, ‘ Quzedam de Ossibus fossilibus animalis,’ p. 27 (Leipzig). 1810. G. A. Goldfuss, ‘ Die Umgebungen von Miiggendorf’ (Erlangen). 1812. G. Cuvier, ‘ Recherches sur les Ossemens fossiles,’ tom. iv, iv, pp. 5—9. 1822. W. Buckland, “ Account of an Assemblage of Fossils—Kirkdale Cave,” “Jeli, Weanns., Gam, Ps Is. 4. 26 PLEISTOCENE MAMMALIA. 1823. W. Clift and J. Whidbey, “On Some Fossil Bones—Oreston,” ‘ Phil. Trans., cxill, p. 88. 1823. G. A. Goldfuss, ‘Saugethiere der Vorwelt,’ p. 401. 1824. W. Buckland, ‘ Reliquize Diluviane ’ (London). 1825. G. Cuvier, ‘ Recherches sur les Ossemens fossiles,’ 3"° ed., tom. iv, pp. 457—467. 1829. J. Rutter, ‘ Delin. County Somerset,’ p. 156. 1833. P. C. Schmerling, ‘ Recherches sur les Ossemens fossiles—Cavernes de Liége,’ tom. 11, pp. 22—46. 1839. J. C. Bellamy, ‘ Nat. Hist. S. Devon,’ p. 439. 1839. M. de Serres, J. M. Dubrueil, and B. Jeanjean, ‘ Recherches sur les Ossemens humatiles des Cavernes de Lunel Viel,’ pp. 72—79 (Montpellier). 1841. G. St. Hilaire, ‘ Hssais de Zoologie Générale,’ 11 (Paris). 1844. H. M. D. de Blainville, ‘ Ostéographie, Carnassiers,’ p. 101. 1846. R. Owen, ‘A History of British Fossil Mammals and Birds,’ pp. 123— 137. 1853. F. J. Pictet, ‘ Traité de Paléontologie,’ tom. 1, p. 202. 1856. R. Owen, “ Description of some Mammalian Fossils from the Red Crag of Suffolk,” ‘Quart. Journ. Geol. Soc.,’ xu, p. 227. 1859. H. Denny, “The Skull of a Dog exhumed from the Alluvial Gravel of Norwich,” ‘ Proc. Yorks. Geol. Polyt. Soc.,’ m1, 1857 (1859), p. 538. 1859. J. MacHnery, ‘ Cavern Researches.’ 1859. W.R. Wilde, “ Animal Remains belonging to the Academy,” ‘Proc. Roy. IrishiAtcade evans eos. 1862. L. Ritimeyer, “ Fauna der Pfahlbauten der Schweiz,’ ‘Neue Denkschr. Schweiz. Gesell. gesammten Naturwiss.’ 1862 and 1863. W. Boyd Dawkins, “ On a Hyena Den at Wookey Hole near Wells,” ‘Quart. Journ. Geol. Soc.,’ xvii, pp. 124 and 125, and abid., xix, p. 267. 1864. W. Boyd Dawkins, “On the Caverns of Burrington Combe,” ‘ Proc. Somerset Arch. and Nat. Hist. Soc.,’ xii, pp. 161—176. 1868. H. Falconer, ‘ Paleeont. Memoirs,’ 1, p. 525. 1872. L. H. Jeitteles, “ Die vorgeschichtlichen Altertiimer der Stadt Olmiitz und ihrer Umgebung,” ‘ Mittheil. Anthropol. Gesell. Wien.,’ 11. 1875. G. Busk, “ List of the Mammalian Remains . . . . inthe Rock Fissure Cavern in Creswell Crags, Derbyshire,” ‘ Quart. Journ. Geol. Soc.,’ xxx1, p. 683. 1875. J. B. Bourguignat, ‘ Recherches sur les Ossemens fossiles de. Canidee, pendant la période quaternaire,” ‘ Ann. des Sciences Géol.,’ vi, p. 59. 1876. W. Boyd Dawkins, “On the Mammalia and Traces of Man found in the Robin Hood Cave,” ‘ Quart. Journ. Geol. Soc.,’ xxxui, p. 248. 1876. S77: S77. S78. ILSADe 1880. 1880. 1880. 1881. 1881. 1882. 1883. 1885. 1884. . 1884. 1884. 1885. 1885. 18386. 1886. 1886. CANIS. 27 . Haughton, “ Animal Bones found in Knockninr ave,” ; : Haughton, “A 1 Bones found Knockninny Cave,” ‘Proc. Ro Trish Acad.’ (2), 11 (Sci.), p. 482. . Pennington, ‘ Notes on the Barrows and Bone Caves of Derbyshire.’ . H. Jeitteles, ‘ Die Stammvater unserer Hunderassen’ (Vienna). . L. Adams, “ Report on History of Irish Fossil Mammals,” ‘ Proc. Roy. Irish Acad.’ (2), iii (Sci.), p. 99. . L. Adams, “ Report on the Exploration of the Shandon Cave,” ‘Trans. Roy. Irish Acad.,’ xxvi, p. 221. . L. Adams, “On Recent and Extinct Irish Mammals,” ‘Sci. Proc. Roy. Dublimy Soc, ty) ps Ob: . H. Huxley, “On the Cranial and Dental Characters of the Canide,” ‘ Proc. Zool. Soc.,’ pp. 288—288. . T. Newton, ‘‘ Notes on the Vertebrata of the Preglacial Forest Bed Series of the Hast of England,” ‘Geol. Mag.,’ dec. 1, vu, p. 152. . Woldrich, ‘“ Beitrige zur Geschichte des fossilen Hundes,” ‘ Mittheil. Anthropol. Gesell. Wien,’ x1. . L. Adams, “ Report on the Animal Remains of the Bone Cave of Bally- namintra, Co. Waterford,” ‘Trans. Roy. Dublin Soc.’ (2), 1, p. 205. . IT’. Newton, “ Vertebrata of Forest Bed Series of Norfolk,’ ‘Mem. Geol. Slaves Oa Oe . D. Cope, “On the Extinct Dogs of North America,” ‘Amer. Nat., xvii, p. 243. . Gaborovski-Moindron, “ Les chiens tertiaires de l’Europe, et l’origine des Canidés,” ‘Bull. Soc. Anthrop. Paris,’ vi, pp. 870—884. . Lydekker, “ Siwalik and Narbada Carnivora,” ‘ Palzont. Indica,’ ser. 10, ii, p. 240. . Lydekker, ‘* Notes on some Fossil Carnivora and Rodentia,” ‘ Geol. Mag..,’ dec. i, 1, p. 442. . Nehring, “‘ Uber eine grosse Wolfsihnliche Hunderasse der Vorzeit u. tiber ihre Abstammune,” ‘Sitzungsb. der Gesell. Naturf. Freunde zu Berlin,’ ix. . Ball, “On the Collection of Fossil Mammalia of Ireland in the Science and Art Museum, Dublin,” ‘Trans. Roy. Dublin Soe.’ (2), iti, p. 340. . Lydekker, ‘ Catalogue of the Fossil Mammalia in the British Museum,’ pt. 1, pp. 121—186. . von Pelzeln, “ Kine Studie tiber die Abstammung der Hunderassen,”’ ‘Zool. Jahrbuch,’ 1, pp. 225—240. . Hicks, “On the Ffynnon Beuno and Cae Gwyn Caves,” ‘ Geol. Mag..,’ eee cee M. Wilckens, “ Palzsontologie der Haustiere,” ‘ Biol. Centralbl.,’ v, pp. 719 and 751. il! 28 1888. 1889. 1890. 1890. SOE 1892. 1893. 1894. 1894: 1902. 1908. 1903. 1906. IGQO7. A. M. 15% A. H. IN C. PLEISTOCENE MAMMALIA. Nehring, “ Wolf und Hund,” ‘ Naturwissenschaftliche Wochenschrift,’ ii. Boule, “ Les Prédécesseurs de nos Canidés,’ ‘Comptes Rendus,’ eviii, TOs “AIL C. A. Windle and J. Humphreys, “ On the Cranial and Dental Characters of the Domestic Dogs,” ‘ Proc. Zool. Soc.,’ p. 5. Smith Woodward and C. Dayies Sherborn, ‘A Catalogue of British Fossil Vertebrata,’ p. 324 (London). T. Newton, “ Vertebrata of Phocene Deposits of Britain,’ ‘Mem. Geol. SWE Va5 Do Gaudry and M. Boule, “ Matériaux pour l’histoire des temps quaternaires,” fase. iv, ‘ Les Oubliettes de Gargas,’ pp. 123—129. Grevé, “ Die geographischer Verbreitung der Hyaniden und Caniden,” ‘Zool. Jabrb.,’ f. v, p. 400. - . Vieira, “ Etude comparative du Squelette du Chien et du Loup,” ‘Ann. SGilo Nett. IBOHIO.,° 1, JO. LOY). .T. Newton, “On the Vertebrate Fauna from the Fissure near Ightham, Kent,” ‘ Quart. Journ. Geol. Soc.,’ 1, pp. 188—211. . Studer, ‘ Die Przehist. Hunde in ihrer Beziehung zu den gegenwartig lebenden Rassen,” ‘ Abh. schweiz. pal. Ges.,’ xxvii. . F. Scharff, “ Exploration of the Caves of Kesh, Co. Shgo (Mammals except Man),” ‘Trans. Roy. Irish Acad.,’ xxxu, B, pt. 4, p. 203. . Keller, “‘ Zur Abstammunesgeschichte unserer Hunderassen,” ‘ Viertel- p) io) ~S) jahrschr. Ges. Zurich,’ xlvii. . F. Scharff, “ Exploration of the Caves of Co. Clare (Animal Remains except Birds),” ‘ Trans. Roy. Irish Acad.,’ xxxiu, B, pt. 1, p. 43. . H. Reynolds, “ A Bone Cave at Walton near Clevedon,” ‘ Proc. Bristol Nat. Soc.,’ 4th ser., 1, pt. m1, p. 183 (issued for 1906). . | Nt ee Lag RRR ye inl len tli RS gta Gace RS a acim el PLAIN, IL, PrEistockNeE CaNIDm. Cranium. Wolf (Canis lupus). (Two thirds natural size.) Fic. 1. Dorsal ) 2. Lateral ¢view of a cranium from Kent’s Cavern, Torquay. 3. Ventral This specimen, which was figured by MacHnery (‘ Cavern Researches,’ Frontis- piece) and by Owen (‘ British Fossil Mammals and Birds,’ p. 123), is preserved in the Torquay Museum. Since these authors figured the skull, the mandible has been lost and the cranium has been somewhat damaged. a. Post-orbital process of frontal. b. Occipital condyle. c. Canine tooth. d. Infra-orbital foramen. e. Premaxilla. Anterior palatine foramen. PALZONTOGRAPHICAL SOCIETY.I909. } Reynolds, Pleistocene Camde. Pl] J.Green del. lith.et imp. : WOLF , CANIS LUPUS. Cranium. Ventral ! view of a cranium from Hutton Cave, Somerset. Dorsal Lateral ) Palatal : Fd een \ view of the ec Ventral view of a cranium from Banwell Cave, Somerset. Anterior view of the jaws from Oreston Cave, Plymouth. AAV tae PLEIsvocENE CANIDA. Craniwn and Mandible. Wolf (Canis lupus). (One half natural size.) orrespondinge mandible. All the above are preserved in the British Museum. Occipital condyle. b. Auditory bulla. ¢. Premaxilla. d. Jugal. e. Anterior palatine foramen. jf. Mandibular condyle. Coronoid process. ‘aj[gtpueuw. ® WNIUe dL) “SNdNn1 SINVWD‘A1OM ‘dun ya-ugt Tap wseag'p sl aleh plwuey) suao0jsie] J‘ spjouAey *6O0GI ALBIDOS IWVOIHdVHYDNOLNOAW1Vd ae p iy, + fy : 7 » } i aaa bie , ia) eee i % r . | ie | > I > | | ’ , | | | 1 ‘ | 2 ° i} . | i Vs hh | he i | || . Hh, . i ; Pipa ere’ TPIbVAS) JOE PLEISTOCENE CANIDA. Oranum and Mandible. Common Fox (Canis vulpes). (Natural size.) Fic. ee aterall 2 Dorsal view of cranium. ou) Ventral 4. Left mandibular ramus seen from the outer side. These specimens are from the Pleistocene of the Ightham fissure, near Maid- stone, and are preserved in the collection of Mr. W. J. Lewis Abbott, F.G.S., of St. Leonard’s-on-Sea. a. Parietal. b. Post-orbital process of frontal. c. Jugal. d. Maxilla. Cae Niasalle jf. Nasal process of premaxilla. yg. “ygomatic process of squamosal. h. Angle of mandible. PALZONTOGRAPHICAL SOCIETY. 1909. b Reynolds, Pleistocene Canide. J.Green del. lith, et imp CANIS VULPES. dible. Cranium man COMMON FOX, gy i ‘ % ae PLATE IV. PLEISTOCENE CANIDA. Cramum and Mandible. Arctic Fox (Canis lagopus). (Natural size.) Fic. 1. Dorsal | . Ventral }view of cranium. 3. Lateral 4. Left mandibular ramus seen from the inner side. 5. ‘lhe same seen from the outer side. These specimens are from the Pleistocene of the Ightham fissure, near Maid- stone, and are preserved in the collection of Mr. W. J. Lewis Abbott, F.G.S., of St. Leonard’s-on-Sea. a. Maxilla. b. Nasal. c. Post-orbital process of frontal. d. “ygomatic process of squamosal. e. Jugal. f. Premaxilla. g. Condyle of mandible. PALZONTOGRAPHICAL SOCIETY. 1909. Reynolds Pleistocene Canidee. | Pps 1. OND: 2. d.Green del.lith.et imp. ARCTIC FOX, CANIS LAGOPUS. Cranium & mandible. PLATE V. PLEISTOCENE CaNIDA&. Dentition. (Natural size.) 1. Right upper permanent dentition of Canis lupus. 2. Right lower permanent dentition of Canis lupus. 3. Right upper deciduous dentition of Canis sp. 4. Right lower deciduous dentition of Canis sp. The above teeth, which are all from the Pleistocene of the Torbryan Cave, | Torquay, are in each case seen from the inner side. D.i.3. and d.c. | were not represented in the series figured and are shown merely in outline. Preserved in the British Museum. | 5. Right | 6. Left These specimens, both seen from the outer side, are from the Pleistocene of { mandibular ramus of a young Arctic Hox (Cans lagopus). the Iehtham fissure, near Maidstone, and are preserved in the collection of Mr. W. J. Lewis Abbott, F.G.S., of St. Leonard’s-on-Sea. 7. and 8. Fourth lower premolar of the specimen from Spritsail Tor, Gower, described as Lycaon anglicus. ‘his specimen is preserved in the British Museum. 7. Outer aspect. 8. Inner aspect. a. Anterior cusp. In all the above specimens the roots when shown in outline are not visible, being hidden by the bone of the jaw. dun qo yp [Sp Us945'P op erp 270 | “eepluey) 220048131 * spjoukey ‘A Id "6061 'ALFIDOS IWOIHdVHDOLNOW1Vd | 2 ae | See SeeS EOCE 10. Ik, IW, 13. 14, Lo. ILO, PIA, WAL, PLEISTOCENE CANIDA. Vertebre. (Natural size.) Atlas, Wolf (Canis lupus), Oreston, ventral view. ‘ | 2, Axis; 3, fifth cervical; and 4, first thoracic; Wolf (Canis lupus), Durdham | Down, all seen from the left side. Sacrum, Wolf (Canis lupus), Oreston, ventral view. All the above are preserved in the Bristol Museum. Atlas, posterior view ) | Axis, seen from left side Third cervical, posterior view >— Common Fox (Canis vulpes). Fourth cervical, posterior view Fifth cervical, seen from left side J The above are from the Pleistocene of the Ightham fissure, near Maidstone, and are preserved in the collection of Dr. F. Corner, F.G.8., of Poplar. Seventh cervical, seen from left side ) First thoracic, posterior view | Seventh thoracic, seen from left side + Arctic Fox (Camis lagopus). First lumbar, seen from right side First free caudal, posterior view | The above are also from the Ightham fissure, and are preserved in the collection of Mr. W. J. Lewis Abbott, F.G.S. Sacrum, Common Fox (Canis vulpes), Durdham Down, dorsal view, Bristol Museum. a. Neural spine. b. Neural canal. c. Pre-zygapophysis. d. Post-zygapophysis. e. Vertebrarterial canal. jf. ‘Transverse process. g. Hypapophysis. h. Metapophysis. 2. Surface for articulation with head of rib. ), Nerve foramina. k. Surface for articulation with condyle of cranium. l. Posterior articular surface. m. Odontoid process of axis. PALA ONTOGRAPHICAL SOCIETY. 1909. Canide. 1stocene Reynolds, Ple J.Green del. lith et imp Vertebre.. Palxontographbical Society, 1911. A MONOGRAPH OF THE BRITISH PLEISTOCENE. MAMMALIA WOM Ti, PARE LV. THE MUSTELIDA. BY SIDNEY H. REYNOLDS, M.A., F.GS., PROFESSOR OF GEOLOGY IN THE UNIVERSITY OF BRISTOL. Pagrus 1—28; Piares I—VIII; TITLE-PAGE AND TABLE OF CONTENTS oF Vou. II. FON DON : PRINTED FOR THE PALHONTOGRAPHICAL SOCIETY, 1912. MONOGRAPH ON Mik BRITISH MAMMALIA PLEISTOCENE PERIOD THE MUSTELID‘. Order—CARNIVORA. Famity—MUSTELID A. I. INTRODUCTION. Te Pleistocene Mustelide,’ which form the subject of the present memoir, are an easier group to deal with than either the Urside or Canidz, not showing the variability and inconstancy of character which render it so difficult to come to a satisfactory conclusion about the mutual relationship of the members of the above groups. Including, as the group does, the glutton, badger, and otter, in addition to the Musteline (marten, polecat, stoat, weasel, etc.), it is somewhat too extensive and diverse to be conveniently treated from the historical aspect as a single entity. In the following paragraphs only the literature of a more general character will be referred to, each species being subsequently considered separately. As was the case with the bears and hyenas, the remains from the great Continental caves attracted attention at an earlier date than those of Britain, Goldfuss* recording bones of the glutton from the caves of Gailenreuth in 1818 1 As in a previous memoir dealing with the Pleistocene Canide, the classification and nomencla- ture adopted are those of Flower and Lydekker, ‘ An Introduction to the Study of Mammalia, Living and Extinct’ (1891). The generic name Mustela is employed in a wider sense than is now usual, most zoologists adopting Nilsson’s name Putorius for the polecat, stoat and weasel. * “Nova Acta Acad. Caes. Leop.,’ ix, 1818, p. 313, and ‘Saugethiere der Vorwelt,’ p. 468. ] 2 PLEISTOCENE MAMMALIA. and Sundvig in 1823, and Schmerling’ those of the badger, marten, and polecat, from the caves of Litge in 1833. Marcel de Serres, Dubrueil and Jeanjean” figured an otter’s mandible from Lunel Viel in 1839, while Croizet and Jobert? recorded the same species from the Puy-de-Dome deposits in 1828. The remains of the smaller Mustelidz were naturally not as a rule recognised at so early a date as those of the larger species, but Buckland,‘ as early as 1822, figured musteline teeth from Kirkdale, which he attributed to the weasel, and Goldfuss’ figured a mandible from Gailenreuth, which he attributed to a Viverra. Schmerling,° however, pointed out that the latter bone was musteline. The records to the date of writing were summarised by H. v. Meyer’ (1832), F. Holl? (1829—1831), de Blainville® (1844), and Giebel” (1847), while Owen, in 1842," and subsequently in 1846,” eave a full account of the available information regarding British occurrences. Gervais” (1859) dealt fully with all French records. Five species of Mustelidze were recorded by Falconer™ (1868) from the various caves of Gower, and other records were given by Dawkins” (1869) in his paper on the “ Distribution of British Post-glacial Mammals.” Very little has been written concerning Musteline remains from Ireland, though Adams," in 1881, recorded the marten and badger from Ballynamintra, co. Water- ford, and Scharff the badger, otter and stoat from the caves of co. Clare,’ and the stoat from Kesh,* co. Sligo. The most important records of quite recent date in Hneland are those of the Ightham” fissure, in which, in addition to Mustela robusta, the polecat, weasel and badger were met with. During comparatively recent times a number of important papers dealing with the Pleistocene Mustelide have been published on the Continent. HH. Cornalia, in his ‘ Mammiferes fossiles de Lombardie ’” (1858—1871), described remains of the badger, marten, and polecat, some of the polecat skulls being very large. This ‘Recherches Oss. foss. Cavernes de Litge,’ 1, pp. 158—166 ; 1, pp. 5—15. ‘Recherches Oss. humatiles Cavernes de Lunel Viel,’ p. 70, pl. 1, figs. 14 and 15. ‘Recherches Oss. foss. Dept. Puy-de-Dome,’ p. 89. Phil ianise exp sparle2snpleexcxe > “Die Umgebungen von Muggendorf,’ p. 282, pl. v, fig. 3. 6 « Recherches Oss. foss. Cavernes de Liége,’ i, p. 5. 7 *Paleologica,’ p. 47. 8 * Handbuch der Petrefactenkunde,’ p. 36. 9 « Ostéographie,’ fase. 4. 10 «Fauna der Vorwelt,’ 1, pp. 55—64. ll « Brit. Foss. Mammals,” ‘ Rep. Brit. Assoc.’ (Manchester, 1842), pp. 70—72. 2 « Brit. Foss. Mamm. and Birds,’ pp. 109—122. 13 *Zoologie et Paléontologie Francaises,’ pp. 243—253. 14 «Pal. Mem.,’ p. 525. 1 «Quart. Journ. Geol. Soc.,’ xxv, p. 192. 16 «Trans. Roy. Dublin Soc.,’ (2) i, p. 205. 17 «Trans. Roy. Irish Acad.,’ xxxiii, B, pt. i, pp. 40—43. 18 Ibid., xxxu, B, pt. 4, p. 205. 9 «Quart. Journ. Geol. Soc.,’ 1, p. 200, and lv, p. 425. 20 «Pal. Lomb.,’ ed. Stoppani, ser. 2. mo %w e MUSTELIDA. 3 paper was followed in 1879 by Liebe’s'’ account of the caves at Vypustek, in Moravia, where glutton, marten, stoat, and polecat were met with. In Woldrich’s’ three beautifully illustrated papers (1880, 1882, 1884), on the fauna of Zuzlawitz, near Winterberg, in the Bohmerwald, remains of polecat, stoat and weasel are figured, and in particular some very large skulls, which are attributed to the polecat, closely resemble the form afterwards described by Newton as Mustela robusta. Lastly, in 1886 appeared an important paper by Winterfeld,’ giving a general account of the Quaternary Musteline remains of Germany. Il GENERAL ACCOUNT OF THE VARIOUS BRITISH MUSTELIDA. Mustela martes, 14% Prne Marren. Two British species of marten, the pine marten (Mustela martes, Linn., or abietum, Fleming) and the beech marten (Mustela foina, Erxl.), have commonly been recognised as members of the British fauna. Alston,* however, shows good reason for believing that Mustela martes is identical with Mustela sylvatica of Nilsson, and that Mustela foina is not really an inhabitant of the British Isles. He mentions among others the following points of difference between the skulls of M. martes and M. foina, though many of them appear to be inconstant or inappreciable. M. martes. M. foina. 1. The breadth of the skull across the zygomatic 1. The breadth of the skull across the zygomatic arches is rather more than half the length. arches is much more than half the length. 2. The arches are highest posteriorly, whence 2. The arches are regularly curved, and broadest they slope rather suddenly downwards and and highest in the middle. forwards. 3. The sides of the muzzle are nearly parallel. 3. The sides of the muzzle are converging. 4, The anterior narial opening is oval. 4, The anterior narial opening is heart-shaped. 5. The palate is comparatively narrow. 5. The palate is comparatively broad. 6. The upper premolars are placed regularly in 6. The upper premolars are crowded, and often the line of the series; the fourth has the inner placed diagonally, their anterior extremities cusp large and placed nearly at right angles bemg directed imwards; the inner cusp is to the axis of the tooth. small and placed somewhat diagonally. 7. m. | has a slightly developed inner tubercle. 7. m. | has a well-developed inner tubercle. The marten was not mentioned by Owen in his ‘ British Fossil Mammals and 1 Thid., 1 (1894), p. 201. 6 «Nova Acta Acad. Caes. Leop.,’ ix (1818), p. 311, pl. viii. The mandibular ramus attributed by Goldfuss to a Viverra and figured by him (‘Die Umgebungen von Muggendorf,’ vy, 1810, 3) is assigned by Schmerling to a marten or polecat (‘Cavernes de Liége,’ ii, p. 5) and by de Blainville (‘ Ostéographie—Carnassiers, Mustela,’ p. 53) to the glutton. 7 *Oss. Foss.,’ ed. 2, 1825, pl. xxxi, figs. 23—25. 8 «Recherches Oss. Foss. Cavernes de Liége,’ i, p. 167. 9 “Geol. Mag.’ [2] vu, 1880, p. 424, pl. xv, and “ Vert. Forest Bed” (‘ Mem. Geol. Surv.,’ 1882), jo, U5 jolle ia 10 “Nat. Hast. S. Devon.,’ pp. 89, 94, 102. ll «Trans. Devon. Assoc.,’ iv, 1871, pp. 98, 102. 8 PLEISTOCENE MAMMALIA. 7 Dawkins and Sanford’ (1866) include it in their list of Pleistocene mammals on the evidence of the crowns of three canine teeth obtained from the caves of Bleadon and Banwell, Somerset, and from one of the Gower caves (see T'ext-fig. 1, 6, D, and &, for the Somerset specimens). It is not, however, mentioned by Falconer? in his list of the Gower cave-fauna, and Woodward and Sherborn? do not include Gower as one of the localities where its remains have been met with. Dawkins‘ in 1871 described a left mandibular ramus from the Plas Heaton cave, Cefn, near 8. Asaph, this fine specimen, which is shown in Text-fie. 1, s, bemg now preserved in the | Grosvenor Museum, Chester. Finally, in 1875 Busk* added Creswell Crags to the list of localities, though the determination was based only on two fragments of pelvis (see Text-fig. 2). A comparatively recent continentalrecord of the occurrence of the glutton is | by Liebe® (1879) from Vypustek in Moravia. Winterfeld’ (1886) discussed its distribution, and gave some German records of its occurrence in loess and other deposits. Meles tavus, tHE Bapacer. The remains of the badger were discovered in Pleistocene caves at an early date, and have been recorded from a yery large number, though, perhaps, not from so many as the habits of the animal would lead one to expect. The earliest records of the badger from Pleistocene deposits are by Schmerling® (1833), who gave good figures of the sku!l and limb-bones from the caves of Ligge, and by Mister’ (1836), who described it from the neighbourhood of Baireuth. Both these authors regarded their species as distinct from the modern species, Schmerling referring to Ins as Meles antediluvianus and Minster to his as Meles antiquus. . M. de Serres, Dubrueil and Jeanjean” in 1859 figured a skull and other bones from Lunel Viel, and affirmed the identity of the badger of the caves with the recent species, a point concerning which subsequent writers have been unanimous. 1 « British Pleistocene Mammalia ” (‘ Pal. Soc.,’ 1866), pt. 1, p. 21. | | 2 «Pal. Mem.,’ 11, 1868, p. 525. 3 «Catal. Brit. Foss. Vertebrata,’ 1890, p. 350. | 4 ‘Quart. Journ. Geol. Soc.,’ xxvii, 1871, p. 406. TY oly soo IUS7/3), jo, (OS7- | 6 «Sitzb. k. Akad. Wiss. Wien.,’ lxxix, 1879, pt. 1, p. 476. 7 «Ueber quartiire Mustelidenreste Deutschlands,’ 1886, p. 40. 8 «Recherches Oss. foss. Cavernes de Litge,’ i, 1833, p. 158. 9 ‘Verzeichniss der Versteinerungen . . . zu Buaireuth,’ 1836, p. 87. 10 «Recherches Oss. humatiles Cavernes de Lunel Viel,’ 1839, pl. i. , Se MUSTELIDA. — 9 Nordmann! in 1847 recorded it from Odessa, and McHnery, as reported by de Blainville? (1844) from Kent’s Cave, Torquay. Owen’ (1846) figured mL. Fia. 1.—Glutton (Gulo luseus). A. Left mandibular ramus seen from the outer side. B. The same seen from the inner side. From the Pleistocene of the Plas Heaton Cave, Cefn, near St. Asaph (Grosvenor Museum, Chester). ©. Crown of left upper canine. From the Pleistocene of the Bleadon Cave, Somerset (Taunton Museum). [D. and E. Crowns of left lower canines. From the Pleistocene of Banwell Cave, Somerset (Taunton Museum). All three teeth seen from the outer side. Nat: size. Fie. 2.—Glutton (Gulo luscus). Busk’s figure of a fragmentary innominate bone from the Pleistocene of Creswell Crags. Reproduced by permission of the Council of the Geological Society. Nat. size. 1 «Découv. Gites riches en Oss. foss. Odessa,’ p. 4. 2 «Ostéographie,’ fase. 4. 3 « Brit. Foss. Mamm. and Birds,’ p. 109. 10 PLEISTOCENE MAMMALIA. McEnery’s specimen—a well-preserved mandible, now in the British Museum. He also recorded the badger from Berry Head. H. von Meyer’ (1859) described remains of the badger from the neighbourhood of Weimar, giving a number of references to French and other records. Dawkins and Sanford’ (1866) recorded it from Banwell and Wookey Hole, and Falconer® (1868) from a number of the Gower caves. Other records are from the caves or fissures of Durdham Down,* Uphill,’ Ightham,® Cefn’ (near St. Asaph), and Hoe Grange.® Pleistocene deposits other than caves have yielded bones of the badger at Newbury, Berkshire, and Grovehurst, Kent, though the age of the latter deposit is somewhat doubtful. Adams’ records it from Ballynamintra, co Waterford, and Scharff from the Edenvale, Newhall and Barntick caves, co. Clare. The remains of the badger were remarkably abundant in the Langwith cave, near Mansfield, and included the remarkably elongated skull figured in PI. V. LTutra vulgaris, THE Ovrer. Owen" referred to a mandible of this animal as having been found in the Norwich Crag at Southwold, and to a humerus found in the same beds at Aldborough, but Newton” was unable to see the specimens and verify the record. He, however, recorded” it from the Forest Bed of Hast Runton. He further believed that an otter occurred in Britain in Red Crag times, referrine™ to de Blainville’s species Lutra dubia, a specimen from the Red Crag of Foxhall, near Woodbridge, which differed from Lutra vulgaris in having the carnassial tooth longer from before backwards, and narrower than in the recent species, while ‘Paleontographica, vil, 1859, pp. 41—40. “ British Pleistocene Mammalia” (‘ Pal. Soc.,’ 1866), pt. 1, p. xxu. ‘Pal. Mem.,’ 11, p. 525. 4 «Proc. Bristol Nat. Soc.,’ n.s., v, 1885—88, p. 44. > Thid., w.s., ix, 1898—1900, p. 159. 6 « Quart. Journ. Geol. Soc.,’ lv, 1899, p. 428. 7 «Geol. Mag.’ [3] ii, 1886, p. 571. 8 «Quart. Journ. Geol. Soc.,’ lxi, 1905, p. 50. 9 «Trans. Roy. Dublin Soc.’ [2] i, 1881, p. 208. 10 «Trans. Roy. Irish Acad.,’ xxxiii, B., pt. 1, p. 42. 1 « Brit. Foss. Mamm. and Birds,’ p. 119. 12 «Geol. Mag.’ [3] iv, 1887, p. 145. 13 «Quart. Journ. Geol. Soc.,’ xlvi, 1890, p. 444. 14 “ Vert. Pliocene Deposits ” (‘Mem. Geol. Surv.,’) p. 12. MUSTELIDA. U3 the inner tubercle is also smaller. The specimen further differs from Lutra vulgaris in that each of the premolars has the posterior root much longer in proportion than the anterior. In the same memoir (p. 13) Newton described a new J]. TaspLe sHowine tHE Disrripurion oF Musretip® In tHe Preisrocenn Derpostts oF THE BririsH Isnns. | & 2S SS = 3 : _~ ~ = 5 2 » 5 3 ol a o s © As | es | 2 | Se | Ss | SS | Be ae S| a5 S 3 Ses | as 3 ye) 2s eed) es) oS a EMOCENERMEPOSItS eran. s..cphcun henecnta<- Se BA ie a a an Sr + + PLEISTOCENE CavEs| AND |FISSURIES. Bacon Hole, Gower......... BN ARs ckallit tes ae + = == Ballynamintra, co. Waterford _ PRS cesta Spe ies ae ie ve hg + ae TBaniTelllly ccs eae coeltees eae on wenn nie a So we af Fe te Sees + IS AEMT Ck com Clancum ae vale a + oe se ae fon ae enchiepeteee vcr ets he Nad. 8 ia ee rc ey: + eraavaiilicadiirr oe: actrees eek Aone cee: fe = + + oe ch + ae BCA OMG ee det ye AMM Med ol. Os ope HE WA ay se oe oe ae cheese Bees + HS GIO NTO et MA Nek yeh ve cease + + +? “ts ae en WeimeneaniSt., Asaph Yo cfsies lec cs | se + + Creswell Craigs 801 ise SPE RL AM ey, + + set Dog Holes, Warton Crag SHUR Ohare | de ale Dardkan IDO Waa ee atertaen oe ee eee wae me : at ais Hidenyalesco: Olare .. 6.2.24. 8). es ~ co ol + ee f at Gmays Whurrock, Hssex 2...) eee. se * a Oe a a oe aF Happaway, Torquay ... .. Fidei Pea | ap ee + Pera a + Me Hoe Grange, near Matlock sae ss a soe oy ee et: + Thelaytlingninil= 52), 3 fa te a, ara fi + ae a ae + are Ipswich 2d Dc Se aR see a Beas ie ng tl a ; sie - rt. oF tenis Cane, MOrOMayi: y-pec.s2 000 taser Ls at es + + be ae + TOSI; COs SINE), ae a NO era ee Se ee 2 it + ba $e: ee TACTILE Wee) pose Us ae a ea Ie eee ee we ea See 4) +? oe Langwith Bassett, near Mansfield... af ce + es + ate SIs Womomole (Gower eee! hee. dee ee ae ce a aioe oe & + + Newhall, co. Clare dc ieee + ey Temes + + Orestonwreten meet w hte FN. a a ay +? Hlymrouuln Geaisedulbeach)\mype 8h. .s6| | oe + ¥ oe vayenselidteaG owen sees. ose aas se aif: 4. S rearia cl Onin cya? eo eh Ne tu aoe a4 A + SpritsalleRomaGower 6.2. fo 5 es. + oe + + Be MNCS Gall Cee etsy. nn i aa et rs al ee sey ee a + ACO jiolantl aera gi Oe pt VSR AP oka Maret. af VAY CDI a es eee Moe ars eee ee e. . ne ee ae =F WealmyBridce: Devon) 4) c-s eee ae ae oe ee Ge ar Note.—Some of the records from co. Clare are probably Prehistoric rather than Pleistocene. 12 PLEISTOCENE MAMMALIA. species (Lutra reevest), founding it on a right lower carnassial tooth which had not cut the jaw, from the Norwich Crag of Bramerton. As is natural from the habits of the animal, the remains of the otter are scanty in caves, but more abundant in river-gravels and similar deposits. Marcel de Serres, Dubrueil and Jeanjean' described and figured a lower jaw from Lunel Viel, which they referred to a new species, and Croizet and Jobert* described bones from the Puy-de-Dome district. The only Post-Phocene specimens referred to by Owen in his Bean Fossil Mammals and Birds’ are from the peat and its associated marls of Cambridgeshire, and belong to the Prehistoric rather than to the Pleistocene fauna. Dawkins and Sanford’ (1866) stated that the only Pleistocene remains of the otter with which they were acquainted were from Kent’s Hole, Torquay, and Banwell Cave, and from the brick-earth of Gray’s ‘Thurrock, Essex. In Dawkins’ paper* on the “ Distribution of the British Post-glacial Mammals” (1869) the additional cave-localities of Durdham Down and Lone Hole, Gower, were given, with Ipswich, as a river deposit. The otter occurs rarely in the Langwith Cave. Scharff? recorded the otter from the Newhall Cave, co. Clare, this being the first notice of its occurrence in Ireland in Pleistocene strata. Ill. DESCRIPTION OF THE SKELETON. The Mustelidz form a large and somewhat heterogeneous group of Carnivores, and are grouped with the Urside and Procyonidee in the section Arctoidea. The cranial part of the skull tends to be considerably elongated and somewhat sharply marked off from the facial portion. The glenoid cavity is relatively far forward. The Mustelide agree with the Felidee and Hyzenidee, and differ from the ereat majority of the Urside, Viverride, and Canidx, in havmeg no alisphenoid canal. The auditory bulla is not as a rule much inflated. The palate is generally considerably produced behind the last molars. The hamular process of the pterygoid is prominent. The infra-orbital foramen is generally very large, and the orbit communicates widely with the temporal fossa. The post-glenoid process tends to curve over the mandibular condyle, and sometimes holds the mandible attached to the cranium. The dental formula in the great majority of cases is 1. $,c. t, pm. 324,m. 3. In 1 * Recherches Oss. humatiles Cavernes de Lunel Viel,’ p. 70, pl. u, figs. 14, 15. 2 “Oss. foss. Dept. Puy-de-Dome,’ p. 89. ’ « British Pleistocene Mammalia” (‘ Pal. Soc.,’ 1866), pt. 1, p. xxi. 4 «Quart. Journ. Geol. Soc.,’ xxv, 1869, p. 198. 6 «Trans. Roy. Irish Acad.,’ xxxii, B. pt. 1, p. 41. q ai ee Ty 2 MUSTELIDA. 15 rare cases the molars may number + or 3. ‘The upper carnassial, pm. 4, differs from that in Urside and resembles that in Felide, Hyzenide, and Canide, in possessing a more or less antero-internally placed inner tubercle supported by a distinct root. ey ee as eUED- Mustela.—The cranial portion of the skull is not so sharply marked off from the facial as in Meles and Lutra. The sagittal and superciliary crests are less developed than in Meles; the occipital crest on the other hand is commonly very strong. The post-orbital processes of the frontal and jugal are fairly prominent, and sometimes approach one another, especially in Mustela ermiiea. The foramen magnum is of relatively large size. The auditory bulla is considerably inflated. The infra-orbital foramen is smaller in proportion to the size of the cranium than in Lutra and Meles, and the post-glenoid process is not sufficiently recurved to hold the mandible attached to the cranium. Gulo.—The cranial portion of the skull is more strongly marked off from the facial than in Mustela, but less so than in Lutru. There is a greater development of ridges, especially of the sagittal crest, than in any other British member of the Mustelide, except Meles. The jaws are very powerful. The foramen magnum is of relatively smaller size than in Mustela, and the auditory bulla, though variable, is less inflated. The paroccipital process of the exoccipital is prominent, while the post-orbital process of the jugal is very slightly developed. The post- glenoid process is much incurved, and holds the mandible firmly attached to the cranium. Meles—The skull of the common badger bears a very close resemblance to that of the glutton in general form, development of ridges, strength of jaws, and relative size of the foramen magnum. Also in the development of the paroccipital process of the exoccipital and of the post-glenoid process, which attaches the mandible to the cranium perhaps even more firmly than in Gulo. The superciliary ridges are somewhat stronger than in Gulo, and the zygomatic arch is rather stouter. Inutra.—The skull of the otter is of a peculiar character, broad and depressed, with the cranial portion, which is much expanded posteriorly, sharply marked off from the facial portion by a strong constriction behind the orbits. The sagittal and superciliary ridges are but slightly developed. The infra-orbital foramen is very large, and the post-glenoid process is not so much recurved as in Meles. The ventral surface of the cranial portion of the skull is notably broad and flattened, and the auditory bulla is very httle inflated. The mandible is small and weak in comparison with that of Meles. . PLEISTOCENE MAMMALIA. C66 c9G 9-€ LL LIL "SUIT YOUMSpoag) Udy [eMang ‘or107 Yoid ‘s1pbpna vaynyT “ST 68 6&1 “(TIED SUTIN) Plegsavyy IBIT “YIIMIUVT] ‘99009 -SI0[q “AIBA 8X} sajzaTy ¢8 IL RGi@e) SUTIN) PISGSARIN | 1Beu ‘YIIMouBT ‘aued -OFSI9[q ‘snxn} sa7ayy GSG-01 G9-T gos c9-F &-& £9 [6-6 6-9 ST-T Sé-§ “ g[Apuoo Jo pus 09 pens “voll SHUIeL TupNdIpuvut fo Yjsuey “TT Teeeiee eae Aten t ara cnsaaecee sen a -SVUE WSUIBTOF FO TOPOUIVIP OSCOASURTT, “OT AURROssasacteccost cccaconenncareaccens TT Jo sny 7OOATE OF penis aqered jo yycueTy <6 ses =" Sntuedo [elMvu 04 [2] Woy Fo sosso90 ad yVq1q.10-gsod waem4oq, AvMpttue gutod wor voVE JO YYouey gs Sopaopeddenedsnaoc[S “+ 7esourenbs Jo spot oreutosdz puryod ATeyeIpeULUAL YFPI MA © POFHO UISC RAG OD KOS ACO sessanoad [wAIq.t0- are pulyeq wmiueto Jo [ZpIM wWniutury, 9 ee ee we tee et eee eee eee yequo.az jo sosson0ard. [e}tqt0-ysod jo spue woaajoq YIPIM “S “ Toulvaog [VUTATTPORT 92 YIPLMA “P Pes ceeietia aise sear Haba apanr on ie son taeee qgoxo [e} -jrsus jo doq 07 prousyds-tsvq pur jeqrd -1090-IS¥q WIM} IINJNS WOT] FY SIOFT “g 5 ee MER enc RTeR Ine sunt cemou emer coaetebe soyoue oeuoshz ssolov YPM osuUleI4xm “Z Sui a dhenanahens tens TMs Jo puto zortojwe 0} Yoj}ou «V[Apuodte4UL WoAT Y}cUSeT “T “CSUN IMG) F880 TT “ON ‘ouImMoqg.culyyIg *quimqaAory ‘oued -OFSTO[q ‘sry? 89797 “CSN “HIIG) “19S LL OL “ON “ABAION ‘pansio.o] ‘quadad ‘ P snaRn) ON ‘(TOD Leusr0oH) am04s -PIBIT TvoU ‘ULLByI yoy “QUDIOISLO[ qT “Mynurw “IBA S2.UDb] na MjaIsnN TL “CHLOO 99040 F STMOT) BUOISPIey, Iva “WIBYAYST ‘aued04 -SId[q ‘seep hyna npapsnyye “CSD “9Ltg)) ‘pray Aalog ‘aue004 “SIO[_ “vaurma vpapsnqr ts Core) IIUIOD) PUWOJSPIBI IVIU ULBYIY.S]T ‘OILOISIY -Olg ¢ ‘sniwoynd vjapsn qe “CSU "FIIE) ‘T6010 ON ‘ourysaesd -1plBVepN “YABITOg 4Yuedet “9 snasopnd npaysnzr “C1109 oad y SIMAT) OUOISPLET 1BaU *MBYIY.ST ‘99004 -SIO[_ ‘Mysnqot wjaysnyy “CSU FI) “€66ZL ON ‘AuBULIay *g “Quaded‘ Pouro/ mjaysn ye “(aad “SNL YBN) atRIQ “09 ‘SOABD [[BUMON “OLLO9SUT “Old d ‘sapimul vjapsnyqy ‘STING ATNITHLSAP JO SUNGINGUASVA TAILVAVaWOH xo aTavy, (T) MUSTELIDA. 15 B. Tue DENTITION. Mustre.a.—Dental formula—i. 3, c. +, pm. 2-4, m. 4. Permanent Dentition of the Upper Jaw.—I. 1 and 2 are very small, one-rooted, simple teeth, 1. 3 is a larger and caniniform tooth. The canine is a relatively very large tooth with a lon o@ shehtly recurved crown. Pm. 1] is a very small one- rooted tooth which, while present in Mustela martes and other representatives of the genus Murtes of Nilsson, is absent in MZ. putorius, M. robusta, M. erminea and M. vulgaris, representatives of the genus Putor/us of Nilsson. Pm. 2 and 3 are simple conical teeth with two roots and a rather well-marked cingulum. Pm. 4, the carnassial, is a large tooth with a prominent blade consisting of a larger anterior and a smaller posterior lobe. The cingulum is well developed, and there 1s a prominent inner tubercle near the anterior border of the tooth. M. 1, which is as large a tooth as pm. 4, 1s short, but very wide, with a raised outer portion bearing several small, ill-defined cusps, and a depressed and more flattened inner portion terminated by a raised semicircular inner border. Permanent Dentition of the Lower Juw.—l. 1, 2Q0and 8 are all very small teeth, their crowns being only from a quarter to a third as long as that of the canine, which is somewhat sharply recurved. Pm. lisa small, simple, one-rooted tooth, and like pm. 1, is absent in Putorius. Pm. 2, 3 and 4 are all very similar teeth with two roots and conical crowns. In pm. 3, and still more in pm. 4, there are indications of a slight additional cusp on the posterior edge of the main cusp. M. 1 is a large tooth with a bilobed trenchant blade and a depressed posterior portion or talon only half the length of the blade. M. 2 is a small one-rooted tooth with a rounded crown. Guto.— Dental formula—i. 3, c. +, pm. 4, m. 4. Permanent Dentition of the Upper Jaw.—I. 1 and 2 are relatively powerful teeth, rather sharply curved downwards, with edges of a somewhat chisel-shaped character and indications of slight lateral cusps. I. 3 isa large caniniform tooth with a strongly marked cingulum, which passes obliquely ¢ along the inner face of the tooth, ending in a slight cusp not far from the point. C.1is of the usual character, and powerful, but not specially large. Pm. 1 is a simple, conical, one-rooted tooth. Pm. 2 is two-rooted, and has the apex of the crown placed far forward. Pm. 3 is a very powerful two-rooted tooth with a rather low conical crown. Pm. 4, the carnassial, is a large tooth with a prominent bilobed blade supported by two roots, the anterior lobe being the larger. The inner tubercle, which is supported by a third root, is small and depressed, but very sharply marked off from the rest of the tooth. There is a fairly promiment cingulum which is raised into a slight cusp at the anterior end of the tooth. M. 1 is a rather large tooth transversely placed. A depression divides it into an outer. portion supported by two roots and 16 PLEISTOCENE MAMMALIA. raised into three ill-defined cusps, and an imner somewhat larger portion with a low cusp and a raised inner border. Permanent Dentition of the Lower Jaw.—lI. 1, 2 and 3 are relatively powerful teeth not differing greatly in size, though 1. 3 has the crown somewhat expanded ; i. 2 arises from the jaw at a point behind i. I andi. 3. C. is a powerful tooth with a somewhat prominent cingulum, which often gives off a ridge running along the inner face of the tooth to the apex. Pm. 1 is a small one-rooted tooth with a circular crown. Pm. 2, 3 and 4 are powerful two-rooted teeth increasing pro- gressively in size. The crown is conical, and the apex, central in pm. 4, is further forward in pm. 8 and still further forward in pm. 2. M. I is a powerful tooth with a large blade, consisting of two equal-sized trenchant lobes and a small depressed talon. M. 2 is a small tooth with an oval crown not raised into any prominent cusps. Merzs.—Dental formula—i. 3,c. +, pm. 4, m. ¢. Permanent Dentition of the Upper Jaw.—I. | and i. 2 are simple teeth with somewhat chisel-shaped blades, 1. 3 is more caniniform. The contrast in size between i. 3 and c. is not so greatasin Dutra. Pm. 1 is a very small tooth which almost always falls out at an early period. As a rule its alveolus is completely closed in old animals. Pm. 2 and 3 are simple, conical, two-rooted teeth. Pm. 4, the carnassial, has a prominent blade with a large conical anterior lobe and an ill- defined and often scarcely recognisable posterior lobe. The inner tubercle is large and depressed and not so much anteriorly placed as in Mustela, or so sharply marked off as in Gulo. M. 1 is a very large tooth with a broad surface covered by a series of small tubercles, which rise to form two rather prominent cusps at the antero-external border. This tooth has three roots. j Permanent Dentition of the Lower Jaw.—The lower incisors are simple teeth of the same character as those in the upper jaw. The canine has a thickened base to the crown, which is somewhat sharply recurved. Pm. 1 is very small, and early falls out; pm. 2, 3, and 4, are simple conical two-rooted teeth. M. 1 is a long and relatively very large tooth. The anterior half has a rather ill-defined bilobed blade, with a cusp placed internal to the posterior lobe. The posterior half has a some- what depressed middle portion surrounded by a series of low cusps. M. 2 is a small one-rooted tooth, bearing several slight elevations on the crown. Lurra.—Dental formula—i. 3, c. +, pm. ¢, m. >. Permanent Dentition of the Upper Jaw.—l. 1 and 1. 2 are small cylindrical teeth ; i. 3 is somewhat larger and more caniniform, but the canine, which is a rather long and slender tooth, contrasts strongly in point of size with the incisors. Pm. 1 is a very small simple tooth, and often falls out early. Pm. 2 and 3 are simple conical two-rooted teeth. Pm. 4, the upper carnassial, has a trenchant blade, with one very prominent principal lobe and a somewhat smaller posterior lobe. The MUSTELID A. iz inner tubercle is large and depressed, with a sharp raised edge. M. 1 1s a large, somewhat irregular tooth, broader than long, with two cusps on the outer edge, divided by a depression from two on the inner border. Permanent Dentition of the Lower tie 4s very small; 1.2 and 3 are slightly larger, but are very simple one-rooted teeth. The canine, as in the upper jaw, is greatly larger than the incisors. Pm, 2, 3 and 4 are simple, conical, two- rooted teeth, the cone in pm. 2 being obliquely truncated in front. M. 1, the carnassial, is a relatively large tooth, somewhat variable in character. The posterior half, or talon, is depressed; the anterior half bears two trenchant lobes, with a large tubercle internal to the posterior lobe. The cingulum is prominent in m. 1 and all the lower premolars. M. 2 is a rather small square tooth, with a flattened crown and a single root. o. THe VERTEBRAL CoLuMN. The numbers of the vertebre are as follows: Cervical. Thoracic. Lumbar. Sacral. Caudal. Mustela 7 14 6 3 18—33 Gulo a ‘ 15 5 3 . 14orl1d Meles . a 15 5 3 18 Intra 7 144—15 . 5-6 . 3 . 25—26 There is little in the vertebral column of the Mustelidze which demands special comment, but allusion may be made to the following points : (1) The length of the tail in Lutra. . (2) The relatively large size of the atlas vertebra in Lutra and Mustela. (5) The length of the spines of the anterior thoracic vertebree of Gulo. (4) The elongated character and shortness of the neural spines of the lumbar and posterior thoracic vertebre of Mustela. (5) The expanded character of the transverse processes of the lumbar vertebrze of Lutra. p. THe Limp GIRDLES. The Pectoral Girdle.—The scapula shows a considerable amount of variation in shape and in the character of the acromion, which is always strongly developed, while the coracoid process is scarcely defined. In Mustela, Gulo, and Lutra there is a very large pre-scapular fossa and the coracoid border is gently curved. The supra-scapular border in Gulo and Lutra forms an angle not much less than a right angle with the spine. In Mustela the supra-scapular border is very short, and the @) 18 PLEISTOCENE MAMMALIA. es | iy \ PLA \\ y fh Fic. 3.—The left scapula seen from the outer side. A\ Polecat (Mustela putorius), from the Pleisto- cene of Ightham (Corner Coll.). B Badger (Meles taxus), from the Pleistocene of the Langwith Cave (Mullins Coll.). (© Otter (Lutra vulgaris), from the Prehistoric peat of Burwell fen (Sedgwick Mus.). Natural size. 1, glenoid cavity ; 2, spine; 3, acromion; 4, coracoid process; 5, pre-scapular fossa ; 6, post-scapular fossa ; 7, supra-scapular border ; 8, coracoid border. von s, a) Fie. 4.—A Left innominate bone of a weasel (Mustela vulgaris). B left innominate bone of a stoat (Mustela erminew). Both from the Pleistocene of Ightham near Maidstone and preserved in Dr. F. Corner’s collection. © right innominate bone of a giant polecat (Mustela robusta) from the Pleistocene of Ightham (Lewis Abbott Coll.). [D right innominate bone of a marten (Mustela martes) from the ? Prehistoric deposit of the Edenvale Cave, co. Clare (National Mus., Dublin). Natural size. All bones seen from the ventro-external aspect. 1, acetabulum; 2, obturator foramen ; . 3, ischium ; 4, sacral surface of ilium: 5, iliac surface of ilium; 6, tuberosity of ischium ; 7, pubis. MUSTELID A. 19 post-scapular fossa is only about half as wide as the pre-scapular fossa at its widest. The scapula in Meles differs considerably in shape from those of the other members of the group, especially as regards the post-scapular fossa, which is about equal in size to the pre-scapular. The supra-scapular border forms an angle of about 60° with the spine, and the coracoid border is sharply angular. The development of the spine 1s greatest in Meles and Lutra, less in Gulo, and slight in 3 Mustela. The acromion is prominent and sharply recurved in all four genera, but less in Gulo than in the others. The Pelvic Girdle.—In Mustela and Lutra the pelvis is relatively weak and the ilium is little expanded. In Gulo, and still more in Meles, the ilium is considerably expanded. In Gulo the junction between the supra-iliac and acetabular borders of the ium is gently rounded, while in Meles, it forms a prominent projection. The ischial tuberosity is more prominent in Meles than in Glo. 20 PLEISTOCENE MAMMALITA. (2) TasnEes or Comparative MEASUREMENTS or THE Limes Girpuus. Meles taxus, Lutra vulgaris, Pleistocene, Lang- , Prehistoric, Burwell with (Mullins fen (Sedgwick Coll.). Mus.). SCAPULA. 1. Length from coracoid process to end of SPINE pa surasetisiagec suisse solos sete aoseanion goatee: 9°45 7 | | 2. Length measured along glenoid border ... | ' 17-55 6:0 3. Maximum diameter of neck .................. 2-0 1:8 | eee foe |\Mustelarobusta,| >, y fe Lutra vulgaris ustele martes,” Pistooenc, | Mustle pute | Matele cul | yretestaxun, | “Prehistoric, ; > | Iehtham, nr. | , § gante, Pleistocene, Reach fen, * Newhall, Co. 5 | cene, Ightham,| cene, Izhtham, fe Olare (Nat. Maidstone Take, WLNGISo NA || sane, RiaAélsii@raye Langwith Cambs, (Sedg- Mus. Dublin). | pe eas (Corner Coll.). | (Corner Coll.). (Corner Coll.) Were: Prnyic GIRDLE. 1. Maximum length ............... 61 5°35 39 18 11:0 104 2. Length from acetabulum to | supra-iliac border of ihum ... ow 2°8 2°2 1-1 6:05 48 3. Dorso-ventral measurement of ilium at widest point......... 12 | 11 ) 35 2'8 1:95 4. Antero-posterior diameter of ‘ acetabulum ieee ret eerie tener 95 ‘8 65 25 17 1:3 5. Length from acetabulum to | posterior border of ischium ... 2:05 | 18 1:35 i) 30 4:3 |6. Maximum diameter of obtu- vator foramen ..................+5+ U7 1°35 1:0 a) 2°2 2:3 ee gk. Tae Limes. These show a progressive decrease in relative length from Gulo, in which they are longest, through Meles, Lutra, and the larger members of the genus Mustela to Mustela erminea and vulgaris, in which they are very short. The limbs are sub-plantigrade in Gulo, Meles and Lutra, digitigrade in Mustela. The claws are strong (except sometimes in Dutra), and in Mustela are semi-retractile. Anterior Iimb.—The humerus in Meles and Lutra is a very powerful bone with strong deltoid and supinator ridges. An ent-epicondylar foramen is present in Meles, Gulo and Mustela, and may or may not be present in Lutia. In Meles the radius and ulna are also very short powerful bones, the lower end of the ulna bearing a prominent ridge on its inner side. The metacarpals tend to be shorter than the metatarsals, especially in Lutra. Posterior Limb.—In Mustela this is considerably longer than the anterior, but the difference is less marked in the other genera. In Meles and Lutra the femur does not show such conspicuous ridges for the attachment of muscles as does the humerus of these animals. The fibula is slender, and stands somewhat widely away from the tibia except at the extremities. The metatarsals and digits are of greater relative length in Gulo than in Meles. ; MUSTELIDA. Fig. 6.—Limb bones of a polecat (Mustela putorius). A\ rightfemur; B left tibia; © right fibula (anterior view); D right humerus; E right radius (antero-external view); F right ulna (inner aspect). All from the Pleistocene of the Brixham Cave and preserved in the Hritish Museum. In Text-figs. 6 to 10 the anterior aspect of the femur and tibia is shown, the posterior aspect of the humerus. Lettering of Text-fies.6 to 10. 1, head of femur; 2, great trochanter ; 3, third trochanter ; 4, internal condyle of femur; 5, external condyle of femur; 6, cnemial crest; 7, head of humerus; 8, trochlea; 9, internal condyle of humerus; 10, external condyle of humerus; 11, ent-epicondylar foramen ; 12, humeral articulating surface of radius; 13, carpal articulating surface of radius; 14, siemoid notch; 15, olecranon. All fioures of the natural size. Fie. 7.—Limb bones of a giant polecat (Mustela robusta). A right femur; B right tibia; © right fibula (posterior aspect) ; D left humerus; F right ulna (inner aspect). All from the Pleistocene of the Ightham Fissure. The femur, tibia and humerus are preserved in the collection of Dr. F. Corner, the fibula and ulna in that of Mr. W. J. Lewis Abbott. Fig. 8—Limb bones of a weasel (Mustela vulgaris). A\ right femur; B right tibia; DPD left humerus ; F left ulna (outer aspect). All from the Pleistocene of the Ightham fissure. The femur, humerus and ulna are preserved in the collection of Dr. F’. Corner, the tibia in that of Mr. W, J. Lewis Abbott. 21 (3) Tasies or Comparative MuasuremEnts or THe Bones or THE AnTERIOR Lime. — to Fe ; 5 ; P Mustela vulgaris, ae ‘ Pian aeE | Dieist Claas, Mustela putorius, var. minuta, ; Telesis, LTE SD = 2 Pleistocene, Pleistocene, — om Ightham, nr. Brixham, No. Ightham, nr. | Happaway, No. Maidstone | 43919 (Brit. Mus.).| Maidstone M, 5811 (Brit. (Corner Coll.). (Corner Coll.). Mus.). Homerus. Ns DESOVEtH nse sdoeaacowcconbacce S00 o7 42 2:0 115 2. Maximum diameter at distal OTe asi faaciuctieines sages ROSES 15 1:05 “4, * 3:3 | i Fiptiee, wali Mustela putorius Meles taxus nou Cait, Pleistocene, | Pleistocene, re nsorie. iislaaaa, Noa |) Hampaway Gamba Gedeurer ‘ 48920 (Brit. Mus.). (Brit. Mus.). Mus., Cambs.). | | 4 . i. Ravivs. : ne j MP) ile! Dye yo¥en dlc Wen menaowece sé apeei dpeerpaandae sqeoads 30 9:1 6°4. | : 2. Longer diameter at carpal articula- y GLOMS 7: Sees een ee Cer CEC ERR Renova 6 19 1°35 He BA : our ee . se 5) Se eee Mustela robusta 5 Mustela vulgaris deles tarus, Pleistocene, : cee aa Pleistocene, f Pleistocene, Ightham, nr. TBaeasara N A Ightham, nr. Happaway, No. Maidstone (Lewis 48897 (Brit arene ) Maidstone M. 5808 (Brit. Abbott Coll.). 7 = uS+/-| (Corner Coll.). Mus.). Una I Bf saYeq nl llaagrepntcsh aesane ee namaaneidescee oda 5:05 3°95 17 10:05 (4) Tapiys or Comparative Murasuremenrs or BonEs or Posterior Lime. | Mustela robusta, Hustelal putonius: Mustela valde Meles tarus, Pleistocene, Pleistocene, Pleistocene, Pleistocene, Ightham, nr. Brixham, No Ighiham, ur Happaway, No. Maidstone (Corner 48921 (Brit. Mine, raaalciion a M. 5808 (Brit, (Clo) (Corner Coll.) MER. i FEMUR. i Tl, IDG TAYE M, nonodpno 200009 o00nGG90A0oNNGODHO 6:4 47 2°05 12°3 q 2. Longer (right to left) diameter Ph Or yNES), Snnconnocccos0sne00900c0000n¢ 13 1-0 4, 27 ii i | i a ee ee “2 le” seed ral. | he ” MUSTELIDA. 25 TABLE OF COMPARATIVE MEASUREMENTS—continued. | Mustela robusta, | | Mustela vulgaris Meles taxus Se os Pleistocene, > ate pu as Wus, Pleistocene, Pleistocene, piece culgant D Ightham, nr, Boca No Ightham, nr. Happaway, No. Bene itn Maidstone 48922 (Brit, Mus.) Maidstone (Lewis M. 4933 (Brit. | (Sedewick Mus.) (Corner Coll.). = peter PALO ODULOOLI =)» Mus.). 5 igre TIBIA eMenothin ance ysccien: eects caeesl | 67 475 1°85 10°95 | 10°4 2. Right to left diameter at proxi- na UNE TA Cire mesh set aw scmieceacodeteanle suse 1-25 5) *35 29 2°25 | | Ce lade Mustela putorius, | Lutra vulgaris, I hers aD Pleistocene, Prehistoric, MeO GieaaS Brixham (Brit. Burwell Fen mennats Coll Nagel = Mus.). (Sedgwick Mus.). | i} | FIBULA | | | | JURTINGA G1 va SesaaeisctareeGs (yon aaa eae aa AN EEN ee eee | 52 4:65 9-4. | IV. CONCLUSIONS. The present memoir has afforded little scope for critical treatment and is almost purely descriptive in character. No novel conclusions have been reached. Of the eight species of Mustelide described, only the marten, glutton and badger appear in British Pliocene deposits, and only Mustela robusta and the elutton are no longer found in Britain. No Musteline remains are recorded from Scottish Pleistocene deposits, and only the marten, stoat, badger, and otter from those of Ireland. Sincere thanks are tendered to Mr. W. J. Lewis Abbott, Dr. F. Corner, and the Rey. H. H. Mullins for kindly placing their large collections of Musteline bones from Ightham and Langwith at my disposal for examination and figuring. J am further indebted to Mr. H. St. George Gray, Mr. A. Newstead, Professor T. McKenny Hughes, Dr. R. F. Scharff and Dr. A. Smith Woodward for the loan of specimens preserved respectively in the Taunton Museum, Sedgwick Museum (Cambridge), Grosvenor Museum (Chester), the National Museum of Ireland and the British Museum. I further wish to thank Dr. C. W. Andrews, Mr. Oldfield Thomas and Mr. H. Woods for help and information, and Mr. J. Green for the care he has taken with the illustrations. 26 1810. 1812. 1818. 1822— PLEISTOCENE MAMMALIA. VI. BIBLIOGRAPHY. G. A. Goldfuss, ‘ Die Umgebungen von Muggendorf.’ G. Cuvier, ‘ Recherches sur les Ossemens fossiles.’ G. A. Goldfuss, “ Beschreibung eimes fossilen Vielfrass-Schidels aus den Gailenreuther Hohle,” ‘ Nova Acta Acad. Caes. Leop.,’ ix, p. 313. 23. J. F. Krueger, ‘Geschichte der Urwelt,’ p. 851. . . . Quedlinburg and Leipzig. 1823. G. A. Goldfuss, ‘Siugethiere der Vorwelt.’ 1823. 1824. 1825. 1528. 1829— 1832. 1853. 1834. 1836. 1839. 1839. 1842. 1844. 1846. 1847. 1847. 1866. 1868. G. Cuvier, ‘ Recherches sur les Ossemens fossiles,’ ed. 2, iv. W. Buckland, ‘ Reliquiz Diluviane,’ p. 15. G. Cuvier, ‘ Recherches sur les Ossemens fossiles,’ ed. 3, iv. J. B. Croizet and A. C. G. Jobert, ‘ Recherches sur les Ossemens fossiles du Département du Puy-de-Déme,’ p. 89. 31. F. Holl, ‘ Handbuch der Petrefactenkunde.’ H. von Meyer, ‘ Palzologica,’ p. 47. P. C. Schmerling, ‘Recherches sur les Ossemens fossiles—Cavernes de Liége.’ G. Cuvier, ‘ Recherches sur les Ossemens fossiles,’ ed. 4, vil, pp. 484—516. G. von Minster, ‘ Verzeichniss der Versteimerungen . . . zu Baireuth,’ 1s SM. J.C. Bellamy, ‘ Natural History of South Devon,’ pp. 86—101. M. de Serres, J. M. Dubrueil, and B. Jeanjean, ‘ Recherches sur les Ossemens humatiles des Cavernes de Lunel Viel,’ p. 67. R. Owen, “‘ Report on the British Fossil Mammalia,” a Brit. Assoc.,’ Manchester, pp. 70—72. H. M. D. de Blainville, ‘ Ostéographie,’ fase. 4. R. Owen, ‘ A History of British Fossil Mammals and Bir dye pp. 109—122. C. G. Giebel, ‘ Fauna der Vorwelt,’ 1, pp. 55—64. A. Nordmann, ‘ Découverte de Gites riches en Ossemens fossiles faite en 1846 a Odessa.’ . F. J. Pictet, ‘ Traité de Paléontologie,’ 1, pp. 214—220. . P. Gervais, ‘ Zoologie et Paléontologie Frangaises,’ pp. 243—253. . H. von Meyer, “Meles vulgaris aus dem diluvialen Charen-Kalke bei Weimar,” ‘ Palzeontographica,’ vu, pp. 41—49. W. Boyd Dawkins and W. A. Sanford, “ British Pleistocene Mammalia,” PIER, SOGay ty JO]V> IM, SOR, H. Wankel, ‘‘ Die Slouper Héhle und ihre Vorzeit,” ‘Denkschr. k. Akad. Wags Wiens: Scxyaili wt 2 aps: ~MUSTELIDA. 27 1869. W. Boyd Dawkins, “On the Distribution of the British Post-glacial Mammals,” ‘ Quart. Journ. Geol. Soc.,’ xxv, p. 192. 1870. EH. Cornalia, “Monographie des Mammiféres fossiles de Lombardie,” ‘ Pal. Lomb.,’ ed. Stoppani, ser. 2, pp. 20—31, pl. xi. 1871. W. Boyd Dawkins, “On the Discovery of the Glutton in Britain,” ‘ Quart. Journ. Geol. Soc.,’ xxvu, p. 406. 1871. W. Pengelly, ‘Trans. Devon Assoc.,’ iv, pp. 98, 102. 1875. G. Busk, “List of the Mammalian Remains ... in the Rock Fissure Cavern in Creswell Crags, Derbyshire,” ‘ Quart. Journ. Geol. Soc.,’ xxx1, p- 685. 1878. A. Nehring, “Die quaterniren Faunen von Thiede und Westregeln nebst Spuren des vorgeschichtlicher Menschen,” ‘ Archiv fiir Anthropologie,’ Ke [Od Ok 1879. HE. R. Alston, “On the Specific Identity of the British Martens,’ ‘ Proc. Zool. Soc.,’ pp. 468—474; and ‘ Zoologist,’ 11, pp. 441—448. 1879. T. Liebe, “ Die fossile Fauna der Héhle Vypustek in Mihren,” ‘Sitzb. k. Akad. Wiss. Wien.,’ Ixxix, pt. 1, p. 472. 1880, 1882, and 1884. A. Woldrich, ‘ Diluviale Fauna von Zuzlawitz bei Winter- berg im Béhmerwald,” ibid., 1xxxu (1880), p. 32; Ixxxiv (1882), p. 194; and Ixxxvi (1884), p. 993. 1881. A. L. Adams, “‘ Report on the Animal Remains of the Bone Cave of Ballynamintra, co. Waterford,” ‘Trans. Roy. Dublin Soc.,’ (2) 1, p. 208. 1881. R. F. Hensel, ‘“‘Craniologische Studien,” ‘Nova Acta Acad. Caes. Leop.,’ xli, p. 125. 1882. H. T. Newton, “ Vertebrata of the Forest Bed Series of Norfolk,” ‘Mem. Py Geolssurve,. pa 20: 1885. R. Lydekker, ‘Catalogue of the Fossil Mammalia in the British Museum,’ i, pp. 176—191. 1885. EH. Wilson, “ The Bone Cave or Fissure of Durdham Down,” ‘ Proc. Bristol Nat. Soc.,’ N.s., v, p. 44. 1886. H. Hicks, “On the Ffynnon Beuno and Cae Gwyn Caves,” ‘Geol. Mag.,’ (3) i, p. 566. 1886. F. Winterfeld, ‘Ueber quartire Mustelidenreste Deutschlands.’ Berlin. 1890. H. T. Newton, “On some New Mammals from the Red and Norwich Crags,” ‘Quart. Journ. Geol. Soc.,’ xlvi, p. 444. 1890. A. Smith Woodward and C. Davies Sherborn, ‘A Catalogue of British Fossil Vertebrata.’ London. 1891. H. T. Newton, “Vertebrata of the Pliocene Deposits of Britain,” ‘Mem. Geol. Sury.,’ p. 10. 28 1894. ISOM 1899. 1900. 1903. 1906. 1906. TOMO: PLEISTOCENE MAMMALIA. H. T. Newton, “The Vertebrate Fauna of the Ightham Fissure,” ‘ Quart. Journ. Geol. Soce.,’ 1, p. 200. . M. Boule and G. Chauvet, “Sur l)’Existence d’une Faune d’Animaux arctiques dans la Charente 4 ’ Epoque quaternaire,” ‘Comptes Rendus,’ Grocynils [Os Iekerss EK. T. Newton, “‘ Additional Notes on the Vertebrate Fauna of the Rock Fissure at Ightham (Kent),” ‘Quart. Journ. Geol. Soe.,’ lv, p. 425. H. Wilson and 8. H. Reynolds, “ Uphill Bone Caves,” ‘ Proc. Bristol Nat. SOGo, MeSky 1, In 152), R. F. Scharff, “ Exploration of the Caves of Kesh, co. Shgo (Mammals except Man),” ‘Trans. Roy. Irish Acad.,’ xxxii, B, pt. 4, p. 202—205. K. F. Hamy, “Le Gulo borealis dans la Grotte de la Grande Chambre a Rinxent, Pas de Calais,” ‘ Bull. Mus. Hist. nat. Paris,’ xu, p. 137. R. F. Scharff, ‘‘ Exploration of the Caves of co. Clare (Animal remains except Birds),” ‘Trans. Roy. Irish Acad.,’ xxxili, B, pt. 1, p. 205. J. W. Jackson, “ On the Vertebrate Fauna found in the Cave-earth at Dog Holes, Warton Crag,” ‘ Lancashire Naturalist,’ Feb., 1910, p. 330. eS ae C=“ 1 a ey te ee PLATE I. PieistoceNE Mustecipa. “Cramum and Mandible. Polecat (Mustela putorius), and Giant Polecat (Mustela robusta). (Natural size.) Mustela putorius. Fig. 1. Dorsal \view of a ? Prehistoric cranium from Ightham, near Maidstone | 2. Ventral) (Corner Coll.). 3. Lateral view of a cranium from Brixham (Brit. Mus.). 4. Palatal view of the corresponding mandible (Brit. Mus.). 5. Outer : : = aspect of the same mandible. | 6. Inner Mustela vobusta. . Dorsal ; + oi [view of a cranium from I¢htham, near Maidstone (Lewis Abbott . WVentra Coll. | 9. Lateral J on | 10. Palatal view of the corresponding mandible (Lewis Abbott Coll.). ak tex | et *Jaspeet of the same mandible. 12. Inner} a. Occipital condyle . b. Auditory bulla | d. Post-orbital process of frontal ;Plates I to VI. | e. Infra-orbital foramen j. Mandibular condyle PALAZONTOGRAPHICAL SOCIETY. IQI]. Reynolds, Pleistocene Mustelidas. Pl. I. J.Green del. lith.et imp. POLECAT:MUSTELA PUTORIUS. & GIANT POLECAT: M.ROBUSTA. Cranium & mandible. Fic. tS aie Go NI SP fea S PLATE II. PLEISTOCENE MUSTELIDA. Cranium and Mandible. Marten (Mustela martes), Stoat (Mustela erminea), and Weasel (Mustela vulgaris). (All except fig. 12 a natural size.) Mustela martes. Dorsal ) . k ; view of a ? Prehistoric cranium from the Edenvale Cave, co. Clare Ventral eet (National Mus., Dublin). Lateral Inner) view of a left mandibular ramus from the Pleistocene of the Langwith Outer) Cave, near Mansfield (Mullins Coll.). Mustela erminea. Dorsal Latera Outer aspect of the right ramus of the corresponding mandible (Brit. Mus.), Ventral view of an imperfect cranium from the Pleistocene of Kent’s Hole, Torquay (Brit. Mus.). ipview of a cranium from the Pleistocene of Berry Head (Brit. Mus.). Mustela vulgaris. Dorsal ) view of a cranium from the Pleistocene of the Ightham Fissure, near tec Maidstone (Lewis Abbott Coll.). Left mandibular ramus seen from the outer side, from the Pleistocene of the Brixham Cave (Brit. Mus.). The same two and a half times natural size. Lettering as in Plate I. PALZONTOGRAPHICAL SOCIETY. 1911. Reynolds, Pleistocene Mustelide. PI. IL. pm2. pred M2. J.Green del.lith.et imp. MARTEN: MUSTELA MARTES, STOAT: M.ERMINEA, & WEASEL: M.VULGARIS. Cranium & mandible. Je iGuadEde JULI, PLEIstoceENE Musraipa. Cranium and Mandible. Weasel (Mustela vulgaris, var. minuta), and Badger (Meles tavus). (Natural size except figs. 2a and 4.) Mustela vulgaris (var. ivinuta). Fig. ee Dorsal ee : 9 Bos is view of a cranium from the Pleistocene of Ightham, near Maidstone Sal (Corner Coll.). 2a. Anterior part of fie. 2 two and a half times natural size. 4. Associated right mandibular ramus seen from the inner side (Corner Coll.). 4a. The same nearly three times natural size. Meles tavus. x Left mandibular ramus of a young individual seen from the inner side. Left mandibular ramus of a young individual seen from the outer side. Both the above specimens are from the Pleistocene of the Happaway Cave; 5 1s preserved in Dr. F. Corner’s Collection; 6 in the British Museum. 7. Left side view)of a cranium and mandible from the Langwith Cave, Mans- Anterior a. field (Mullins Coll.). : 9. Inner view of a left mandibular ramus from the same locality (Corner Coll.). or) ge Lettering as in Plate I. PALA ONTOGRAPHICAL SOCIETY. IQI1. Reynolds, Pleistocene Mustelide. PL UL 4a. x3 pree pm 4 ml i d.Green del. lith. et imp. ) WEASEL:MUSTELA VULGARIS (VAR.MINUTA), € BADGER: MELES TAXUS. Cranium & mandible. PLATE IV. i . PLEIstocENE MusTELIDz. Oraniwm. Badger (Meles taeus). (Natural size.) Fie. veDorsal eine ¥ ME AOE eet se view of a cranium from the Pleistocene of Grovehurst 2. Ventral (Brit. Mus.) . a 3. Lateral eri” a Lettering as in Plate I. J.Green del. lith.et imp. oO > = Lu 5 ‘ wn e) 5) e a g O (ep) — Ww 2 x= aie z zat oc Ae © oc fe) LJ kK oO Z < Oo m Ss Sie ey a 8 a 5 istel at Emme > Reynolds, Pleistox OUEST TE. JUNIE Piuistocens Musrunma. Cranium. Badger (Meles taxus), var. (Natural size.) Fic. ie Dorsale a. k ; i: ae view of a remarkably elongated cranium from the Pleistocene of the 2. Ventral 3 . ; Langwith Cave, near Mansfield (Corner Coll.). 3. Lateral Lettering as in Plate I. PALA ONTOGRAPHICAL SOCIETY.IQII. Reynolds, Pleistocene Mustelide. Sy ve ay: BADGER: MELES TAXUS. (vaAR.) Cranium. d.Green del -lith et Pil, VW, imp. | | | Fie. oe to IPN) WIE. Piuistocene Musrsipa. ‘ Cranium and Mandible. i a a ae eb Otter (Lutra vulgaris). ae FN i: (Natural size.) rhs Dorsal ae of a cranium from the Prehistoric ery of Ventral (Sedgwick 1 Mus., Cambridge). oe The same cranium with the associated mandible seen from the Inner view of the left ramus of the above mandible. Lettering as in Plate I. PALZONTOGRAPHICAL SOCIETY.IQII. Reynolds Pleistocene Mustelidee. Pl. WIE i ras) Sinclair yeas d.Green del.lith.et imp. OTTER: LUTRA VULGARIS. Cranium & mandible ee cue be. 1 r é ie 1 ) y PLATE VII. Puierstocenr MustrEe1ips. Vertebre. Badger (Meles tawvus). (Natural size.) Fie. 1. Atlas, dorsal view | 2. Axis, left side view From the British Museum Collection. 3. Third cervical, posterior view E ae be eae From the collection of Dr. F. Corner, F.G.S., 6. Fifth cervical, posterior view BseOlen. 7. First thoracic, left side view 8. Dorso-anterior view of the same bone | Ou Mitta thoracic. deb meron cw From the British Museum Collection. ‘10. Dorso-anterior view of the same bone | 11. Thirteenth thoracic, posterior eee the collection of Dr. F. Corner, 12. Fifteenth thoracic, left side view ! F.G.S. 13. First lumbar, left side view 14. ‘Third lumbar, posterior view TeOmeeeoee aes 15. The same bone seen from above rom the British Museum Collection. 16. Sacrum, ventral view All these vertebre are from the Pleistocene of the Happaway Cave, Devon. a. Neural spine b. Neural canal c. Pre-zygapophysis | dai Giost-7ygapopliys!s —'\ miares aysllitenaimraaa | e. Vertebrarterial canal | jf. ‘Transverse process g. Anapophysis h. Metapophysis PALA ONTOGRAPHICAL SOCIETY.IQII. Reynolds, Pleistocene Mustelidee. J.Green del. lith.et ump. : MELES TAXUS. Vertebre. BADGER =~ eh ee Le | PLATE Va PretsroceNns Musveripa. Vertebre. Polecat (Mustela putorius), and Otter (Lutira vulgaris). (Natural size.) Mustela putoris. Fic. 1. Atlas, dorsal view 2. Axis, left side view 3. Fourth cervical, left side view} All from the Pleistocene of the Brixham Cave, 4. The same bone, anterior view| and preserved in the British Museum. 5. Sacrum, dorsal view 6. Sixth lumbar, dorsal view Lutra vulgaris. 7. Atlas, ventral view. 8. Axis, seen from the right side. 9. Third cervical, posterior view. 10. Fifth cervical, anterior view. 11. Sixth cervical, seen from the left side. 12. Seventh cervical, anterior view. 13. First thoracic, anterior view. 14. Ninth thoracic, seen from the left side. 15. ‘Twelfth thoracic, posterior view. 16. ‘Thirteenth thoracic, seen from the left side. 17. Fourth lumbar, posterior view. 18. Fourth lumbar, seen from the right side. 19. Sacrum, dorsal view. All the Otter vertebra figured are from the Prehistoric peat of Burwell Fen, and are preserved in the Sedgwick Museum, Cambridge. Lettering as in Plate VII. ————— PALZONTOGRAPHICAL SOCIETY. /I9II. Reynolds, Pleistocene Mustelide. 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