HENRY FAIRFIELD OSBORN WorLp. DistrRiBUTION OF STEGODONTS, He LS Ple an E toc oO Pli ? -fathom Present coast l 100 ine y AE { COLOR iy) AMMONTINAE © Sh MM 9 TEGODONT/ oo 8 Archidisko NAE 0 Sfegolophodon Removed to the Mastodontiddé too SITEGOLOPHODONT/ don = 73 > o @ <2) x ee oe = = o = S 3 a. i ae, q)- < 5 OTHS, LOxODONTS, AND ELEPHANTS I35° 120. 105° 90W 75 60° iskodon ® Loxodonta @ Elephas as Ben h @ Paladeoloxodon @® Hlypselephas sk © Hesperoloxodon @® Platelephas ie DF.Levett Brad Et oO BOS CLD BX RESTORATION OF THE JEFFERSONIAN MAMMOTH (PARELEPHAS JEFFERSON!) After a painting by Charles R. Knight in 1909, under the direction of Henry Fairfield Osborn The skeleton on which this painting was based is mounted in the American Museum of Natural History, Hall of the Age of Man. It was found near Jonesboro, Indiana, on the farm of Dora E. Gift, in 1903; purchased for the American Museum with the Jesup Fund in 1904; mounted in 1906; first described and figured by the present author in 1907. As found the skeleton was embedded in a muck deposit of late Pleistocene age, fifteen feet below the surface. This deposit is probably of post(?)-Wisconsin age (according to the geologic time scale having been laid down about 15,000 years ago). Representatives of the Parelephas phylum appeared in Europe in the early Pleistocene and persisted into the Third Interglacial. This Third Interglacial period may mark the time of migration across Asia into North America. In fact, it is suggested by the present author that such migration might have occurred in Second or even in First Interglacial time. The most striking features of this individual are the complete incurvation and crossing of the tusks, indicating that it 7s an old bull, and the relatively small size of the head. It is here represented with a hairy covering, as Parele- phas vs characteristic of the north temperate region, both of Europe and the United States. PROBOSCIDEA A MONOGRAPH OF THE DISCOVERY, EVOLUTION, MIGRATION AND EXTINCTION OF THE MASTODONTS AND ELEPHANTS OF THE WORLD BY HENRY FAIRFIELD OSBORN A.B. Princeton, 1877; D.Sc. Princeton, 1880; Honorary LL.D. Trinity, 1901; LL.D. Princeton, 1902; Sc.D. Camsripceg, 1904; LL.D. Cotumsia, 1907; Px.D. Curistranta, 1911; D.Sc. YAtz, 1923; D.Sc. Oxrorp, 1926; D.Sc. New York, 1927; LL.D. Union, 1928; Docror OF THE UNIversity oF Paris, 1931; Doctor or NATURAL SCIENCE, JoHANN Wo trcGanc GortHe University, 1934 REsEARCH Proressor OF ZOOLOGY, CotumsiA Universiry; Honorary Curator-In-Cuizr OF VERTEBRATE PALAEONTOLOGY, THE AMERICAN Museum or Naturat History; SENIOR PALA}ONTOLOGIST, Untrep States GEoLocicaL Survey; Honorary Presipent, THE AMERICAN Museum of Naturat History; Honorary PRresipDent, Tue New York ZOOLoGIcAL Society EDITED BY MABEL RICE PERCY VOLUME II STEGODONTOIDEA ELEPHANTOIDEA pp S i: AY VS Was Nog e\poR Ves ¢c a Naf s\\RC ‘el INCORPORATED z ) Xu :) 1S cA SN G \ as S PUBLISHED ON THE J. PIERPONT MORGAN FUND BY THE TRUSTEES OF THE AMERICAN MUSEUM OF NATURAL HISTORY THE AMERICAN MUSEUM PRESS NEW YORK, 1942 PUBLICATION NOTE Volume I of this work was issued August 15, 1936. The present volume, containing the Stegodontoidea and Elephantoidea as well as tables, conclusions, and general index, has been prepared from the materials left by the late author (see Publication Note to Volume I). Copyricut, 1942, By Tue AmpricAN Museum or Naturat History CONTENTS PAGE AT Sola ©) Em ITEN UUs pL Fe AWTS OINISH eqecetecreiece tec exetene each cresusy ste cebsucscucyorawete clas a ney PPtaSU exe Royce Voy SCENE MAUL PU(oRe CETL EIEREVoEMC ACN eae oni VPS eat XV CHAPTER XLV. THE ROOF-TOOTHED STEGODONTS; SUPERFAMILY STEGODONTOIDEA.................:.......--. 805 Classificationsom the:zeneray Stegolophodon and) Stegodoname ae een neon ace eee ae ase 807 History of discovery of the subfamily Stegodontine. Principles of type revision of species.................. 815 itheistez odontinssiandelMastodontins ofl Ching sean ene race een ho eerste eee ae ee errr 816 Pliocene to) PlerstoceneyeroboscidesonJapanen ieee uence een eee nearer ice eeaaee aaa aen ace 818 Phylogenetic discussion of the thirty described species of Stegodonts and Stegolophodonts................... 819 Probable European—Asiatic origin and migration of the primitive Stegodonts..........................0.. 822 Type revision of the species in order of original discovery and description.............. Bs eis ne ae ee 822 Birstitworstezodontsdiscovereduneburmar 928) pees er eet en ele eae 825 Mastodonlatidens: Clitt.1828) |=Stegolophodenlatidens| een nas sateen ae aera eee 827 Mastodon elephantoides Clift, 1828 [= Stegodon elephantoides]............ 6-060 c ccs 828 Discoveriessinclndiay and (Urmay e ieee eles occ sitcocs eet RO eee en SE ee ee eet ota ci cay peor 829 Stegodonts of China, India, Java, the Philippine Islands, Austria, Japan, and Burma......................-. 831 @haractersiofethe subfamily; Sterodontinse yee aemeer eee cor erie riers retiree iterate ree 837 Stegolophodorschlesingery 1917,, zeneric definitions a--eee eile ere tae eter reve ern 839 TDI ia Ke (eq yn) bots eines iene Den an Ane On nO Om aa ae ew OOD ModuDHaS prow ep oo. cciccc 840 itis Page Oita cP: ane ne eee et eee aren men So aloo ino Maka wannmOn ne cm ab. icra woteowiog doc hal 842 Sublatidens Schlesinger) 1 ON2 sraverse «can arene ica: Mme eaves tee Roe eee er tae ear vet nen te 846 APP edies Weikaatiny, WONG ooo oacaccpaasosaeagengcsepaebanosooudonageocedo0d saoddosodeccsujacue 846 UO INOTAR OR ody ICP Vhs pemleooonddsocncnsdo conn oocK DOH oORoDeDeuAaUHOUnOUOS ondoobavcucdoasuds 847 CAULEY sepnOgnessitss OSHOKNe1 O29 hem taet ree Ieee ere eee eae e 848 decker Osborn 193Glc ery. see ees ee he Oe ers inc eae eee roe ee 851 Stegodon Falconer and Cautley, 1847, 1857, generic definition. ...... 20... 2.6... eee eee eee eee eee eee 853 PPS Oy, col e(DE oman seb weld oboe Acad eadaooSpecccondodbancpopauob somoboomeDocHougmsonHen Ge 860 elephanioides Clift VI828) 22 «ai iar wes ions ote ie eels vate PYF eS oie eee Sra 861 bombéfrons Falconer’and Cautley, 1846... 22.027 see rie oe ee ee ie 863 insignis Falconer and Cautley, 1845, 1846................... Ppa corp aacepmoma a oesantomance> ss: 866 ganesa. Falconer and Cautley, 1845, 1846... ..... 5.2... cee cece ce nent eee eee ee cetera sees 869 UDSULOSE See Gag Oe Pose One HAD Hoops tcioob cuabooDDn DHUe GEC UDOOObOD Senco A boCKdEGEDbD DOOD Ic 874 msignis birmanicus Osborn, 1929... . 2.2.2... e ice nce ee cre ne es Hone ote ne eee cneneewnee 874 orientalis grangert Osborn, 1929)... 6.2 0... 5.0. oun wise sous «2 Sere Seed sage es ree shee fesies peo) elena 875 mingorensis Osborn, 1929... ie. 5 ie ie nse sine a eae eee nee See Ne Ae Nc eRe edhe ee 883 orientalis’ Owen 1S70ls. os. neste Sere es nes eee nese tee eee Con ete oe 884 airdwana Martin, 1890). 2 2..0 see ects oa riers ote aioe eee ern ao ce 885 ganesa var. javanicus Dubois, 1908. ... ...... 2. 1. e eee een een ent ete scent anes 889 trigonocephalus Martin, 1887... 2.0... 6.0.06 cece cece cee ne ce ee ne een ee eee nee ee eee tsetse se tee 890 mindanensis: Naumann, 1890); . commen ae cis eme tts tec ict yeti ee etre stat ee ekanet vers ae enter 892 qQurore Matsumoto, 1905; VOUS sae ee scrcuetenel aera tater elt ee eee een ee Eee el ee 892 orientalis shodoénsis Matsumoto; 19245 2. fees ote ere cles = ect rete et alee eee ely eet ocho t= chek b= elon 893 bondolensis van der Maarels 1932). 5.02 o.oo tee eres oe mem eee ele ete = tied dele tensed elo ete lelielitcl =r 894 trigonocephalus praecursor von Koenigswald, 19383........... 2... 6s esse eee e ceteris 896 (Parastegodon?) kwantoensis Tokunaga, 1934... . .... 2.0.6. eee cette eee eee eens 897 ylishensis Young, 1935). 05. 15 o.com epee ee settee ei eee toe ete ee 897 officinalis Hopwood, 1935... < . 2. sce ce aan see eee ole ia ne ele ele ele ese ae oles ota ete 898 zdanskyt Hopwood, L935... 2. (ele ea a overload ce es fcrns svete tet eet evry) -V to el pened lst tae orc = ee 899 (Parastegodon) sugiyamai Tokunaga, 1935... .......-.-.- 12. eee ee eee eee teens 899 Matsumoto on the phylogeny and classification of the Japanese Mastodonts, Stegodonts, and Elephants... ...901 Osborn’s comments (1929) on Matsumoto’s phylogeny and classification of 1924-1927. ....-.............55- 908 XV. CLASSIFICATION OF THE ELEPHANTOIDEA BY THEIR DIVERGENT AND HARMONIC CRANIAL ANID DEN TAY CHARACTD RS i secant cecil ce asec reesei ie ar eee ee othe ee 911 Elephantoidea Osborn, 1921, superfamily definition. ...........-- 0. ++ 022s este eee eee e seen tenes 912 Elephantide Gray, 1821, family definition. ............ 2... 2-2... een n eee rete eee rece n scree 912 i OSBORN: THE PROBOSCIDEA APTER xv—Continued PAGE Hailunelofsprevioussdentallicl asstticeitlo ns spt sp onstrate neh eee tate tee eet ee aoe cee eo eee 914 @lasciticationspyrcranialian dycentellech ana cers meget itty cktonee te cre oie eye teste eters ete ele eee 914 @ranialsmechanicsof Hlephas (Weithoter| Osborm, Grerory,) ia ee yeteroeer tiie ee 915 Comparative craniallsectionsiofrelep ham) SkcUI IS tregetereitee treet tetera ete eee ea ete ere 918 Ontozenetie cranialuchanges une Hep has yin G7 Cis erate ear Tore ete ele ea eee ee 919 Dental and cranial adaptation to prevailing feeding habits the key to phylogenetic classification. ............ 927 Ridge-plate formule of primitive and progressive genera in adaptation to prevailing habits of feeding... .. 927 Food of the Indian and African elephants and of the mammoth...........................22-.2+.--05 927 Seps@anll Goanrgasiin rorecl Oi wee wa. oo dab sacs vena aon HaoadeBD Abd FOoANSadosudnwousu ose gadoS 929 Summary of progression from browsing to grazing dentition...................----- esse eee eee eee 929 Vertebral distinctions of Elephas, Lorodonta, Mammonteus, and Parelephas...............5...000 00 cee e eee 930 Vertebral formless. oaks ocass come 07 aie ong hiccne comeing series s os. exctsiay uatecte ire pees agnor Rt Rear Ce ees cua 930 Synopsis of subfamily classification of the Elephantoidea.....-... 2.2.2.6... eee e eset see eerste ee 932 XVI. THE GENUS ARCHIDISKODON (SUPERFAMILY ELEPHANTOIDEA), MOST PRIMITIVE MEMBER OF ASHE) SUiIBIAIVIDIGY: IMUAIMIMO NGMINAR. Cn oes ec a a neieieye coe cease cet See erent tre pearance bree 935 SHOAy Ot WA sWlotenthis WEVA ONO. oo coodooneceseosoo poo eougG Heo dso eo soa sun seg seauaDSoo2a0d0DS: 935 Manmmontine Osborn 1921 ssubtamnilive de timiti ome ery-e reteset cte eters rete tel tte ene eee et eet eae el etel ene te en eet 937 TEINS? ONL Wa xe AAMT ooc6 oo co bacda ros ananeoohamoneosoonondoNoDoDDDONO UO UUS OSA HATOEE 939 Order of discovery and description of twenty-two species of Archidiskodonts........................--2005. 942 Archidiskodonts of Murasia.andeAmenican naa. ciae se ise eine alot iis etree ere ie ec ieee 943 INewaAnchidiskodonts and sluoxodOntslonmAtricayeseerter vere casi neseaees rei perrer eter meet eerste ease ret ea 944 Approximate phylogenetic order of succession of species of Archidiskodon and Parelephas (1928).............. 946 Ag. chadiskodor ohligal SSo nl ssonceneric) Gelinicl Olean eee a cee tee ratte reer ire rele Oe ner iats 947 planunonsualconerands Cautleyeel S460n( S40 | pert ysteieie aera eke toca tee tetott tet e tee ete edt tere teeta Teter 950 Measurements of twenty-seven specimens collected by Barnum Brown in the Siwaliks.............. 954 Wevth=Adamsia svwalekienstsaias aoe eee eee es ne TE Er yao erate 959 DUCNUIRONSUMON USS LELANESC Up O 2A eet teyel ten Var were yer ver IIe cio eRe ert ene leer 968 OCH OL GS ING nal Piel ae ee og too On Ute pon GG Od fon SOR OMESaR GUO DB. Goob m>inodolonccodeoonoo omnes 969 Lyrodoril—Asvmendvonalis female]arpeerr ite clern ns stich stevia seek ere tere tte acre eet tegen 977 VD Yb Ss (0) a tags) oe (2) 10) 0 phe Pe PO rs 10nd nicotene oc UOe ubio M0'6u0 Cc 977 MeETaLonalisrenomenensis»Deperetyands Mayet L923 qe cis sass se secre ieee etter shel tenia rare yeee 980 Archidiskodonts and Metarchidiskodonts of South Africa.................-- eee ee ee ee eee tees 983 Archidiskodon (continued) jinoullmajodes Osler, WBE 6 > omnsandoadsoasuus juecaccagoepoddbouEuuUoOddacupdooDonU DoD OUnODSD COSC 986 coli ihnaans Oslo, IWS. oo aduonsmesouadods aoouussoqbo dD GnG OD MOOD OHOCHOOU SCO UUd OD DdZBONDDD ORS 987 RECTOR One, PISA oe hans armenn onand enue sb dec du GaOdmoOOmmne ots E ame abEmooanin esos sod oegd bao 989 Regaine Dent, UEP; Game cose neue nae dcdan do soe J4sor PoGdigtoUn olor ptaucnp.s mondentosdodccon oc 990 main Dents ICP Me ceine cab ete a abnor cl soon benods hn saNsdeo toe aman cduns aOR toons sta ona é nob cole 991 LOZ OUGTILOLA CSE Dart Ail O2OUE we tes cant crated edbcteys nunrat RRA ge muter meme ER Sean hatetsestc Ureasiet ices MoE H cee ecelea uke eee ame 991 HO DEM, WO) eS abo a doen ner A Goes ooodg Non AA OG CORRE De deoIa aoc ocdtd ota omar Sadi oo. HbIsg cobb « 992 Mietarchidiskodoni@sborn.934.;eeneric definition.) sav ais ele ne ite rts es el trometer 994 ajptagiias lelnurd mutoiy IOP EE ane oe penbon enn CoM Onae bens cd nbs Koodo spy GoULoUsRoRuCoCG no boodaok bueete: 994 (Anchiciskodonts of theiUmited states anduvlexieor --a2 a see sy teeta: crete iste Sisters) ele =) erclcie ile evcle nisi iencels 996 Archidiskodon (continued) Gygratter IUCN ew, Cok a o oad ROMO pe UGS 428 anoneicoiny mp obcdeHaden boutroopodise sto durodsudstanone 998 Miyano ALCS TES aval ors IPP Sao wannesboobesdees sun euoon do bas 00ema cn colby acdsooeducUbeaT. 1015 imiantlan lle aie Vb euolnoore IPS Oo naboduecentv ees auoGuGoUoonuooO obocdnoccoBadnonocGn GaaKd 1016 TAL CO Map EMS Ci uaratop Ouaealofy-al YP a5 ao olnd toh dno nbuecovoudcGdon0oHooGueedgGocuBuHOUe 1017 moment: (skeletal characters) \0) 15 yccas vs. caste enero tals hirer velecke eae re Rarer rereelehe sterol 1019 payebarbour, GUS Riaiete at GUS h. Mee wean sttenh se cenarete otic ie et snohene te Be wissen Ranma aiaer eve eres ye cr 1023 Giananatt Keni, Iarqooybie, ICP). g do ann Cooma Dann Sdounod sop osc oo0 doo oodmadOnon Nd ncOnaas So0egr 1025 UP En ALOT MALDEN AL DOUT pi OQ bg. cc \yolcvo elo tens here er Adele oho) ia ake sh cael ee ere 1027 iin yas hig Cyl nk let en may dadveon foGecone accu aouc oro cmc sutmad aan 1029 alotsiayd erie binuiy cre ICP sese Goya motion aot ah eld omaenon oopacUOr coun. Ghomeducatiap so auac 0st SONOTLENSIS OSDOIN 1929). . sesso ssercloe cleic tence eiele sitiuerccetetenebere cake Ee shee) eer Ae ee Re Re ROOT aCe ere te 1033 METLOLOMAIISINEDT ASCETSIS! OSDOLD PL OG2 oo ere) e eels cio sversie 1s aie ietenetoletel Wee ore =) «nek onero ter rete tsi aaa es Taneteletar=neuecrnys 1033 CONTENTS ix CHAPTER PAGE XVII. THE GENUS PARELEPHAS (SUPERFAMILY ELEPHANTOIDEA), OF THE SUBFAMILY MAMMON- TINE, INTERMEDIATE BETWEEN ARCHIDISKODON AND MAMMONTHEUS, DISTRIBUT- ED IN THE NORTH TEMPERATE ZONE OF EURASIA AND NORTH AMERICA........... ..1039 European north temperate origin. History of separation from other extinct proboscideans.............. ..1039 Order of discovery and description of species of Parelephas..........0. 00sec cece eee ct tte eee 1047 Parelephas Osborn, 1924, generic definition... ........ 0... eee e cee eee tree eee tee etter teen e nee ee 1048 regoninenvonlest Ait andy, ol QUS yoniy. ts, S cele tela Sa vA ee and atete m ehe eo eke tate eno -P-Fe ett el syne 1055 trogonthentt, Rohligs 1885; 1888189 Ue yy aye jese anew ntti cts toi s © lel aleve ecole ieee tennfeted) = chs es cians seagate 1056 pogorsheri esti Bobligs (SOU) so. yes. ois nlovapn sprees dole ole cosye ciel sinseaa) eveveyeds = es Ucuay 1c ase o late heres eyes 1059 HMENTACHS Hal CONCI PL SH Camere elicits critter eae TTT neta eaten ete ea 1060 TOA VOM EI UDI 6500 cobboudoH ooddRod oe coos nHoSosooe dar soos ooboUcEAssdEnEDccsogauodsoe 1062 neon, IQ0Ys oscoueasodunvoceugudosoesadcuondocs sconce oundorouncRsouobaquoodQdaIeosDAaNK 1065 iINorthwand | GouthwAmericane species\OLemarelephase me erect oer tne iste eter tte ed seated arti et 1067 ARGO INE) ore JIGS) salen on BEBE AAO phoebe SSlclode (eb Ond panera cherin Isotope peu otb wold aor 1068 (@imississupprensiavHOster wlSi2eya cialis ss rete cere ecco t seers ore heel retces i-th ele eve 1070 @olumbianeMammoths(Barelephas\colwmbs)e ae ecrsecr tetera ke iter ee eet tener 1070 Columbipbal concer ml Soiap S033 SOSner meer ree a rieeie ie cee ie tae se ie ee 1071 texianus{=columbi] Owen, 1859, Blake, 1861, 1862...... 2.2.22... ee eect eee ee eee ee ee eee 1073 Cohen Collection (Phosphate Beds of South Carolina)...........-.--2 22sec eee reece tet e eee 1075 JAial Hara CEG, ave oct eG Saco eine ooo nen Goced unues coon abbonanmcodmeuccim acco WoodAclsso 1079 columbs felicia Wreudenberg, 1922 0.2.0.6. obec ise oe i ee megs te ens ee ae en en le ners olele 1082 columbi cayennensis Osborn, 1929... . 2.0... ee eee ete nt nett see ener estes teae 1083 HAAR ci Os 01014014) PP RAM GO OOM Rone ce Soon nod. cide uaopr onan sou SH eeo sm euaor a4 ea crid cic 1083 Elephas roosevelti [= Parelephas jeffersonii] Hay, 1922.......... 65-62 eee eet ett tere ene 1095 prognessus Osborn, 1924. . 2... <2. se ces commer tae sane ees como, ene ee ge ace cleo ens eta ceed 1097 mashingioni Osporn, 19239 0.0. 2 aso os oils erp se wheat arenes Veta 1101 CITA S De SCP ie ekoe Ben eaeD ae BEenmenne Sus aore nn o.c00ddd omen pron un aton OSD On AS oD ahem cuccs 1104 floridanus Osborn, 1920)... eee ere.s cio cue ele eee eyes ge 2 eee y= rie le ele le ele ef ale lero l a pie ede ea 1105 XVIII. THE GENUS MAMMONTEUS (SUPERFAMILY ELEPHANTOIDEA), OF THE SUBFAMILY MAM- MONTIN, THE TRUE NORTHERN WOOLLY MAMMOTH ...............--.0.-0- sees eee 1117 Sceleto Elephantino Tonn, 1695, confused with the mammoth, Blumenbachl9 9 eemeeree eer eer 1118 Breyne’s description (1735) of the Elephas primigenius of Siberia ys 5 ious ters gee ER a Reyer at devant Sekar 1119 Names successively applied to the mammoth... ........... 00. see eee eee eee tenet tet eee ees 1120 Native Siberian origin of the word Mammut...... 2.0... 60-1 see e eee eee eee teen eee eens 1124 iG dt (0) )) ARORA See oron Sn caaddager cco fbaumnaniocotosegunpoot dese Ss 1124 Hloworth (ES82)is-..ca cs = ote eocc btn stevens ctor ort otal Sete eaeketer thet ealer sete Reno lo een. a) eke irate eater Necro 1125 Mammonteus Camper, 1788, Osborn, 1924, generic definition... ......- 2... +22. esse eee eee teen ees 1126 External characters and feeding habits. .............. esse eee bette eee ee eee eee c eee liz Skeletal characters of Mammonteus primigenius... 0.0... 0.0 cece ete eee eee ete eee 1129 Historical order of naming of species of Mammonteus exclusive of species which are now known to belong tocParelephas trogonthentt, UG csi tro aya terry tao areas cele rE 1136 Aurignacian mammoth hunters of Moravia... .....--- 22... 025s sere enter ere eee sete eet ete e sete: 1139 Typical progressive Eurasiatic stages of Mammonteus.........-- +. +--+ 500 be veers resent eens tsetse es 1140 primigenius Blumenbach, 1799, 1803... . 25. -. +. 2. ee reenter ne eee eee ete enna 1141 Elephas odontotyrannus [= M. primigenius] Eichwald, 1835....... 0... 2 202200 s sere eee settee 1146 Primitive European stages of Mammonteus primigenius......-.-- 0-0-0 eee teense tenets cs 1149 primigenius leith-adamsi Poblig, 1888... ........ 2-0... secret ee ee eee tents ee nese 1150 primigenius hydruntinus Botti, 1891... .......--.. +2... eee e eee e ene eee nent ener e esses 1150 primigenius fraast Dietrich, 1912-202 eee oie einer ee eee eid otis asain 1152 primigenius astensis Depéret and Mayet, 1923...........--.. ++ sees eee eet ttc e tenet ees 1154 BorestiBed jor Cromerians faunsie oo. beseeie eels 2 crt se ieee yogi cane nee a 1155 American stages of Mammonteus. ... 22.00.2021 e cece eee eee e ee eet as ree see nmn recs ener eres sacs 1156 primigenius americanus DeKay, 1842.........------2 seen eee rene eet ee eet eee e neers ccs sceeee 1156 x OSBORN: THE PROBOSCIDEA CHAPTER XvilI—Continued PAGE prumigentvius compressus: OSHOIN, VOLE sie. clove os ore er hee ay eee atuar Yai NOT vn Or erate Ces Sara aietnte ee ere 1157 PEUMNAGENTUS ALASKENSTS) SPs TOW slau m cosets nic tsps cacr snes ote faut suckarshape «, eVeeusts lew ekabeacsons oem eis aii ak seae oeusasI sree 1159 ETGZENMAMMOth, OL SIDES Nac eereece tet oles nee oe cree eras sae TR TR ROE TA cide crave sit eee 1162 Summary of the discovery and natural history of the woolly mammoth......................0000 0000005 1163 XIX. THE GENERA LOXODONTA, PALAHOLOXODON, AND HESPEROLOXODON OF THE SUPERFAMILY IDJODIBEVAINA NOW IBVA, SLOPE VAI UNLNS IL ODOIDXOINIMONUAD)s 5 os 00 eunadagooo boos utsoncanadonaoosdcobaDSS 1171 @lassification and history, of discovery, of thesoxodontinses- errno aaa rae eines cere 1173 Difhculties of generic nomenclature. + sas. ceca Sorc HOO ne: eee meit es aon onus ciearercneceieieitie ite 1174 Order of discovery and description of the fifty-three type species of the extinct Loxodontine............... 1187 Moxodontins. Osborn lols asublamilyadenni tone eee reer eer een ener nents 1191 [pocodontoe Cuvier elo el S2(preenericrdennitionwemnnee are cere aee or cere rie cee ei see eo eece etre 1191 Order of description of eighteen living African species and subspecies. ..............00.00 see eee eee eee eee 1192 Loxodonta (continued) africana Blumenbach fA7 90s sac omptachinon uterus ol enenacortieiakcretorels oko rocnieks cee RIE La ceteest t xe OT eee 1197 africana) (?)cottony Bales: 19261929 aio, cieuctesderertynays aula cic ol ie neue sini ue ernie oko Siete ener 1202 cornaliae Arad as IB TO se acs ccavansyconcinr et olonies aides whens oreo osiete eiots asiGlais. & ake oun Sek Os cusses eM eeeleuc eave eee 1204 iRalzolorodon Matsumoto 241 generic) definition... 4+) os ose ae ree cierto aeteitaete 1207 mamadicus Halconer.andiCautleywl846, 1847, 265.2 .se mig sone acne cele eiees ase Oe erect 1211 Hesperolocodon) OSbomm 951-72 enenicidennitlon eerie ei tea ete oe eee velit etree 1217 aniiguus Waleonerand Cautleys 18471850 ac, 6 eco ccs 2 sen rd eee ells eokoe oo eee ee rier eee 1217 Wpnoriskeletonvofs: antiquws a trcysere oye ce =o tiee ee euauein wspiarsttnge Wisesuesteieus enslO rusbes seemed er ereasas ote eke 1222 Coadaptation of the vertebral column with the superior incisive tusks of the elephants..................... 1228 Hesperoloxodon (continued) antiquus nanus ACCOnciy 880 \aa 7 bos occ vie ic cesce se clio sna S)die ofa ten rsiasa ely e rues ciao eeeiael rete set eheedetee rE 1230 UNL GUUS PLAT ny NCHS) PACS MSO Talo, aye, cejaie et pe oxo overs Oe ale Otekan Poors aie RTOS aI mole ac ockeioe) eee 1231 antiquus ausonius Major, 1875, Verri, 1886, Depéret and Mayet, 1923........................ eee eee 1232 ONUGUUS GER MONICUS SUCLANESCU LODE cyeyere sate) feisheielaieeeieyera oe eet ctr otc eee eo Reece teria terrane 1233 Hesperoloxodon antiquus italicus of Southern Italy and H. antiquus germanicus of North Central Germany... .1238 ONUGUUSALICUSSOBDOMMs, LOB eye ore 6c. mycrvincs oe ieee; fe 0) tekers as 6 witeray suede Sieiste) uslead or elecsuetonrenckel ste Neree eee roa 1245 NUT UUSIG ERMANTCUS OAS UENO ono 3 oye) soe Siste Chains oO ator ele ie cary yetsl cies oP ohe (eee odeae heh meee oko em eter eae 1253 Extinct warredsspecies/ othe Mediterranean) Islands ci cere ese era acta sede teeter stre deren ert 1257 Palzoloxodon (continued) ANGLULETLSES MH AIC OMENS sl Sass 5 asco eet ysis sr ers ov sue WONG URES A Septal tarat aah akenameeen evens fe tel Syatet eaten otololbepoity hr oueeeie tenon een 1262 HL LLCOMENCMSUSK LOO (ire sare eos tex 21-1 seers. $s+) Gr ep allekn foksiah taped aes Monee atc adnibrto) uate reeeta olsen iotey tia Pen onete ey here cloraee eae 1263 CTE TEE Lao Enh IIGY Das dig On ang eeeeaamoe cinaceicar acca mcecro an odacceia \adansomdas Bch 5. 1265 lammanmonacmLorsy the Ma] Or SSB ei. csc art ce cs exoeiet tence evelan eve Sie ener ened aero enetaita eM icbeistorcithateyetotterehaatcte 1266 (puree nies ICE eo sounder aso RoMe Eame or Gorn ae Cou acne aac ow cod coo EDDA Tan CG HOOE Oa IY Mo10'0 C 1266 CRELLCUS IES AUC LOO (pte ec nteyre ee ccs (oa do at's aici vats coe edPepepele Neve enews Uaea AMON eee hays lets ea iane te ley fe whe) Nelnlebelenorecerens 1267 The dwarfed elephants of the Mediterranean Islands and the question of Pleistocene isthmuses (Vaufrey, 1929) , 1268 Ancestral istagesi ot Ralcolorodon Mo Atri Cai ajo: acc, hrc diols atatoneley Pek ake Piste ever ae elolDy ene) tated alia lovee) oie) eleyeke] = eeveuerane 1273 GilANLICUSPEOMEL ALSTOM coy. ceys co orgs ovals Seonsiss #e eT ROHR NO NO PR rect stole s lakh usenet ecole 1274 OLENSTS OME] sgl SOO pater atal. & ereve wie elolone oe re isc Sus HIRE Clean Ree HR LTR RSS NSN ENC Ie neePe Ketones neta Fel itera shots 1274 Ha ID Ghiuts Neal Ne aca ee er eee Metso che oct os c.do.oOoReo comet acq Aare Te AaCer come ary ac 1275 (ei limealip Cravol lbpyapes byited (hi felowvads) bby (oga | booaun ocuddoodo doa booed UoODeOsO DODOUOUNOUDDOONGoU nO DOC 1277 Palzoloxodon (continued) Aye Db Ta 4 eee pes oIgnS SECT eme do ro Orson COOme neo eS OO Cueb Roonc0 a HO cman omndc 1278 Oe ODE Wht PAU ees Gat HOC OPEST cols 70005000 Gdnm DU COUT OOOC OUT DONOR ADE EMO ODUCE 1279 Olean OE att (4) ees, 6 ate Neate PIC OR DCR MeteEs oS 75.5 5:c OG COA Yoo TODO OU OA SO COU aH GOMdUemOOt GO SOO 1280 iL ELCON Di hatha 0)" Eee aan ae eee ee MET at oi bre. apie CON Veo OT OnE DOOD UUDIOO GO co OIE 1280 Tn OF con 71 ie er Mneb rds Meenas nico soo ols Gdnn dono Oo ABU oocn Seem ome To oom 1281 archidiskodontordes: Haughton, 1932 secre sauce ie & oleerclerels) ate eae netahe teks ale Pele voneialere etchetey Merete retells teholoael stoi oiolote 1282 transvaalensis: Wart. VO2 (52. scsiercca osc Sassou ole oss ohaiessuhe tale olel eRe ae Rete RSet ter es Ret Roney NAREN Tee Mefol orale Pelee) sKekekeyeas 1284 C]ivay ny at 12a) DEY ad A) 4 Pe En RIS INL APIOREAT G.7.0.0, 6b ocd On TOR DCOGOD SoU do Gnu GORD ODD OCOD UOHK 1285 CONTENTS xi CHAPTER XIX—(Continued) PAGE Loxodonta (continued) ZULU SCO Ube OU deegevecs Naicg= 25 Heo alae at oycos tat ale pies aly oon, 5 salto ARE ee Peto a RT MODE Cicaee ra cv e eT aan ae alge SO 1286 GUO STEN DEW ete P Lapel ast mechs Be Accra CR AERA Re SR I eT A RIES BA Ae SR ea emo eric ete | 1287 ALTPICONGEV AT KOULLQUL OP ANb yl 92 Ohare rene ott Sa eet Een A eT SEAT Pt eat en ak 1287 SUDCNIGUCs ELAN OM COM LOS ia. cucu a aie ofa eis Siete (ae RGM PEI SC USHERS COIS IE TREE 1288 oxodontinesyoleJapantand +) ava, strce is so rin ohh ls Peete ee Ie ee ee 1289 Summary of Matsumoto’s final observations and theories of 1924 and 1929........................... 1290 Osborn’s summary (1930) of the observations of Makiyama (1924) and of Matsumoto (1924-1929)... ... 1292 Two Japanese subspecies described by Makiyama (1924)................. 000 e cece eee eeceecseeeeees 1293 Palzoloxodon:namadicusmaumannt Makiyama, 1924°,-. 0.5... 00.. ones oes ee ese ce eee ee 1295 paleolorodoninamadicusmnamanuNiakiyamas1 924 eee eee ania ene 1296 Hive Japanese loxodontines described by Matsumoto. «4440 20422 eee aan eee ean eiee 1297 Palzoloxodon protomammonteus Matsumoto, 1924, 1926.............0... 02.0 - eevee eevee eee esaes 1297 maleoloxodongtokunagaNviatsuino tol 929 ears eee 1298 Palzxoloxodon protomammonteus proximus Matsumoto, 1926............0. 0. e eee eee 1298 laaleoloxodonmamadicusly abersNViatsumOtLowl92 9) eee an eet ener ee ea ee 1299 Palzoloxodon (Archidiskodon?) tokunagai mut. junior Matsumoto, 1929.......................0.. 1299 Japanese species described by Saheki and Tokunaga (1931, 1934)...............0. 00. c cece eee cece eee 1300 Rarelephasyprotomammonteus matsumotor saheky, 1930 )9..2 42 oe eee eee aaee 1300 Ralzolocodonyokonamanusmhokunag as] 934 ee ee ee 1301 Javanese species) described by Duboist.. anette eer eee en ee en ee ae eee 1302 (Raleolonodongny SUG NAvCUSMD UD O18 lO OS rere reiterate eet eee ee 1302 Geographic) distribution along theseasterncoastiot Asian.) sesso cece ate ence erieaene aeiie eee 1304 XX. THE SUBFAMILY ELEPHANTINA (SUPERFAMILY ELEPHANTOIDEA), OF EASTERN ASIA, IN- CLUDING THE RECENT ELEPHAS AND RELATED LOWER PLEISTOCENE SPECIES....... 1307 Enistoricallintroductionyandinomenclaturen(So0?)b.C:—_1936) sae eee eee aoe eee eee eee 1308 Falconer (1868) on the specific unity and vertebral formule of the Asiatic elephants....................... 1312 Corse, de Blainville and Falconer on characters of the geographic varieties....................00-.000000e 1313 Hlephasindicusicodactylusetlod 80 newer nya etter eta eae terete ee 1313 Hilephasmndicuseactenodacty us lod fSOnh em nee ne ae ner OnE rene ner erent 1313 Elephaswndicusiceylanicuside Blainvalley nan nae ore eee ic een eer eee ene eee 1313 Bilephasiindicusivengalensiside) blainvilles ere pee ee epee Oe ee nee ee oor eee ete 1313 Klephas andicus (Dauntela var:)): nance ee deh oer eek eo en ee eee eee eerie 1314 Blephas:indicus) (Mika svar:)io cis to4 a aca sersbace Ge ric ao oie eee EEC Cn nn eer 1314 Iniving specific orsubspecific forms, continental and ansularsee yee eee eee eerie 1315 Hossil'formsimorejor less closelysrelated itopHlephaswndzcusmese pice hentai eee eta 1318 Names of species and subspecies of the subfamily Elephantine in order of description..................... 1319 iHlephantinse!@sborn, 1910subtamilysdefinitionemcrnce tse eee ee rete ee ee 1320 Hlephas Linneusy 135-58. ;2eneric definition ae eee caer ere eee eo ee eee rE eee eer Creer errs 1322 indicus Linn eus lio (O41 COleCtiVe|ISPECIES ayae eccrine tee eee eee 1323 indicus ceylanicus de: Blainville; 1845512 (Reieees ON CLARE e I ee ee eee 1450 pleistocenc? Of B Umm ate ew yah gk hed oases ere ioe eee eee 1450 Pleistocenesor Southey @ bina s coe sovc costes sce neat seratevor te haemo ete ee ere ree 1452 Pleistocene ofelndo-Ohrinaers Ac tege tec s,s oho Oe cress meee Tee Tey ene oe ee ae 1453 Pleistoceneiofatherbiastula diesel ke skated so eteve te ome a Te a REET Gree Pe 1453 | DU 0) oY Reine toe io cian, Pian cco A ROR ACE RO Se Ie Ca eon 6 Raper c aes Wad hin Hobo aoe Se Gans 6c 1456 TowersMiocene eS Urdig alia nyse forces cite eae te le ae Or Oe creche ee eee 1457 Middle Miocene: Helvetian and Tortonian—Vindobonian...................00 cece eee eee e cence 1460 WpperMiocenesm Sarma tianecrs .\f. fers ame cit rie eater een nte ie Ee oe ae eee eae 1464 Omer Pelt OCeMe ee OMAN Cys icte .aie eX syoved oe See teae ed eet ehese eC TE eee ON ect eco EE 1466 Middletbliocenest bl aisaneian cys. cccys) costs, ersrancs scot eRe Se ae reise en eee eae 1469 Wipperr Pliocene smeA's tlarictars... et aactsten ere avo eats cuustsver Poa Sree aR Ge hae Se ee Ie ue suave Oem eee 1470 JPA ATSROVTS OVS. O'S chp 6 a-c chee CERO ROR BL CRORE OPERONS OR ETOP oir UA A Aerts he A AN tN RRR 8 3's 1472 INTENSE 40s de Ora Sar OTE TRUER EERE TS CEE ASO GAT RE gee PEE, Ot A A Oa SURES OL RE 1477 Miocencote ong oliavand! CembraleAsiayys. oe sna ssclerie ae orci srstancvateeterces Sobre mateiey sisi nrarse alc einoreiel erases 1477 INTO CENELOTm NOt © im aint ote tre cy Se ei ite yes ee ce rome acto se ei ee raek Arn Aro en tea eeak OLS Ai ree 1479 bhiocenetore Wong oli arecites.s cree od Secheete elt ie YO OG eh Roe oe eee ioe ace 1481 VEIbCoLersr eS). Che, INMoyA Net O) abbey pee OM Re en Boe eG ed Nita A Sie A ae ee on LA OR Ae nh Sto a A mnO Id He 1481 Pleistocene of North China.......... Par fir SECRET OF RIO IC EO TIE aD aR Dearne nh oat 1483 NiioceneKcoEblelstocene Ol aa paiin tcc soe steer ese A CI EEO SRS REL cee eerste te 1490 IN Ox te ATTIC LI CA Mice La ot Str fe Te Seco Palio A AIEREG OU eA cticicums AHR Vas eer aR EI Re ro Ser tito hobs nears 1490 Uppers Miocene sa B arstovasin'. a-pren ta cricnse asia eS nena seh Oe eho eee VC Pee a oe ee a ll ciake apna vee 1491 ltowersbliocene-s Clarendonianvacaite ces. certo kere oor ee Te ae ee ane OEE ocr 1495 MiddleyPliocene-sfemphilliaticy i. .v.ranta ve eas ca iar ce aati et chniue ave Porn siael sere leven srae crore ceorenceea 1502 Wippenieliocenes!Blancam s 2%: Oies.2 Aaa Aoe eee eee, Jk cits nn: HRN Sie a SORA Ra a tr, oP ea 1503 IP UTORENSTOLE VLC RA COT eases co ieee ae es aa ren NPs Se ETI ce oP Saale yc one Bea ee 1506 Proboscideans from undetermined levels in the Miocene and Pliocene of North America................ 1507 North American Tertiary horizons containing fragmentary proboscidean remains....................+ 1508 Pleistoceneioft North Ameria teach. syatieers oe sa teotees Rhos eo re Ste Me oe Sey er acer 1510 PlGIStOCEneTOLe VICKICOA Cee yee ee inte eee ore a ener te ae Ne Joly EC CE NP te tee cect arene err eyetineee 1515 Centralvands SouthAmertegicc.t-rccaconac werers sateretetncl stein to NERC He RA ae ate ae a eet earn ode Soar AT Rea tern 1516 Pliocene tof CentralhAmmierica spac et. sors ste. A « 1024 Archidiskodon hayi, type mandible, compared with A. imperator ref............ 6.6 ee eee ee eee 1025 Archidiskodon imperator, referred juvenile jaw, subsequently made by Barbour the type of Hlephas [A rchidiskodon] scott. 7 TUPLE SUELO HEE) hy tem 0] eno Ren ARs Os ois en cia ns Sion iio Car Oe akon ooo iO ice Nore acre iio} ari 1026 Comparison of type mandibles of Archidiskodon imperator maibeni and A. imperator scotti; also enlarged views of right second hariauton TAME CO UC IAAge Ol Zhe SS a gauee oadoes su cons Goosu ge ood andeos on odcunD oe sn aohoanagdaegroneasec 1027 Archidiskodon imperator maibeni, type superior and inferior dentition. After photograph. ............-..-......55-5. 1028 Archidiskodon haroldcooki, type mandible with third right molar in situ. After Hay and Cook....................45. 1029 Archidiskodon exilis, type, compared with A. imperator ref. After photograph. ............. 6.005 e eee eee eee 1030 Archidiskodon evilis, type. Facial portion of skull, with tusks and lower jaw. Restoration. After photograph......... 1031 Map showing location of some occurrences of fossil elephants on Channel Islands. After Stock. ..............-.-.-5. 1032 Archidiskodon sonoriensis, anterior portion of type mandible and maxilla, showing third superior and inferior molars... . 1033 Archidiskodon meridionalis nebrascensis, type mandible. After Osborn... ........... 0... e eee ee eee 1034 Archidiskodon meridionalis of Durfort, referred superior and inferior molars, found associated with skeleton. After casts... 1035 Archidiskodon meridionalis of Durfort, second and third superior and inferior molars of skeleton. After casts.......... 1035 Archidiskodon meridionalis nebrascensis, type. Restoration by Flinsch................... 00sec eee eee ees 1036 Archidiskodon meridionalis of Durfort and A. meridionalis nebrascensis of Nebraska. Restorations by Flinsch......... 1037 Parelephas trogontherii of Mosbach. Restoration by Flinsch. .........-..-. 660 e eee eee eee eee teenies 1038 Parelephas jeffersonii, type. Restoration by Knight. ............. 60.6 e cece n eee eee eee eee n ene enee 1040 Parelephas jeffersonii, type skeleton. Second figure after Osborn. ..... 0.2.66... eee eee ee eee tenes 1041 Map of chief Lower to Upper Pleistocene localities in which occur species of Archidiskodon, Parelephas, Mammonteus, Loxodonta, and Palvolocodons eAttern! OSDOrI irs ce csies ieee ee bere teleraitatetietona tstelcaverers:leltel olell lotevieke fedtiestite its laQelle aitelataMyieteane)/-s1 1042 Map showing geographic distribution of species of Parelephas, types and referred specimens... 0.66.62 eens 1047 Mammonteus primigenius and Parelephas trogontherti, cranial profiles. After Pohlig and Falconer... ........-...+.5. 1050 Left lateral profiles of seven species of Parelephas with progressive ridge formul®.... 0.0.0... 5.000000 e eee eens 1051 FIGURE 936. 937. 938. 939. 940. 941. 942. 943. 944. 945. 946. 947. 948. 949. 950. 951. 952. 953. 954. 955. 956. 957. 958. 959. 960. 961. 962. 963. 964. 965. 966. 967. 968. 969. 970. Ale 972. 973. 974. 975. 976. Oe Ele 978. 979. 980. 981. 982. 983. LIST OF ILLUSTRATIONS XX1 PAGE Parelephas of Europe and America in comparison with Hlephas indicus bengalensis. Restorations by Flinsch.......... 1052 Cranial profiles of Archidiskodon imperator, Mammonteus primigenius, Parelephas jeffersonii, and P. washingtonii......... . 1053 Parelephas trogontherioides, lectotype and cotype molars. After Zuffardi.............0...c0c0ecc cect ee eeeeeeveeeee. 1054 Parelephas trogontherii, type third superior and inferior molars. After Pohlig.................... 0200000 cece eee eee. 1057 Parelephas trogontherii, referred molars from Siissenborn and Weimar. After Wiist...........................-22-.. 1058 Elephas antiquus Nestii Pohlig |= Parelephas(?) trogontherii nestii], cotype or syntype left third superior and inferior molars. PACE TAT HO, LO RT APNG bah shog ae See SRN Aa i RU a SRN Cie eA Na etl iy aS Ph aca Nn oa ds Voit. yale sant baal Pee O59 Rarelephasianmentacus, type left third!superior molars eAfter Halconers. 454.2208. . 0stoee sos eee eee oss elle ee eee 1061 ponelephaswiiermedius, reternedemolarss, eAti ter photographs saan ares e eee ee ae tence ne aan 1063 Parelephas intermedius, restored skeleton in Lyons Museum. After Lortet and Chantre............................. 1064 Rarelephaswwust), cotype molars, from’ Tiraspol; Russia. After/Pavlow......9.20.+.-.2.se0-02ese0e0.. cee ee se ee ... 1066 Parelephas jacksoni, type juvenile jaw, and referred Hlephas [Mammonteus] primigenius jaw. After Mather........... 1068 Diagrammatic cross-section of type locality of Parelephas jacksoni. After Mather............................-..-.. 1069 Parelephas columbi, type third right lower molar (middle portion) longitudinally and vertically bisected. After Falconer... 1071 Parelephas columbz, restored type molar, redrawn for present: Memoir................---e--c cece cc cee veecseeeeeee. 1072 Elephas texianus |= Parelephas columbi ref.|, type right third inferior molar. After Blake........................... 1073 Parelephas columbi, Falconer’s type third right inferior molar and Osborn’s neotype third left inferior molar. After Osborn. 1074 Parelephas columbi, key to superior and inferior grinding teeth selected from the Cohen Collection, phosphate beds, Charles- WGI Ish (Chata clo ct ene eee een eee As netin hey crea ape ee AL Re ORM fe Me POT Or en Wome cers thre Ee 1076 Map showing distribution of Parelephas jeffersonii, Mammonteus primigenius, Parelephas columbi, and Archidiskodon mipenaionminaunesumitedi statesyand CanadaseeAttersH aye eerreee ernie. ieee een ee eee iar ieee 1078 Parelephas columbi, superior and inferior molars found in incomplete skull of Amherst skeleton. ...................... 1079 anelephasycolu moma herstiSKCletOn eA acy tras Sen eee es Te TI ya fs Sere Sepa eer ee 1081 Parelephas columbi felicis, type third right superior molar. After Freudenberg................... 000.00 ee ceeeeeeees 1082 Parelephas columbi cayennensis, type fragmentary third right superior molar. After cast...................0..00000. 1083 Parelephas jeffersoniz, showing ridge-plate compression at three levels of third right inferior molar.................... 1085 Parelephas jefferson, aged type third superior and inferior molars; also same superposed on type molars of Elephas TO OSEVELLIR ER ree oe heen, oaks hE ara Site Shs Bees ee Del aes Mee Canin ny, De See tpn RL a 6 ae eee 1086 Parelephas jeffersoni, type and paratype [ideotype] grinding teeth and jaws..................000 0.00 e cece eve eeeeee 1089 Hrontaliviews ofcraniaiof Parelephas jefferson and PP: washingtonia.2.) eee eee eae eee 1090 Profile views of type and referred crania of Parelephas jeffersonii, P. washingtonii, and Mammonteus primigenius; also {OMUBVAC WHOL MM DTUINLG ENTS a. -< ca 3 y2506. 5 Sestak eventos Be tN So Nota eS) Cal eR Foe 1091 Parelephas jeffersonii (Franklin County Mammoth) and EHlephas indicus bengalensis crania...........-..0...02.0 0000. 1092 Parelephas jeffersonvi (Franklin County Mammoth) skull in Nebraska State Museum. After photograph............. 1093 Parelephas jeffersonvi (Franklin County Mammoth), aged. Diagram showing ridge-plates of second and third superior MOTs wandyportion of crown) of thirdinfenor molar —AftersBarboury ase ode eee oe eee eee 1093 Parelephas jeffersonii, type skeleton in American Museum. First published type figure by Osborn.................... 1094 Parelephasvetiersonim paratype lideoty pel] awe... cusveeet career eters een eee ater ee eee 1096 Elephas roosevelti (syn. Parelephas jeffersoniz), type third superior and inferior molars......................0..000005 1096 Parelephas progressus, type third superior and inferior molars from Zanesville, Ohio, side views, originally figured by Oshormbasipanatypes of Hlephasjeffersoniz. Atter Osbornh jeer ieee eerie eee iene eee 1098 iRanelephasipnogressusy uy pevmolars.wcrowny Views a) Aditer OSbOnmiererteeera ara aes oer eee eae ee 1098 Porelephaswvashingtonia, referred (cranium: & 42s. eed tats eet ere hon eee ne eee eee 1100 Rarelephasswashingtoni, type adult jaw. aciss. «owe sec Soe G eeOnios chee eee ne Serie ae ee eter ae 1101 Parelephas washingtonii, referred young adult male skull, right lateral aspect... 2.2... 2.02. cc eee 1102 Parelephas washingtonii, referred young adult male skull, left lateral aspect, combined with type jaw................. 1102 Parelephas washingtonw, type jaw compared with that of P. jefferson... ... 2.2... 0221 ee ett 1103 Parelephas washingtonii, referred cranium with second and third superior grinders in situ... 2.0... eee 1103 Parelephas washingtonii, referred second left inferior molar. After Peterson.................... 0. cee eee cece cece eee 1104 iRanelephas()neellsestyperskulletragmaent. eAtitery Eel aiyzerepatseeetcic eat wey ta teeta ee eee ee eee en ey ae 1104 Parelephas floridanus, type and paratype crania and jaws compared with type molar fragment of P. colwmbi cayennensis.... 1105 Parelephas floridanus, type and paratype palates, with molars im situ... 2.2... 2.22 ee ee ee 1106 Parelephas floridanus, type right and left superior and inferior molars. After Osborn. .........................0000, 1108 Parelephas floridanus, type third left superior molar (detailed photograph) and drawing showing method of measuring length of superiommolaricrown: After’ @sborn......; = 2422 aaae some aii a/ Sies 2 ean Oren ee tener eerie 1109 Ronelephasmonaanus sy peaandibles! AtterOsbornemee eater eee eee etter 1110 XXil FIGURE 984. 985. 986. 987. 988. 989. 990. 1030. 1031. 1032. OSBORN: THE PROBOSCIDEA PAGE Parelepias onaanus | pataby pe mandi pleseAtter, Osborne serie ene tet test teehee er eter ee 1110 Parelephas jlojdanis type wtenionmolars: Auten Osborne rier ryote eet t steele ete eetetetet et este eaten eee 1110 Parelephas flomdanus, reconstructed type cranium. -Atter Osborn se ees] ete eerie elie 1111 Parelephas floridanus, referred inferior mandible and milk dentition..........................-52.2-..0....5200-50-- 1111 New standard method (1930) of proboscidean skeletal measurement, illustrated on type skeleton of Parelephas jeffersonit. AES rt OSWOLI ee srs Arse ae PR TN TS eI ONC NTO 3 AEC Cer eee REG SI nS rE 1112 Parelephas floridanus, referred right third inferior molar, showing method of ridge-plate measurement................. 1115 Mammonteus primigenius (Woolly Mammoth). After painting by Knight............................-....--.0005. 1116 Messerschmidt cranium of the mammoth of Siberia. After Breyne.................. 00sec cece eee tee eens eee 1119 Comparison of crania of Mammonteus primigenius, Elephas indicus, and Loxodonta africana. After Cuvier............ 1121 Mammonteus prumgenius, lectotype molars: “Afitericastss-.-- oo. 2-4 >] 4s. oo ee eer ae sic teen iene 1122 Migrating Woolly Mammoth (Mammonteus primigenius) as it appeared on the river Somme, northern France. Restoration [on ca) <0 td 01 eee eee tegen, Ori ae DARA ica Cero an DOGMA RR Ob thin mooie nine Abb. o is scien. e 1126 Comparison of the tip of the trunk of the mammoth (Mammonteus primigenius) with that of Hlephas indicus and Loxodonta CO [18s 0 eae SUAS el Ol (210) eee een ps eee loin Co Micking SST ma ane Seni Cana SUD bn Oye Go4 Duet iD mn bins 5 ¢ 1128 Mammonteus primigenius, skeleton, from Kolyma-Beresowka River, Siberia. After Salensky........................ 1130 Mammonteus primigenius fraasi, skeleton, from Steinheim on the Murr, Wurttemberg. After Abel................... 1130 Mammonteus primigenius, skeleton, from Borna, Germany. After Abel................ 0 0c cece eee eee eee ee eee 1130 Mammonteus primigenius, skeleton, from Lierre, Belgium. After Dupont..................5 20. e eee eee eee eee 1130 Restoration of the Woolly Mammoth sketched on the wall of the cavern of Les Combarelles aux Eyzies (Dordogne), Brances -Atten Capitan Breuiliand) Peymronys,5.oes.cxctiernce che sceete cies are rake ac aere erste vee oe tee te Nene wee 1131 Outlines ol theawoolly; Mammoth iromithe!Grottoot@ombarellessee ey. eaeeieeeeeree 1131 Charging mammoth incised on a tusk of Mammonteus primigenius discovered in the rock shelter of La Madeleine (Dor- dogne)“France:, After. artetiand! Christy. oicrecuscu otis Svs ern eles aici eer eae ele aad eens ores 1132 Map showing geographic distribution of Mammonteus and Parelephas in North America. After Hay................. 1133 Map showing location and principal discoveries of fossil mammalian fauna of Alaska-Yukon to the year 1929.......... 1134 Diagram showing geographic range of Mammonteus and Parelephas. Now superseded by figure 795 above............ 1135 Map showing geographic distribution of principal species of Mammonteus, types and referred. ...................---, 1136 Mammonteus primigenius, skeleton from Moravia. After photograph.................. 0.20. e cece eee cee eee 11389 Mammonteus prumigenmisyretermed molars ofyAlaskars aie cia ctiecisc cr isle crete oes seco seis ocialle eetenanet el elelcnel = ielieteeeetetsticte- es 1142 Mammonteus primigenius and Parelephas jeffersonti and P. washingtonii crania compared.............-...00 0.00025 1144 Mammonteus primigenius, male cranium, from the Yukon. After photograph.....................0 00. 0cee eee eee. 1145 Mammonteus primigenius from Alaska, showing growth stages in the jaws and teeth. .......................-...-5-. 1145 Type of Elephas odontotyrannus [=third superior molar of Mammonteus primigenius]. After Eichwald................ 1146 Mammonteus primigenius, Elephas indicus bengalensis, and Loxodonta africana oxyotis. Restorations by Flinsch. ......... 1147 Mammonteus primigenius (“Adams skeleton”) from the Lena River, Siberia. Subsequently made by Brandt the type of LEPRGStDTAChy RAM PhUs PALLCTPUILERIUS ebay tebe aetels esa is eerae eee ei patie etoa pin Aes e eskmiebal al shals Cages phatase cnn ene 1148 Mammonteus primigenius leith-adamsi, type third left inferior molar. After Pohlig.................. 2.0.0.0 22.0 e eee 1150 Mammonteus hydruntinus, type first left superior molar. After Botti. ............. 0... ccc cece eet 1151 IGT GEES (EMAL TES OC, oe Genie, ANiieye IDEAS Gag soos ecano Go maeeecsso5 950005400445 50005 0s0R5C 1152 Mammonteus primigenius fraasi, type mounted skeleton. After photograph... 2.0.2.0... 50000. c cee ee eee eee 1153 Mammonteus primigenius astensis, type and paratype molars. After Depéret and Mayet........................055- 1154 Mammonteus primigenius (?)astensis, referred molars, compared with (?)Parelephas, Archidiskodon or Hesperoloxodon Molar PAT temdinecscale tracings) byatheypresent/sUluhOms .eicaary-cie chae eienetmeele peers cle raves Bess char teemeieie teterhe 1155 Mammonteus primigenius americanus, type. Portion of upper molar from near Rochester, New York. After DeKay... .. 1156 Mammonteus primigenius compressus, type second and third superior molars. After Osborn... ........--.-..-.000055 1157 Mammonteus primigenius compressus, type female skull. After Osborn................. 00. e ee cee eee eee eet 1158 Mammonteus primigenius compressus, paratype third right superior molar. After Osborn. ...............6.0 005 eee 1159 Mammonteus primigenius alaskensis, cotype crania, superposed outlines. .............0 00. e eee tee eee 1159 NUCHTIMOTREUSEDT UIT GCNTUSIQLASKKENSTS NCOUY, DP CxCLAIL Dina eo) gele aie ches teve eine ne ieee etna a tet reat eee eae ene teen 1160 Discovery sites of frozen carcasses of the Woolly Mammoth and Rhinoceros. After Tolmachoff..................... 1162 Map showing geographic distribution of mammoths in Upper Pliocene and Pleistocene times. After Osborn.......... 1164 JehersomanmViammo thsof Indian asiyaons trvecrs ta ieace teh ere eeu ie te te ee en ote Roth MARIE NC oo ele Ree es 1165 imperiale Mammo thot Nebraska: is cys eserves cicero eee Tee eee Neel eis rete Pes liey tote tanto to tetettemey svete Wree Fore ctiys 1165 Archeditusks ofthe African elephan tis 7azt. size \ STEGOLOPHODON CAUTLEY! PROGRESSUS 24 =—————_ e 2 — ae ————— Fig. 726. Type of Stegolophodon cautleyi progressus Osborn, 1929. Detailed study of right superior dentition, Dp*-M!, combination drawing of two sides. Coronal ridge-crests intermediate in formula and pattern between the typical Mastodon [=Stegolophodon] caulleyi Lydekker and M. [=Stegolophodon| latidens Clift; similar in the regular disposition of the cones and conelets. 849 Fig. 727. Type skull (Amer. Mus. 19446) of Stegolophodon caut- leyt progressus Osborn, 1929, collected by Barnum Brown twelve miles east of Chinji Bungalow, summit of the Lower Chinji horizon, 2,000 feet above the base of the Lower Siwaliks, India. This skull, containing the right and left superior tusks, third and fourth superior deciduous premolars, and the first superior molars (partly erupted), shows the laterally compressed tusks with enamel band, 4+ ridge-crests on Dp‘, 4+ ridge-crests on M!; it is somewhat more progressive than the type of Mastodon { =Stegolophodon| cautleyi Lydekker. Left lateral, palatal, and right lateral aspects, one-fifth natural size. $50 eC. One tet OLDOIN SSNS z ay ero Big S.LYDEKKERI TYPE TYPE PACS Coy RNS . S.CAUTLEY! S.LATIDENS after Clift Ty ae ZS C S. NATHOTENSIS S.SUBLATIDENS TYPE PLATE XIII STEGOLOPHODONTIN.®, STEGOLOPHODON: PrimitTIvVE LONGITUDINAL SuLcus, SHADED (A, B), Persistine rn I-IV AnTEeRtorR RinGe-crests (HK, E1), tn L-III Anrertor Ripce-crests (F), VesTiGIAL IN RipGE-crests I-III (G, H, 1). Constets Rounpep, INcREAsING FROM 4-5 (A, D), 5-7 (C, H, I, G). Posrertor RipGce-crests [V-VI Proaressive, with ConEeLEts 5-7 (G), ConeLets Nor Excrerepinea 5 (H, I) Mio-Pliocene B, Stegolophodon cautleyi progressus type, r-M'. Amer. Mus. 19446, near Chinji Bungalow, India. Summit of Lower Chinji horizon, 2,000 feet above base of Lower Siwaliks. C, Stegolophodon nathotensis type, r.M3. Conelets 5-6. Amer. Mus. 19455, near Nathot, Lower Middle Siwaliks, India. Lower Chinji horizon. Lower Pliocene! D, Stegolophodon latidens cotype, r.M3. Ridge-crests I-VIII, conelets 4, sulcus on ridge-crests I-IV. After Clift, 1828, Pl. xxxvin, fig. 1, near Yenangyaung, Burma. E, Stegolophodon latidens cotype, r.M*. Ridge-crests I-VI, sulcus on ridge-crests I-IV. E 1 (section), ridge-crests coalescent at base (I-III). After cast (Amer. Mus. 21978) of Clift’s type, 1828, Pl. xxxvu, fig. 1, near Yenangyaung, Burma. F, Stegolophodon latidens ref., r.M'. Sulcus on ridge-crests I-III, conelets 4-6, ridge-crests 445. After Matsumoto, 1926.1, PI. y, figs. 1 and 3 (Prostegodon), Shiwogama, Miyagi District, Province of Rikuzen, Japan. Pliocene (?) G, Stegolophodon lydekkeri type, 1.M®. Ridge-crests I-VI, conelets 4-7, sulcus on ridge-crests I and II only. After Lydekker, 1886.2, fig. 19 (as M. latidens), Borneo. Middle(?) Pliocene A, Stegolophodon sublatidens type, r-M®. Conelets 4-5. After Schlesinger, 1917, Taf. xv, fig. 2, Teschen (Schlesien), Austria. Middle Pliocene H, Stegolophodon cautleyi lectotype, |.M*. Ridge-crests I-V, sulcus on ridge-crests I and II, conelets 5. After Lydekker, 1886.1, p. xv, fig. 6, Perim Island, India Upper(?) Pliocene I, Stegolophodon stegodontoides Pilgrim, type, r.M®. Ridge-crests I-VI, sulcus on ridge-crests I-III, conelets 5. After Lydekker, 1880, Pl. xxxrx, Lehri, Punjab, India, possibly Upper Siwaliks. CoMPARATIVE OBSERVATIONS (1935) Molars of Stegolophodon are readily distinguished from those of the Mastodon, Zygolophodon, and Turicius phyla by the following characters: First, by the persistence of the median sulcus separating the inner and outer pairs of cones of all the crests (A, B, D), of the three to four anterior crests (EK, F, I), of the two anterior crests (G, H). Second, by the rounded, bunoid conelets separated by median sulcus (A-H). Third, by the closure of the enamel in the base of the transverse valleys, as seen in section (El), very characteristic of Stegodon. The second and third of the Stegolophodon characters enumerated above link this genus with the genus Stegodon. But we must remember that Stegolophodon cautleyi is of Middle Pliocene age (Perim Island), contemporary with the true Middle Pliocene Stegodon bombifrons (Dhok Pathan). [See note on page 824 above regarding the Lower Pleistocene rather than Lower Pliocene age of Stegolophodon latidens.—Editor.] THE STEGOLOPHODONTIN A: evidence in the structure of the superior grinding teeth of the affinities of this tetralophodont type with the much more primitive trilophodont mastodonts of the Lower Miocene of France (cf. also Figs. 725 and 727). Stegolophodon lydekkeri Osborn, 1936 Figure 728, Pl. x1 From vicinity of Bruni, northwest coast of Borneo. [In an intensive reexamination of various species of Stegolo- phodon, Professor Osborn discovered wide differences between the Fig. 19. Mastodon latidens.—The third left upper true molar of a small individual in a partially-worn condition ; from the Pliocene (?)of Borneo. $. The lower border of the figure is the inner border of the specimen. (From the ‘ Palxontologia Indica.’) Fig. 728. Stegolophodon lydekkeri Osborn, 1936, Volume I, p. 700, fig. 660, of the present Memoir. Third left superior molar exhibiting six ridge- crests anda talon. Figured by Lydekker, 1886.2, fig. 19, as Mastodon latidens. Compare Lydekker, 1885.2, Pl. xtvin, also Pl. xm of the present Memoir. Two-thirds natural size. STEGOLOPHODON 851 types of Stegolophodon sublatidens Schlesinger, 1917 (Pl. xur, A) and S. latidens Clift, 1828 (Pl. xu, E) and a molar tooth from Borneo described and figured by Lydekker as ‘Mastodon latidens’ Clift (1885.2, 1886.1, 1886.2). This molar Professor Osborn selected as the type of a new Pliocene species, namely, Stegolo- phodon lydekkeri (Vol. I, p. 700). Tyrr.—Third superior molar of the left side, 1.M’, original in the Zoological Society of London; cast in the British Museum (M.2498). Horizon AND Locauiry.—(Lydekker, 1885.2, p. 777): “The specimen forming the subject of the present notice was forwarded from Borneo to the Secretary of this Society by Mr. A. H. Everett, C.M.Z.S., who stated that it was found during the early part of the present year [1885] by a Kadayan in the jungle in the vicinity of Bruni, on the north-west coast of Borneo.” Pliocene (?). Tyrer Figurn.—Lydekker, 1885.2, Pl: xivmt, figs. 1 and 2; see also Lydekker, 1886.1, fig. 7, 1886.2, fig. 19, as well as Pl. xin and figures 660 and 728 of the present Memoir. men is the crown of the last left upper true molar of a tetralopho- dont Mastodon, and agrees so closely with Indian teeth of the Siwalik Mastodon latidens, Clift [Footnote: ‘Trans. Geol. Soe. ser. 2, vol. ii, pt. 3, p. 371 (1828).’], that it may be safely referred to that species, although it indicates a very small individual... . It will be seen that the Borneo specimen agrees [with MM. latidens of the Punjab—made by Pilgrim in 1913 the type of Mastodon stegodontoides| in the number of ridges (although the hind talon is considerably smaller), but is of greatly inferior size, the dimen- sions of the two specimens being as follows, in inches :— Punjab Zorneo. ixtremes en mt hina ann Se O 6.3 Width of first ridge... . . Bite: coaches Gay 2.95” 70. 75. 80. SS. Tlurdoo a on mings ~T 2 ef ortope tp. eeliengeceane Lake be Cape Comorin 70 75 80. LED 90. Fig. 729. Falconer’s map of the geology of India (see ‘‘Palzontological Memoirs,” 1868, Vol. I, description of Pl. m). The red fossiliferous areas, here colored black, are de- seribed by Falconer as follows: “The red [black] stripe represents the Sewalik Hills, stretching from the Hydaspes to the Gunduck River, 800 miles. The small red [black] patch behind the Himalayahs repre- sents the ossiferous plain of Tibet about 16,000 feet above the sea. The other red [black] patches represent the Nerbudda [Pleistocene], and the [black dot within the circle the] Gulf of Cambay [Perim Island—Middle Pliocene] fossil tracts.” The following is Falconer’s interpretation of the geology of India: “The great mass of light shading represents the supposed insular form of the continent of India at an early period of the Tertiary epoch, the island forming a sort of triangle, of which the eastern and western Ghats formed the sides and the great Vindhya range the base, with an irregular patch of mountainous country stretching north forming the Aravalli range.” “The dark shading represents the plains of India, forming the valley systems of the Ganges and Indus drainage, which were formerly narrow ocean straits. These straits were the recipients of the silt and alluvium washed out of the Himalayahs, and were at length elevated above the sea, so as to form the existing continent. The Sewalik Mauna then spread over the continent, from the mouth of the Irrawaddi to the Gulf of Cambay 2,000 miles, and north to the Jhelum 1,500 miles. After the long establishment of the Sewalik Fauna, a great upheavement took place along the line of the Himalayahs, elevating a narrow belt of the plains into the Sewalik Hills, and adding many thousand feet to the height of the Himalayahs.” 852 IV. SUCCESSION OF SPECIES OF THE GENUS STEGODON SurpRFAMILY: STEGODONTOIDEA Osborn, 1935, 1936 FaMILy: STEGODONTIDA Young-Hopwood, 1935 SUBFAMILY: STEGODONTIN& Osborn, 1918, 1921 Genus: STEGODON Falconer and Cautley, 1847, 1857 Original reference: Falconer and Cautley, “Fauna Antiqua Sivalensis,” 1846 [1847, Pl. xxi]; also Falconer, Quart. Journ. Geol. Soc. London, 1857, Vol. XIII, pp. 314, 318, and Synoptical Table opposite page 319. Genotypic species: Elephas Cliftii, EZ. bombifrons, E. ?Ganesa, E. insignis.1 Syn.: Emmenodon Cope, 1889; Parastegodon Matsumoto, 1924 (in part). Generic Derinrtion.—(Falconer, 1857, p. 318): ‘‘Dentium molarium 3 utrinque intermediorum coronis complicata colliculis hypisomeris (e.g. 7+7+8), mammillatis, tectiformibus. Praemolares non- dum observati.”’ GeneRIC CHARACTERS.—Ridge-crests intermediate between Stegolophodon and Archidiskodon planifrons types, progressively multiplying from six to eleven in the intermediate molars, from nine to fifteen and a half in the posterior molars. Cones rapidly subdividing by binary or ternary fission into multiple conelets. Ridge-crests elevating from brachyodont (Stegodon sinensis) to subhypsodont (S. airawana and S. aurore stages). Cement developing in the valleys. Crania of mastodontoid (S. bombifrons) to extremely abbreviated, female? (S. insignis), more elongated, male? (S. ganesa), more triangular (S. trigonocephalus) form. Tusks attaining great dimensions (S. ganesa). Phylum parallel to that of the true Archidiskodon and Elephas, not directly ancestral, readily distinguished by cranial and dental characters. The generic name Stegodon was first printed Pig. 730. After Osborn, 1910.346, p. 323, fig. 154. Compare figures 729 and 826 of present Memoir. in the “Fauna Antiqua Sivalensis” of Falconer and Cautley, 1846 [1847, Pl. xtu]. It was ten years later that it appeared as a subgenus (Fal- coner, 1857, pp. 314, 318, table opp. p. 819) to include the following species, Elephas cliftii, E. bombifrons, E. ganesa(?), and EF. insignis, and has subsequently been used as a genus by all the principal authors, except Lydekker who clung to the Cuvierian division of the mastodonts and elephants into the two genera Mastodon and Elephas. Pohlig in 1888, p. 252, wrote the genus in this way: Stego(lopho)don. Schlesinger, 1917, p. 115, separated the species M. latidens as the type of a new genus Stegolophodon. Stegodon (omitting the six’ species described above under Stegolophodon) is readily definable —Chief Miocene and Pliocene fossil mammal deposits of Asia. oD Ou Le O = = Persia. 2. Perim Island. 3. Manchhar Beds of Sind. 4. Siwaliks of the Punjab. 5. Sub- and readily distinguishable fr om any genus of himalayan Siwaliks (River Brahmaputra to River Jhelum). 6. Valley of the Lower Irawadi, ° Burma. 17. Miocene and Pliocene deposits of China (Provinces of Shan-si, Shen-si, Sze-chuan, the Mastodontide, on the one hand, Or of the Kwang-Tung, Ho-nan, Hu-nan, Hu-peh). 8. Miocene and Pliocene deposits of Japan. Elephantide, on the other (see Vol. I, p. 25), by the brachyodonty to subhypsodonty of its ridge-crests, in contrast to the hypsodonty in all the species of the Elephas and Loxodonta phyla. The relative height attained is shown in figures 687 and 688. While the ridge- Hopwood in his Memoir of 1935 on the “Fossil Proboscidea from China,” p. 72, remarks: “According to the modern school of priority-purists this necessitates the selection of H. cliftii as genotype as a matter of course. Fortunately, we can do so and adhere to the author’s original intention, for, in his later writings Falconer expressly says that Stegodon corresponds ‘with the forms collectively designated Mastodon elephantoides by Clift’, (1857, p. 314) —Editor.] *ITo these six species should be added the Stegolophodon lydekkeri Osborn, described in Volume I of the present Memoir (p. 700). —Editor.] 853 854 crests are composed exactly as we know that we shall find them in the direct ancestors of Hlephas, they remain short crowned, even in the progressive species Stegodon insignis and S. ganesa, and in the still more pro- gressive S. orientalis and S. airawana. This is undoubtedly an adaptation to browsing on leaves and softer kinds of food, which leads us to believe that the Stegodonts were persistent browsers rather than grazers, as in OSBORN: all the phyla of Hlephas and of Loxodonta. TABLE V. Valeoner (1868), Lydekker (1886), Martin (1890), Matsumoto (1918), Osborn (1929) THE PROBOSCIDEA Species tN APPROXIMATE ASCENDING OrpDER OF COLLECTIVE MAXIMUM AND Minimum RipGe-crests | Dp 2 Dp 3 Dp 4 M 1 M 2 M 3 STEGOLOPHODON cautleyr Y-B-¥6 Y-4-¥ 4 5-4 cautley? progressus | 3 aad vedo Ee 2 Vira 41-5, Ye -I6 latidens 4 2 aa a lydekkeri | B+ sublatidens 2645 stegodontoides Gays STEGODON SImensis y-4 ‘ cena > 5- -6% % 648 elephantoides (=cliftii, fide Fale.) = bee EO GH & elephantoides Clift | 8% 8% | 10 bombifrons zh == Tos == % 7 = ae ie 5-818 = trigonocephalus 4 mindanensis (Archidiskodon?) Frag | ments, far more progressiv | ein the direction of Archidiskodon than either | insignis or ganesa. | PEA 2 5-6 mie 6-7-7 4-8 _1-¥e-7--Yh- 8 9-1 0-1 1-11)4-4-1 1-48 UNSIGNLS 2 Ton 7-9 4-7-7 4-10 7-8-M-9-M-212 | 9-1 0-4-1 1-}-12-1 2413 : 2 5 a _BY-7 7-7% 6}-1 0-1 0-38 ganesa - 7 BMT 8 7H-8 7Mit-8-Vi- 8-18 es . 144-15 pinjorensis 2 eae ae xd = —— 2 = insignis birmanicus 12-4 S a5, eS Ye-2-16 5-4 M- 6-44 4-6-4 Mes-i Y-1 1-4 orientalis grangert 2 +54 7 8 ()-9-45 W138 , | AUrOre | | ue | eee et os E a | 22) bes Lee 2S = rd | 2 | 1-6 | 7-9 7-9 9-11 12-14 arawana 13-15-14 THE STEGODONTINA: STEGODON 855 The order of ridge-crest addition and development (Table V) corresponds approximately with the phylogenetic and geologic ascending order as shown in Table IV above of the present chapter, in which the species are grouped partly by geographic distribution. The ridge-crest formule of certain species, e.g., Sfegodon sinensis, S. ganesa javanicus, S. orientalis, and S. orientalis shodoénsis, are too imperfectly known to determine precisely their phylo- genetic position. In general, S. senensis appears to be the most primitive, while S. azrdwana appears to be the most progressive and geologically recent. STEGODON GANESA 3050MmM..10’e STEGODON PINJORENSIS 2745mm.,9’e INDIA SIWALIKS INDIA SIWALIKS 2 TEGODON INSIGNIS 925MM. 6'47%e' INDIA SIWALIKS STEGODON GRANGERI 2250MmM.,7/5’e CHINA TEGODON TRIGONOCEPHALUS 1428 mm.,4’8/4"e JAVA Peal 2148mm., 7/%%e STEGODON AIRAWANA 176mm. 3\0/%8"e SIWALIKS JAVA (a Fie. 731. Species or SrEGODON FROM INpIA, CHINA, AND JAVA Restorations BY MarGret FLINSCH, UNDER THE DIRECTION OF Henry FairrieLp OSBORN, ONE ONE-HUNDREDTH NATURAL SIZE SKULLS OF STEGODON BOMBIFRONS, S. INSIGNIS, AND S. GANESA IN THE BritisH AND INDIAN MUSEUMS Before considering in detail the succeeding species of Stegodon, it is necessary to examine and compare the known crania of various species with each other, as assembled in figure 732 from Falconer and Cautley’s beautiful plates, with the crania of other Stegodonts from the East Indies, e.g., Stegodon airawana (Fig. 773) and S. trigono- SPECIES OF STEGODONTS FROM THE SIWALIKS, INDIA. (PLATES AFTER FALCONER, 1847.) All figures one-twentieth natural size ELEPHAS GANESA Ref PI. XXII, Fig. 3 ELEPHAS GANESA Ref. Pl. XXIN, Fig. 2 ELEPHAS INSIGNIS Ret ELEPHAS INSIGNIS Ref. PI. XLII, XV. B Pi, XLII, XV. A <<< — j = tik i ELEPHAS GANESA Ret. é ) ( = L ) H | ~ ae = _~ \ t Noa ¢ 2S p Pl. XXII, Fig. 1 (rev. Cae! YE } é = rr my hy 4 \ | Arik i i\ / NOAA Nee | Len { ‘) | i f \ q | ELEPHAS GANESA Ref \ iH atin Pi. XXI ae Ih KT he 9! Wye ‘ . ELEPHAS INSIGNIS Ret : Pl. XVI, Fig. 3 ELEPHAS INSIGNIS Ret ) Pi XV. if See a Pe aaa = ELEPHAS INSIGNIS Ref. ELEPHAS INSIGNIS Ref. ; re Cs a PI. XVI, Fig. 2 PI. XLV, XV. B [rev.) ( i ey --~ \\e & : nie os \ ey ELEPHAS BOMBIFRONS Cotype ep a PI. XXVIN, Fig. 2 | ELEPHAS BOMBIFRONS Lectotype Lyd ELEPHAS BOMBIFRONS Ref, PI, XXVI PI. XLIN, XII ELEPHAS GANESA Ref. PI. XXII, Fig. 2 Vig. 732. The three species of Stegodonts of which the crania were known to Falconer are illustrated in this comparative plate in such a manner that their distinctive characters may be readily contrasted. Throughout the Stegodon crania are of relatively small size, lacking the cancellate structure characteristic of Elephas. All one-twentieth natural size. Compare figure 777. Elephas | =Stegodon| bombifrons, a Middle [to Upper] Pliocene stage of evolution, is illustrated in the lower line. [The cranium of E. bombifrons at extreme right should read: Pl. xiv, xu (rev.).] Elephas | =Stegodon] insignis, a Lower to Middle [Upper] Pleistocene stage of evolution, is illustrated in the middle of the diagram. Elephas | =Stegodon| ganesa, a Lower to Middle [Upper] Pleistocene stage of evolution, is illustrated in the top lines. It is extraordi- nary that tusks of such enormous length and width should be supported by a cranium of such small size. The extremities of the tusks should probably be turned inward, instead of outward as figured by Falconer and Cautley; the left tusk is complete; a portion of the right tusk is omitted in this drawing. [Compare figure 733 for revised restoration of the tusks.] 856 THE STEGODONTINA: STEGODON 857 cephalus (Fig. 776) of Java, also with the newly found S. orientalis grangeri (Fig. 763) of China. These Stegodont crania should be compared with the large cranium of Archidiskodon planifrons (Fig. 830), one of the most primitive of the true elephants. In general the cranium of Stegodon bombifrons (Fig. 732, bottom row, also Fig. 742 and Fig. 744) is more generalized and subelephantine in character, whereas the crania of Stegodon ganesa and S. insignis are very highly specialized, of relatively small size, and bear little resemblance to the crania of the true Elephantide. Quite different are the small, triangular crania of Stegodon trigonocephalus and S. airdwana of the East Indies. SexuAL DIvERGENCE vERY Markerp.—The referred giant skull of Stegodon ganesa obviously belongs to a full-grown male Stegodont (as shown in four aspects in Fig. 732). On the contrary, the crania referred to S. ELEPHAS GANESA Ref. Pl. XXIII, Fig. 2 Vig. 733. Cranium of Elephas [Stegodon| ganesa, after skull referred to this species by Falconer. Drawings made from original plates in Falconer and Cautley’s “Fauna Antiqua Sivalensis’’ reduced to one-sixteenth natural size. See figure 732 opposite for frontal, lateral, and palatal views, reproduced to a one- twentieth scale. The frontal profile is seen to differ profoundly from that of Stegodon insignis and less profoundly from that of S. bombifrons. In an attempted restoration of Stegodon ganesa from the tusks as originally represented by Falconer and Cautley, it was found impossible to lower the proboscis between the closely appressed tusks; it was also observed that the extremities of the tusks turned outward, unlike all other proboscideans. Accordingly on receipt of the gift of the cast from the British Museum (Jan. 23, 1931) the tusks were readjusted in a position with the extremities turned inward, allowing a sufficient space for the descent of the proboscis between the butts. The present figure represents the specimen according to this conception of the position of the tusks, in contrast (Fig. 732) with Falconer and Cautley’s restoration. [The most painstaking examination of the original specimen by Professor Osborn, Doctor Gregory, and Doctor Colbert failed to reveal any evidence either that the tusks have been transposed or that they are twisted in the alveolus by post-mortem changes. Nevertheless all felt that the space between them is insufficient for the trunk as restored by Falconer and Cautley and that the whole arrangement looks abnormal.—Editor.] 858 OSBORN: THE PROBOSCIDEA insignis (Fig. 732) obviously belong to small-tusked females, because the alveolar processes for the insertion of the tusks are extremely small and narrow. If, as seems possible, all the crania referred by Falconer and Cautley to Elephas [= Stegodon| insignis are females, and the great cranium referred to Hlephas {= Stegodon| ganesa is a male, there is in Stegodon a far greater sexual disparity and difference than prevails between the female and male crania of either Hlephas indicus or Loxodonta africana, as figured below in the present Memoir. If this be true, the sexual disparity in cranial charac- ters constitutes an important specific distinction of Slegodon insignis and S. ganesa. In the present Memoir we treat the two species separately but agree with the theory suggested by more than one author, especially Lydekker and Matsumoto, that the crania of S. insignis represent the females of the same collective species as the referred male cranium of S. ganesa. British Museum, W. D. Marruew, SepremBer, 1920.—The skull of Stegodon bombifrons is essentially elephantine, the shortness of the enamel plates being the chief distinction; the skull is apparently shorter than in Loxodonta; the jaw is deeper; the symphyseal process may be a little heavier. In contrast, the skull of S. insignis has a supranarial region with a great thickening of the cellular tissue which appears to round back into the occiput, the occipital crest being very little developed; a very curiously shaped head and very small tusks. Fig 2: CHARACTERS OF REFERRED SKULLS OF INDIAN STEGODONTS, AFTER FALCONER, 1868 (See Figs. 732-736, 754, 777) Skull of Hlephas |= Stegodon| bombifrons (Falconer, 1868, Vol. I, p. 458, Pl. xxvu): “Very fine and perfect skull, anterior view. Four other views of same skull are given in Plate xxvii. This head is very marked; it is convex from occiput to front and also across, and is very narrow at the temporal contraction. The bounding ridges sweep round by a A GENERALIZED CRANIUM A SprcraAuizep CRANIUM A Specratizep Mate Cranrum Fig. 734. Hlephas [=Stegodon| bombi- Pig. 7385. Hlephas [=Stegodon] insig- Fig. 736. Restoration: Elephas frons. Vront view of cranium, one-twelfth nis, one-twelfth natural size. After [=Stegodon| ganesa, one-twelfth natural natural size. Brit. Mus. M.2979; cast Falconer and Cautley, 1846 [1847, Pl. xv]. size. Brit. Mus. M.3008. After Falconer Amer. Mus. 10378. After Falconer and and Cautley, 1846 [1847, Pl. xxi]. Cautley, 1846 [1847, Pl. xxvu]. THE STEGODONTINA: STEGODON 859 bold curve into the post-orbitary processes, as in #. |= Archidiskodon| meridionalis. There is a deep furrow between the tusks. The nasal opening for the trunk is above the line (or nearly so) of the post-orbitary processes of the frontal bone. Above the infra-orbitary foramen on the right side there is another smaller opening.”’ Skulls of Hlephas [= Stegodon| insignis (Falconer, 1868, Vol. I, p. 448, Pl. xv): ‘This is the most remarkable of all the Indian fossil Elephants. The cranium is as singular and grotesque in construction as that of the Dino- therium giganteum. ‘The cranium is seen to differ remarkably from that of H. Ganesa (Plates xx1. and xxm.) notwithstanding that the molars of the two species agree so closely. That of #. insignis is flattened at the top, elongated from side to side and singularly modified, so as to bear an analogy to the cranium of Dinotherium giganteum, while that of H. Ganesa does not differ much from the ordinary type of the Elephants.” (Op. cit., p. 449, Pl. xv1, fig. 1): “This head is very cubical in form, is old, very concave in front and vertically ; teeth broken. Interval between incisive sheaths deep. No tusks.’ (Op. cit., p. 449, Pl. xvu, figs. 8 and 4): ‘‘Anterior and lateral view of another cranium. Both zygomatic arches are missing, and the left side of the cranium is deficient. Shows the great length of the incisive sheaths.”’ Skulls of Hlephas |= Stegodon] ganesa (Falconer, 1868, Vol. I, p. 453, Pl. xxi): ‘Large skull, with fragment of left incisive 7n stlu, and corresponding fragment of right incisive detached. The incisive alveoli are remark- ably elongated, as in E. primigenius. The plane of the incisives is continuous with that of the frontal, but with a tendency to obliquity forwards. The skull is very imperfect on right side.”’ (Op. cit., p. 454, Pl. xxi): “Fig. 1.—Hlephas Ganesa. Lateral view of large skull figured in Pl. xx1.—B.M. Fig. 2.—KH. Ganesa. Palate view of same skull. The right incisive is seen in section. The posterior true molar is seen on either side of palate. It has ten plates and a heel behind, and a small talon in front; the hind heel has few denticles; the four front ridges are worn. The alveoli are parallel as in the Mammoth.—B.M. Fig. 3.— E.Ganesa. Sketch showing restoration of skull, with tusks, of H.Ganesa, profile view,one-thirteenth of natural size.”’ (Op. cit., p. 454, Pl. xxur): Restoration. “‘Sketch showing restoration of skull, with tusks, of H. Ganesa, oblique antero-lateral view, one-thirteenth of natural size.”’ (Op. cit., p. 455, Pl. xxiv.a.): “Figs. 1 and la.—Hlephas Ganesa. Fragment of skull with palate and back molars on both sides. This is a most remarkable specimen. I have called it H. Ganesa (H. F.), and it much resembles the molar of the big Ganesa specimen (Plate xxi. fig. 2) in form and in the compression of the ridges, but the ridges are few... . B.M.” STEGODONT CRANIA OF CHINA AND OF THE East INDIES (See Figs. 763, 776, 773, 777) A most fortunate discovery from the pits along the Yangtze River near Wanhsien, Province of Szechuan, China, is the extensive series of Stegodont crania in all stages of development, infantile, juvenile, young adult, and mature adult. This priceless collection made by Walter Granger of the Central Asiatic Expedition of the American Museum illustrates in a perfect manner (Figs. 759, 761-763, 686) the complete ontogeny, dental succession, and metamorphosis of the cranial form. The young adult and mature adult crania, together with the inferior mandible (Fig. 763), apparently resemble the cranium of S. bombifrons more closely than the crania of S.insignis and S.ganesa. This material was described by the present author as belonging to a new subspecies, namely, Stegodon orientalis grangerv. The cranium of the East Indian species Stegodon airdwana (Fig. 773) of Java belongs to a very advanced mid-Pleistocene stage, resembling S. bombifrons and S. orientalis grangeri in profile but differing in the frontal aspect, which is flattened. The cranium of S. trigonocephalus (Fig. 776), as its specific name indicates, has a 860 OSBORN: THE PROBOSCIDEA triangular rather than a rounded superior profile, and, although of the same geologic age as S. insignis and S. ganesa, it is entirely different from the Siwalik species in its profile and proportions. Consequently it appears that both in cranial and dental characters the East Indian species represent a distinct and somewhat dwarfed side-branch of the northerly continental species. SYSTEMATIC DESCRIPTION OF SPECIES OF STEGODON Stegodon sinensis Owen, 1870 Figures 687, 702, 737 Alleged to be “from marly beds in the vicinity of Shanghai,’ China. This animal (ligs. 687, 702) is probably of Upper Miocene [Lower Pliocene] age, for it is more primitive than Slegodon bombifrons, and possibly ancestral to the S. bombifrons stage. Speciric CuHaracrers.—This little-known true Stegodont, Stegodon sinensis (Dp 3 =*), is somewhat more primitive than S. bombifrons (Dp 3+), since the ridge-crests are less elevated or hypsodont as compared (Figs. 687, 688) with S. bombzifrons, S. orientalis grangeri, and S. insignis. Estimated number of cone- lets on fourth ridge thirteen to fifteen. Falconer, Lydekker, and Martin ascribe (see Table V above) four ridge-crests to Dp* of S. elephantoides (=cliftii), S. bombifrons, and S. trigonocephalus. History.—(1) After comparing the type deciduous molar with all the Siwalik specimens in the British Museum, Owen concluded (1870, p. 420) that the above Chinese tooth was most parison is strengthened by the fact that Koken (1885) referred the Stegodon sinensis of Owen to the species Stegodon cliftii of Faleoner. (3) Lydekker (1886.2, p. 79) also referred S. sinensis Owen to Elephas cliftii Fale. (Op. cit., p. 80): Brit. Mus. 41925. “A third right upper milk-molar, provisionally referred to this species; from the Pliocene near Shanghai, China. This specimen is the type of Stegodon sinensis, and is described and figured under that name by Owen in the Quart. Journ. Geol. Soc. vol. xxvi. p. 417, pl. xxvii.; it is also figured and provisionally referred to the present species [Elephas chftii] by the writer, in the ‘Palzontologia Indica,’ ser. 10, vol. i. p. 257, pl. xlv. fig. 2. There are four complete ridges, and a large anterior talon, which is reckoned by Owen as a fifth ridge. The median longitudinal cleft is very indistinct. Presented by Prof. Sir R. Owen, K.C.B., 1870.” (4) Osborn (1924) prefers to retain the name Stlegodon sinensis until further local material can be secured for comparison. Fig. 737. closely related to undetermined Siwalik specimens which he com- pared with the M. elephantoides of Clift. These two undeter- mined specimens are described by Falconer (1868, Vol. I, p. 460) as follows: Plate xxrx. a, “Tig. 5.—H. bombifrons? Fragment of molar, from lower jaw, right side, with four ridges—B.M. Length, 5.8in. Width, 4.5in. Fig. 6.—E. bombifrons? Fragment of molar with three ridges and a heel. ‘Doubtful what figs. 5 and 6 are.’— H. F. Length, 4.4. in. Width, 4.5 in.” This shows that Falconer regarded these Siwalik teeth as doubtfully related to L. bombifrons and that Owen also indirectly compared them with #. bombifrons. Stegodon bombifrons lectotype is of Middle Pliocene age. (2) One should also compare Owen’s type tooth with figure 700 above of Mastodon |Stegodon| elephantoides (=cliftii). This com- Type of Stegodon sinensis Owen, 1870, Pl. xxvur, figs. 1 and 2, natural size. Alleged to be from ‘‘marly beds” near Shanghai, China. (Op. cit., p.417): “The tooth in question is the second upper molar (d3 of the type series) from the right side. Its crown, in a length of three inches, is divided into five transverse ridges.” Falconer, Lydekker, and Martin (Table V above) ascribe four ridge-crests to Dp* of Stegodon elephantoides (=chftii), S. bombifrons, and S. trigonocephalus. Stegodon sinensis Owen, 1870. ‘On Fossil Remains of Mammals found in China.’’ Quart. Journ. Geol. Soe. London, Vol. X XVI, p. 417. Typrn.—(Op. cit., p. 417): “. . . Second upper molar (d 3 of the type series) from the right side [Dp*].” Brit. Mus. 41925. Horizon anp Locaurry.—Alleged to be “from marly beds in the vicinity of Shanghai,” China. Probably Upper Miocene [Lower? Pliocene]. Typr Ficurr.—Op. cit., Pl. xxvii, figs. 1-3. Typr Descriprion.—(Owen, op. cit., pp. 417, 418): ‘“Stegodon sinensis, Ow. The tooth in question is the second upper molar (d 3 of the type series) from the right side. Its crown, in a length of three inches, is divided into five transverse ridges, the proportions of which, as to height and basal breadth, with the ridged and THE STEGODONTINA: wrinkled character of the enamel, suffice for its reference to a species of the group of Proboscidians discovered by Crawfurd in the Irrawadi Tertiaries of Ava, and described by Clift in the second volume of the second series of the Transactions of the Geological Society (p. 369, pls. 36-39, 1828). . . . In the present tooth the first or foremost ridge (Pl. xxvit. figs. 1 & 2, 1) is defined by a cleft on the outer side of the tooth, but not on the inner side, fig. 3; here the abraded surfaces or ridges 1 and 2 are blended by wear into a common hollow field of smooth dentine (fig. 1, a). There is a slight constriction near the part where the worn surface of the first ridge blends with that of the second; and this constric- tion, which may be detected in the succeeding ridges, I take to be a trace of that stronger one which more completely divides the WAY doe 6 Schasy” lchoy 1. Taght Lower Jaw, Mastodon lakdens. 2. Lot Lower Jaw, Mastodon Liliphantndes Fouad by J Crawfurd Esq] 25 STEGODON 861 Stegodon elephantoides Clift, 1828 Figures 683, 686, 695, 696, 700, 701, 738, 7389-741 Lower Pliocene, lowest levels of the Irrawaddy Series (fluviatile),' near Yenangyaung, 250 miles below Ava, Burma. Syn.: Elephas cliftii Falconer, 1846 [=Stegodon elephantoides (=cliftit) of the present Memoir.] Speciric CHARACTERS (CuLiFtr, 1828, Osporn, 1929).—Third inferior molar, M3, with nine complete ridge-crests and a well- developed half ridge, equaling ten; five to eight conelets on each ridge-crest; length 11 inches=280 mm., est. breadth 315 inches = 90 mm.; no apparent cement. Superior molar, 1.M?! (= Falconer’s type of Hlephas cliftii) with six and a fourth ridge-crests, length 155 mm., breadth 83 mm., ten to twelve conelets on each; cement in the bottom of the valleys. Palate with 1.M?, r.M?, with six and a fourth ridge-crests; length of 1.M?, 186 mm., breadth 102 mm.; From the three specimens ridge-crests worn; traces of cement. Printed by € Fudliarsded, Nad. size ) Miles below Ava LECTOTYPE OF STEGODON ELEPHANTOIDES Fig. 738. Lectotype left Me, M3, of Mastodon elephantoides Clift, 1828, Pl. xxxviut, fig. 2, one-third natural size- (Op. cit.,p. 372 and Explanation of Plates): ‘Left side of the lower jaw of Mastodon elephantoides. The remains of the anterior molar tooth [].Me] are seen, and behind it, the posterior tooth which was advancing, and which, in consequence of the jaw-bone being broken away, is seen through its whole length. This tooth is eleven inches long and three and a half broad.” transverse coronal ridge in the molars of better Mastodons into an inner and an outer part. A well marked tubercle (figs. 1 and 2,f) projects at the outer side of the base of the first ridge, 1, near the interspace between that and the second ridge. . . . Never- theless in the number of ridges in a given tract of the grinding- surface, in their height and breadth of base, and in the absence of intervening cement, the conformity of the Chinese molar with the grinders of the Mastodon elephantoides is close. ‘The enamel also shows the same vertical linear impressions and ridges, by which we may reckon that the summit (say, of the fourth ridge in the tooth here described), if it were unworn, might be cleft into from thirteen to fifteen small mamille.’”? Owen did not compare this specimen with Clift’s original type lower molar of Mastodon elephantoides, but with specimens referred by Clift to that species. enumerated below the lectotype and cotype ridge formula is compiled as follows: M 1°% M 2°* M 3,5. Osborn, 1927: This Stegodont is similar to S. bombifrons in ridge-crest formula. Hisrory.—As fully explained above (pp. 855, 856), Stegodon elephantoides Clift is the second species of Stegodont based by Clift on a lower jaw (Pl. xxxvutt, fig. 2) and on an upper molar (PI. XXXxIx, fig. 6). Unfortunately Falconer was led to abandon ‘Mastodon elephantoides’ and to substitute ‘Elephas cliftw’; conse- quently all the literature subsequent to Clift’s original description and most of the reproductions of his illustrations appear under the specific name Hlephas cliftii, which is actually a synonym of Stegodon elephantoides. Faleoner’s mistake partly arose through Clift’s error in entitling the palate of Mastodon [= Stegodon| 'ISee note on page 824 above regarding the Lower Pleistocene age of Stegodon elephantoides.—Kditor.| 862 elephantoides (figured in Pl. xxxv1) as ‘Mastodon latidens.’ Consequently Clift’s species M. [=Stegodon| elephantoides rests upon three specimens: Lectotype: “Left lower jaw, Mastadon Elephantoides’’ (Clift, 1828, Pl. xxxvin, fig. 2=figures 696 and 738 of the present Memoir). First left superior molar, |.M', ““Upper molar of W. Hlephantoides” (Clift, 1828, Pl. xxxrx, fig. 6= figures 700, 739, and 740 of the present Memoir), sub- sequently made the type of Elephas cliftii by Falconer and Cautley. Palate, with 1.M? and r.M?, “Upper jaw of Mastodon latidens”’ (Clift, 1828, Pl. xxxv1), not figured in the present Memoir. Mastodon elephantoides Clift, 1828. ‘On the Fossil Remains of two New Species of Mastodon, and of other vertebrated Animals, Cotype: Referred: OSBORN: THE PROBOSCIDEA cene, lowest levels of the Irrawaddy Series (fluviatile).2- Lecro- TYPE Wigure.—Clift, 1828, Pl. xxxvim, fig. 2. CoryrPr I'tcurE.—Clift, 1828, Pl. xxxrx, fig. 6 [=Stegodon elephantoides (=chfti2).| REFERRED PaLatTe.—Clift, 1828, Pl. xxxv1. Currr’s OricinaL Description oF ‘MaAsropoNn’ ELEPHAN- roinrs.—We refer to Clift’s clear and consistent description quoted in full above (p. 827) together with the beautiful lectotype figure reproduced in our figure 738 herewith. Clift’s specimen comes from Burma and his figureagrees exactly with his deserip- tion, namely, M 370, with five to eight conelets on each ridge. Cotyrr or STEGODON ELEPHANTOIDES Curr (=cLirrit FALCONER) Fig. 739. Falconer and Cautley, 1846, first figured by Clift, 1828, Pl. xxxrx, fig. 6, as an “Upper molar of MW. Elephantoides.” Reproduced herewith from cast (Amer. Mus. Warren New figure of type (I.M!) of Elephas cliftii Coll. 10382) one-half natural size. Original in British Museum (Brit. Mus. M.10520). From near Yenangyaung, Burma. Vig. 740. Cotype1.M! of Stegodon elephantoides (=cliftei), after Falconer and Cautley, 1846 [1847, Pl. xxx, figs. 2, 2a], left M!', said to be the same molar (see Pal. Mem., 1868, Vol. I, p. 461) as that figured in Clift, 1828, Pl. xxxrx, fig. 6, although the drawing does not agree well with Clift’s original illustration nor with the reproduction from the cast, (Fig. 739 opposite). Observe rudimentary anterior ridge (pro-protoloph) and rudimentary seventh ridge (heptaloph), also the six ridge- crests (proto-, meta-, trito-, tetarto-, penta-, and hexalophs), the ridge-crest summits each crowned with from ten to twelve conelets. found on the left Bank of the Irawadi.” ‘Trans. Geol. Soc., London, (2), II, Pt. III, 1828, pp. 372, 373. Lucrotyrpr.—(1) Left side of lower jaw with Moe, M;; original in Museum of Geological Society of London; cast Brit. Mus. 7393 [referred by Faleoner and Cautley in the “Fauna Antiqua Sivalensis,” PI. xx, figs. 9, 9a, Pl. xx.a, fig. 6, to H. insignis].' CoTyPE.— (2) First upper molar of the left side, 1.M' (original in the British Museum (Brit. Mus. M.10520), cast Amer. Mus. Warren Coll. 10382); same tooth was selected by Falconer and Cautley as the type of Elephas cliftiz. Rererrepd (Osborn).—Palate with |.M? and r.M? (figured as Mastodon latidens by Clift). Hor- IZON AND Locauiry.—Left bank of the Irrawaddy River, near Yanangyaung, 250 miles below Ava, Burma. Lower Plio- COTYPE CHARACTERS OF STEGODON ELEPHANTOIDES (=CLIFTIT) FALCONER AND CAUTLEY, 1846, PAGE 47, BASED ON CLIFT’S SPECIMEN AND FIGURE Valconer’s designation of this type (?left M*) and of this species is as follows: “The same group comprises a fourth extinet Indian species, named in this work, #. Cliftii, which furnishes the next link in the chain of forms presented by the molars of the Klephantide. . . . In our view, the tooth represented in pl. 39, fig. 6, of Mr. Clift’s memoir in the Geological Transactions [Clift, 1828], under the name of Mastodon Elephantoides, and the palate specimen represented in pl. 36 of the same memoir, under the name of M. latidens, belong to this species. . . . The penultimate and antepenultimate molars in the upper jaw have only six transverse See Chakravarti, D. K., Quart. Journ. Geol., Mining, Metallurg. Soc. India, 1937.1, p. 34, who referred it to S. elephantoides.—Editor.] *[See note on page 824 above regarding the Lower Pleistocene age of Slegodon elephantoides.—Kditor. | THE STEGODONTINA: ridges, continuous, and chevron shaped, with numerous mammil- le, as in H. insignis and EB. Ganesa; but the cement does not fill up the interspaces of the ridges, being reduced to a comparatively inconsiderable quantity in the bottom of the hollows. HE. Cliftii, in the reduced number of the coronal ridges, and in the other characters of the teeth, appears to constitute the dental link which forms the immediate passage from Hlephas into Mastodon.” ADDITIONAL CHARACTERS OF THE COTYPE SPECIMEN OF STEGODON ELEPHANTOIDES (=CLIFTII) (Faleoner, 1868, Vol. II, p. 84): “But the detached tooth {Fig. 740] on the upper jaw is seen entire, and beautifully preserv- ed, in the specimen fig. 2 of the same plate [i.e., Pl. xxx], presenting six ridges and a small hind talon. The same tooth is represented by fig. 6 of Pl. xxxrx. of Mr. Clift’s Memoir (Geol. Trans., vol. ii. 2nd series). It is there described as an upper molar tooth of Mastodon Elephantoides, under which title Mr. Clift included specimens that are referred in our arrangement to two distinct forms. ... The Elephantine affinities of this tooth are indicated by the absence of a longitudinal line of division along the crown, and by the great number of points (about eleven in each) that enter into the composition of the ridges.” Of the same cotype molar, Lydekker (1886, p. 81) observes: ‘7388. Cast of the first (?) left upper true molar in an early stage of wear (woodcut, fig. 20). The original was obtained near Yenankhoung, on the left bank of the Irawadi in Upper Burma, by Crawfurd in 1826, and is pre- served in the Museum of the Geological Society;!!! ... There is scarcely any trace of the median cleft, the cement is slight, and there are numerous cusps. Mantell Collection. Purchased, 1836.” FALCONER’S NOTES OF 1868 ON STEGODON ELEPHANTOIDES (=CLIFTII) Falconer, ‘‘Pale#ontological Memoirs,” Vol. I, 1868, pp. 461, 462, Plate .xxx of the ‘Fauna Antiqua Sivalensis.” It is very important to observe that the four specimens described by Falconer, including the cotype of S. elephantoides from Burma, agree with each other both in the number and char- acter of the ridges and in the number of conelets on each ridge. Uprrr JAws.—Burma, left bank of the Irrawaddy River, 250 miles below Ava. Plate xxx, figs. 1, la, EF. cliftii F. & C., palate, Dp‘, ? ridges; figs. 2, 2a, type,"! Burma, 250 miles below Ava, 1.M!, ridges 6%, beautifully preserved, ‘‘six ridges and a small hind talon . . . as many as eleven to twelve denticles. . . . Its elephantine affinities are indicated by the . . . great number of points [denticles]’’; fig. 3, superb palate, M?, ridges 6}, little cement (referred by Clift, 1828, Pl. xxxv1, to M. latidens); figs. 4, da, 4b, fragment of M2, right side, 5 ridges, cement moderate in quantity, from near Yenangyaung, upper Burma. LOWER Jaws.—Burma. Plate xxx, figs. 5, 5a, 1.Ms, “eight ridges and a talon,” little cement. Coryrr CHaracters.—From the above it appears that the four specimens described by Falconer from near Yenangyaung, 250 miles below Ava, Burma, exhibit the following characters: (1) Denticles very numerous, eleven to twelve on each ridge; (2) ridges moderately high with little cement; (3) ridge formula as follows: Stegodon elephantoides (=cliftii): M 1°°*M 25" M 35x. STEGODON 863 PHYLETIC CONCLUSIONS Osborn, 1927: Falconer considered Clift’s Lower Pliocene?’ type of Mastodon |= Stegodon| elephantoides as close to his Upper Pliocene S. insignis, but S. elephantoides proves rather to be close to the Middle Pliocene S. bombifrons, as shown in the comparative ridge formule table above, the two formule abbreviated being as follows: S. bombifrons: Dp 3% Dp 4 ony Mi Mie MS S. elephantoides: Dp 33 Dp 4? M14 M24 M3 ,5. Elephas cliftii [= Stegodon elephantoides (=cliftir)], fide Vale. and Caut.: Dp34Dp42M1°4#M2°4* M3 &. Fig. 1 REFERRED STEGODON ELEPHANTOIDES (=CLIFTI) Fig. 741. Referred 1.M3 of Elephas cliftii, after Falconer, 1868, Vol. II, Pl. v, figs. 1, 2, ‘‘Views in plan and profile of last true molar, lower jaw, left side, rather less than one-fourth (two-ninths) of the natural size.’’ See also the “Fauna Antiqua Sivalensis,” Falconer and Cautley, 1846 [1847, Pl. xxx, figs. 5, 5a]. From Burma, presented to the British Museum by Colonel Burney (Brit. Mus. 14759). The ridge formula of 1.M3, namely, gg, assigned by Falconer and Lydekker, actually occurs in an aged specimen from Burma, Falconer’s figure of which (Falconer and Cautley, 1846 [1847, PI. xxx, figs. 5, 5a] is reproduced herewith in our figure 741; this grinder is so old that the two anterior ridge-crests may have worn off, consequently the ridge formula is uncertain; the dimensions (length 12.7 in. =323 mm., breadth 4.5 in. = 115 mm.) considerably exceed those of the type of Stegodon elephantoides given above. There is no substantial basis, therefore, for the assignment to the synonymous Elephas [= Stegodon| cliftii of a ridge formula inferior to that of Mastodon [= Stegodon|] elephantoides Clift. Stegodon bombifrons Falconer and Cautley, 1846 Figures 686, 687, 699, 731, 732, 734, 742-746, 777, Pl. xx Lectotype: Siwalik Hills (fide Falconer), probably from the Dhok Pathan horizon, Lower [Middle] Pliocene, India, in which this species is very abundant and characteristic (Pilgrim-Brown). 'Now in the British Museum (Natural History), M.10520 =Stegodon elephantoides (=cliftit). *[See note on page 824 above regarding the Lower Pleistocene age of Stegodon elephantoides.—Editor.| 864 This is an early Lower [Middle] Pliocene stage of the true Stegodon based on types from the Dhok Pathan zone which also contains Stegolophodon latidens. From this species some of the earlier authors believed that the genus Hlephas originated. Re- ferred Stegodon bombifrons occurs in the Tatrot horizon, Middle [Upper] Pliocene. A superior molar from the Dhok Pathan horizon (Fig. 746) exhibits very clearly the cone and conelet structure of the crown. E. [Elephas] bombifrons Falconer and Cautley, 1846. “Fauna Antiqua Sivalensis” . . . letterpress, 1846, pp. 46, 47. Lrcro- TYPE AND Coryprs.—(Op. cit., p. 46): “. . . several crania con- taining perfect teeth,” of which the lectotype is figured in PI. XXVI. Horizon aNp Locaurry.—Siwalik Hills, India, prob- ably from the Dhok Pathan horizon, Lower [Middle] Pliocene. Lecroryer AND Coryrn Ficures.—Op. cit., 1846 [1847, Pls. xxvr (leetotype), xxvu, xxvuu], figures 742 and 744 of the present Memoir. Falconer and Cautley, 1846, p. 46 (see above p. 858) based this species on ‘‘several crania containing perfect teeth,” described as “from the Sewalik Hills.’ Of these Lydekker (1886.2, p. 83) designated as the lectotype Brit. Mus. M.2978. The exact locality is not given. According to Pilgrim (Vol. I, fig. 413, also Chap. XXII below) this species occurs abundantly in the Dhok Pathan horizon of Lower [Middle] Pliocene age, a horizon which also contains Tetralophodon punjabiensis, Synconolophus corrugatus, and S. hasnoti. Falconer assigned to this species a low ridge formula, namely: M3433. CHARACTERS (FALCONER AND CauTLEy, 1846, Pp. 46); SEE OsBORN ABOVE, PP. 855-859.—(1) Type based on several crania containing perfect teeth. (2) Crown divided into transverse ridges, composed of numerous mammille [conelets] of chevron-shaped section. (3) Interspaces occupied by a thick coat of cement. (4) Principal ridges of third upper and lower molars, M 35, in contrast to [S.] insignis. (5) Third upper molar measures 11 inches (279 mm.) in length by 434 inches (114 mm.) in width. (6) The lower third molar of the left side, with nine [9}4] ridge-crests, measures 13.4 inches in length (=340 mm.) by 4.2 inches in breadth (= 105 mm.), considerably exceeding the dimensions of the lectotype of Pl. XXVI ELEPHAS BOMBIFRONS Ref. Pi. XLIIL, XIII OSBORN: THE PROBOSCIDEA S. elephantoides, namely, length 280 mm., breadth 90 mm. (7) The collective (Falconer, Lydekker, Osborn) ridge-crest formula (not including the unexposed ridge-crests) is as follows: Stegodon bombifrons: Dp 34 Dp 4 isacgueg M1 1oE 9 8-7 yy 9 74-8-81-9-915 M 23-70 M 3 secon an: FALCONER’S NOTES OF 1868 ON ELEPHAS [=STEGODON] BOMBIFRONS Falconer, ‘Palswontological Memoirs,” Vol. I, 1868, pp. 456, 458-461, Plates xxv-xxrx.B of the “Fauna Antiqua Sivalensis.”” {Errors of determination indicated by Osborn in square brackets.| Elephas bombifrons. Valeoner and Cautley, 1846 [1847, Plates XXVI, XXVII, XXVIII]. Lrecrorypr AND Coryprs.—Cranium represented in Plate xxvr (Lydekker’s lectotype); Plate xxvir, very fine skull, M%, ridges 9}, length of molar, 10.2 in.=257 mm., width 3.7 in. = 93 mm., and Plate xxvimt, fig. 2, same skull. Fig. 743. Stegodon bombifrons. Much more primitive than S. insignis. Observe transverse ridges of trunk to elevated nasals, extreme broadening of summit of occiput. Ears conjectural. Limbs given the same proportions as the Stegodonts throughout, without knowledge of skeletal material. Iemale to right, directly after lectotype skull; no inferior tusks. Male to left. Restoration by Margret Flinsch, 1930. One-fiftieth natural size. Upper Jaws.—Plate xxrx, fig. 1, broken cranium, M®, ridges 84, length of molar 10 in. =253 mm., width 4 in.=101 mm.; figs. ELEPHAS BOMBIFRONS Cotype Pi. XXVIII, Fig. 2 ELEPHAS BOMBIFRONS Ref. Pl. XLV, XIII (rev.) Srrcopon CRraAntiA, AFTER ALCONER AND CauTLey’s ILLUSTRATIONS IN THE “I'AUNA ANTIQUA SIVALENSIS” Fig. 742. Crania of Stegodon (Elephas) bombifrons, lectotype, cotype, and referred. Outlines assembled from original plate drawings of this species in Falconer and Cautley’s “Mauna Antiqua Sivalensis,” frontal, palatal, and lateral aspects. All one-sixteenth natural size. THE STEGODONTINA: 2, 2a, cranium, M°, 8 ridges; figs. 4, 4a, r.M*, 8+ ridges; figs. 5, 5a, upper jaw, r.M°, 7% ridges; figs. 6, 6a, palate with r.M', 915 ridges, length 10.9 in. =276 mm., width 3.8 in. =97 mm. (at ends), 4.3 in.=112 mm. (in middle). Lower Jaws.—Plate xxv, figs. 3, 3a, lower jaw, r.M3, 9!6 ridges, enamel very thick, scanty cement. Plate xxrx.a, figs. 1, la, lower jaw with Dp,, ridges 6%, “probably the third [fourth] milk molar”; figs. 2, 2a, lower jaw, r.My,, ridges 7}, “certainly the first true molar”; figs. 3, 3a, lower jaw, 1.Me, ridges 7}; figs. 4, 4a, lower jaw, Mz, ridges 714; figs. 7, 7a, lower Jaw, My, ridges 7. Plate xx1x.B, figs. 5, 5a, lower jaw, r.Dp, “‘with 5 ridges and an anterior and posterior talon’’; figs. 6, 6a, lower jaw, r.M,, ridges 76; figs. 7, 7a, lower jaw, |.Ms, ridges 9!5 [11+]. In the two plates (xxrx.a, xxrx.B) of the ‘T’auna Antiqua Sivalensis” about sixteen specimens of inferior molar teeth are beautifully figured; they exhibit six to ten conelets on the unworn crown and a maximum of eleven conelets on the worn crown; five of the conelets double by dichotomy. Thus the number of STEGODON 865 conelets is approximately the same as in Stegodon elephantoides (=cliftiz). From the above observations we deduce the ridge formula of Stegodon bombifrons practically as above under ‘‘Characters.”’ LYDEKKER’S NOTES OF 1886 ON ELEPHAS [=STEGODON] BOMBIFRONS “Catalogue of the Fossil Mammalia in the British Museum (Natural History),” Part IV, 1886, pp. 82-88 Lydekker (1886.2, p. 83) designates as his lectotype Brit. Mus. M.2978: “The cranium, showing the third true molars of both sides in an early stage of wear. This specimen is the type, and is figured by Falconer and Cautley in the ‘Fauna Antiqua Sivalensis,’ pl. xxvi”; reproduced in figure 742 of the present Memoir. Lydekker’s notes are based on forty-four specimens in the British Museum referred to this species, chiefly from the Cautley Fig ? CoryrE oF STEGopON BoMBIFRONS Wig. 744, Brit. Mus. M.2979; cast Amer. Mus. 10378. Irom the Siwalik Hills, India. Valconer, Pal. Mem., 1868, Vol. I, p. 458: (Pl. xxv) “Very fine and perfect skull, anterior view.” Cotype of Elephas bombifrons Falconer and Cautley, 1846. After Falconer and Cautley, 1846 [1847, Pls. xxv, xxvini], one-sixth natural size. (Pl. xxvur) Fig. 1. “Lateral view of same skull, as figured in Plate xxvu.—B. M. Tig. 2... Palate view of same skull, showing sections of tusks, and last ? true molar on either side, with 9 ridges and a heel; the 8 front ridges worn. The interval between the molars in front is very narrow; behind they are extremely divergent.—B.M.” Fig. 5. another skull. Occipital view of 866 Collection (1842); the specific references therefore are based on l'aleoner’s determinations. Speciric CuHaracturs.—(Lydekker, 1886.2, p. 82): ‘The ridges are rather taller, somewhat wider apart, and more numerous than in H. clifti [=Stegodon elephantoides (=cliftii)|, and the valleys are generally completely filled with cement; it is, however, sometimes very difficult to distinguish between the hinder teeth of the two species, while in the opposite direction it is often difficult to distinguish between those of EH. bombifrons and EF. insignis. The teeth figured by Falconer and Cautley under the name of E. ganesa cannot be distinguished from those of the present species, and are therefore provisionally classed under the same head. The teeth are frequently very large, and the ridges are often Rererrep Rigur Tarrp Superror Mouar or StEGODON BOMBIFRONS Fig. 745. size. curved; a trace of the median longitudinal cleft can often be observed in the first two or three ridges, and the inner columns of these ridges occasionally show accessory tubercles near the longi- tudinal cleft, where they assume a Mastodon-like shape. The plane of wear of the teeth of this and the following species [Z. ganesa] is similar to that of the true Elephants. The mandibular symphysis is produced into a spout-like termination, as in FE. indicus. The cranium has the fronto-parietal region very convex, the constriction of the frontals by the temporal fossee being more marked than in the other species. Hab. India (Punjab to Siwalik Hills) and (?)China [Footnote: ‘Koken, Pal. Abhand. vol. iii. pt. 2, p. 12 (1885).’]. The species may perhaps also occur in Java.” The ridge formula of Lydekker (op. cit., 1886.2, p. 82), namely, Mm. [Dp] 72-2, M $(2-2-¢-"), is less precise in not referring to the half-ridges but does not otherwise differ excepting in the supe- rior ridges (*) of Dp 3 from the type ridge formula gathered from Falconer’s observations above. Lydekker’s observations fully substantiate Falconer’s formula of 1868, and, omitting the half- A partly worn right superior molar, r.M®, referred by Falconer and Cautley to Blephas bombifrons and corresponding very closely in all details to Falconer’s description of the cotypes of this species, namely: M 3 with nine and a half ridges; conelets—six divid- ing into eleven. After Falconer and Cautley, 1846 [1847, Pl. xxrx, fig. 6], one-third natural OSBORN: THE PROBOSCIDEA ridges, enable us to write the standard maximum formula as follows: Maximum ridge formula of Hlephas {= Stegodon| bombifrons: Dp 2: Dp 3t Dp 4# M1$M2, M38. Stegodon insignis lalconer and Cautley, 1845, 1846 Figures 686, 688, 697, 731, 732, 735, 747-753, 756, 760, 764, 766, 776, 819, Pl. xx Siwalik Hills, India, Upper Pliocene, Pinjor horizon, to Lower Pleistocene, Boulder Conglomerate (fide Falconer and Pilgrim), to Upper Pleistocene. In the present Memoir (see Fig. 413) the upper levels of the Pinjor horizon are of Lower Pleistocene age (fide Barnum Brown). ‘alconer’s types of this species agree in the ridge-crest formula with the type which he subsequently described as Stegodon ganesa Fig. 746. Stegodon bombifrons collected by Barnum Brown in 1922 in the Dhok Pathan horizon, Lower [Middle] Pliocene, two and a half miles northeast of Hasnot, Referred fragment of superior molar of India. This specimen is wrongly numbered; it should read Amer. Mus. 20044. Three anterior ridge-crests of a right third superior molar, r.M® (compare Fig. 759 of Stegodon orien- talis grangeri type). but they exhibit profoundly different characters in the cranium, as shown in a comparison of figures 735 (S. insignis) and 736 (S. ganesa), or in a comparison of figures 752 (S. insignis) and 732 (S. ganesa). This profound difference, as explained above and below, is attributed to the fact that all the crania referred by Falconer and Cautley to Stegodon insignis represent small-tusked and probably female individuals, while crania referred to S. ganesa represent large-tusked and probably male individuals. Faleoner originally described Hlephas insignis in 1846, p. 37, and in the same communication (p. 45) he named a fourth species Elephas Ganesa, describing a third superior molar (which had been figured in 1845, Pl. m1, fig. 7a) and remarking that the tooth bears the closest resemblance to the corresponding tooth in Ff. insignis. This doubt always remained in Falconer’s mind, for in his notes of 1867, p. 4, and of 1868, Vol. I, p. 424, he remarked: “Tn fact, there are no good characters by which the teeth of these two species can be satisfactorily distinguished, although the erania are so remarkably different.” In Faleoner’s mind, therefore, BLlephas {= Stegodon] insignis possessed a cranium of the type he THE STEGODONTINE: figured in Pl. xv of the ‘Fauna Antiqua Sivalensis,’”’ reproduced herewith in our figure 735, in contrast to Hlephas [= Stegodon| ganesa which possessed a cranium of the type figured in PI. xx, reproduced in our figure 736. SpexvuaL Dispariry.—As explained above in the discussion of the erania, Falconer apparently selected the female crania as refer- able to Elephas |= Stegodon] insignis and the male crania as refer- able to H. [S.] ganesa. A detailed comparison of all the referred specimens has failed to establish any true specific distinction be- tween these two species. Consequently we may regard the lecto- type and referred specimens from the Pinjor, Boulder Conglomer- ate, and Godavari, Narbada Alluvium, as ‘collective species’ including a number of ascending mutations or subspecies which will be recognizable by profound monographic research; it is not STEGODON 867 Horizon anp Locauiry.—Siwalik Hills, India, probably Pinjor horizon, Upper Pliocene or Lower Pleistocene. LECTOTYPE FigurE.—Op. cit., 1846 [1845, Pl. 1, fig. 6a]. CorypE.—Op. cit., 1846 [1845, Pl. mu, fig. 60]. Description.—Falconer’s original description of 1846, p. 37, quoted in part above in this Memoir (p. 829) was restated in 1868, Vol. I, p. 423, fig. 6a of Pl. 1, as follows: “Elephas insignis, from the Sewalik Hills [I ig. 747 of the present Memoir]. Vertical section of last upper molar. The four anterior ridges are affected by wear; the six posterior ridges are entire, the fangs are fully developed, and their mode of implantation in the jaw is distinctly shown. The white mass in the centre represents the body of ivory, which is projected upwards in ten angular lobes, terminating in a sharp edge. The height of these lobes does not much exceed the width of their base, and closely Fig 6b LECTOTYPE AND CoTyPE OF STEGODON INSIGNIS Fig. 747. Lectotype left superior molar, 1.M*, of Elephas insignis Falconer and Cautley, 1846, from the Siwalik Hills, India, after Falconer and Cautley, 1846 [1845, Pl. u, fig. 6a], also cotype third inferior molar, M3, after Falconer and Cautley, 1846 [1845, Pl. u, fig. 6b]; both figures one-third natural size. (Falconer and Cautley, 1846, p. 37): ‘Fig. 6a, pl. 2 [lectotype 1.M’—Brit. Mus. M.3015], represents a vertical and longitudinal section of the last upper molar of an Indian fossil species, which we have named Elephas insignis in this work.” (Falconer, ‘Paleontological Memoirs,” 1868, I, p. 423): “The four anterior ridges are affected by wear; the six posterior ridges are natural position. entire, the fangs are fully developed, and their mode of implantation in the jaw is distinctly shown.” Length of this tooth 10.3 inches. Inverted to show Fig. 6b (cotype M3—Brit. Mus. M.3039) is a vertical section of a third lower molar of Z. insignis, one third natural size (cf. Falconer, 1867.1, p. 4; 1868.1, I, p. 424; also caption to figure 697 above). probable that a single true specifie stage passed from the Pinjor into the Narbada Alluvium horizon. In the present Memoir we shall first treat Falconer’s descrip- tions of the types and referred specimens of these two species separately and then unite them under the collective species name Stegodon insignis-ganesa. Crantum.—The lectotype cranium of the third species of Stegodont, described as Elephas insignis by Yaleoner and Cautley, 1846, is from the Cautley Collection but is without record as to its exact geologic level. The specimen is now in the British Mu- seum. Lydekker (1886.2, p. 91) designates it as “M. 3015. An imperfect cranium, showing the third true molar of either side.”’ Blephas insignis Yaleoner and Cautley, 1846. ‘Fauna Anti- letterpress, 1846, p. 37. Lecrorypr.—A third superior molar of the left side, 1.M’, in an imperfect cranium (Brit. Mus. M.3015) containing the third true molar of either side. qua Sivalensis,’’ applied over them is a thick layer of enamel reflected up and down in a continuous zig-zag plate. The interspaces of the five posterior ridges of enamel are completely filled up by a mass of cement much exceeding the enamel in thickness (vide Plate vi. fig. 7). This is the best illustration of the intermediate type of a proboscidean molar tooth, from which those of the other species diverge in opposite directions. It belongs to the Mastodon Elephantoides of Clift. The dark granulated shade below the portion of the ivory nucleus sustaining the five posterior ridges indicates the hollow of their common fang, which in the fossil is occupied by a core of sandstone.—B.M. (Reproduced in PI. ty. fig. 1.) Length of tooth, 10.3 in.” CHARACTERS OF HLEPHAS [=STEGODON] INSIGNIS.—(Lydekker, 1886.2, p. 89): “The apparent impossibility of distinguishing the dentition of this species from that of 2. ganesa has been already mentioned. ... The ridges of the cheek-teeth are usually rather dm? foreshorlened ; Game foreshartened +3, P| Ry <7 ae wy - Fig. 748. Stegodon insignis ref. (Amer. Mus. 19869) superior and inferior molars probably belonging to the same individual as the mandible shown in figure 750. Pinjor horizon, Upper Siwaliks, near Siswan, India. One-third natural size. A, Al, A2, Imperfect second superior molar, 1.M2, with +544 ridge-erests much worn; a complete third superior molar, 1.M%, with }5-8-}5 ridge-erestis, of which the two anterior are slightly worn. B, Right third inferior molar, r.Ms3, displaying 46-9 + ridge-crests, of which the three anterior are slightly worn (compare Fig. 750). The 1.Mg, now completely exposed, displays a total of 12 ridge-crests. 868 THE STEGODONTINA: taller and narrower than in HL. bombifrons, their average number greater, and the cement still more abundant. It is, however, not always easy to distinguish between the two. The third molar is usually narrower posteriorly, and the enamel frequently thinner. The taller and more numerous ridges indicate that the present species is intermediate in respect of dental characters between L. bombifrons and EH. planifrons. The adult cranium is remarkable for the great depression of the fronto-parietal region, although this feature is less marked in some specimens than in others [Footnote: ‘Compare “Fauna Antiqua Sivalensis,” pl. xliii. figs. 15, 15b.’]; but in the young cranium the contour is indistinguishable from that of the adult HL. ganesa [Footnote: ‘It is of course self-evident that these young crania (like detached teeth) might equally well have belonged to F. ganesa.’|.” REFERRED MATERIALS IN THE AMERICAN Muspum.—A typical individual (Amer. Mus. 19869—Tligs. 748, 750), in the primitive S. cnstgnis-ganesa stage, was found by Barnum Brown near Siswan (Pinjor horizon), India, in which the upper and lower jaws are fortunately associated; this specimen exhibits the following ridge-crest formula: +54(= 7) 9 1-8-6 Mi ESD yg, Also recorded from Siswan are two other specimens, namely, a second superior molar, r.M? (Amer. Mus. 19804—Tl'ig. 749), in Vig. 749. Referred Stegodon insignis (Amer. Mus. 19804), a second superior molar, r.M?, taken from palate found below the conglomerates, three miles northeast of Siswan, Upper Siwaliks, India. Ridge-crests, namely, —-7—-}s, agree closely with S. insignis, somewhat too progressive for S. bombifrons. which the ridge-crests (“"*) agree closely with the typical formula of Stegodon insignis, and a third left inferior molar, 1.M, (Amer. Mus. 19859—Fig. 753) with ;;o, ridge-crests. A juvenile right lower jaw (Amer. Mus. 19858—Fig. 751) exhibits Dp, with 7 ridge-crests; a young adult lower jaw (Amer. Mus. 19964—T'ig. 752B) shows |.M> in full use, 1.M3; coming into place; the ridge- crest formula of 1.M. is 7. Still another specimen from the same locality of Siswan is a lower jaw, also aright upper tusk complete (Amer. Mus. 19773—Fig. 766). The grinding teeth of these three individuals in the American Museum collection (see Figs. 748, 749, 750, and 753), all recorded from the Pinjor horizon, Upper Pliocene [or Lower Pleistocene], exhibit progressive variation in the ridge count, as indicated by numerals in these figures; according to Doctor Brown, it is not certain that they belong in the Pinjor horizon proper, but may have come in by erosion from the conglomerates above, STEGODON 869 A. 7. /9869 Vig. 750. Referred inferior mandible, associated with two upper teeth, of Stegodon insignis (Amer. Mus. 19869), collected by Barnum Brown in the Pinjor horizon, Upper Siwaliks, near Siswan, India; it contains both second inferior molars, Mg, well worn, also right and left third inferior molars, Ms, little worn, with 4-9+ ridge-crests; the 1.M3, after removal of the bone, displays 12 ridge-crests (compare Fig. 748, probably of the same individual). Stegodon ganesa Falconer and Cautley, 1845, 1846 Figures 686, 698, 731-733, 736, 754, 755, 757, 760, 766, 819, Pl. xx Siwalik Hills, India, Upper Pliocene Pinjor horizon, to Lower Pleistocene, Boulder Conglomerate (fide Falconer and Pilgrim), to Upper Pleistocene. In the present Memoir (lig. 413) the upper levels of the Pinjor horizon are of Lower Pleistocene age (fide Barnum Brown). This species (fide Falconer, Lydekker) exhibits the same dental characters (lig. 757) as Stegodon insignis but is distinguished by dpz STEGODON INSIGNIS Ref- crown view 4 nal. SIZE, FEY. Al Weel (2) r. Ih y. 4 Amer Mus. /9858 “A Juventle jaw, rey. % nal. sxze Tig. 751. Referred lower jaw, left aspect, of Stegodon insignis (Amer. Mus. 19964), mature adult, collected by Barnum Brown in 1922 in the upper clays below the conglomerates, three miles north of Siswan, Pinjor horizon, Upper Pliocene, India. The juvenile right lower jaw of S. insignis (Amer. Mus. 19858), collected by Dr. Brown also in the upper clays but two miles north of Siswan, summit of the Pliocene, is inserted for comparison. The ramus (Amer. Mus. 19964) exhibits a fully adult form, with abbrevi- ated symphysis, 1.Me in full use, 1.M3 coming into place. The ridge-crest formula is M 22, M3 z. One-sixth natural size. Observe Dp4 of Amer. Mus. 19858 which exhibits seven ridge-crests as compared with the typical Dps of S. insignis with seven to nine ridge-crests (ef. Table V, p. 854 above). One-fourth natural size. (/ ELEPHAS INSIGNIS Ref. ELEPHAS INSIGNIS Ref PL XU, XV. B Pi. XLII, XV. A a : ene 8 ae) \ ~ Le / & Sep ee 4 ee \ ) ( \@) Fi | — oy aes» {) ' thd! ELEPHAS INSIGNIS Ret. PI. XVI, Fig. 3 ELEPHAS INSIGNIS Ref, PI. XV. 4 neat. size B Young adult jaw y 2), HY <) LOFT) ELEPHAS INSIGNIS Ref. ELEPHAS INSIGNIS Ref. Pi, XVI, Fig. 2 Pi. XLV, XV. B rev.) Vig. 752. Referred crania of Elephas [=Stegodon] insignis. Specimens reproduced in outline from original plates in Falconer and Cautley’s “Fauna Antiqua Sivalensis,’”’ as indicated in the printed legends. Frontal aspect of four crania, lateral aspect of one cranium, and posterior aspect of one cranium. All one-twenticth natural size. Compare figures 732 and 777. 870 THE STEGODONTINA: profoundly different cranial contours, profile and facial aspects (compare Figs. 732, 752). Falconer, immediately after describing Elephas insignis (1846, p. 37), described Elephas ganesa (1846, p. 45); apparently he was very much puzzled by the exact similarity in the structure of the teeth as contrasted with the profound difference 5] in the character of the skull, for in the ‘Paleontological Memoirs’ of 1868 (Vol. II, p. 84) appears the following statement: “Regarding the specific distinetness of 2. (Steg.) Ganesa I am by no means so well assured; this species is chiefly founded on a huge cranium in the British Museum with long tusks, presented by Colonel Baker. I have not been able to reconcile the form of this cranium with either that of #. (Steg.) insignis or E. (Steg.) bombifrons; but at the same time I must confess that I have failed in tracing its dentition satisfactorily as a distinct form through different ages.” E. [Elephas| Ganesa Falconer and Cautley. ‘Fauna Antiqua Sivalensis,” letterpress, 1846, p. 45. Lecrotryprr.—A last upper molar, M*. Brit. Mus. 18489. Horizon AND LocaLity.— Siwalik Hills, India, Pinjor horizon, Upper Pliocene or Lower Pleistocene. Lrecroryre FicurEe.—Op. cit., 1846 [1845, Pl. 1, fig. 7a—see figures 698 and 757 of the present Memoir]. LrctrotyPe Drescriprion.—(Op. cit., 1846, p. 45): ‘The crown consists of ten principal ridges, with a subordinate ‘talon’ ridge in front and behind. The anterior seven ridges have their summits worn, the two in front being ground down to the common base of ivory, the tooth having been a considerable time in use.” Cuaractrers.—Falconer (1868, Vol. I, p. 424) remarks of this Fig. 754. This is the famous and oft-repro- duced referred skull and tusks of Hlephas [=Stegodon] ganesa Falconer and Cautley, 1845-1847, reproduced from Plate xx, fig. 3, of the ‘Fauna An- tiqua Sivalensis.” Same skull figures 732, 733, and 736. Original in the British Museum STEGODON 871 molar, M*: “Plate 1m, Fig. 7a.—Elephas Ganesa, a fossil Indian species. Vertical section of last upper molar. The crown consists of ten principal ridges, with a subordinate talon ridge in front and behind. The anterior seven ridges have their summits worn. A small portion is broken off at the anterior end. The disposition Fig. 753. Referred Stegodon in- signis (Amer. Mus. 19859), collected by Barnum Brown in 1922 two and a half miles south of Charnian, near Siswan, India, below the conglomerates, probably Pinjor horizon, Upper Pliocene. A third left inferior molar, 1.M3; length 291 mm., width 93 mm.; ridge- crests of M 3 yyen. One-third natural size. and relative proportions of the ivory, enamel, and cement bear the closest resemblance to those of the corresponding tooth of FH. insig- nis, and the number of ridges agrees. In fact, there are no good characters by which the teeth of these two species can be satis- factorily distinguished, although the crania are so remarkably different —B. M.... Length of tooth, 9.25 in.” Lydekker (1886.2, p. 89) erroneously selects Brit. Mus. M.3008 as the type of H. ganesa: “The imperfect cranium, showing the partially-worn third true molar of either side, the base of the left and the greater portion of the right incisor; from the Pliocene of the Siwalik Hills. This specimen (the missing portions of which have been restored in wood) is the type, and is figured by Falconer and Cautley in the ‘Fauna Antiqua Sivalensis,’ pls. xxi., xxil., xxiii., and xliii. fig. 14. It is remarkable for the enormous size of the incisors. Presented by Gen. Sir W. EB. Baker, K.C.B., 1848.” Hailmmae & Walton Lithographers Seale of Feet | (M.3008). Pal g! ELEPHAS GANESA Fig. 755. Restoration (1930-1933) of Stegodon ganesa, to a one-thirty-fifth scale, by Margret Flinsch, under the direction of Henry Fairfield Osborn. Based on Falconer’s figures of the skull and tusks [see also Fig. 733.] Vig. 756. Restoration (1934) of Stegodon insignis, by Margret Flinsch, under the direction of Henry Fairfield Osborn. All figures about one thirty- sixth natural size. Restoration based on crania figured in the “Fauna Antiqua Sivalensis” by Falconer and Cautley. Aged individual at left (Pl. xv1, fig. 1), center (Pl. xm, fig. xv.a, ef. Pl. xv), right (Pl. xm, fig. xv.B, ef. Pl. xvit, figs. 1 and 2). 872 THE STEGODONTINA:: STEGODON 873 FALCONER’S NOTES OF 1868 ON STEGODON INSIGNIS AND 8S. GANESA Taleoner, “Paleontological Memoirs,” Vol. I, 1868, pp. 428, 424, 452, Plates 1—xxv of the ‘Fauna Antiqua Sivalensis”’ Slegodon insignis. Lucrorypnr of Stegodon insignis from the Siwalik Hills. Plate 11, fig. 6a, M* with 10 ridges, the height of these ridges does not much exceed the width at the base, closely covered with a thick layer of enamel, interspaces of five posterior ridges of enamel completely filled with cement. (I alconer and Cautley, 1846.1, p. 37): “This tooth belongs to one of the forms which have been included under the name of Mast. Hlephantoides, by Mr. Clift; and which Professor Owen names ‘Transitional Mastodons.’ ” Stegodon ganesa. Luctoryrn. Plate mr, fig. 7a, M*, length 9.25 in.=235 mm.; “. . . ten principal ridges, with a subordinate talon ridge in front and behind. ... The disposition and relative proportions of the ivory, enamel, and cement bear the closest resemblance to those of the corresponding tooth of #. insignis, and the number of ridges agrees. In fact, there are no good characters by which the teeth of these two species can be satisfactorily distinguished, although the crania are so remarkably different.—B.M.”’ . Upprer JAWS AND GRINDING TEETH STEGODON INSIGNIS.—Plate x1x, figs. 1, la, upper jaw with Dp? and Dp*, Dp? with 6 ridges; figs. 4, 4a, skull, M! with 7}; ridges; figs. 6, 6a, skull with M?*, M¢ with 11}; ridges, abundant cement, length of M* 11 in. =280 mm., width 3.8 in. =97 mm. Plate x1x.a, figs. 2, 2a, palate with M*, M*; M 2 ridges 7, M3 ridges ¥-11-'. Plate xxrv, figs. 6, 6a, M! ridges 744, M? ridges \-8, ridges very high and compressed, ten points or denticles on 5th ridge. Plate xxtv.a, fig. 2, skull fragment, M! ridges 6%, M? ridges }5-7-', little cement. STEGODON GANESA.—Plate xxl, fig. 2, fine large skull, M* ridges 10}5, small talon in front. Plate xxrv, fig. 1, upper jaw, M! ridges 6); fig. 2, upper Jaw, (?) M? ridges 7; fig. 3, upper jaw, M? ridges 7}5; fig. 4, upper jaw, M® ridges 615, last ridge with 7 denticles; fig. 5, imperfect upper molar—6 ridges with 9 denticles on fourth ridge. Lower JAWS AND GRINDING TEETH STEGODON INSIGNIS.—Plate xvim.a, figs. 3, 3a, lower jaw, M3 with 124: ridges, 9 denticles on the 7th ridge; figs. 4, 4a, lower jaw with My, Me, My with 4 ridges, Mz with 7 ridges, plates very deep; figs. 5, 5a, lower jaw, 1.Ms, with 11}: ridges. Plate xx, fig. 6, 6a, lower jaw, Me with 9 ridges and front and back heel; figs. 7, 7a, lower jaw, M3 with 12 or 13 ridges; figs. 9, 9a, lower jaw, Me with 9s ridges. Plate xx.a, fig. 6, lower jaw, M27 to 8 ridges. Plate xxrv.4a, fig. 3, Me [M3] ridges 12s, length 11.5 in.=291 mm., width 4in.=101mm. Plate xxv, fig. 4, lower jaw, M, ridges 7}. STEGODON GANESA.—Plate m1, fig. 7b, M3 [?Mo»| “appears to have consisted of eight principal ridges, with a talon ridge behind, and a subordinate ridge in front. . . . It bears a close resemblance to the corresponding inferior tooth of EH. insignis in the form of the ridges, thickness of enamel, and proportion of cement.—B.M.” Plate xx.a, figs. 1, la, lower jaw, My with 5% ridges, Me with 8 ridges; figs. 2, 2a, Me with 7/5 ridges. Plate xxv, fig. 1, M3 ridges +8. Plate xxv.a, fig. 1, lower jaw, M; ridges 74+. Plate xx1x.p, figs. 2, 2a, lower jaw, Dp, ridges 7%. SKULLS OF S. INSIGNIS AND 8S. GANESA Stegodon insignis. Plate xv. ‘The cranium is seen to differ remarkably from that of #. Ganesa (Plates xxi. and xxII.) notwithstanding that the molars of the two species agree so closely. That of #. insignis is flattened at the top, elongated from side to side and singularly modified, so as to bear an analogy to the cranium of Dinotheriuwm gigantewm, while that of #. Ganesa does not differ much from the ordinary type of the Elephants.” Plate xv1, figs. 1, 2,3. “Fig. 1.—Hlephas insignis. Broken cranium, oblique antero-lateral view. Left orbit, &e., gone. This head is very cubical in form, is old, very concave in front and vertically; teeth broken. Interval between incisive sheaths deep. No tusks. A black specimen in Cautley’s collection.— B.M.” Fig. 4, skull with M*, number of ridges +104, length of M* 9.4 in.=239 mm. Plate xvu, figs. 1, 2, same skull as Plate XVI, fig. 3. Extreme length from occipital bulge to plane of molars 23 in. =583 mm., extreme width of occiput 25.5 in. = 647 mm., width of naso-maxillary opening 11.3 in. =288 mm., occipital condyles to anterior end of palate 22 in. =560 mm. See also Plate xvn, figs. 3, 4, and Plate xvi, figs. 1, 2, 3, 4. ; Summary oF Rince ForMuLa The above observations on ridge formule include teeth in different stages of development and of attrition, hence the many discrep- ancies. The adult ridge formule are indistinguishable in the two species. Consequently we may combine the ridge numbers observed by Falconer (in Murchison) in the following collective ridge formula of Stegodon insignis-ganesa. Stegodon insignis-ganesa: Dp 38 Dp 47m M17} M2 #3" M 3 18 1248-13 This collective ridge formula includes the maxima numbers only; whereas that given below by Lydekker (1886.2, p. 89, under /. insignis) includes both maxima and minima. In a collective ridge formula we include ascending mutations, i.e., successive geologic stages in the evolution of the ridges, some more primitive, some more progressive, all constituting a collective species. LyprKKER’s (1886) CoMPARISON OF STEGODON BOMBIFRONS, 5. INSIGNIS, AND 8S. GANESA “Catalogue of the Fossil Mammalia in the British Museum (Natural History), Part IV, 1886, pp. 88, 89 CHARACTERS OF EF. [=STEGoODON] GANESA.—(Lydekker, 1886.2, p. 88): “The third true molars in the type cranium of this species contain ten ridges, and thereby agree with the corresponding teeth of #. insignis rather than of H. bombifrons, a conclusion confirmed by a second cranium, in which there appear to be either ten or eleven ridges in the same tooth [Footnote: ‘See Ree. Geol. Sury. Ind. vol. ix. p. 48 (1876).’]. This close resemblance between the last molar of this form and of HL. insignis renders it apparently impossible to draw any distinction between the earlier teeth of the two forms [Footnote: ‘The majority of the teeth figured in the “Fauna Antiqua Sivalensis”’ under the name of F. ganesa have the low ridge-formula of #. bombifrons (q. v.).’], and all such teeth are therefore referred to the latter. Falconer [Footnote: ‘See ‘“Palseontological Memoirs,” vol. ii. p. 84.’] had considerable doubts as to the specific distinctness of the present form, and as the resemblance between the type cranium and the young cranium of Z. insignis (Footnote: ‘See “Fauna Antiqua Sivalen- 874 OSBORN: THE PROBOSCIDEA sis,” pl. xliii. figs. 14, 15.’] indicates that the two are closely related, it is possible that 2. ganesa may be the male form of H. insignis. The adult cranium does not differ very widely from the type of 2. ¢ndicus, although the frontal constriction is less marked.” Lydekker’s ridge formula of Hlephas [=Stegodon] insignis-ganesa, namely, Mm. [Dp] the half-ridges; it also gives a higher ridge formula to M 1 and M 2. 2.(5-6)-7 (7-8).(7-8).(9-11) . : 1) 2.5.(7-9)) M (7-1 0).(8-12).(9-13) omits Consequently the following maximum ridge formula according to Lydekker is higher than that of Falconer; this ridge formula, however, is not sustained by Lydekker’s own observations on fifty-nine specimens of the Cautley Collection in the British Museum, many of which are described and figured by Falconer. Lydekker’s combined ridge formula of Hlephas |= Stegodon| insignis-ganesa may be written as follows: Dp 23 Dp 33 Dp 45M 17M 2 Stegodon insignis-ganesa Valeoner and Cautley (1846)—Osborn (1928) Having now reviewed in detail the observations of lalconer and of Lydekker, we appear to be forced to the conclusion that Stegodon insignis and S. ganesa are respectively female and male representatives of a single ‘collective species’ which first appears in the Lower Pleistocene, Pinjor horizon, extending into the Boulder Conglomerate, while specimens at present referred to the same species occur in the Middle [Upper] Pleistocene Godavari, Narbada Alluvium, contemporaneous with Hlephas (Hypselephas) hysudricus and Palzoloxodon namadicus of the Godavari. Doubtless we are dealing with a series of ascending mutations which in time may be distinguished by valid constant specific or sna M34. presence of Stegodon airdwana in the same Kendeng-Schichten or Trinil horizon [Middle Pleistocene] of Java with Palxoloxodon hysudrindicus, which, according to Stremme (1911) and Janensch (1911) is comparable to Elephas antiquus (i.e., P. namadicus). See note on the fauna of the Kendeng-Schichten below. In summation, the phylum Stegodon insignis-ganesa of India and the phylum Stegodon airdwana (=javanicus)' of Java repre- sent the last surviving Lower and Middle Pleistocene [to Upper Pleistocene] members of the southern forest-browsing Stegodonts. These two Stegodont phyla were geologically con- temporaneous with the forest- and plains-browsing Loxodonts, and with the grazing ancestral true elephants, such as Archi- diskodon. Sectrronep Mo.uars OF STEGODON GANESA AND STEGOLOPHODON LATIDENS Fig. 757. 1846 [1845, Pl. m1, fig. 7a], less than one-half natural size. bombifrons). From Siwalik Hills, India. subordinate ‘talon’ ridge in front and behind. down to the common base of ivory, the tooth having been a considerable time in use.” (Left) Vertical section of lectotype left third superior molar, 1.M%, of Blephas [=Stegodon] ganesa Falconer and Cautley, Brit. Mus. M.18489 (erroneously referred by Lydekker, 1886.2, p. 84, to 2. (Falconer and Cautley, 1846.1, p. 45): “The crown consists of ten principal ridges, with a The anterior seven ridges have their summits worn, the two in front being ground Molar inverted to show natural position. (Right) Vertical section of two last upper molars of the right side, r.M?*, of Mastodon [=Stegolophodon| latidens Clift, after Falconer and Cautley, 1846 [1845, Pl. m1, fig. 8], less than one-half natural size. From Irrawaddy River, Burma. Compare figure 719 above. Xxxvil, fig. 1. subspecifie characters; meanwhile we regard these ascending mutations as a collective species Stegodon insignis-ganesa. ‘The collective ridge formula is probably as follows, indicating as minima the ridge-crests characteristic of the more primitive stages and as maxima the ridge-crests characteristic of the more progres- sive stages so far as known: Dp 23 Y-1 1-46 M 3 wa 1-}i-1 2-1 214-13" Dp 352 Dp4—aes MIs; S. insignis-ganesa: M 9 ¥o-7-Y4-Y4- 8 Y-9-Ya The survival in India of referred Stegodon insignis-ganesa in the same Middle [Upper] Pleistocene levels with the true elephants and Loxodonts, such as Elephas (Hypselephas) hysudricus and Paleoloxodon namadicus, is paralleled in the East Indies by the Same molars as those figured by Clift in 1828, PI. Stegodon insignis birmanicus Osborn, 1929 Figures 758, 760 Mingoon, opposite Mandalay, Burma; upper levels of the Irrawaddy Series, Upper Pliocene.” This left ramus (Amer. Mus. 20002), collected by Barnum Brown in 1922, agrees with Stegodon insignis in the number of the ridge-crests of the third molar but greatly exeeeds this typical Siwalik species in size; the jaw and inferior grinder appear to be one of the largest of the Stegodonts heretofore described (Figs. 758 760); the tooth indicates a higher degree of specialization than the typical S. 7nsignis-ganesa and a specialization different from that of S. orientalis grangerz; the S. insignis birmanicus molar is of gigantic Synonym of Stegodon airawana or S. trigonocephalus (see footnote on page 889 below).—Editor.] *[See note on page 824 above regarding the Lower Pleistocene age of the upper levels of the Irrawaddy Series.—Editor. |] THE STEGODONTINA: size, there are many ridge-crests, far apart, with a medium number of conelets, and considerable cement; the S. orientalis grangeri molar is of medium to large size, there are many ridge-crests, approximated, many conelets, and moderate development of cement. Stegodon insignis birmanicus Osborn, 1929. “New Eurasiatie STEGODON 875 second pair, are partly worn, ridge-crests slightly open; this is followed by ridge-crests pentalophid to octalophid (Sth to Sth), partly worn; conelets still separate; the ninth to twelfth ridge- crests are entirely unworn. The conelets are few and stout; they vary in number from four to twelve on each ridge-crest. is present all the way back. Cement Vig. 758. inferior molar, 1. Mg, in the same ridge-crest stage as Slegodon insignis-qanesa. Mandalay, Burma. and American Proboscideans.’? Amer. Mus. Novitates, No. 393, Dee. 24, 1929, pp. 15, 16. Horizon anp Locauiry.—Upper Pliocene [now regarded as of Lower Pleistocene age]. Mingoon, opposite Mandalay, Burma. Tyrr Ficgurr.—Op. cit., 1929.797, p. 16, fig. 16. Typr.—Amer. Mus. 20002, a very large and massive left inferior jaw containing the left third inferior molar, 1.M3. The ridge-crests of 1.M3, namely, ;52;, are the same in number as in Stegodon insignis, but the elongation of this inferior molar and the open character of the ridge-crests are quite distinctive from S. insignis; the jaw is more massive and the inferior grinding teeth surpass in length measurement those of any other Stegodont type known; the grinders are larger and exhibit fewer conelets. The principal measurements (in millimeters) are as follows: Total length of third inferior molar, 1.M; At the crown 333 At the base 362 Total maximum breadth of 1.Ms, 97 Breadth-length index of 1.M; 29-27 Length of mandibular ramus from base of |. Ms, to tip of symphysis 560 Horizontal thickness of mandibular ramus 204 Height of ramus at symphysis 208 In detail, the entire jaw and posterior grinder are very mas- sive. Of the total of 12—s ridge-crests, the anterior pair are deeply worn and widely open; the tritolophid and tetartolophid, or Type of Slegodon insignis birmanicus Osborn, 1929 (Amer. Mus. 20002), a twelve and a half crested third left One-third natural size. Irom Mingoon, opposite Stegodon orientalis grangeri Osborn, 1929 Figures 682, 684, 686, 687, 731, 759-764, 777, 1231, Pl. xx Upper Pliocene [Lower Pleistocene] of Yenchingkou, near Wanhsien, Province of Szechuan, China. The subspecies Slegodon orientalis grangeri, as shown in fig- ure 687 is more primitive than the type of S. orientalis, also from a cave in Szechuan; the ridge-crests are less elevated and wider apart at the base and seem to be even more primitive than those of the S. znsignis type (Vigs. 688, 747); the cranium is much smaller and simpler than that of S. 7nsignis-ganesa (Vigs. 735, 736) and resembles in its contour rather that of S. bombifrons (Figs. 132, 134, 777). Stegodon orientalis grangeri Osborn, 1929. ‘New Eurasiatic and American Proboscideans.”” Amer. Mus. Novitates, No. 393, Dec. 24, 1929, pp. 16 and 17. Typr.—Amer. Mus. 18714, a left third superior molar, 1.M*, and right and left third inferior molars, r.M3, 1.Ms3, of the same individual. Horizon AND Locauiry.—Upper Pliocene [now regarded as of Lower Pleistocene age]. Yenchingkou, near Wanhsien, Province of Szechuan, China. Typr Fraurn.—Op. cit., Osborn, 1929.797, p. 16, fig. 16. Typr AND RererrepD Fiaures.—Aside from the type (Figs. 759, 760, 762) there is very abundant referred material from the same locality, including the complete superior and inferior den- tition, Dp 2-Dp 4, M 1—M 8, in various stages of attrition and dental succession, as illustrated in figures 687, 759, 763, 761, 762 of the present Memoir, constituting the most complete material of dental suecession known in the fossil Proboscidea. STEGODON ORIENTALIS GRANGER! PERMANENT MOLARS gy nal. Size A. 17. (8636 A./7. (8642 Z m2 ZmZ 7 7p (ren) 77m 7 (rev) 9+ crescs GS crests A./7. 1871/4. Type outer side enner stde A./7 /87/4 Zmz +73 crests STEGODON ORIENTALIS GRANGERI MILK DENTITION % 7aZ. szze A171. 18711 TOPE - Zdox doz A.M. /86302a Fig. 759. Type (Amer. Mus. 18714) and referred specimens (Amer. Mus. 18705, 18711, 18630a, 18636, 18642) of Stegodon orientalis grangeri, part of the collection from Yenchingkou, Province of Szechuan, China, made by Walter Granger during the winter of 1920-1921. All figures one-fourth natural size. (Cf. Pigs. 761 and 762 for crown views, giving ridge-crests and conelets.) The specimens bearing number 18642 have been sent to Peking, China, in exchange. Left lateral aspect of: R.Dp’, r.Dp* (rev.)—1.Dp», 1.Dp3 (Amer. Mus. 18705) L.Dp*, ].Dp* (Amer. Mus. 18711), and 1.Dp4 (Amer. Mus. 18630a) L.M}, r.M, (rev.) (Amer. Mus. 18636) L.M?, also r.Me (rev.) (Amer. Mus. 18642) L.M®, 1.M; (Amer. Mus. 18714)—type. 876 Crown view of: Type third left inferior molar, 1.M3, strongly concavo-convex, out- wardly arched. Amer. Mus, 18714. Type third left superior molar, 1.M®, strongly convex, parallel sides. Amer. Mus. 18714. THE STEGODONTINA:: STEGODON MATERIALS OF STEGODON ORIENTALIS GRANGERI, TYPE AND Rererrep.—Collected by Walter Granger in 1920-1921, all from the same pit; estimated specimens: Separate crania, more or less complete 6 Separate crania, more or less fragmentary a Right and left mandibular rami and complete in- ferior jaws 28 Separate superior and inferior grinding teeth in various stages of succession 49 Skeletal bones, very few, not exceeding 3-4 CHARACTERS.—An ascending mutation or a subspecific stage regarded as somewhat more primitive than the type of Stegodon Upper Miocene [Pliocene] stage. Owen’s type of Stegodon sinensis Brachyodont. Lophs compressed, far apart. Compare Stegodon bombifrons. Amer. Mus /87/4 Type STEGODON ORIENTALIS GRANGER) Amer /7us. 200. STEGODON INSIGNIS BIRMANICUS STEGODON INSIGNIS — GANESA Lower Pleistocene stage. Osborn’s type of Stegodon orientalis grangeri Subhypsodont (first stage). Lophs more compressed, more elevated. Compare the typical Stegodon orientalis. 877 orientalis Owen (Fig. 769) which is recorded from a cave in the Province of Szechuan, northwest China, and much more progres- sive than Owen’s type of Stegodon sinensis which appears to be comparable to S. bombifrons. The ridge-crest formula, beautifully shown in figure 759, is as follows: Dp 2*3" Dp3 24%, Dp4*3# M3 4-11-46 M1*3~ M2or0% M3 4avg COMPARISON WITH OTHER CHINESE STEGODONTS.—On closely comparing Owen’s types of Stegodon sinensis (Figs. 687, 702) and S. orientalis (Figs. 687, 769) with the Stegodon orientalis grangeri teeth (Figs. 687, 759, 761) collected by Doctor Granger at Yen- chingkou, it is certain that we have to do with three distinct spe- cific and subspecifie stages broadly distinguished as follows: (?) Lower Pleistocene stage. Owen’s type of Stegodon orientalis Subhypsodont (second stage). Lophs still more compressed, still more elevated. Compare the referred Stegodon orien- talis and S. airdwana. “reg LY nat. Suze wnner View Lmz inner view es Tez ComMPaARrIsON OF STEGODON INSIGNIS-GANESA, S. INSIGNIS BIRMANICUS, AND S. ORIENTALIS GRANGERI Vig. 760. Left third inferior molars, to a one-third scale, from the internal aspect, showing the eruption line of the grinder and exposing the six anterior ridge-crests in Stegodon orientalis grangeri (upper) and S. insignis ganesa (lower). After Osborn, 1929.797, p. 16, fig. 16. Stegodon orientalis grangeri, type (Amer. Mus. 18714). buried in the jaw. Large size with 13+ ridge-crests of which 6+ anterior were erupted while 7-13 were Stegodon insignis birmanicus, type (Amer. Mus. 20002), with 5-6 ridge-crests erupted, 6-12 buried in the jaw and encased in cement (dotted). Stegodon insignis-ganesa ref. (Amer. Mus. 19869), with +-1-6 ridge-crests erupted, 7-11 buried in the jaw, mostly lacking cement. This comparative figure illustrates the gigantic size attained; the largest (S. insignis birmanicus) slightly exceeds in length the correspond- ing grinder of S. orientalis grangeri. we ‘ee ie 3 s a Fig. 761. Superior and inferior second, third, and fourth premolars and first molar; (Dp 2 —M 1) of Stegodon orientalis grangert (Amer. Mus. 18636, 18711, 18630a, 18705), side and crown views, one-half natural size (cf. Fig. 759). (Upper pair). First superior and inferior molars (Amer. Mus. 18636), 1.M! with +6-+ ridge-crests, conelets 10 maximum, r.M, (rev.) with 8 ridge-crests, conelets 9. (Middle pair). Fourth superior and inferior deciduous premolars, 1.Dp* (Amer. Mus, 18711) with 3-615 ridge-crests, conelets 12-13, 1.Dp4 (Amer. Mus. 18630a) with 7—!4 ridge-crests, conelets 12 maximum. (Lower pair). Second and third superior and inferior deciduous premolars (Amer. Mus. 18705), r.Dp” (rev.) with +-3-+ ridge-crests, r. Dp* (rev.) with 6+ ridge-crests, conelets 15 maximum, 1.Dp» with 2 ridge-crests, 1. Dp3 with 6+ ridge-crests, conelets 16 maximum. 878 Vig. 762. Type and referred superior and inferior molars of Stegodon orientalis grangeri. Crown views (cf. Fig. 759). After retouched photographs, one-half natural size. (Upper pair) Type (Amer. Mus. 18714), left third superior molar, 1.M3, with -++-1145 ridge-crests. Left third inferior molar, 1.Ms, with 4-13 ridge-crests; inner side strongly convex, outer side strongly concave, in contrast to the parallel sides of the upper molar. (Lower pair) Amer. Mus. 18642. Left second superior molar, 1.M2, with +8-+ ridge-crests. Right second inferior molar, r.Mg (rev.) with 9+ ridge-crests. These molars have been sent to Peking, China, in exchange. 879 880 OSBORN: CRANIAL CHARACTERS OF STEGODON ORIENTALIS GRANGERI (Figs. 759, 761, 763, 777) The infantile cranium of Slegodon orientalis grangeri (Amer. Mus. 18638, 18702), as represented in the composition drawing (Vig. 763), exhibits the small deciduous superior incisors and the deciduous premolars, Dp 2-4 (for crown views of these deciduous THE PROBOSCIDEA to lower grinders, Mj-2, with ridge-crests 8 and 9 respectively (cf. Vigs. 759, 762 lower); these ridge-crests, partly bathed in cement, of moderate hypsodonty, exhibit the characteristic convex coronal curvature of the upper ridge-crests, the concave coronal curvature of the lower ridge-crests, a feature much more strongly marked in the third molars (Fig. 759). In the mature adult cranium (Amer. STEGODON ORIENTALIS GRANGERI Resse Pie eS. iis Ee Re ve fi aS | at : ‘ ‘ ‘ JUVENILE a \ a EH \ — C4 ‘ AM.18702 a : a7 , P \ 4 (l( s YOUNG ADULT a : MATURE ADULT /“ / N A174. 1/8630 < Awwieyos | i \ J : 1 \ ‘ A/./8705, /8640 lig. 763. Infantile, juvenile, young adult, mature adult crania showing eruption of Dp*—M?®, Dp3—Ms. Young adult cranium (Amer. Mus. 18630) showing M! and M?. nacesee ‘6. fy Le Yenchingkou, Province of Szechuan, China. Infantile cranium (Amer. Mus. 18638) showing succession of Dp?—Dp’, also Di’, together with jaw (Amer. Mus. 18640) belonging to this skull, showing Dp2—Dps. Juvenile cranium (Amer. Mus. 18702), jaw (Amer. Mus. 18711), showing eruption of Dp3, Dp4. Jaw (Amer. Mus. 18636) with Mo» in situ, sent to British Museum. Mature adult cranium (Amer. Mus. 18708), jaw (Amer. Mus. 18629—sent abroad in exchange), showing M?, M*, and Msg in situ. premolars, compare Fig. 761); the rounded profile, of subtriangu- lar form, should be compared with the infantile cranium of Mas- todon acutidens (Vig. 131) and that of Hlephas indicus (Fig. 799). The juvenile cranium (Amer. Mus. 18702, skull, 18711, jaw— a composition drawing) has lost the deciduous incisors, also Dp 2, but retains the functional Dp 3—Dp 4 (ef. Figs. 759 and 761). The young adult cranium (Amer. Mus. 18630, skull, 18636, jaw) exhibits the rounded and greatly elevated dome of the brain case; all the deciduous teeth, Di?, Dp 2-4, have disappeared and the grinding function is now assumed by the first and second true molars, M 1-M 2 (ef. Figs. 761, 763). It would appear that the grinders M'”, with ridge-crests 6 and 8 respectively, are opposed Mus. 18708, skull, 18629 jaw) the second superior molar, M?, is disappearing, while the third superior and inferior molars, M®’, Ms, are fully functional, their reciprocal convexo-concave relations being beautifully shown in figure 763 (right). The upper portion of this mature adult cranium is fractured, leaving the exact profile contour in doubt. The third superior and inferior molars of this subspecies grandly represented in the type specimen (Amer. Mus. 18714) seen in both lateral and crown views in figures 759 and 762, add a great deal to our knowledge of the Stegodont dentition and show very marked mechanical contrasts between the superior and in- ferior grinders as follows: THE STEGODONTINA: STEGODON 881 Tuirp Superior Morars A very careful study of these four ontogenetic stages of the Lateral profile strongly convex. cranium, namely, infantile, juvenile, young adult, and mature Posterior ridge-plates elongated, partly or wholly bathed adult, reveals resemblances and contrasts with the Mastodon and in cement. Elephas erania which will probably prove of phyletie or generic Sides of crown parallel. value. Similarly, the remarkable ridge-crest succession, the mechanical warping of the superior and inferior grinders, the Turrp Inrertor Monars gradual increase in height (hypsodonty) of the superior ridge-crests Lateral profile horizontal. from Dp 2-M 3, indicate that the highest and most progressive Anterior and posterior ridge-plates of uniform height; ridge-crests are reserved for the extremely adult stages in which the cement chiefly in median plates. posterior ridge-crests of the eleven and a half crested third superior Sides of crown strongly arched, convex inner side, concave molar grind against the less hypsodont ridge-crests of the thirteen outer side. crested third inferior molar. STEGODON PINJORENSIS Amer. us. 19772 Jype STEGODON AIRAWANA Vig. 764. Comparative sections of third left superior molars of Stegodonts of India, Java, and China, all to a one-third scale. A, Stegodon pinjorensis Osborn, type. A 14+ ridge-crested 1.M*, from near Siswan, India. Amer. Mus. 19772. B, Stegodon insignis Falconer and Cautley, lectotype. A 11 ridge-crested ].M’, from the Siwalik Hills, India. After Faleoner and Cautley, 1846 [1845, Pl. 1, fig. 6a]. Brit. Mus. M.3015. C, Stegodon airawana Martin, ref. Portion of an 1.M® of the Middle Pleisto- cene of Trinil!, Java, exhibiting 7 of the 12-14 ridge-crests; much more recent in structure than the molars of Stegodon insignis of the Lower Pleistocene. ‘This tooth was presented to the American Museum by the Geological-Palacontologi- cal Institute and Museum of the University of Berlin, Germany, through the courtesy of Geheimrat Pompeckj and Doctor Dictrich. Amer. Mus. 22636. D, Stegodon orientalis grangert Osborn, type. An 1.M8 with +11 }5 ridge-crests, from near Wanhsien, Province of Szechuan, China. Amer. Mus. 18714. Amer: Mus. 19772 One-twelfth natural size. s nat. stze Fig. 765. Type of Stegodon pinjorensis (Amer. Mus. 19772) from three miles north of Siswan, India. Compare Osborn, 1929.797, p. 17, fig. 17. A, Right lateral view of skull, with r.M® in situ, ridge-crests 14}5-15. A1, Palatal view, showing both right and left third superior molars. A2, The top view of this skull is an orthogonal projection on the plane indicated by the arrow-tipped line shown in side view (A.) A3, Posterior view of skull. ¢ a I> di B STEGODON GANESA Ref (Rev) After Falcorer STEGODON GANESA ef Amer Mus. 19773 Me Natural haf Fig. 766. Referred lower jaw and right superior tusk (A) of Stegodon ganesa (Amer. Mus. 19773), collected by Barnum Brown in 1923 in the Upper Original in British Museum (Brit. Mus. M.3008). Both figures one-sixteenth natural size. Siwaliks, below the conglomerates, three miles north of Siswan, India, compared with skull and tusks (B), after Falconer, 1846 [1847, Pl. xxt1, fig. 3]. 882 THE STEGODONTIN#: Stegodon pinjorensis Osborn, 1929 Figures 711, 731, 764, 765, 767, 768, 777, 817, 1217, Pl. xx “Three miles north of Siswan, India.’’ Upper levels of the Pinjor horizon, Lower Pleistocene. Stegodon pinjorensis Osborn, 1929. “New [Eurasiatic and American Proboscideans.”” Amer. Mus. Novitates, No. 393, p. 18. Typr.— ‘Amer. Mus. 19772. A male cranium, rostrum wanting; portions of right inferior tusk preserved.” Horizon AND Locauiry.—“‘Recorded by Barnum Brown as follows: ‘Skull. Just below Conglomerate beds, Upper Siwaliks, three miles north of Siswan, India.’ This progressive cranium does not belong in the Pinjor (Upper Pliocene) horizon, as the specifie name pinjorensis suggests, but was probably deposited from the overlying Boulder Conglomerate beds of Lower Pleistocene age.” Tyre Fraure.—Osborn, 1929.797, p. 17, fig. 17. Tyrer Descrierion.—‘‘Superior grinding teeth distinguished from those of Stegodon insignis-ganesa by their superior size, much more numerous ridge-crests, progressive hypsodonty; the comparative ridge formule of M 3 are as follows: a ee : 44-15 Stegodon pinjorensis: M 3 **%*° Stegodon insignis birmanicus: M 3 yea; : Ss Y-11-%4 Stegodon insignis-ganesa: M3 yor This type male cranium resembles that of the male type of Stegodon ganesa Fale., namely, with small rounded _parieto- YE Smash. ag Fig. 768. STEGODON 883 occipital crest, lofty and greatly abbreviated frontonasal surface, anterior nares correspondingly elevated, grinding surface of the large molars very strongly arched, but the cranium is relatively more depressed or bathycephalic than in S. ganesa.” Ye Naturat size S. PINJORENSIS Type A./4 19772 S. GANESA Rex After Faiconer Fig. 767. Type skull (Amer. Mus. 19772) of Stegodon pinjorensis (A), also skull of same individual (A) superimposed (dotted lines) on referred skull of S. ganesa, after Falconer (B). One-sixteenth natural size. Observe the extreme bathycephaly in the skull of Stegodon ganesa in comparison with the more typical Stegodont form, as exemplified by the skull of S. pinjorensis. Front view of Stegodon pinjorensis before present arrangement of the tusks, which now are regarded as possibly turning inward, as in new restoration of Stegodon ganesa (see Fig. 733, p. 857 above); otherwise head true to present proportions. TF orelimbs entirely conjectural, drawn in pro- portion to size of head. Ungues 5 and 5. Restoration by Margret Flinsch, May, 1930. One-fiftieth natural size. Stegodon orientalis Owen, 1870 Figure 687, 703, 769, 770 Szechuan, northwest China, near the city of Chungkingfoo. Swinhoe Collection of 1870. The imperfectly known Stegodon orientalis, as shown in com- parison of figures 687, 688, 697, and 769, is apparently more progressive, with more elevated ridge-crests, than the type of S. insignis (Fig. 697); the ridge-crests (Fig. 769) are more acute and more compressed at the base, and the summits are more closely approximated; with numerous conelets and strong layers of cement in the valleys. It is, however, impossible clearly to define and separate this species until further material is found in the type locality. It is somewhat more progressive than S. orzentalis grangert. This species described by Owen from an imperfect type figure (Fig. 769) is regarded by Lydekker (1886.2, p. 97) as a synonym of Probably Lower Pleistocene. OSBORN: THE PROBOSCIDEA city of Chung-king-foo, in the province of Sze-chuen.’”’ TYPE FigurEe.—Op. cit., Pl. xxvii, figs. 1-4. Tyrer Description.—(Op. cit., p. 421): “The dentine retains its original white colour, . . . the enamel also has its recent pearly tint; a thick mass of cement appears to have been retained in the intervals of the coronal ridges. One of these ridges, with the con- tiguous halves of two others, form a molar two inches nine lines in breadth (Pl. xxvin. figs. 1 & 2); a portion of a posterior ridge with a low basal heel, from the same, or the same-sized tooth, and the last two ridges, with a terminal half ridge or talon, of a milk- molar, one inch and a half in breadth (ib. figs. 3 & 4), represent the present species. ... The condition of the fragments agrees with the statement, viz. that they were from a cavern. .. . The ridge (ib. fig. 1) a a runs straight, or nearly so, across the tooth; the entire ridge is cleft at the summit into about a dozen mamille by as many vertical grooves; the dentine rises into the base of each 3 Pig. 769. Original in the British Museum (41926-7). Vig. 3. ‘Hind end of milk-molar, d 3, grinding-surface.” Tig. 4. Type of Stegodon orientalis Owen, 1870, from a cavern in Szechuan, northwest China. (Op. cit., p. 433, fig. 1): “Portion of true molar, grinding-surface.” Fig. 2. Same, ‘“‘side view.” After Owen, 1870, Pl. xxvun, figs. 1-4. Same, “side view.” Locauiry.— (Op. cit., p. 421): “These fragments form part of the scries of teeth obtained by Mr. Swinhoe, and said to be ‘from a cave, near the city of Chung-king-foo, in the province of Sze-chuen.’ The condition of the fragments agrees with the statement, viz. that they were from a cavern. . land—he believed, on the course of the Yang-tse-kiang.”” Stegodon insignis (see citation below). Koken (1885) also regarded S. orientalis as a synonym of S. insignis, Schlosser (1903) agreeing with Koken. In Osborn’s opinion, the imperfect character of the type molar (lig. 769) and associated milk molar renders it difficult to determine whether these specimens are in the Stegodon insignis stage; in both specimens the lophs are elevated. From another cave locality, 140 miles distant, near Yenchingkou, comes the superb material collected by Granger, which proves to be somewhat more primitive than Owen’s type of Stegodon orientalis and is described as Stegodon orientalis grangeri (see p. 875 above). Owen’s type description is in part as follows: Stegodon orientalis Owen, 1870. “On Fossil Remains of Mam- mals found in China.” Quart. Journ. Geol. Soc. London, Vol. XXVI, Pt. 1, p. 421. Typer.—Molar fragments (Brit. Mus. 41926-7). Horizon AND LocaLity.—(Op. cit., p. 421): “... obtained by Mr. Swinhoe, and said to be ‘from a cave, near the .. [p. 434] Mr. H. Woodward stated that Mr. Swinhoe had himself obtained a series of these fossils from a cave many miles in- mamilla. The enamel (e) averages two lines in thickness.” “From the above-defined characters it is plain that we have here, also, parts of a ‘transitional Mastodon,’ in other words, a species of Stegodon, Vr. In the straight, or nearly straight, direction of the coronal ridges, and the absence of any trace of mid cleft, these molar fragments more resemble the teeth of Stegodon Cliftii, St. insignis, and St. ganesa of Falconer than does the St. sinensis; and in the apparent quantity of coronal cement (ib. fig. 2c) as well as in the evidence of a hinder talon (ib. fig. 34), they are more like St. insignis than St. Clifiii. Yet the two hinder ridges, with the terminal talon of the tooth (ib. figs. 38 & 4), which, in breadth, corresponds with the second upper deciduous molar of St. insignis and St. sinensis, clearly differ from both. The last two ridges run straighter across, are of the same extent, and are divided by more numerous vertical grooves into smaller and correspondingly numerous apical mamille. The second of THE STEGODONTIN&: these ridges is cleft in the middle. From the alleged conditions of discovery, and the little-altered condition of the above-described portions of proboscidian molars, one would be led to deem them to be of as comparatively recent geological age as our ordinary British Cave-fossils.” LYDEKKER’S NOTES OF 1886 ON STEGODON ORIENTALIS “Catalogue of the Fossil Mammalia in the British Museum (Natural History),” Pt. IV, 1886, p. 97 Lydekker (1886.2, p. 97) treated this species as follows: “(Brit. Mus.] 41926-7. The last two ridges and talon of an un- worn fourth lower milk-molar and portions of two other cheek- STEGODON Stegodon airawana Martin, 1890 Figures 686, 688, 707, 731, 764, 771, 773, 774, 777, 779, Pl. xx Kendeng-Schichten, Pithecanthropus erectus zone, Middle(?) Pleistocene, Alas-Tuwa, Trinil, Java. This is the most progressive Stegodont known, surpassing Stegodon orientalis and greatly surpassing S. insignis-ganesa in the elevation (hypsodonty) and number of the ridge-crests. As shown in figures 687 and 688, the fourteen to fifteen ridge-crests of the third right inferior molar, r.M3, are almost columnar in side view and much more elevated than in S. orientalis (Figs. 687, 779, and 769). In front view the cranium (Figs. 773, 777) resembles that of 10 +43 Fig. 770. RrrERRED STEGODON INSIGNIS(?) =ORIENTALIS(?) AND TyPE OF SERRI- DENTINUS LYDEKKERI. AFTER SCHLOSSER, 1903, Tar. xtv, Fras. 7-10 Fig. 7. Tooth referred by Schlosser to Mastodon aff. latidens Clift, from the rothe Thone =Schansi; fig. 10, a left inferior molar, ?M3, from ?Fokien, referred by Schlosser to Stegodon insignis. Fig. 8. Type of Mastodon [ = Serridentinus] lydekkeri Schlosser, from the rothliche Sande = Tientsin, Honan, north China, presumably a left M? (cast Amer. Mus. 10374) ; fig. 9, supposed inferior premolar referred by Schlosser to Mastodon lydekkert { = Serri- dentinus lydekkeri of the present Memoir]. teeth, provisionally referred to the present species. These specimens, which were obtained from a cavern in Sechuen, north- west China, are the types of Owen’s Elephas (Stegodon) orientalis, and are described and figured by him under that name in the Quart. Journ. Geol. Soe. vol. xxvi. pl. xxviii. figs. 1-4. They show, however, no characters by which they can be distinguished from the teeth of the present species [i.e., Stegodon insignis], as the writer has already observed in the ‘Palewontologia Indica,’ ser. 10, vol. i. p. 269. Purchased from R. Swinhoe, E'sq., 1870.” S. trigonocephalus (Fig. 776) and differs widely from that of S. insignis-ganesa or S. bombifrons. This is a dwarfed insular form, very progressive in molar structure. History.—This species was named by Martin in 1890 and regarded by Janensch (compare Janensch, 1911, p. 171, fig. 12, also Taf. xx, fig. 3) as differing in skull structure both from Stegodon insignis and S. ganesa while similar in jaw structure. The assigned ridge formula is; Dp 32 Dp 44 M 14M 2*M3*4,>. Dubois in his Trinil-Fauna (1908, p. 1256) observes that this 886 species of Stegodon is highly characteristic of the Trinil Pzthecan- thropus beds, also that the third lower molar rises to fourteen ridges, i.e., M 3 rz, and is more progressive than that of insignis. This would tend to place Pithecanthropus erectus as of Middle Pleistocene age. Dietrich (letter, March 10, 1924, and notes), as cited above in this Memoir (p. 813), on morphological grounds, regards this Javanese species as more recent than any of the known continental species of Stegodonts. Stegodon Airdwana Martin, 1890. ‘Ueber Neue Stegodon- Reste Aus Java,” Verh. Kon. Akad. Wetensch. Afdeel Natuurk., Amsterdam, Deel XXVIII, p. 4. Typr.—Incomplete man- dible with third molar of either side in place. Horizon AND Locatiry.—Kendeng-Schichten, Pithecanthropus erectus zone, Middle(?) Pleistocene, Alas-Tuwa, ‘Trinil, Java. TYPE Ficure.—Martin, 1890, Tab. 1, figs. 1 and 2 (mandible), also Tab. 11, figs. 3 and 4 (type Ms). Tyre DescripTion.—(Op. cit., p. 4): “‘Nur von Java in einer unvollstéindigen Mandibel und darin steckenden Molaren bekannt. Letztere mit 9 Jochen und 2 Talons. Durch den mastodonartigen Charakter der Kronenspalte und die geringen Cementmengen schliesst sich die Art an St. Cliftzi und St. bombifrons, durch die hdheren Joche an Sf. insignis und St. ganesa an; sie nimmt somit, gleich dem ebenfalls auf Java beschriinkten St. trigonocephalus [Footnote: ‘Vgl. Sammlgn. Ser. I, Bd. 4, pag. 102.’], eine Mittel- stellung zwischen beiden Gruppen ein.” Janensch, 1911 (‘‘Die Proboscidier-Schidel der Trinil-Expedi- tions-Sammlung’’) concludes his detailed description of the skull, dentition, and skeleton of Stegodon airdwana with the statement (p. 192) that while Stegodon insignis and S. ganesa are closest in their dentition to S. airdwana, yet in certain details they are some- what more primitive; also in skull structure S. airdwana differs both from S. insignis and S. ganesa, while in jaw structure the three species are similar. Stremme (“Die Siugetiere mit Ausnahme der Proboscidier,” 1911, p. 143) observes: ‘‘Stegodon Airdwana Mart. ist nach Janensch mit Stegodon ganesa und Stegodon insignis am nichsten verwandt, die im indischen Pliociin und Pleistocin vorkommen.”’ Janensch (1911, p. 192) determines the Trinil Stegodon as Stegodon airdwana Martin, and observes regarding the ridge formula (op. cit., p. 187): ‘Die Jochformel [Footnote: ‘In dieser Formel be- deutet x den Talon.’| lautet, soweit bis jetzt bekannt: Mm * eee Dubois gibt neuerdings (Trinil-Fauna §. 1256) an, dass die Zahl der Joche der letzten unteren Molaren bei dem Stegodon von Trinil bis zu wenigstens 14 gehe, doch ist aus seiner Angabe nicht zu ersehen, ob er die Talons etwa mitzihlt.”” He also observes that this species of Stegodon is highly characteristic of the Trinil Pithecanthropus beds. The other Javan species, Stegodon trigono- cephalus Martin, does not oceur at Trinil. Osborn, 1922: Our knowledge of this very progressive Javan type has been greatly extended by the researches of Janensch (1911) on the rich collection discovered by the Selenka-Blancken- horn Expedition in the Trinil beds in which the type of Pithecan- The fauna of these beds, as described by Stremme (1911, pp. 82-150) includes (pp. 141, 142) the following. thropus erectus occurs. OSBORN: THE PROBOSCIDEA STEGODON ATRAWANA FAUNA OF THE KENDENG-SCHICHTEN LAYER, JAVA, AS LISTED BY STREMME, 1911 This fauna was first regarded as of Lower Pleistocene age (=Boulder Conglomerate zone of Pilgrim), or transitional to Middle [Upper] Pleistocene (=Godavari Alluvium, Nerbudda of Pilgrim) whieh contains Stegodon insignis ref., S. ganesa ref., and Palzoloxodon namadicus. Stremme remarks (1911, p. 144): “Hin wichtiges Leitfossil wiire eventuell Hlephas, dessen Zahnbruchstiiek Janensch dem Hlephas antiquus am niichsten stellt. Das Stiick Pig. 771. 2, one-fourth natural size. The type right inferior molar, r.Msg, is also figured Type of Stegodon Airawana Martin, 1890, Tab. 1, figs. 1 and 9 in Tab. m, figs. 3 and 4, two-thirds natural size. (Martin, op. cit., p. 4): “Nur von Java in einer unvollstiindigen Mandibel und darin steckenden Molaren bekannt. Letztere mit 9 Jochen und 2 Talons.” THE STEGODONTIN: stammt nicht von Trinil; auch Dubois hat keine Hlephas-Reste yon Trinil in seiner grossen Sammlung.” STEGODON 887 The last word by Dietrich (1926.1, p. 139) makes it still more recent, namely: ‘‘Selbst wenn das Entwicklungstempo rascher ge- worden ist, kommen wir fiir Azrawana zu einem sehr viel jiingeren Primates: Pithecanthropus erectus Dubois. Alter als bisher fiir ihn und damit fiir die Trinilschichten fest- Macacus nemestrinus saradana, related to exist- gesetzt wurde, nimlich zu Jung- bis Jiingstpleistociin. Diese Auf- ing Zati of Sumatra and Nemestrinus of fassung des geologischen Alters der Pithecanthropusschichten Borneo. bahnt der von Dubois 1923 gewonnenen Erkenntnis, dass Pithe- Ungulata: Stegodon ganesa javanicus Dubois=|Stegodon canthropus ein Glied der Hominiden ist, den Weg zu dem weiteren Airdwana or S. trigonocephalus Martin]. Schritt, das P. bereits zur Gattung Homo gehort.” Elephas hysudrindicus Dubois = Elephas sp. an- Osborn, 1928: It now seems probable that Stegodon airawana tiquus [namadicus] Falconer (fide Stremme, is of lower Middle Pleistocene age, somewhat more ancient than 1911, and Janensch, 1911). the Godavari, Narbada Alluvium, and with a more early fauna, Cee ee Tafel KXXTK Rhinoceros sivasondaicus Dubois, intermediate 12°0L.+..Gr between R. sivalensis and R. sondaicus."! Rhinoceros kendengindicus Dubois. perhiwan Tapirus pandanicus Dubois. y “Bodjonegoro Artiodactyla: Sus brachygnathus Dubois, related to recent Vinggarg =p Sus verrucosus of Java. nS N on Sus macrognathus Dubois. 47 Sarco res Bae N sort 5 , Sree : ru 5 ““o(Nsawi o@ ¢ &o SON ay Al Hippopotamus (Hexaprotodon) sivajavanicus ra} ae A y Dubois, related to Hexaprotodon sivalensis. Baper® Tas se Cervulus kendengensis Stremme, related to asus = q Pee S.Br existing muntjac. 2 eR WU 2s 6Madiun: : Cervus (Axis) liriocerus Dubois = Cervus (Axis) Ge earns: UND: tie 64 Lydekkeri Martin. ate Se aN Cervus (Rusa) kendengensis Dubois. es Cervus (Rusa) paleomendjangan Dubois. Duboisia (Tetraceros) Kroesenti Dubois, re- Ese lated to existing Boselaphus and Tetraceros. Leptobos Groeneveldtit Dubois. Leptobos dependicornis Dubois. Bibos palzosondaicus Dubois, related to the “= Mnochertihrende Tufhe ra Vulkanmantel g. Guartar existing Bibos sondaicus."! ante Bibos protocavifrons Dubois. a. Kartenskizze der Umgebung von Trinil. Masstab I : 1 500000. Buffelus (Bubalus) paleokerabau Dubois, re- lated to existing Javan Buffelus."! Fig. 772. Sketch of the Kendeng horizon (vertical shading), Trinil, Java, u containing the type of Pithecanthropus erectus Dubois and the type of Stegodon Rodentia: Hystrix. ganesa var. javanicus (=S. airawana or S. trigonocephalus), also referred specimens Edentata: Manis paleojavanica Dubois. of Stegodon airawana. After Dubois, 1908, Taf. xxxrx. Middle Pleistocene. Carnivora: Mececyon trinilensis, related to recent Mececyon javanicus. Felis oxygnatha Dubois. Felis trinilensis Dubois. Felis microgale Dubois= Feliopsis paleojavanica Stremme. Hyena bathygnatha Dubois. Lutra palxoleptonyx Dubois. More recently (letter, 1924) Dietrich determined that the species Stegodon airdwana, which the Kendeng horizon contains, is somewhat more progressive and consequently more recent geo- logically than S. insignis-ganesa. 1(Maarel, 1932, p. 193) “. . -we do not doubt that Stremme would have come to a pleistocene age, had he including the rhinoceroses related to the F. s/vasondaicus and lacking the R. wnicornis and Equus namadicus forms as well as the Palxoloxodon namadicus of the Godavari, Narbada Alluvium. Comparison of the beautiful figures of Janensch (1911, figs. 1-16, Taf. xx1-xxv) together with casts (Amer. Mus. 6835) indi- cates that Stegodon airdwana is a much more progressive species than the S. insignis type of the Upper Pliocene [to Lower Pleisto- cene], and referred of the Lower and Middle [Upper] Pleistocene; it is nearly as progressive as the Stegodon aurorx type from Mt. Tomuro, Japan, as shown in the comparative figure (Fig. 688). Consequently we are justified in accepting Dietrich’s recent con- clusion that Stegodon airdwana and Pithecanthropus erectus are as we—arrived at the conclusion that the Trinil fauna contains at least three still living species viz., Bibos sondaicus fossilis, Buffelus bubalus yar. sondaicus fossilis, and Rhinoceros sondaicus fossilis.’’| 888 of Middle rather than of Lower Pleistocene age. While the vertical heightening or hypsodonty of the molar crown approaches that of the Archidiskodon planifrons type, the entire conformation of the S. airdwana cranium (Fig. 773) is totally different and demon- strates afresh that the most progressive Stegodontine are parallel with, rather than ancestral to, the Elephantide. Selenka-Trinil- Expedition np, Tafel XXI. Fig. 1-2. Stegodon Airawana Maxt. Verlag von Wilhelm Engelmann in Leipzig oo : : : : Beh F Vig. 773. Front and side views of referred Stegodon atrawana Martin, as figured by Janensch in his memoir of the Selenka-Blanckenhorn Trinil- Expedition, 1911, Taf. xxt, figs. 1 and 2. One-sixth natural size. COMPARISON OF STEGODON AIRAWANA MARTIN WITH STEGODON INSIGNIS-GANESA Translated from the German of Dr. W. O. Dietrich with interpolations by H. F. Osborn and corrected by Doctor Dietrich, April 18, 1924 Stegodon airdwana Martin from the Pleistocene of middle Java (Pithecanthropus beds of Trinil) manifests evidences of insular dwarfing; it is in all its parts, both of teeth and skeleton, smaller than the continental west Asiatic and Indian races and species. Special features of the S. azrdwana teeth include the following char- acteristics: (1) Shallow longitudinal cleavage of the crowns; (2) relatively slender and narrow ridge-crests; (3) strong division of the ridge-crests into numerous mamille [conelets], relatively limited cement covering; (4) small, thin, and strongly folded enamel on the abrasion surfaces. (5) While Stegodon airdwana is a dwarfed tropical form as compared with the giant S. 7nsignis- ganesa, it exhibits more numerous, more elevated, and more com- pressed ridge-crests. OSBORN: THE PROBOSCIDEA It is in these respects the most specialized Stegodont of all known species, a terminal form based upon a Stegodont foundation. The dental formula of S. azrdwana runs as follows: Stegodon airdwana: Dp 23 Dp3*;" Dp 47° M14° M2234 M 3 fsisn The greater or lesser development of the ridge formula de- pends upon the anterior and posterior half-ridges or talons, as in the reckoning of Falconer and Lydekker. In contrast [Osborn] the maximal ridge formula of the best-known continental species, Stegodon insignis, is as follows: 74% M 1 ze 7TY%-9 - 4-10 [Stegodon insignis-ganesa: Dp 3 € Dp 4 IN|) ERD yp Gy ee Va- 9-Y4 12}-131° Lenetu or Morars.—In the following table the characteristic maximal and minimal molar tooth length is expressed in millimeters: Upper Molars Dp? Dp? Dp* M! M2 Mé Stegodon airdwana 18 54 90 126 189 221 Stegodon insignis 18 68.6 121.9 246 280 310 Stegodon elephan- toides (=cliftrt) 70.7 124.4 155 208 236 Stegodon bombifrons 150 188 250.5 Consequently Stegodon airdwana is two-thirds the size of S. insignis-ganesa and is also inferior in size to S. bombifrons. The construction of a specialized stegodont molar arises through the simple outgrowth of the anterior and posterior half ridge-crests. The lengths of the molar teeth in the above table are taken from Falconer and Lydekker and from original measure- ments by Dietrich. The maximal length measurements of M3; of the lower jaw compared with the width give us the indices of the third lower molar which is the longest in the series: Stegodon airdwana ap. so4 Tq, mm.= 22 length-breadth ridge index tr. S y } s gqnis §& 5 : 2 Sieqodan insignis ee S*° mm. = 28 length-breadth ridge index Stegodo ftir ap. ze , : F Stegodon clifii = 5“ mm. =36 length-breadth ridge index Ste, bombi- ap. 33: : : Biegada bom I s8* mm. =33 length-breadth ridge index frons tr. By this means we may calculate the length-breadth ridge quotient. Such indices may be calculated also for the third upper molar, M*®, when we secure similar figures which will enable us to Mitnatt. Mee Vig. 774. Juvenile cranium with small tusks. Elevation of narial openings as in S. trigonocephalus. Observe the peculiar straight front of the forehead and great breadth across the top of oecipitals. Skeleton conjectural as in other Stegodonts. Restorations by Margret Flinsch (1930), under the direction of Henry Fairfield Osborn. One-fiftieth natural size. Stegodon airawana. THE STEGODONTINA: determine positively the progressive development within the Stego- dont series. In order to measure the specialization of the molar teeth, including anterior addition and reduction and _ posterior addition and reduction, a relative proportion may be established between the development of the two anterior teeth and the two posterior teeth. It is noteworthy that the proportion between the first and second molars in the Chinese and Javanese species is the same. From careful calculations it follows that M? in S. airdwana is more strongly reduced than in the corresponding tooth of the S. orientalis of Owen. On all these structural grounds Doctor Dietrich concludes that S. airdwana is more recent than either of the Asiatic species of the continent and that its newer Pleistocene age is rendered certain. STEGODON GANESA VAR. JAVANICUS DUBOIS, 1908 Middle Pleistocene, Trinil, Kendeng-Schichten, Java This subspecies, Stegodon ganesa javanicus, belonging in the same Kendeng beds as S. airdwana Martin, 1890, is to be regarded, as remarked by Stremme, as a synonym of S. aivrdwana.! In his paper of 1908, entitled ‘‘Das Geologische Alter der Kendeng-Oder Trinil-Fauna,” Dr. Eugen Dubois, the discoverer of Pithecanthropus erectus, assigns to Pithecanthropus and the accompanying Kendeng-Trinil Fauna at the most a Lower Pleisto- cene age (“‘alt-diluvialen Alters’). The remains occur in the fossil-bearing tufa known as the ‘‘Kendeng-Schichten,” in the center of which lies Trinil, as shown in Plate xxxrx (Fig. 772 of the present Memoir). Recently Dietrich has assigned a Middle Pleistocene age to the Kendeng Trinil fauna. Dubois (1908, p. 1257) erroneously remarks that the two skulls described by Martin as S. trigonocephalus really belong specifically to Stegodon ganesa in their cranial and dental characters, but in order to distinguish this smaller Kendeng species, as an insular variety, from the giant continental form of ganesa, he proposes the subspecific name Stegodon ganesa var. javanicus. He considers it probable (1908) that this Javan subspecies be- longed to the Upper Pliocene fauna. Stremme, “Die Siiugetiere mit Ausnahme der Proboscidier,”’ in “Die Pithecanthropus-Schichten auf Java,” 1911, p. 142, regards this subspecies as closely related to Stegodon airdwana Martin, 1890. According to Dubois (1908) the two skulls described by Martin as Stegodon trigonocephalus belong to this species.' It is important to note that the true S. trigonocephalus of Martin does not occur in the same level as Pithecanthropus erectus. Stegodon Ganesa var. javanicus Dubois, 1908. ‘Das Geolog- ische Alter der Kendeng-Oder Trinil-Fauna.” Tijdsch. Konink. Neder. Aardrijks. Genoots. Amsterdam, Tweede Serie, Deel xxv, STEGODON 889 No. 6, p. 1257. Typr.—Material from Kendeng. Horr ZON AND Locaurry.—Trinil, Kendeng Schichten, Middle Pleis- tocene, Java. Description (Op. cit., 1908, pp. 1256, 1257).—‘‘Die Untersuch- ung eines sehr reichlichen Stegodonten-materiales aus dem ganzen Kendeng, unter welchem auch mehrere Schidel von jungen und alten Tieren, hat mich nun zu dem Ergebnisse geftihrt, dass alle diese Ueberreste einer einzigen Art angehdren. Diese unterscheidet sich von Stegodon ganesa kaum anders als durch ihre viel ger- ingere Grdsse. Der Schidel besitzt dieselbe weite Temporal- grube, ist ebenso charakteristisch stark brachycephal und auch ubrigens sind beide Formen sehr ihnlich. Nur altere Schadel weichen in einigen Beziehungen etwas von der typischen Ganesa- Form ab. indem nimlich die Frontal- und Occipitalteile sich gegeneinander abflachen und mehr oder weniger scharf von einander getrennt sind. Das braucht uns aber nicht davon abzuhalten diese Formen derselben Art zuzuschreiben, denn es kann ja bei den Elephanten die Schiidelform einer und derselben Art, innerhalb gewissen Grenzen, betrichtliche Verschiedenheiten zeigen. Der Ganesa-Typus bleibt bei unserer Form doch immer erkenbar.” “Auch durch ihre Molaren ist sie von S. ganesa spezifisch nicht zu trennen, nur geht die Lamellenzahl im Unterkiefer bis zu wenigstens 14 anstatt 13; die Zahl der Lamellen hat aber, wie wir durch Poutia wissen, fiir die Unterscheidung der EKlephan- tenarten nicht die grosse Bedeutung welche man ihr friiher zu- schrieb.” “Zu dieser Art geh6ren auch die zwei von Martin als Stegodon trigonocephalus beschriebenen Schidel. Waren diese gut erhalten gewesen, so hitten sie erkennen lassen, dass die dreieckige Form keine urspriingliche und der Art eigen- tiimliche ist. Der darauf Bezug nehmende Name ist also zu kassieren. Ich schlage nun vor, diese kleinere Kendeng-Form nur als Varietiit von der riesigen Festlandsform, der sie deutlich sehr nahe steht, zu trennen, fiihre sie also als Stegodon ganesa var. javanicus ein.” CuHaRAcTeErs (Op. cit., 1908, p. 1257).—‘‘Wenn man nun auch nicht zugeben kann, dass Stegodon ganesa und Stegodon insignis einer Art angehéren, so sind beide jedenfalls einander sehr nahe verwandt (die Molaren sind nicht oder kaum zu unterscheiden), Stegodon insignis aber kann man, mit M. Scuuosspr .. . ), geradezu als das Leitfossil der jiingeren Pliocinfauna Ostasiens ansehen. is ist schon hierdurch wahrscheinlich, dass unsere javanische Stegodon ganesa der gleichen jungpliociinen Fauna ange- hort. Unzweifelhafte Ueberreste von Stegodon ganesa sind nun aber noch nicht in jiingeren als Pliociinen Schichten angetroffen worden, denn der in den Narbad&-Schichten gefundene Stoss- zahn kann man dieser Art nicht mit Sicherheit zuschreiben .. .” The following note was prepared by Dr. George Gaylord Simpson (October, 1937): ‘“Stegodon trigonocephalus Martin, 1887, pp. 27, 36, 41, was founded principally on two skulls, Professor Osborn selecting the younger as type. Dubois briefly characterized abundant material from Kendeng, referring it to Stegodon ganesa but as a new variety javanicus Martin, 1887.1, pp. 25, 27). (Dubois, 1908, pp. 1256, 1257). He also referred S. trigonocephalus to S. ganesa. but he did not explicitly say so and the clear implication is that they have different types. He implies that S. ganesa var. javanicus is the same as S. trigonocephalus and S. airawana and S. ganesa but the same as S. airawana (1933, pp. 103-105). These were probably from the vicinity of Surakarta (Solo), Java (see He probably considered trigonocephalus as the same as his variety javanicus Von Koenigswald considers S. trigonocephalus as distinct from applies to the species the oldest of the three names, S. trigonocephalus. Maarel, however, considers S. trigonocephalus to be distinct from S. atrawana, and this was Professor Osborn’s opinion. It appears, in any case, that S. ganesa var. javanicus Dubois, 1908, is synonymous with S. trigonocephalus, S. airawana, or both, and as both names antedate it, Dubois’ name is invalid.’’—Kditor.] 890 OSBORN: Stegodon trigonocephalus Martin, 1887 Figures 705, 731, 775, 776, 777, Pl. xx Probably vicinity of Surakarta, Java. Dubois notes that Stegodon trigonocephalus does not occur in the Kendeng-Schichten of Trinil; Matsu- moto regards the geologic level of S. trigonocephalus as equivalent to the Lower Pleistocene, Boulder Conglomerate beds of India. Stegodon trigonocephalus is a dwarfed insular Stegodont, based chiefly upon an immature cranium (Fig. 776) which has sub- stantially the same characteristic insular form as the immature cranium of S. avrdwana (Fig. 773), but which is very different from Rai (ede, .< Ge * Oi i SS. Fig. 775. Stegodon trigonocephalus. Rather young cranial profiles drawn directly from type. Frontal line much longer than in S. pinjorensis. Cranium extremely abbreviated. Skeleton conjectural, as in other Stegodonts. Restoration by Margret Flinsch (1930), under the direction of Henry Fairfield Osborn. One-fiftieth natural size. the crania of S. bombifrons or S. insignis-ganesa. This juvenile type represents the immature condition of the Stegodont cranium and consequently is very interesting and important. It is now re- garded as in a Pleistocene stage of evolution, but the extremely immature condition of the teeth renders its specific determination very difficult. From the following description and figures of Martin, it is difficult to distinguish this immature type from Stego- don airdwana; it is apparently a somewhat more primitive Lower Pleistocene form. THE PROBOSCIDEA Stegodon trigonocephalus Martin, 1887. ‘Tossile Siiugethier- reste von Java und Japan,” Sammlung. Geolog. Reichs-Museums, Leiden, Beitriige z. Geolog. Ost-Asiens und Australiens, Ser. I, Bd. IV, Heft 2, pp. 27, 36 (1887); also Jaarb. Mijnw. Neder. Oost-Indié, 1887, Wetensch. Ged. 16, Paleeontologie van Neder- landsch-Indié. Verhandeling No. 21, pp. 3, 12 (1887). Typr.— Young skull with third and fourth deciduous premolars in the Geological Museum of Leiden. Horizon anp Locatiry.— Probably from vicinity of Surakarta, Java; said to be associated with remains of proboscideans referred to Stegodon bombifrons, Buelephas namadicus, and Huelephas hysudricus. TYPE Fieurre.—(Op. cit., Martin, 1887, Sammlung. Geolog. Reichs- Museums Leiden, Tab. 11, figs. 1, la, and Tab. 1m, fig. 1. Descrivtrion.—The original description by Martin (1887, pp. 36-41, with figures) gives the characters of the immature type skull and teeth in great detail, and on page 45 offers the follow- ing distinctions from Stegodon cliftii and S. bombifrons: ‘“Zunichst ist es von Bedeutung, dass die Zihne von denjenigen des Stegodon Cliftti so vollig verschieden gebaut sind, dass eine Vereinigung der javanischen Art mit der genannten, von der ein Schiidel noch nicht Ebenso_ be- stimmt unterscheiden sich aber (soweit unsere Kenntniss bis jetzt reicht) auch die Zihne von St. trigonocephalus durch die Jochzahl der Praemolaren von allen anderen, bis jetzt bekannten Arten von Bezeichnend fiir den 2“" Praemolaren von St. trigonocephalus ist ferner seine ovale Form, und ebenso fiir den 3°" Praemolaren die kaum merkliche Convergenz seiner Seiten- fliichen nach vorne zu, Merkmale, die von allen bekannten, ent- sprechenden Ziihnen der iibrigen Stegodontenarten abweichen. Durch die hohen, schmalen Joche und vor allem durch die feine laltelung diirfte auch bei Bruchstiicken der Molaren von St. trigonocephalus eine Unterscheidung von St. bombifrons bisweilen bekannt ist, von vornherein ausgeschlossen wird. Stegodon, Fig. 776. natural size; the crown view of the tooth is reproduced one-third natural size. Taf. 1, figs. 3 and 4 (Naumann, 1887). Type of Stegodon trigonocephalus Martin, 1887, Tab. u, figs. 1, la, and Tab. m1, fig. 1. The side and front views are reproduced one-eighth A single crest from Java, distinet from S. trigonocephalus and doubtfully referred by Naumann to S. insignis Fale. and Caut. (op. cit., p. 9): ‘Das Zahnbruchstiick kann nur zu St. insignis oder zu St. Ganesa gehoren.” THE STEGODONTINA: 3 Son | mature adult S. PINJORENSIS S. BOMBIFRONS Cotyoe Type Tig. 777. S. ORIENTALIS S. ORIENTALIS GRANGERI Aef STEGODON 891 young adult “ ; | guvenile ' ' Juvenile GRANGER] Ref S. TRIGONOCEPHALUS 7yse —S. AIRAWANA Ref ‘ L (ngs ‘ PL ) chile & \\ E| All 4 nal srze 2ngantile Front and side view outlines to a one-sixteenth scale of Stegodont crania of: A, Al, Stegodon pinjorensis type, mature adult (Amer. Mus. 19772), from near Siswan, India. See also figure 765. B, B1, Stegodon bombifrons cotype, mature adult (Brit. Mus. M. 2979, cast Amer. Mus. 10378). See figure 734. C, Stegodon orientalis grangeri ref., young adult (Amer. Mus. 18630) collected by Walter Granger, Wanhsien, Province of Szechuan, China. See figure 763. D, Stegodon orientalis grangeri ref., infantile (Amer. Mus. 18632) collected by Walter Granger, Wanhsien, Province of Szechuan, China. E, El, Stegodon trigonocephalus type, juvenile, from vicinity of ?Surakarta, Java. I, F1, Stegodon airawana ref., juvenile, Java. See figure 773. ermoglicht werden; dagegen ist die Trennung solcher Reste yon St. insignis und ganesa mit sehr grossen Schwierigkeiten verbun- den.” Ridge formula of S. trigonocephalus: Dp 3+ Dp 4+ as compared by the author with S. cliftii: Dp 3* Dp 4*; with S. bombifrons: Dp 3’ Dp 4°; with S. ganesa: Dp 2* Dp 3* Dp 4*. The skull, however, is quite different in form from that of S. insignis. Martin inclines to compare S. trigonocephalus with S. insignis rather than with the Lower [Middle] Pliocene stage of S. bombifrons. It is important to note that these characters relate both to the Sammlung. Geolog. Reichs-Museums, Leiden. See figure 776. immature type skull and to a second older skull which is a referred specimen, namely: (op. cit., p. 41) “b. Alter Schddel. (Tab. rv u. Val Eii oo) ee A specimen from northwest Mindanao, Philippine Islands, referred to this species is described in detail by Dr. Edmund Nau- mann of Munich University in his ‘“Fossile Elephantenreste von Mindanio, Sumatra und Malakka,’”’ Abhand. u. Berichte des K6nigl. Zoolog. Anthrop.-Ethnog. Museums zu Dresden, 1887, pp. 5-8. He subsequently (1890) made it the type of Stegodon mindanensis. STEGODONTS OF THE PHILIPPINE ISLANDS AND JAPAN We owe to Naumann (1890) and to Matsumoto (1915, 1918) the discovery and description of three very progressive species, originally referred to Stegodon, which in part or wholly seem transitional in structure to Archidiskodon or to Hlephas. The true affinity of these types rests upon (a) the still unknown structure of the cranium and (b) the ridge formula and height and form of the ridge-crests as now revealed in the case of the species Stegodon aurore (Fig. 781). In brief, until the cranium is known, we cannot be certain whether these animals are progressive Stegodonts or primitive Archidiskodonts. The history of discovery and description is as follows. 892 OSBORN: Stegodon (Archidiskodon?) mindanensis Naumann, 1890 Figures 706, 778 Mindanao, Philippine Islands, Lower(?) to Middle Pleistocene. Stegodon Mindanensis Naumann, 1890. “‘Stegodon Minda- nensis, eine neue Art von Uebergangs-Mastodonten.” Zeitschr. deutsch. geol. Ges., Bd. XLII, Heft I, pp. 166-169. TYPE.— Fragment of a molar tooth. Horizon AND LocaLiry.— Mindanao, Philippine Islands, (?)Lower to Middle Pleisto- cene. Typr Ficurre—Naumann, 1887, Taf. 1, figs. 1 and 2. Tyre DescripTtion.—(Naumann, 1890, p. 167): ‘. .. so bleibt doch das friiher erzielte Resultat, nach welehem durch die Type or STEGODON (ARCHIDISKODON?) MINDANENSIS lig. 778. Type incomplete inferior molar of Stegodon mindanensis Naumann, 1890, from Mindanao, Philippine Islands (originally figured by Naumann in 1887, Taf. 1, figs. 1 and 2, as Stegodon trigonocephalus). As compared with Stegodon airawana (Vig. 779), the ridge-crests in S. mindanensis type are more vertically placed, as in the Stegodon aurore type (Pig. 780). Untersuchung der beiden Zahnbruchstiicke von Mindanao ‘die Verbreitung der Siwalikfauna tiber das Gebiet der Philippinen bewiesen und die enge Verkniipfung einer wahrscheinlich Jungter- tifiren Sdugethierfauna auf Java und den Philippinen durch eine in der Entwicklungsreihe der Stegodonten und Elephanten hochwichtige Art’ constatirt sein sollte, zu Recht bestehen. Merkwiirdig ist ferner ein medianer Einschnitt der Krone, der jederseits von einem secundiren Kinschnitt begleitet wird. Durch diese Spaltungen werden die Mamillenreihen in Gruppen zerlegt.”’ The type of this species was originally figured by Naumann in 1887 as Stegodon trigonocephalus; it was regarded by him as an outlyer of the Siwalik proboscidean fauna. A single ridge-crest from Java (see lig. 776, lower), distinct from S. trigonocephalus, Vig. 779. Referred by Janensch to S. airawana. of a third superior molar, M°, after Janensch, op. cil., p. 174, fig. 13. (Left) Lateral view. (Right) Ridge-crest section. THE PROBOSCIDEA was doubtfully referred by Naumann to S. insignis or S. ganesa Faleoner and Cautley. (Naumann, 1887, p. 9): “Das Zahn- bruchstiick kann nur zu St. 7nsignis oder zu St. Ganesa gehoren.” If a true Stegodont, Stegodon mindanensis (Fig. 778) is even more progressive than S. avrdwana, because the valleys between the ridge-crests are entirely closed up; it compares somewhat more closely with Stegodon aurore (Figs. 780, 781), also an imperfectly known species possibly referable to Archidiskodon. Osborn, 1924: It is very difficult to determine the characters of this type (Fig. 778) without examination of the original speci- men. The type lower molar is far more progressive in the direction of Archidiskodon or Elephas than the types of either Stegodon insignis or S. ganesa; it is, in fact, a true-crested tooth in which the ridges are closely compressed and the valleys closed. Conse- quently this tooth should be compared with that of a primitive species of elephant or of Archidiskodon. Stegodon aurore Matsumoto, 1915, 1918 ligures 688, 709, 780, 781, 819 Upper(?) Pliocene, Mt. Tomuro, Kaga, Japan. Like Stegodon mindanensis, Elephas (Prostegodon, Parastego- don) aurore is either a highly progressive Stegodon or a primitive Archidiskodon, a point to be determined positively by the discovery of a cranium. ELEPHAS (PROSTEGODON, PARASTEGODON) AURORH Marsu- moto.—In describing the type specimen, determined as a second right superior molar, r.M?, Matsumoto remarked (1918, p. 52): «".. this specimen represents a species new to science, being... a transitional form from Stegodon to Elephas.’”’ Originally described as Hlephas aurore Mats., 1915, also 1918, p. 52, Pl. xx, it was after- ward referred by Matsumoto (in Osborn, 1923.601) to Prostegodon, the genotype of which is Mastodon latidens, and finally (1924.2) was made the genotype of Parastegodon Matsumoto, to include the Stegodon mindanensis-E. aurore group. Elephas aurore Matsumoto, 1915, 1918. Preliminary reports of this species were published in the Scientific Gazette, Tokyo, Vol. III, No. 5, 1915, pp. 308-315, and in the Journal of the Geological Society, Tokyo, Vol. XXIII, No. 275, 1916, p. 294 (both in Japanese). Also (in English) “On a New Archetypal Prot ery il %, aS Te 0 vy 2 12 Nine ridge-crested second superior molar, M®, after Janensch, 1911, p. 171, fig. 12, and ridge-crest section One-half natural size. Both figures inverted to show the molars in natural position. It will be observed that the convexity of the ridge-crests of the second molar crown, resulting in the wide divergence of the crests, together with the more numerous ridge-crests, tends to relate this animal to Stegodon rather than to Archidiskodon, in which the ridges are more vertical and more closely compressed. THE STEGODONTINA:: STEGODON 893 TyPE OF STEGODON AURORA Fig. 780. Type of Elephas (Prostegodon, Parastegodon) aurore Matsumoto, after Matsumoto, 1918, Pl. xx, figs. 1 and 3. One-half natural size. Determined by Matsumoto (1918) as a ten and a half erested second right superior molar, r.M?. Compare figures 699 of S. bombifrons and figure 762 of S. orientalis grangeri showing convex external side and flat internal side; also figures 686, 7, and 688. FARCHIDISKODON =AURORAE | AR rate Fig. 781. Vertical section of type molar, r.M*, of Stegodon aurore (compare Fig. 780, crown and side views of type of Hlephas (Prostegodon, Parastegodon) aurorx, also Fig. 825, lectotype of F. [= Archidiskodon| planifrons). One-half natural size. After photograph sent by Doctor Matsumoto. Univ., (2), XIII, No. 1 (Matsumoto, 1929.3).—Kditor.| Fossil Elephant from Mt. Tomuro, Kaga.” Sci. Rept. Tohoku Imp. Univ., 1918, (2), III, No. 2, p. 52. Typr.—aA right M? with ten and a half ridges, thus exceeding the ridge formula of M2 in Archidiskodon planifrons, namely, M 2*°. Original in Geological Institute, Tokyo. Horizon AND Locauiry.—Mt. Tomuro, Kaga, Japan, (?)Upper Pliocene. Typr Ficgurr.— Op. cit., 1918, Pl. xx, figs. 1-3. SuppLneEMEnTARY DEscriPTion (Marsumoro, 1924.2, pp. 256, 257, 262)—Made the genotype of the genus Parastegodon (see p. 9038 of the present Memoir). Type Description (1918, pp. 52-55).—‘‘The unique type specimen corresponds to an upper, probably penultimate, molar of the right side. It measures 180 mm. in length, 75 mm. in maximum width and 48 mm. in the maximum height of the crown.” The reasons why the author referred this new species to Hlephas and not to Stegodon are as follows: The ridge formula of M? corresponds to 10}, while that of the known species of Stegodon corresponds to 6 to 9. The author gives six additional reasons for separating the animal from Stegodon, and six additional reasons for separating it specifically from FL. planzfrons, to which he regards it as allied generically. The author concludes (p. 55): ‘The present species appears to the present writer to be more archety- pal and more Stegodon-like than EH. planifrons in the second distine- tive characteristic, but just the opposite in the third, fifth and sixth distinctive characteristics. At any rate, the discovery of the present species is worthy [to] be considered as an additional datum to prove the very intimate alliance of Stegodon and Elephas.” Stegodon orientalis shodoensis Matsumoto, 1924 Middle Pleistocene. Island of Mitsugo (Mitsugo-shima) and Island of Shédo, Inland Sea, Japan. Stegodon orientalis shodoénsis Matsumoto, 1924. ‘Preliminary Notes on the Species of Stegodon in Japan.” Journ. Geol. Soe. Tokyo, XX XI, No. 373, pp. 333-335. Typr.—A fragment of skull—left and right upper jaw bone with third molar attached. Uyeno Museum of Tokyo, No. 2194, presented by Mr. Tomekichi Ozaki. Horizon AND Locauiry.—Mitsugo-shima, Yoshima- mura, Nakatadotsu-gun, Province of Sanuki, Japan. Milazzian- Tyrrhenian. Middle Pleistocene. Tyre Fiaurre.—Type figure not seen by the present author.’ Description in Japanese by Matsumoto; translation by Messrs. Shoichi Ichikawa and Ushinosuke Narahara.—(Matsumoto, 1924.3, pp. 333-335). ‘Materials: A fragment of skull; left and right upper jaw bone with third molar attached (found at Mitsuko- shima, Yoshima-mura, Nakatadotsu-gun, Province of Sanuki. Specimen No. 2194, Uyeno Museum of Tokyo, presented by Mr. Tomekichi Ozaki). A small fragment of third molar (found at Shédo-shima and the property of the Tokyo Imperial University). Third upper left molar (found at Shédo-shima, the cast is kept at Kyoto Imperial University). A fragment of an upper palate with third molar of left and right attached (found at Shédo-shima, the property of Kyushu Imperial University and the cast kept at Ixyoto Imperial University).” MSee also Matsumoto’s article (in English) “On Parastegodon Mastumoto and its bearing on the Descent of Earlier Elephants,” Sci. Rept. Tohoku Imp. *[Dr. Jird Makiyama has recently reviewed the Japanese Proboscidea (see Makiyama, “Japonic Proboscidea,” 1938, Mem. Coll. Sci. Kyoto Imp. Univ., (B), XIV, No. 1, pp. 1-59), in which he figures (p. 18, fig. 7) the holotype palate of Stegodon shodoénsis, with right and left third molars in situ. On page 19 he states that “The material of Matsumoto was not enough to give full details of the species, but it was named before Parastegodon akashiensis Takai, 1936, which seems to me to be in the nearest relation, as it may have a new name form S. shodoensis akashiensis.’ three figures (Figs. 10-12).—Editor.] ’ This subspecies he describes on pages 21 to 27, including 894 OSBORN: “Ridge formula of molar: left, a+9X [=-9—4]; right, at+t8x [='%-8-\4]. The exact measurement of ridges cannot be determined. The length as it is, left 226 mm., right 215mm. The greatest width, left 97 mm. at 2nd ridge, right 96 mm. at 3rd ridge to 6th ridge. The height of crown, left 52 mm. at 6th and 7th ridges, right 49 mm. at 7th ridge. The 6th ridge of left, and 7th ridge of right shows the beginning of wear. . . . There are four ridges within 100 mm. The length of each ridge is 25 mm. and this is rather narrow as a width for third molar of Stegodon. ‘The valley is narrow, and the entrance of inside of valley has supple- mental cusps. The fold of enamel is very regular and fine. The cement is well developed. The interlocking of opposing surfaces of the molar is of Mastodon type.” “Second example: Ridge formula may probably be X11X. Greatest width, 93'4 mm. at 6th ridge, height, 55 mm. at 7th ridge. There are four ridges within 100 mm. Judging by the measurement, it looks like a second molar, but by the shape of its being attached THE PROBOSCIDEA a+8x. The exact measurements of ridges cannot be determined. Length, left 205 mm., right 200 mm. Greatest width, 85 mm. at 6th ridge from back (left), 84 mm. at 5th ridge from back (right). Height, 45 mm. at 3rd ridge (left), 50 mm. at 2nd ridge (right). There are four ridges within 100 mm. The interlocking occlusion of the molar is elephant type.” “A small fragment of third molar, which shows three ridges. I cannot determine whether it belongs to the upper or lower jaw. The length from inside to outside of each ridge is 90 to 96 mm. The width of front to back is 24 to 25 mm.” ““Among these specimens, the first has comparatively low crown and the second and third have very high crowns. The fourth specimen is rather low as absolute value but as a ratio it is high. The molars are different in sizes, and this differentiation (or varieties) is the one which occurs in the last period of the existence of the species.” “These specimens resemble the Stegodon insignis which is to the palate is a third molar. Ridge formula, left a+9X; right found in the Narbada Valley of India.” V. RECENTLY DESCRIBED STEGODONTS FROM JAVA, CHINA, AND JAPAN (The following species were described respectively by Dr. Franciscus Hendricus van der Maarel (“Contribu- tion to the Knowledge of the Fossil Mammalian Fauna of Java,” 1932), by Dr. G. H. Ralph von Koenigswald (“Beitrag zur Kenntnis der Fossilen Wirbeltiere Javas,” 1933), by Dr. Shigeyasu Tokunaga (“Fossil Elephant teeth found at Yokohama and Kakio, Kanagawa Prefecture,” Journ. Geog. (Tokyo), XLVI, No. 546, 1934, and “4 New Fossil Elephant found in Shikoku, Japan,’’ Proc. Imp. Acad. Tokyo, XI, 1935), by Dr. Chung-Chien Young(‘‘Miscellaneous Mammalian Fossils from Shansi and Honan,” 1935), and by Dr. Arthur Tindell Hopwood (“Fossil Proboscidea from China,” 1935). While the species of Stegodonts in these publications (other than Parastegodon? kwantoensis and Parastegodon sugiyamai of Tokunaga) were noted by Professor Osborn, they were not studied intensively by him, consequently they are listed here together with excerpts from the original descriptions, including type figures, but without comment or determinations by the author of the present Memoir. Doctor Hopwood states on page 103 of his Memoir that ‘Insufficient is known of the Stegodontide of China to make a comparison with those of India very profitable. At present it seems as though they occur earlier in India than in China, where they are a late invasion from the South. Palaeoloxodon, on the other hand, may be more recent in India than in China.’’—Editor.] Stegodon bondolensis van der Maarel, 1932 Vigure 782 Bondol near Kuwung, district Randublatung, Regency Blora, Residency Rembang, Java. Stegodon bondolensis van der Maarel, 1932. ‘Contribution to the Knowledge of the Fossil Mammalian Fauna of Java,” pp. 158-164. Typr.—“‘Fine fragment of the mandible, com- prising the whole of the horizontal ramus and the lower portion of the ascending ramus of either side; containing on either side a molar which is inferred to be the M; from the absence of the indication of any other tooth behind.” Horizon AND LocaLiry.—Bondol near Kuwung, district Randublatung, Regen- cy Blora, Residency Rembang, Java. Tyrer Fiagurrn.—Op. cit., Pl. xiv, figs. 1, 4, 5, also text figures 24 and 25. Derscription.—(Van der Maarel, 1932.1, pp. 158-164): “The mandible is broken into two in the symphysis, but both parts match exactly. The alveolar border of both molars is damaged.” “The diastema descends obliquely forwards at an acute angle of about 35° with the inferior border of the ramus. The mandibu- lar symphysis is produced into a relatively broad, spoutlike termination. Its lower border is damaged. ... The outer surface of the horizontal ramus presents two foramina mentalia, the one situated some 8 em. below the anterior extremity of the alveolar margin, the other about 8 em. in advance of the former and nearer to the inferior border of the ramus.” “Only the lower portion of the coronoid process has been preserved.” “\feasurements of lower jaw in em. See text fig. 24. I greatest height =16 greatest width = 7.8 ereatest height =12 greatest width =17 Length of BC Alii, In plane In plane II THE STEGODONTINA: “As the front side of both molars is damaged, it seems—at first sight—impossible to ascertain the true number of ridge-crests carried by each of them. I ortunately, however, in the 1.M; a small portion of the front surface of the fangs has been preserved. It is situated below the anterior side of the foremost ridge-crest, and proves with absolute certainty, that this ridge-crest is really the first one. That is to say: not a single ridge-crest has been lost. The ridge formula is, therefore, 8.” STEGODON 895 can, however, only be seen in the last three ridge-crests. In this connection attention may be drawn to the fact that in the molars of the present specimen enamel, cement and the greater part of the dentine had the same irregularly spotted appearance. As a result the crenulation of the enamel was very indistinct, and it was only possible to make it distinctly visible in the photograph by black- ening the dentine of the tooth. Another result is, however, that ‘Stufenbildung’ though present, is in general not clearly exhibited.” Fig. 782. Type mandible and enlargement of third left inferior molar of Stegodon bondolensis. van der Maarel, 1932, Pl. xrv, figs. 1, 4, and 5. one-seventh natural size. From Bondol, Java. After Molar about three-fifths natural size; mandible about (Upper) Crown view of third left inferior molar, 1.M3, with 8+ ridge-crests; same molar as in ac- companying lower jaw. (Lower, right) Superior view of fragmentary lower jaw with right and left third molars in situ. (Lower, left) Right lateral view of same jaw. “The tooth is distinctly curved outwards. The base of ridge- crests 1-7 are more or less of the same length; from ridge-crest 8 inner and outer side of the molar converge suddenly back- wards.” “The base of the crown and the grinding surface are strongly concave. The grinding surface slopes obliquely from the outside inwards.” “The enamel is moderately thick and consists of two layers (see the buccal cusps of ridge-crest 7). A distinct ‘Stufenbildung’ “There is no indication of the presence of a median longitudi- nal cleft. Cement is present, though in small quantities. The valleys between ridge-crests 1, 2, 3, 4 and 5 possess but little cement. More backwards the amount is somewhat less small, not only the base of the valleys being filled to a greater extent, but also the lingual and buceal sides of the ridge-crests, and the posterior side of the hind talon being covered by a thin coat of cement. Noteworthy is the stronger, local development of cement near the buccal side of ridge-crest 7.” 896 OSBORN: ““\IBASUREMENTS OF R. AND L.M3 IN MM. Total length of r. Mj; 190 mm.\measured along median line Ditto “ 1,M3 197 ~ of crown.” “As a matter of fact all the species of Stegolophodon may be left out of consideration, being all very primitive forms which in my opinion may as well be reckoned to the family of the Masto- dontide.” “On the other hand, most of the remaining species [of Stegodon| are considerably more progressive than the form to which the lower jaw in question belongs.” “Tn my opinion we may, therefore, conclude, that the present Javan form is distinet from all other species. I propose the specific name of bondolensis, indicating the locality from whence it has been procured.” THE PROBOSCIDEA scheinend weniger an HGOhe zunahm, als es bei der typischen Form der Fall ist. Die letzten drei Joche und der Talon liegen noch dick unter Zement.” “Diesen letztaufgefiihrten Zahn bilde ich Taf. xxvir ab, neben dem entsprechenden Zahn von St. t. trigonocephalus von Pitoe, nahe Trinil. Dieser letztgenannte Zahn ist ca. 270 mm lang und gehdrte einem mittelgrossen Tier. Zwei letzte Molaren von Trinil sind (nach Janensch) 304 und 303 mm lang, auch solehe von Watoealang und Ngandong erreichen 300 mm, (da sie z. T. mit der Hinterhilfte noch im Kiefer stecken, kann ihre Grosse nur geschitzt werden.) Dem gegeniiber miissen die Zihne von Boemiajoe mit ca. 240 mm klein genannt werden. Dabei haben sie dieselbe Breite und dieselbe Biegung, wie die anderen Zihne. Die Verkiirzung kommt daher, dass die Zihne von Boemiajoe nur X 11 X Joche besitzen gegeniiber X 13 X bei der jiingeren Form. Fig. 783. Type left third inferior molar, 1.M3, of Stegodon trigonocephalus praecursor, after von Koenigswald, 1933, Taf. xxvu, fig. 2, two-thirds natural size. From Bumiaju, Java. Stegodon trigonocephalus praecursor von Koenigswald, 1933 Figure 783 Untere Schichten, Kali Glagah, Bumiaju, Java. Stegodon trigonocephalus praecursor von Koenigswald, 1933. “Beitrag zur Kenntnis der Fossilen Wirbeltiere Javas.’’ Wetens- chappelijke Mededeelingen, Dienst Mijnbouw Nederl.-Indié, I Teil, No. 23, pp. 104, 105. Typr.—Lower jaw with third molar of both sides complete. Horizon anp Locauity.— Untere Schichten, Kali Glagah, Bumiaju, Java. TYPE FigurrE.—Op. cit., Taf. xxvu, fig. 2 (third lower molar of the left side, 1.M3). Description.—(von Koenigswald, 1933.1, pp. 104, 105): “Ts liegt ein Unterkiefer vor, dessen beide Ms; noch komplet sind. Mz rechts ist isoliert, ein Teil der starken Zementlage ist abgewittert, so dass auch die letzten noch nicht angekauten Joche und der kleine Talon hervortreten. Der linke Mz sitzt noch auf dem leider etwas gequetschten Kieferast, der im vorderen Teil an- ” Dubois (1908, pg. 1256) gibt fiir das Stegodon der Kendenglagen sogar 14 Joche an, doch ist aus seinen Ausfiihrungen (Abbildungen hat er bisher nicht verdffentlicht) nicht zu ersehen ob er nicht etwa einen Talon mitgerechnet hat. Die Exemplare der SeLENKA- Expedition haben alle, und auch die mir vorliegenden bis auf eines, nur 13 Joche. Dieses eine von Lepen Alit bei Tingang zeigt nur X 11 X Joche (vergl. v. d. Maarel, pg. 143) und gehdrt aber dennoch einem typischen St. t. trigonocephalus. Dieser Fund zeigt, dass noch unter den Stegodonten der Kendengschichten als Ausnahme Formen mit weniger Jochen auftreten konnen. Es ist eine bekannte Erscheinung, dass oft bei spezialisierteren Arten auch noch Riickschlige auftreten.” “Bin weiteres Kennzeichen der Zihne von Boemiajoe ist, dass der dicke Schmelz viel weniger intensiv gefiltelt ist als bei der typischen Art (Taf. xxvii). Das lisst auch ein Vergleich mit den Abbildungen bei Martin und JANeNnscH deutlich erkennen,” “Yon Boemiajoe stammt auch noch ein auffallend grosser Stosszahn von etwa 3,50 m Liinge, der seiner ganzen Form nach wohl nur einem Stegodon gehOrt haben diirfte.” THE STEGODONTINA: STEGODON 897 Stegodon yushensis Young, 1935 Figure 785 From Yiishe, China. Stegodon ytishensis Young, 1935. ‘‘Miscellaneous Mammalian Fossils from Shansi and Honan.” Pal. Sinica, (C), IX, Fase. 2, 1935, pp. 26-28. TyPr.— ‘““\ well preserved upper left third molar.” Hortr- ZON AND Locauity.—‘‘Pontian ‘violet sands’ of the Yiishe formation from Yishe,’” China. TYPE FigurE.—Op. cit., Pl. v, figs. 1, la. Description.—(Young, 1935.1, pp. 26-28): “With the exception of the fourth ridge and of a large part of the cingulum, the tooth is well pre- served, including the roots. Crown composed of five ridges, not including the posterior heel and the Fig. 784. Type lower jaw with right second molar, r.My, in situ, of Parastegodon? anterior cingulum. . . . The ridges are rather closely kwantoensis Tokunaga, 1934, Pl. rx, fig. 1, one-half natural size. set; the valleys not very deep; only slight traces of cement can be observed. Cingulum is very weakly indicated in the preserved parts.” Parastegodon? kwantoensis Tokunaga, 1934 Figure 784 Kakio, Kanagawa Prefecture, Japan. Upper Pliocene (fide Tokunaga). [The description in Japanese by Professor Shigeyasu Tokunaga of his species Parastegodon? kwantoensis, 1934, was not reviewed by Professor Osborn. However, as the present author regarded Parastegodon as either a progressive Stegodon or a primitive Archi- diskodon, his generic reference cannot be given in the present Memoir. The following résumé in English is by Mr. Ushinosuke Narahara of the American Museum of Natural History.— Editor. ] Parastegodon? kwantoensis Tokunaga, 1934. ‘Fossil Elephant teeth found at Yokohama and Kakio, Kanagawa Prefecture,” Journ. Geog. (Tokyo), Vol. XLVI, No. 546, July, 1934, pp. 365-369. Typre.—Portion of lower jaw with right second molar, r.Moe, in situ. Horizon AND Locariry.—Kakio, Kanagawa Prefecture, Japan. Upper Pliocene (fide Toku- naga). Tyre Figure.—Op. cit., Pl. rx, figs. 1-3. Derscription.—Portion of a lower jaw with right second molar, r.Ms, in sitw; eight and a half ridge-crests, probably ten if perfect; molar valleys deep, narrow, and filled with cement. Length of jaw bone 253 mm. Second molar, r.M» Length 190 Breadth 76 Height (outside) 35 Height (inside) 31 5,4 ridge-crests in 100 mm. The specimen is primitive like Parastegodon |Stegodon] aurore Matsumoto, but with certain differences, which he [Tokunaga] believes warrants its assignment to a new species, namely, Fig. 785. Third superior molar of the left side, 1.M®, of Stegodon yiishensis, after Young, 1935, Pl. v, figs. 1, 1a, two-thirds natural size. "rom Yiishe, China. Parastegodon? kwantoensis. (Upper) Crown view. (Lower) External view. OSBORN: “With the exception of the first ridge which has three roots (external, internal, median) and of the last ridge, under which the roots build a continuous transversal lamella, each ridge has two distinct roots. The four posterior ones are laterally fused along the external side of the tooth, and the three posterior ones along the internal side.” “Dimensions:—[with some omissions] Maximum length of the tooth......... ee loom. Breadth at the third ridge [maximum]........ 83 mm. Height of the crown above the cingulum at the STOMA SCNE eye ee etre oo eae ene eee 36 mm.” “Both Stegodon officinalis Hopw. and zdansky: Hopw. are distinetly larger, and they have a higher number of ridges at the third molar. In size Stegodon orientalis Owen (1870) and Stegodon orientalis grangeri Osborn stand closer; but they also have one ridge more at M* and the cement of their teeth is more developed. S. sinensis Owen and S. aff. bombifrons are described from frag- ments only. The former one is hardly separable from S. orientalis: and the latter one is too large.” “We therefore have to deal with a new species, for which the name Stegodon yiishensis (sp. nov.) is proposed.” Stegodon officinalis Hopwood, 1935 Figures 786, 787 Said to have come from Szechuan(?), China. Horizon unknown. Young in his memoir ‘‘Miscellaneous Mammalian Fossils from Shansi and Honan,” 1935, p. 27, compares Stegodon officinalis Hopwood (ex MS.) with his new species S. ytishensis. 787. “Last unworh ridge and upper(?) molar,” referred by Hopwood to Stegodon officinalis, one-half nat- ural size. Cast Amer. Mus. 21879. After Hopwood, 1935.1, Pl. vu, fig. 4. Vig. Fig. 786. talon of an nalis Hopwood, consisting of the “first Type of Stegodon offici- two ridges of an unworn lower molar” (cast Amer. Mus. 21878), one-half natural size. After Hopwood, 1935.1, Pl. vu, fig. 3. Stegodon officinalis Hopwood, 1935. “Fossil Proboscidea from China.” Pal. Sinica, (C), [X, Fase. 3, 1935, pp. 73-75 (Hopwood, 1935.1). Typr.—‘The first two ridges of an unworn lower molar.”’ Cast Amer. Mus. 21878. “Bought in a medicine shop, Hanchow. Szechuan.” Horizon 1935.1, Pl. vu, fig. 3. Horizon AND LOCALITY. Said to have come from unknown. Tyre Fieurr.—Op. cit., THE PROBOSCIDEA Mareritau.—(Hopwood, 1935.1, p. 73): “The unworn last ridge and talon of a third molar. Probably from the upper jaw.” Cast Amer. Mus. 21879. Description.—(Hopwood, 1935.1, pp. 73-75): ‘A Stegodon with the ridge-crests widely spaced, with four to five mamille on each ridge; first and second ridges of lower molar divided by a median cleft; conules of upper molars not united into ridges.” “The anterior ridge is divided into two cusps by a deep median cleft. The post-trite cusp has two large cones with a third, smaller one, between them. The tips of the cones of the pretrite cusps have, with one exception, been broken off. The second ridge is also divided by a median cleft, but this is not so prominent as in the first ridge. The pretrite cusp has three cones, and the post- trite has two. The ridges are much narrower at the top than at the base, this is especially true of the second hence the labial and lingual surfaces of the tooth slope inwards to a very marked extent. The anterior surface of the second ridge is nearly perpen- dicular, whereas the posterior surface slopes away at a fairly steep angle. This difference in the slope of the two surfaces gives the ridge the appearance of being tilted forwards, a sign of deri- vation from a lower tooth. ... There are no lateral cingula, nor are there cingules at the entrances to the valleys. No cement is present.”’ “The second fragment consists of the last ridge and talon of a large tooth, which is doubtfully regarded as a third upper molar. The ridge consists of five cones which are not fused to one mass, but are separated by definite clefts. One cleft is deeper than the others; hence the cones are divided into two groups, one with three cones and the other with two. A large part of the talon is covered by cement, above which there rise the tips of four cones. . . The ridges are upright, with their anterior and posterior surfaces sloping at about the same angle; the valley between them is narrow, and partly filled with cement.” Discussion by Hopwood.—‘There is no proof that these two specimens are correctly associated. At the same time it is evident from their preservation that they are both derived from the same deposit, which appears to have been of a lignitic, or peaty nature. They indicate the existence of a species which has hitherto been unknown and it is preferable to assume an unproven connexion, instead of making two ‘species’, one of which might have to be relegated to the synonymy at a later period.” “Tn placing the species with Stegodon rather than in Stegolo- phodon, | have had regard to the characters of the ridges of the holotype. One of the chief differences between the two genera is that whereas in Stegodon the cones are united into ridges, in Stegolophodon they remain discrete. Admittedly the cones in the fragment of an upper tooth are still divided by deep clefts, but they show the first stages of union, and the cones in each half of the ridge are more closely joined to each other than the two halves of the ridges.” “This species is the most primitive yet recorded from China. In the division of the lower ridges into two cusps, as well as in the division of the upper ridges and the slight displacement of the two halves, it retains characters which derive from a mastodont ancestry. ‘Traces of this inheritance are never entirely lost among the Stegodontide and they persist with more or less regularity among the elephants. There are several species in a similar stage THE STEGODONTINA: of evolution among the fauna of the Siwalik deposits of India, but they have not yet been described. So far as is at present known, the Indian species have blunter cones than the Chinese form, and their ridges are not so high.” “The trivial name refers to the chance that these specimens were purchased in the shop of a Chinese druggist.” Stegodon zdanskyi Hopwood, 1935 Figure 788 Horizon unknown. The name Stegodon zdanskyi Hopwood first appeared (ex MS.) in Young’s memoir on “Miscellaneous Mammalian Fossils from Shansi and Honan,” 1935, pp. 27 and 28; to this species Young has referred ‘‘an upper left maxillary with M?, M* in situ and... an isolated right M?* evidently belonging to the same individual, both found in the Pontian sands of Yiishe. A fragment of molar and some milk teeth may also belong to the same species.”’ Stegodon zdanskyi Hopwood, 1935. ‘Fossil Proboscidea from China.” Pal. Sinica, (C), IX, Fase. 3, 1935, pp. 75, 76 (Hopwood, 1935.1). Tyer.—“The first four ridges of a right third lower molar.” Cast Amer. Mus. 21872. HorizoN AND Locatiry.—‘Bought in a Medicine Shop, Shanghai.”’ Horizon unknown. Types Figurr.—Op. cit., 1935.1, Pl. vn, fig. 5. Vig. 788. ridges of a right third lower molar” (cast Amer. Mus. 21872), after Hopwood, 1935.1, Pl. vit, fig. 5, one-half natural size. Type of Stegodon zdanskyi Hopwood. ‘“TVirst four Derscriprion.—(Hopwood, 1935.1, pp. 75, 76): “A Stegodon of very large size, with eight to ten mammillz on each ridge-crest ; anterior ridge still preserving the two cusps of the earlier mastodont ancestors, posterior ridges foreshadowing the structure of the plates found in the teeth of the Elephantide.”’ “The first ridge of this specimen is divided into two cusps by a prominent mesial cleft. Each cusp, especially the pretrite one, is divided into two cones by a cleft which is not so prominent as that which separates the cusps. The cones themselves are again divided at the summit into two or three mammille by grooves, which do not proceed very far down the anterior and posterior surfaces. None of the other ridges show this sub-division; they are all divided, but in a very different manner.”’ “ach of the posterior ridges has a large cone at either end, STEGODON 899 with one or two mammille at the summit. Between these cones is a long, roof-like portion with a mammillated crest. The number of mammille varies with the manner of counting. If the main eminences alone are reckoned, there are from four to six; if each tiny sub-division, or indication of one, is counted, this number is increased.” “The crown of the tooth is curved to the right, and, seen from the side, it is very slightly concave. The ridges have a for- ward pitch; that is to say, their anterior faces are steeper than the posterior. All these are marks of a right lower molar.” “Anteriorly there is a relatively slender cingulum, which receives a nodular buttress passing downwards and inwards from the inner cone of the pretrite cusp of the first ridge. There are no lateral cingules at the entrances of the valleys. A small quantity of cement is present.” Discussion by Hopwood. This species is the largest Stegodon yet recorded. Apparently it has nothing to do with the Indian species hitherto described; they are all smaller, have the ridges closer together, and more copious cement. An undescribed Indian species, in which the third upper molar is 243 mm long and 181 mm wide, is of about the same size, but its ridges are more of the type of Stegolophodon stegodontoides (Pilgrim), and it is probably referable to the same genus as that species.”’ “Apart from its size, the most interesting feature of S. edanskyi is the curious mixture of elephantine and mastodontine features in its ridges. In its composition of two cusps, each made up of two cones, and in the relations between the pretrite cusp, anterior buttress, and the cingulum, the first ridge is, fundamental- ly, that of any mastodont of the bunolophodont, or primitive zygodont, type. All the other ridges are essentially elephantine in their structure. They each have a large cone at either end, and a long, mammillated, roof-like portion in the centre. If the two fissures which divide the tooth in this manner are deep enough, the partly worn tooth would show a tripartite enamel figure. This tripartite division is characteristic of the elephants, though it is also shown by some, at least of the Stegodon group (ef. Soergel, 1912 [1912.2], p. 8, fig. 2). In most cases it consists of two outer rings with an ellipse between them; exactly the type of figure which could arise during the wearing down of a lower tooth of S. zdanskyi.” ‘All the other remains of Stegodon from China hitherto de- seribed (Owen, 1870 [1870.1]; Koken, 1885 [1885.1]; Schlosser, 1903 [1903.1]; e¢ ald) belong to more advanced species, and, by a general consensus of opinion, the various writers, other than Owen, have referred them to such species as S. bombifrons, S. ganesa, or S. insignis.” “The trivial name is given in honour of Dr. O. Zdansky of the Egyptian University, Cairo, who is well known for his valuable researches on the fossil mammals of China.” Parastegodon [Stegodon?| sugiyamai Tokunaga, 1935 Figure 789 At Iruhi, in Saida village, Shikoku, Japan. Pleistocene (fide Tokunaga). Upper Pliocene or Lower Parastegodon sugiyamai Tokunaga, 1935. “A New Fossil Elephant ound in Shikoku, Japan.” Proc. Imp. Acad. Tokyo, Vol. XI, p. 434. Typrr.—An upper molar of the left side, 900 probably an M?’. Horizon Locauiry.—‘Mr._ T. Sugiyama recently sent the author [Tokunaga] a specimen of a molar-tooth, obtained during road construction at Iruhi in The stratum from which it was obtained AND Saida village, Shikoku. consists of an alternation of loose shale and coarse-grained quartz- sandstone, and probably referable to either the old Pleistocene or Tyrer Ficure.—Op. cit., p. 433, crown and inner side view of molar. Type Descriprion.—(Op. cit., pp. 432-434): “Since medi- aeval times numerous specimens of fossil bones and teeth of ele- phants have been scooped up from the sea-bottom by fishermen’s nets along the eastern part of the Inland Sea of Japan. How- ever, very seldom are specimens discovered from strata exposed on to the youngest Pliocene.” Typp or PArRASTEGODON [STEGODON?| SUGIYAMAI Fig. 789. Type molar, probably an 1.M?’, of Parastegodon sugiyamai Tokunaga, 1935, text figure, p. 433, one-half natural size. From Shikoku, Japan. the land surface along that part of the sea. The author is aware of only several cases, namely, a few found in situ at Akashi on the north shore of the sea, and at Sue village, Ayauta-gun, and Saida village, Mitoyo-gun, Kagawa-ken, both in the south of the sea.” “The present specimen obviously belongs to Stegodontinae in having a low crown, and in other points, but in shape, size, number of ridges, and especially in the nature of its enamel plication the present fossil is referable to no known species from Japan. It closely resembles Stegodon airdwana Mart., and Stegodon OSBORN: THE PROBOSCIDEA trigonocephalus Mart. from Java; and is closest to the former. The following is the description of the specimen.” “In the crown view, one side of the tooth is nearly straight and the other slightly convex. The grinding surface is oval and slightly convex, and its base is also slightly convex antero-poster- iorly, indicating that it is an upper left molar. The anterior part of the crown is broken off and there [are] preserved a posterior talon, seven ridges and a part of another; most probably the first ridge and the anterior talon are lost. The foremost, namely, second ridge has its anterior portion somewhat damaged and its outer portion broken off, and the next three ridges also lack their outer- most portions. All the ridges are distinctly exposed, not covered by cement. The posterior talon was covered by cement and first showed itself by shaving off the cement; it is half as high as the adjoining ridge. The last ridge is just touched by wear and the other ridges were worn by grinding.”’ “The present specimens, when complete, probably had nine ridges, an anterior and a posterior talon.” “The length of the crown of the present specimen is 120 mm +20 mm, the latter term being the estimated length of the lost part. The width of the grinding surface is 60 mm at the sixth ridge (67 mm at its base). The height of the crest is 34 mm at the worn eighth and 88 mm at the unworn ninth. The eighth and ninth ridge-crests decline forwards and are a little curved in side view, while the seventh but slightly declines forwards and is flat and not curved in the same view; all the other ridges stand up- right. In well worn stage each ridge-crest is narrowly rectangular, not strongly constricted, and has its enamel finely plicated; in a less advanced stage of wearing, it shows a large number of con- strictions. In these features the present specimen resembles in general S. avrawana; however, the detailed enamel plication of its crown-ridges is characteristic when compared with the speci- mens of the latter illustrated by Soergel, Janensch, Stehlin, Maarel and Koenigswald. The unworn last or ninth ridge of ours has seven mammillae on crest and the eighth has seven constric- tions. In the seventh ridge the dentine area is expanded and there are still five constrictions, while in the sixth and all the others forwards there is no constriction. The enamel is finely and rather regularly plicated in the seventh and anterior ridges and never form rough irregular undulations exhibited on grinding surface by Maarel’s Javan specimen, as his photograph shows, which is similar to our molar in the size of crown and grinding degree. The width of enamel measures 4 mm at the thickest portion in grinding surface.” “In this specimen all its ridges except the posterior talon rise higher above the cement and the last or ninth is the highest.” “The property of the enamel plication of each ridge-crest of the author’s specimen agrees with no described Japanese species and differs in several points from the Javan species. Moreover it presents several features different from fossil elephants of other lands. The present specimen evidently needs a new name, and is named, Parastegodon sugiyamai in honor of the discoverer, Mr. Tsurukichi Sugiyama. The author is at present unable to affirm whether it represents the first or the second molar, but it evidently belongs to one of them.” APPENDIX TO CHAPTER XIV MATSUMOTO ON THE PHYLOGENY AND CLASSIFICATION OF THE JAPANESE MASTODONTS, STEGODONTS, AND ELEPHANTS (1924, 1925, 1926, 1927)! The numerous and diversified Proboscidea of Japan prove that this region from Miocene to early Pleistocene times had broad land connections with southern Asia, Burma, and India, with remote relations by migration to Europe and Africa. Subsequent to and independently of the conclusions reached by Osborn on the classification and phylogeny of the mastodontine and elephantine Proboscidea, Hikoshichiré Matsumoto published the systematic and theoretic results of his important revision of the Japanese Proboscidea in six papers, as follows: “Preliminary Note on Fossil Elephants in Japan,’ Journal of the Geological Society of Tokyo, Vol. XXXI, No. 371, September 20, 1924 (1924.2). “Preliminary Notes on the Species of Stegodon in Japan,” Journal of the Geological Society of Tokyo, Vol. XX XI, No. 373, November 20, 1924 (1924.3). “Preliminary Notes on Two New Species of Fossil Mastodon from Japan,” Journal of the Geological Society of Tokyo, Vol. XX XI, No. 375, December 20, 1924 (1924.4). “On Two New Mastodonts and an Archetypal Stegodont of Japan,” Science Reports of the Tohoku Imperial University, Second Series (Geology), Vol. X, No. 1, 1926 (1926.1). “On the Archetypal Mammoths from the Province of Kazusa,” Science Reports of the Tohoku Imperial University, Second Series (Geology), Vol. X, No. 2, 1926 (1926.2). “On a New Fossil Race of the Asiatic Elephant in Japan,’ Science Reports of the Tohoku Imperial University, Second Series (Geology), Vol. X, No. 3, 1927 (1927.1). His results are summed up in his three phylogenetic diagrams reproduced herewith (Figs. 791, 792, 793) and in the following systematic summary or conspectus compiled from notes included in Doctor Matsumoto’s letters of July 14 and November 20, 1924, and from his two more recent papers of 1926 and 1927. Matsumoto’s valuable letters and notes, intercalated in the English text of his published report of 1926, give us a clear statement of his observations and opinions, as compared with those of the present Memoir indicated in square brackets. Osborn (1927) adopts certain of Matsumoto’s generic and specifie terms (e.g., Palzoloxodon Mats., which antedates Sivalikia Osb.), but rejects others (e.g., Hemimastodon Pilgrim, Prostegodon and Parastegodon Mat- sumoto, which are antedated by genera of other authors). A full synopsis of Matsumoto’s work (1924-1927)' is given herewith for monographic purposes, followed by Osborn’s comments (p. 908 below). 1. Genus Hemmastropon Pirerm [=Surna, Votume I, Cuaprer VIII} Hemimastodon Pilgrim. 1. Hemimastodon annectens sp. nov., Journ. Geol. Soe. Tokyo, Vol. XX XI, 1924.4, pp. 401, 405. See present Memoir, Chapter X, p. 457. [= Serridentinus annectens.| 2. Genus TRILOPHODON FALCONER [=TRILOPHODON OF THE PRESENT Memorr, Cuaprer VIII] Trilophodon Falconer. 2. Trilophodon sendaicus sp. nov., Journ. Geol. Soc. Tokyo, Vol. XX XI, 1924.4, pp. 402, 408. See present Memoir, Chapter VIII, p. 280. |= Trilophodon sendaicus.| ‘Por a continuation of Matsumoto’s observations and theories, up to and including 1929, see Chap. XIX, pp. 1289 to 1300 below.—KEditor. | 901 (Gee = : Loam ( Cervus ofr nippor Tyrrhenian Bison occidentalis e Milazzian « PFU BEEP By see & cfr: ae WEN : A 4 orTT ; 5S. orzentalts x (“Mitteldiluvialschotter) 2 A Narita,s.s. Elephas, last mutation aif =i L (=Z4.rnaumeannt) > Sl iat ae -— is ; eri: £ 0 a 7 IO ( ZZephas, : Cromerian -Sictlian lene : | Fide Gtetion oF Toho eee Wy) (“Altdiluvialschotter ) 23 ale Oa Beas Be SiS a \ (=?L. Aysudrindica ) PEO Go pe Payee c Tokyo — | s ates nicl Oxon ceue of ES Fg Zlephas, ?mutation oy aS agate aes || Tt Meeoreateeo aaa Se OPATE = °?L.ausonia) >! (Phocanschotter) x = : >) { Grreffa mecrodan es Calabrian | FS Naganuma F8 Stegodon orientalis ta alas SS eee | Elephas, mutation of /Tinato | F 5 ood Da oeo se a FF? \ (= Parelephas trogonthervt | Astian ee | Ourngt Se) ————— (Eee . C CETES Sa Stegodon orientalis : aoe: ean LIA HE i , 7. Sp. (Merced -Etcheqoin) oi eee Younger Lignites: v2 serra. ae @ @ o fia if EE or Cervid Sp. é RB erat Ke Takamori = we x Iz geo boy 0.0164) (i eee wo < Pontian oD Tatsunokucht F 6{ Zygolophodon, % Sp. = : FO 7 (lpyrenaccus tyoe) o | €Santa Margarita) | Bis al W Lower Lignites fan) 3 << C) (eo) Hanareyama ES Prostegodon latidens QO. i eee _ a or Upper Saboyama 2| Sarmatian = @ : Ll wy) | (:Cierbo) = Taga or Shirado ane z i 4 F 4 Dicrocerus, 7-9P L rs 5 > Tortonian S| ° x= 9 Briones) 0 TJoqgari or fee -F3Kadonosawa 43 JDesmostylus japonicus, type | ards | . in Mammonteus primigenius com- (4) Cranium shortened (cyrtocephalic), deepened (bathycephalic), heightened (hypsicephalic), pointed (acrocephalic), reaching an extreme fore-and-aft abbreviation and elevation in Mammonteus. (5) Habits varying in the three generic phyla. Originally browsing, progressive to extreme grazing type. A. PLANIFRONS Fret Br British Mus. Yq Natural size A. MERIDIONALIS 7Zype G Florence’ Mus. S. PINJORENSIS 7ype D A.IMPERATOR Fref -7. Amer Mus. 19772 va Amer Mus. ,-7 ‘ P47 Ce Fig. 817. Comparative profiles of erania of Archidiskodon (A, B, D) and cranium of Stegodon (C), all one-twentieth natural size. A, Archidiskodon planifrons ref. (Brit. Mus. M.3060). B, Archidiskodon meridionalis type (Florence Mus.). C, Stegodon pinjorensis type (Amer. Mus. 19772). D, Archidiskodon imperator ref. (Amer. Mus. 14476). Observe in A, B, D the progressive increase in size, abbreviation, heightening, and deepening of the cranium, heightening of the occipital crest, and con- cavity of the lengthening forehead. Contrast with the still more abbreviated, hypsicephalie cranium of Stegadon pinjorensis (C). While united by these profound cranial characters, the three genera, on the contrary, are widely distinguished by the specialization of their grinding teeth. It seems hazardous, therefore, to unite them in a single subfamily. The outstanding generic distinctions are as follows: Archidiskodon Pohlig, typified by Elephas meridionalis and EF. planifrons. Archidiskodon tusks and cranium as in other Mammontine. Primitive grinding teeth, subloxodont, subhypso- dont to hypsodont. Ridge-plates ex- tremely broad plated, laminate, slowly progressing in number from M 3 +{¢ (A. planifrons) to +4256 (A. imperator), Enamel thick. Mammoths of the south temperate regions; chiefly browsing. Progressive in size to the gigantic Archidiskodon imperator maibeni. Parelephas Osborn, typified by Ele- phas jeffersonii. Parelephas tusk and cranial charac- ters asin other Mammontinz; cranium less cyrtocephalic than in Archidisko- don or Mammonteus. Convergent with the Elephantine in bathycephaly and multiplication of ridge-plates; grind- ers resembling those of Hlephas indicus; ridge-plates relatively fine, progres- sively hypsodont, multiplying from P. trogontherti (M 3731), to P. jeffer- sonit (M 3 $4), to Parelephas progressus (M 3 3). Enamel medium. Mammoths of the mid-temperate re- gions of Eurasia and North America; browsing and grazing habits, adapta- tions similar to Elephas indicus. Intermediate in size, progressive to Parelephas floridanus. Mammonteus Camper-Osborn, typi- fied by Elephas primigenius. Mammonteus crania as in other Mammontine, but excessively acro- cephalic, hypsicephalic, bathycephalie, and eyrtocephalic. Grinders with ex- treme fore-and-aft compression; ridge- plates progressively deepening and broadening, the most elevated and at- tenuated known, multiplying from M3 +$=39 (M. astensis), to M3 34 (M. primigenius), to M 3 34 (M. primi- genius compressus). Enamel thin. Mammoths of cireumpolar regions; habits chiefly grazing, browsing in un- favorable seasons. Dwarfed in size, with digits in pes reduced to four. THE MAMMONTINA: ARCHIDISKODON AND METARCHIDISKODON 939 2. HISTORY OF THE GENUS ARCHIDISKODON Although long united with Hlephas, the species which are grouped in this genus constitute a very distinct generic phylum for which Pohlig’s name Archidiskodon, referring to the archaic molar ridge-plates, seems very appropriate. As shown in our “Classification of the Elephantoidea”’ (Chapter XV) above, also in figures 805, 806, 816, and 817, Archidiskodon is profoundly related in its cranial characters to Mammonteus and to Parelephas; consequently it belongs in the subfamily Mammontine, as defined on page 937 of this Memoir. A. IMPERATOR Ref. M. PRIMIGENIUS Ref P. JEFFERSONI! 7ype P WASHINGTONII Los Angeles Mts. Naz. (4us. B58O Amer. /7s. 9950 Amer. Mus. 865/ E M. PRIMIGENIUS 3° pid COMPRESSUS - 765432! Type A B D A. PLANIFRONS Type + 10 Yoo natural size Fig. 818 Cc S. AURORAE s 1 Tig. 818. HypsicepHatic CraNniA OF THE MAMMONTIN® Type A, Archidiskodon imperator, juvenile male cranium, from Rancho La Brea, California. B, Mammonteus primigenius, young adult male cranium, from Siberia. C, Parelephas jeffersonii, type cranium, aged male, from Indiana. D, Parelephas washingtonit, adult male cranium, from the state of Washington. Observe (1) the strong fore-and-aft cranial compression and vertical elevation of the occiput (hypsicephaly); (2) the slightly concave forehead; (3) the very deep depression of the mandibular rami (bathycephaly); (4) the sharp downward flexure of the basicranial axis (cyptocephaly); (5) the disparity in size of the M. primigenius skull and jaws. 10 ‘ B Wyee Fig. 819. ComparaTIVE SERIES OF SUPERIOR MOLARS SHOWING EVOLUTION OF THE RIDGE IN THE ELEPHANTOIDEA AND STEGODONTOIDEA. ONE-SIXTH NATURAL SIZE. E, Mammonteus primigenius compressus, type M°, showing extreme compression of 27 ridge-plates. D, Archidiskodon planifrons, type M*, with 9-10 ridge-plates. 10 C, Stegodon aurore, type M?, with 10+ ridge-crests (cf. S. airawana, Fig. 764C). =\ B, Stegodon ganesa, type M*, with 10+ ridge-crests. 4. a A, Stegodon insignis, type M°, with 10 ridge-crests. A ©] Stegodon aurore (cf. p. 892) “is either a highly progressive Stegodon or a primitive Archidiskodon, a point to be SINSICNIE | f I determined positively by the discovery of a cranium.” Type Sa pas wae ee Fig. 819 Pou.ia, 1885, 1888.—To our knowledge Pohlig was the first to separate the elephants with archetypal ridge- plates as Archidiskodonten (1885, p. 1027): ‘8. Ich theile die Elephanten nach Kronenformen und Lamellen- zahlen der Molaren ein in Archidiskodonten (H. planifrons, E. meridionalis), Loxodonten (EF. africanus, ?E. anti- quus) und Polydiskodonten (EF. primigenius, E. indicus ete.), die Stegodonten mit Clift wieder zu Mastodon zihlend.”” Three years later Pohlig (1888, p. 138) defined the genus as follows: ‘1. Archidiskodonten. Typus: E. meridionalis. Uebergang zu der folgenden Gruppe bildet H. planifrons. Tapinodiske, laticoronate, kurze und (i ‘a | UHH ws IN Nn, 4, " J TF @AMRITSAR 4, | «© | @ S MALAHoRE mil Ay elif y a | | INDIA,1857 ts © 190 209 300 400 S00 || 2a* — : 75° AMERICAN MUSEUM OF NATURAL HISTORY ERWIN J RAISZ A.M. 1927 CreNTRAL REGION (SEE BLACK SQUARE) OF THE SIwALik HiLxs, 200 MILES SOUTH AND NORTH OF Srmva, [npr Fig. 820. A flanking Tertiary range, 800 miles in length, on the southwestern base of the Himalay exposures (vertical lines, now divided into Upper, Middle, and Lower in 1831, and determined as of Tertiary age by the bones of crocodiles, and Fig. 826, detail of the Simla region). a Mts., and northeast of the Punjab. Classie “Sewalik” Siwaliks) discovered by Lieut. Proby T. Cautley in 1827, explored by Dr. Hugh Falconer tortoises, and other fossil vertebrates (cf. Fig. 729, after Falconer, Pl. xxv, map of India, 940 THE MAMMONTINA: ARCHIDISKODON AND METARCHIDISKODON 941 pachyganale Molaren. Parsilamellat. (Meist nur 15 Lamellen an M. III.)” In the same Memoir (p. 252) he introduced the generic terms: 4° Archidiskodon 5. Polydiskodon 4°. Loxo(-disko-)don. From these citations we have abstracted the above generic characters. SCHLESINGER, 1912, 1916.—Schlesinger (1912, pp. 98, 99, footnote) confirmed this generic description and amplified Pohlig’s definition as follows: “‘Ausdriicke halte ich fiir einen Missgriff und gebe ihre Erklirung: Tapino- disk = mit niedrigen Jochen (Gegensatz=hypselodisk), latikoronat = breitkronig (Gegensatz=angustikoronat), pachyganal = mit dickem Schmelzblech (Gegensatz = endioganal), parsilamellat = mit wenigen Lamellen (Gegen- satz=densilamellat).”’ In a later paper (1916, Fig. 821. Chief Lower and Upper Pleistocene localities of western Eurasia in which @ = _ =, = y © “ive Ds 102, 103) Schlesinger eites the ter 7 a a occur species of Archidiskodon, Parelephas, Mammonteus, Loxrodonta, and Palzoloxodon. subgenus: “Hlephas (Archidiscodon) meridion- After Osborn, 1910.346, p. 391, fig. 176. See more detailed caption, figure 932, below. alis Nesti,”’ and (op. cit., pp. 112, 113)“Elephas (Archidiscodon) planifrons Fale.’ Schlesinger also in his “Studien iiber die Stammesge- schichte der Proboscidier” of 1912 recognized a resemblance to the broad-plated American species Hlephas columbi and E. imperator, but failed to separate these species into a distinct generic phylum. Osborn (1922.555) distinguished Hlephas columbi from EH. imperator (p. 3) as follows: “We thus find by the characters of the type and neotype specimens that the real Hlephas columbi is not the animal we have been describ- ing under this name; it is a dwarf form, per- haps a dwarf female, of the animal which we Pp 4 PLEISTOCENE EUROPE.—1 Forest Bed of Cromer (Norfolk). Sables de 2 seribi . ‘ KRlephas St. Prest near Chartres (Eure-et-Loire). 3 Malbattu (Puy-de-Dome). 4 Peyrolles (Bouches- have been describing under the pee Hlep : du-Rhone). 5 Solhilac near Puy. Clay deposits of 6 Durfort (Gard). 7% Cajarc (Lot-et-Ga- ronne). 8 Val d’Arno (Tuscany). 9 Leffe near Bergamo (Lombardy). 10 Rizdorf near Pots- dam (Brandenburg). Gravels of 11 Siissenborn near Weimar. Sands of 12 Mosbach in f northern Baden. Freshwater deposits of 13 Clacton (Essex). Sands of Mauer near 14 Hei- He clearly defined (op. Cita, pp. 3 and 4) delberg (western Germany). 15 Chelles on the Marne, near Paris. 16 St. Acheul (Somme). 17 Ilford and Grays Thurrock.(Essex). Lignites of 18 Diirnten and of Utznach, near Zurich. imperator.’”! the characters of the grinding teeth of HE. am- 19 Taubach near Weimar. 20 Wildkirchli cave on Mont Sdntis (eastern Switzerland). Tuffs 5 « c - of 21 the Tiber Valley, near Rome. Caves of 22 Neandertal, near Diisseldorf (western Ger- perator, concluding (p. Sys The cranial char- many), 23 Spy, near Amur (Belgium), 28a Krapina (Croatia), 24 Chapelle-aux-Soints (Cor- réze). Caves and alluvial deposits of 25 Ternifine (or Palikao) near Oran (Algeria), 26 Pointe acters observed in three more or less complete Pescade, near Algiers (Algeria). 27 Prince's Cave (Monaco). Sandy clays of 28 Véklinshofen A (Alsace). 29 Saalfeld (Saxe-Meiningen). Travertines, etc., of 30 Gera, Jena (Saxe-Weimar). skulls referred to Elephas imperator tend to 31 Leipzig Saxony). 32 Solutré, north of Lyons. Loess of $8 Wiirzburg (Bavaria). 34 Thiede : : near Braunschweig (Prussia). Cave of 35 Montmaurin (Haute-Garonne). 86 Chdteauneuf- support the direct descent of this animal from sur-Charente (Charente). Caves of 87 Schweizersbild near Schaffhausen, and Kesslerloch near ace : 9 Thayngen (northern Switzerland). Remains of lake dwellings at 38 Wauwyl (Lucerne), 39 Ro- the EH. meridionalis of the Val d’Arno, Upper _ tenkausen, south of Lake Pfiffikon, 40 Concise on Lake Neuchatel (Switzerland). Peatbogs of : on 41 Hassleben, near Weimar. Travertines of 42 Langensalza (Erfurt) in central Germany. Pliocene of Italy. Caves of the 43 Island of Malta, 44 Island of Crete, 45 Island of Cyprus. Subsequently Osborn (1924.633, p. 2) confirmed Pohlig: ‘‘We therefore confirm Pohlig’s separation of the southern mammoths Elephas planifrons, E. meridionalis, and E. imperator into the distinct generic phylum Archidiskodon,” and in the year 1925 (Osborn, 1925.662) he placed Archidiskodon among the Mammontine, as shown in the diagram of that year (Vol. I, Fig. 7, of the present Memoir), concluding (p. 28) with a redefinition of the family Elephantide and Race XIII, as follows: 'Subsequently (see pp. 997, 1001 )Professor Osborn separated Elephas imperator and E. columbi, assigning the former to the genus Archidiskodon Pohlig and the latter to Parelephas Osborn.—Editor.] FAMILY: OSBORN: THE PROBOSCIDEA ELEPHANTID~, distinguished by plated grinding teeth developing out of the more or less closely compressed, serrated ridges of Stegodon'! into the broadly plated grinders of Archidiskodon, the lozenge-shaped grinders of Loxodonta, and the compressed, finely plated grinders of Parelephas, of Mammonteus, and of Elephas the type genus of the family... . Race XIII. THe SourHern Mammorus, or ArcHIpIskoponts. Excessively broad-plated grinders with abundant cement; first known in India, migrating westward into southern Europe, eastward into America, where arriving in late Pliocene or early Pleistocene time they finally gave rise to the Im- perial Mammoth, Archidiskodon imperator, the last of its race. Fig. $22. Worumerg: i \ ee ee 7 eI} MAMMONTINA Geographic distribution of the principal species of Archidiskodonts. The white dots within the black areas represent the approximate localities where the types of these twenty-two species were discovered; these dots each carry a number in a circle representing the chronologic sequence of type de- scription. 0. The white crosses represent some of the principal referred specimens mentioned in the present Memoir. 3. ORDER OF DISCOVERY AND DESCRIPTION OF TWENTY-TWO SPECIES OF ARCHIDISKODONTS Sree Figure 822 REFERENCE IN PRESENT MerMOIR 1825 Elephas meridionalis Nesti, Val d’Arno, Italy = Archidiskodon meridionalis 1846 [1845] Elephas planifrons Falconer and Cautley, Siwalik Hills, northern India = Archidiskodon planifrons 1855 =F. giganteus Aymard (MS., also in Falconer, 1857, p. 321), France Nomen nudum = Archidiskodon meridionalis(?) {1857-1868 Elephas columbi Falconer, Georgia = Parelephas columbi (see Chap. XVII)] 1858 Elephas imperator Leidy, ?Seneca, Thomas Co., Nebraska (fide Hay, 1924, p. 100)? = Archidiskodon imperator {1859-1861 Hlephas texrianus Owen, 1859, Blake, 1861, Texas = Parelephas columbi (see Chap. XVIT)| 1889-1890 Hlephas lyrodon Weithofer, Val d’Arno, Italy = Archidiskodon meridionalis (female?) {1890 Elephas mindanensis Naumann, Philippine Islands = Stegodon (Archidiskodon?) mindanensis (see Chap. XIV)| 1915 EHlephas hayi Barbour, Crete, Saline Co., Nebraska =Archidiskodon hayi {1922 Hl. |Elephas| Columbi var. Felicis Freudenberg, Mexico =Parelephas columbi felicis (see Chap. XVII)] 1922 Il. [Elephas| Columbi var. silvestris Freudenberg, Ejutla, State of Oaxaca, Mexico = Archidiskodon imperator silvestris ‘(See Chapter XIV, pp. 806, 807, where Professor Osborn’s final views regarding the complete separation of the Stegodontoidea from the Elephantoidea are given.—Editor.| *1Exact locality not cited. Lugn and Schultz (1934.1, p. 373) give Pawnee Loup Branch of Platte River= Middle Loup, probably Hooker County.— Editor.] THE MAMMONTINA: ARCHIDISKODON AND METARCHIDISKODON 943 6. 1922 Hl. [Elephas| Columbi var. Falconeri Freudenberg, Tequixquiac, Valley of Mexico = Archidiskodon imperator falconeri 7. 1922 Loxodonta griqua Haughton, Griqualand West, Transvaal, S. Africa = Metarchidiskodon griqua 3. 1922 EHlephas Columbi var. imperator Freudenberg, Spokam Bar, near Helena, Montana = Archidiskodon imperator 8. 1923 Elephas meridionalis, mutation cromerensis Depéret and Mayet, Kessingland, Suffolk, England = Archidiskodon meridionalis cromerensis 9, 1924 EHlephas antiquus rumanus 8. Stefanescu, Tulucesti (Covurlui), Rumania = Archidiskodon planifrons rumanus 10. 1925 Hlephas scotti Barbour, near Staplehurst, Seward Co., Nebraska = Archidiskodon imperator scotti (or juve- nile A. zmperator) 11. 1925 Hlephas maibeni Barbour, near Curtis, Lincoln Co., Nebraska = Archidiskodon imperator maibeni {1927 Archidiskodon transvaalensis Dart, Lowest Terrace, Vaal River, 8. Africa = Palxoloxodon transvaalensis (see Chap. XIX)] {1927 Archidiskodon sheppardi Dart, Lowest Terrace, Vaal River, 8. Africa = Palxoloxodon sheppardi (see Chap. XIX)] 2. 1927 Leith-Adamsia siwalikiensis Matsumoto, Siwalik Hills, India = Archidiskodon planifrons 12. 1928 EHlephas haroldcooki Hay, Frederick, Oklahoma = Archidiskodon haroldcooki 13. 1928 Elephas exilis Stock and Furlong, Santa Rosa Island, California = Archidiskodon exilis 14. 1928 Archidiskodon subplanifrons Osborn, Sydney-on-Vaal, 8. Africa = Archidiskodon subplanifrons 15. 1928 Archidiskodon broomi Osborn, near Kimberley, 8. Africa = Archidiskodon broomi 16. 1929 Archidiskodon sonoriensis Osborn, near Arizpe, northern Sonora, Mexico = Archidiskodon sonoriensis [1929 Lox. (Pal.) Tokunagai junior mut. Matsumoto = Palzoloxodon (Archidiskodon?) tokuna- gai mut. junior (see Chap. XIX)| 17. 1929 Archidiskodon vanalpheni Dart, Sydney-on-Vaal, 8. Africa = Archidiskodon vanalpheni 18. 1929 Archidiskodon milletti Dart, Sydney-on-Vaal, 8. Africa = Archidiskodon milletti 19. 1929 Archidiskodon loxodontoides Dart, Sydney-on-Vaal, 8. Africa = Archidiskodon loxodontoides 20. 1929 Archidiskodon yorki Dart, near Christiana, 8. Africa = Archidiskodon yorki {1929 Archidiskodon andrewsi Dart, ?Middle Terrace, Vaal River, §. Africa = Palzxoloxodon? andrewsi (see Chap. XIX)] [1929 Archidiskodon hanekomi Dart, Vaal River, 8. Africa = Palxoloxodon hanekomi (see Chap. XIX)] 21. 1932 Archidiskodon meridionalis nebrascensis Osborn, Angus, Nuckolls Co., Nebraska =Archidiskodon meridionalis nebrascensis 22. 1934 Archidiskodon proplanifrons Osborn, Gong-Gong, near the Vaal River, 8. Africa = Archidiskodon proplanifrons 4. ARCHIDISKODONTS OF EURASIA AND AMERICA In the early part of the 19th century there was described by Nesti from northern Italy (1808) the first known species of this phylum, to which he later (1825) assigned the name Hlephas meridionalis, or, as we may now call it, the ‘Southern Mammoth,’ in reference to its remote relationship to the ‘Northern Mammoth.’ In 1846 there was described from India a direct ancestor of H. meridionalis, namely, Elephas planifrons, the name alluding to its flattened forehead. In 1857 Falconer recognized as a remotely related species his Hlephas columbi from Georgia, a species long confused with Archidiskodon. In 1858 Leidy described as a closely related species Hlephas imperator from western Nebraska. The Elephas texianus of Owen and Blake, 1859, 1861, 1862, is a synonym of Elephas [Parelephas] columbi. The Elephas lyrodon of Weithofer, 1889, 1890, from northern Italy, is the female(?) form of E. meridionalis. Then a long interval followed and a very primitive member of this phylum, namely, Elephas hayi, was discovered in western Nebraska and described (1915) by Barbour, indicating that these animals may have migrated into North America directly from India, because EH. hay: appears to be almost as primitive in jaw structure as the Hlephas planifrons of India. 944 OSBORN: THE PROBOSCIDEA In 1922 Freudenberg reviewed the elephants of Mexico and described four subspecies of Hlephas columbi namely, Hl. Columbi var. Felicis, El. Columbi var. silvestris, El. Columbi var. Falconeri, and El. Columbi var. imperator; the first (Hl. Columbi var. Felicis) we regard as more closely related to Falconer’s species Elephas columbi, the remaining three to Leidy’s species Hlephas imperator. In 1923 Depéret and Mayet devoted a volume to their thorough researches of the species of eastern Eurasia, Elephas planifrons and EH. meridionalis, and added the Lower Pleistocene subspecies Hlephas meridionalis, muta- tion cromerensis, of Kessingland, Suffolk, England. The wide geographic distribution of Archidiskodon is indicated by Stefanescu’s description in 1924 of Hlephas antiquus rumanus of Rumania, by the discovery of two American species, Hlephas scotti and EH. maibeni, in Nebraska, described by Barbour in the year 1925, and finally by the most surprising and welcome discovery of all, the Archidiskodon subplanifrons and A. broomi in the Vaal River diggings of South Africa, described by the present author in 1928, as well as Dart’s new species, Archidiskodon transvaalensis and A. sheppardi from the Vaal River gravel terraces, described in 1927.' 5. NEW ARCHIDISKODONTS AND LOXODONTS OF AFRICA After the present author had described and figured, but not published,” the two names above (Archidiskodon subplanifrons and A. broomz), he received Prof. Raymond A. Dart’s ‘‘Mammoths and Man in the Transvaal,” Nature, December 10, 1927, with the welcome figures [Fig. 823 of the present Memoir] and description of the Vaal River terraces at Windsorton and Bloemhof, of which the following is an abstract: The Vaal River valley, near Bloemhof, in the southwestern Transvaal, belongs to a great watershed which has yielded Australopithecus africanus Dart, 1925 [from Taungs district, Bechuanaland], and the Boskop man, besides stone implements, rock engravings, and other evidences of primitive man. The watershed includes three gravel deposits of different geologic age (Du Toit, Ann. Rept. Geol. Comm., 1906) situated on a number of terraces. Upprrer.—The highest and presumably the oldest [Pliocene?] terrace has an altitude of some 200 to 300 feet above the river exposed at a distance of 3! to 6 miles. No fossils have hitherto been described from this most ancient terrace (Dart, op. cit., p. 1). [This is possibly the level at which Archidiskodon subplanifrons Osborn, a very primitive form, was found—Fig. 875 of the pres- ent Memoir.—Osborn. | Mippie.—The middle or second [Pleistocene or Mastodon] gravel terrace is on a lower level at several points west of Barkly West; from this terrace numerous palolithic implements have been described (Hodkinson, 1926); also (Beck, 1906) a frag- mentary tooth of Mastodon (Bunolophodon) sp. Felix, at Waldeck’s Plant, 60 to 80 feet above the river bed. Haughton described Loxodonta griqua |= Metarchidiskodon—see figure 882 of the present Memoir], also a new giraffe (Griquatherium cingulatum) from this 60 to 80 foot terrace. [Doctor Broom holds that the mammoth teeth are washed in from an older geologic deposit (see Nature, March 3, 1928, p. 324) and are not truly associated with the flint implements, in geologic age. ] Lower.—A still lower gravel deposit [Pleistocene], near the level of the present river Vaal, contains Hquus and Hippo- potamus amphibius var. robustus. From this low river bed gravel, 1% miles below The Bend on the Vaal River, Haughton determined as Elephas (ef. antiquus) a portion of a tooth recovered at a depth of 5 feet, but which Dart regarded (op. czt., p. 42) as resembling 2. antiquus recki Dietrich. {Apparently this same specimen is now described by Osborn as Archidiskodon broomi, the label of which bears the inscription, ‘3682 Mus. Kimb. The Bend. H. Else.” Fig. 877 of the present Memoir.| From this level are recently recorded Archidiskodon transvaalensis Dart and A. sheppardi Dart."! 'Archidiskodon transvaalensis and A. sheppardi prove to belong to the genus Palxoloxodon (see Osborn, 1934.925, pp. 2and 14). Since this section of the Memoir was written, four species from South Africa have been described by Professor Dart (1929) and one by Professor Osborn (1934); also two species from North America have been described by Professor Osborn, one from Mexico (1929) and one from Nebraska (1932). A complete list of the species of Archidiskodon is given on pages 942 and 943 above.—Kditor.| 2Subsequently published in Nature (Osborn, 1928.749, pp. 672, 673) under the title “Mammoths and Man in the Transvaal.” THE MAMMONTINA: ARCHIDISKODON AND METARCHIDISKODON 945 Gravet TERRACE 60-80 FEET River BED GRAVEL TERRACE ‘ Dia pase Tue Vaat River GRAVEL TERRACES, SouTH AFRICA, THE SCENE OF THE DISCOVERY OF Mammotus AND MAN IN THE TRANSVAAL Arrer Dart, 1927 Fig. 823. These terraces (upper at Windsorton, middle and lower at Bloemhof) have yielded the types of Boskop man (Homo capensis), of Aus- tralopithecus africanus, of Loxodonta [= Metarchi- diskodon| griqua Haughton, of Archidiskodon [=Palxolorodon| transvaalensis and A. [=P.| sheppardi of Dart, and of A. subplanifrons and A. broomi of Osborn, also of Mastodon (Buno- lophodon) sp. Felix. After Dart, 1927, figs. 1 to 3. Agra covento BY DIAMOND --0--beewn DiGeINCs: ‘ ISLAND J ©. Oxb pry arp ULD RIVER BED DIGGINGS SAND House ON ISLAND NEw River peo ‘ ’ \ ' , GRAVEL BED oF G UNKNOWN DEPTH + Consequently a summary of the discoveries and descriptions of South African specimens (1928) is as follows: ORIGINAL DESCRIPTION REFERENCE IN PreseNT Memoir Loxodonta griqua Haughton, 1922, type, Griqualand West, Transvaal = Metarchidiskodon griqua Elephas (Loxodon) zulu Seott, 1907, type, Zululand = Loxodonta zulu Elephas zulu, referred by Hopwood, 1926, Kaiso Bone-beds, near Lake Albert = Loxodonta zulu Elephas antiquus Recki Dietrich, 1916, type, Tanganyika Territory = Palzxoloxodon recki Elephas aff. meridionalis Nesti, 1825, referred by Hopwood, 1926, from Kaiso Bone-beds, near Lake Albert = Metarchidiskodon griqua (or A. planifrons?) Archidiskodon transvaalensis Dart, 1927, type, middle Vaal River gravel terrace = Palzoloxodon transvaalensis Archidiskodon sheppardi Dart, 1927, type, lowest Vaal River gravel terrace = Palxoloxodon sheppardi Archidiskodon subplanifrons Osborn, 1928, type, middle(?) Vaal River gravel terrace = Archidiskodon subplanifrons Archidiskodon broomi Osborn, 1928, type, middle(?) Vaal River gravel terrace = = Archidiskodon broomi Mastodon (Bunolophodon) sp. Felix, referred by Beck, 1906, Waldeck’s Plant = Trilophodon(?) sp. indet. The above species of Archidiskodon from Africa are fully described below, following the description of the Eurasiatic Archidiskodonts. 946 OSBORN: THE PROBOSCIDEA 6. APPROXIMATE ASCENDING PHYLOGENETIC ORDER OF SUCCESSION OF SPECIES OF ARCHIDISKODON AND PARELEPHAS (1928) Only an approximate phylogenetic order of succession of the species of Archidiskodon may be given at the present time. In this phylogenetic list we may include with some certainty fifteen of the better-known species of Archidiskodon and Parelephas, placed in ascending order as follows: ARCHIDISKODON AND PARELEPHAS OF NORTH AMERICA AND EURASIA Pleistocene? level 1857-1868 Southern United States and Parelephas columbi Falconer, originally described by Falconer as Mexico Elephas columbi, 1857 1859-1861 Texas Parelephas columbi Falconer, originally described as Hlephas texianus by Owen, 1859, by Blake, 1861 1922 Mexico Parelephas columbi felicis Freudenberg, originally described by Freudenberg as Elephas Columbi var. Felicis, 1922 1922 Mexico Archidiskodon imperator silvestris Freudenberg, originally describ- ed by Freudenberg as Elephas Coluwmbi var. silvestris, 1922 1922 Mexico Archidiskodon imperator falconeri Freudenberg, originally de- scribed by Freudenberg as Elephas Columbi var. Falconeri, 1922 Lower Pleistocene 1925 Nebraska Archidiskodon imperator maibeni Barbour, originally described ? Aftonian (fide Osborn) by Barbour (1925) as Elephas maibeni, subsequently (1926) Iowan or Late Pleisto- as Archidiskodon maibent cene (fide Lugn and Schultz, 1934)! Lower Pleistocene 1925 Nebraska Archidiskodon imperator scotti Barbour, originally described by Barbour as Elephas scotti, 1925 Pleistocene 1928 California Archidiskodon exilis Stock and Furlong, originally described by Stock and Furlong as Elephas exilis, 1928 Pleistocene 1858 Nebraska Archidiskodon imperator Leidy, originally described by Leidy as Elephas imperator, 1858 Lower Pleistocene or 1915 Nebraska Archidiskodon hayi Barbour, originally described by Barbour as Upper Pliocene (?)? Elephas hayi, 1915 Lower Pleistocene 1923 England, Forest Bed Archidiskodon meridionalis cromerensis Depéret and Mayet, originally described by Depéret and Mayet as Elephas meridionalis, mutation cromerensis, 1923 Upper Pliocene 1825 Italy Archidiskodon meridionalis Nesti, originally described by Nesti as Elephas meridionalis, 1825 1846 [1845] India and southern Europe Archidiskodon planifrons Falconer and Cautley, originally de- scribed by Falconer and Cautley as Elephas planifrons, 1845, 1846 ARCHIDISKODONTS OF AFRICA Pleistocene 1927 South Africa Archidiskodon |= Palzxoloxodon| transvaalensis 1927 South Africa Archidiskodon {= Palxoloxodon| sheppardi Lower Pleistocene(?)* 1928 South Africa Archidiskodon broomi Osborn Pleistocene 1922 South Africa Metarchidiskodon griqua Haughton, originally described by Haughton as Loxodonta griqua, 1922 Upper (?) Pliocene? 1928 South Africa Archidiskodon subplanifrons Osborn According to the above phylogenetic arrangement, the four species Archidiskodon subplanifrons*, A plani- frons, A meridionalis, and A. hayi? are the only known [1928] Upper Pliocene representatives of this great phylum. See page 1028 below for the present author’s final determination.—Editor. | *{Lugn and Schultz (1934.1, table opp. p. 358) regard Archidiskodon hayi as of Kansan (Grand Island) age, equivalent to the Lower Pleistocene.— Editor.} 3{Dr. C. van Riet Lowe places Archidiskodon subplanifrons in the early Pleistocene and A. broomi in the late Pleistocene (see van Riet Lowe, 1929.1, table opp. p. 682).—Editor.| THE MAMMONTINA: ARCHIDISKODON AND METARCHIDISKODON 947 FamMILy GROUP OF ARCHIDISKODON IMPERATOR MAIBENI ALONG THE PLATTE RIvER, NEBRASKA Restoration by Margret Flinsch Buba in 1935, under the direction of Henry Fairfield Osborn Bull in foreground 1/40th natural size and bull at extreme left 1/140th natural size. Cow and calf in middle foreground 1/60th natural size Fig. 824. The bull in the foreground and the one in the distance at the extreme left were based on measurements of the skeleton of Archidiskodon imperator maibeni in the Nebraska State Museum (Neb. Mus. 5-9-22), from Lincoln County, Nebraska (see Figs. 910-912, 918 of the present Memoir), supplemented by a fine skull of A. imperator in the Geological Institute of Mexico (No. 212) from Tepexpan (see Fig. 902) and of a giant tusk of the same species (A. imperator) in the American Museum (Amer. Mus. 22481), from Post, Texas (see Fig. 894). The height at the withers (top of shoulder blade) is 3826 mm.; adding the usual 6}4% gives a height in the flesh of 4068 mm. or 13 ft. 4)4 in. II. CHARACTERS OF ARCHIDISKODON AND METARCHIDISKODON AND INCLUDED SPECIES Superramity: ELEPHANTOIDEA Osborn, 1921 FAMILY: ELEPHANTID& Gray, 1821 SUBFAMILY: MaAmmMontinaz Osborn, 1921 Genus: ARCHIDISKODON Pohlig, 1885, 1888 Original reference: Zeitschr. deutsch. geol. Ges.. XX XVII, Heft IV, p. 1027 (Pohlig, 1885.1); Nova Acta Leop. Carol., LIII, No. 1, pp. 138, 252 (Pohlig, 1888). Genotypic species: Elephas meridionalis and E. planifrons. Syn.: Parastegodon Matsumoto, 1924 (in part); genotype, Hlephas aurore Matsumoto, 1924. Leith-Adamsia Matsumoto, 1927; genotype, Lezth-Adamsia Siwalikiensis Matsumoto, 1927. GENERIC CHARACTERS (TRANSLATION AFTER PoHLiG, 1888, Pp. 138).—Genotypes E. meridionalis, E. planifrons; transitional to higher groups. Molars with few ridge-plates, not exceeding fifteen [?] lamelle; crowns broad, short, and with thick enamel (i.e., ‘“laticoronate, kurze und pachyganale Molaren’’); ridge-plates tapering to summit in sections. 948 OSBORN: THE PROBOSCIDEA GENERIC CHARACTERS (OSBORN, 1928).—Superior tusks large, incurved, crossing in old males as in other Mammontine. Cranium extremely heightened (hypsicephalic), foreshortened and broadened (brachycephalic), deepened (bathycephalic). Molar ridge-plates extremely broad, enamel border thickened, cement usually very thick. Ridge-plate formula slowly progressing from minimum, M 3 1° (A. planifrons), to maximum, M 3 +235 (A. imperator). Finally attaining gigantic size. This genus is given first consideration in the history of the subfamily Mammontine for the reason that Elephas |= Archidiskodon| planifrons of the Upper Pliocene of India is geologically the earliest of the Elephantide at present known. It is a very striking circumstance that until we reach the Upper Pliocene, Pinjor horizon, of India we discover Mastodonts and Stegodonts only, there being no trace of the Elephantide; consequently Archidiskodon planifrons appears to be a new arrival in southern Eurasia in Upper Pliocene time, probably a de- scendant of ancestors coming from some region of southern Africa, such as Archidiskodon subplanifrons.' Osborn, 1929: Comparing Falconer’s measurements of thirty-nine Siwalik specimens (Table VI, opposite page) with Barnum Brown’s measurements of twenty-seven Siwalik specimens (p.954 below), we find a closely similar range of ‘ascending mutations’ rising from the most primitive ridge formula, M 3 5.24, through intermediate mutations, M 3 ;%;, to the most progressive mutations, M 3 ,4:°%4;. Thus in sixty-six specimens collected in the Pinjor horizon, of the Siwaliks, India, we observe gradual ascending mutations of the Archidiskodon planifrons stage toward the A. meridionalis stage. The ridge-plates throughout in M; are more numerous than those in M*. Summary.—The ascending mutations in the thirty-nine specimens referred by Falconer (1868) to ‘Hlephas planifrons’ are seen to compare very closely with those in the twenty-seven specimens of the collection made by Barnum Brown and referred by Osborn to Archidiskodon planifrons. This renders it probable that the Falconer collection and the Brown collection were from the same geologic horizon. The extremes in both collections are seen in the maximum and minimum heights of the ridge-plates: Upper PLIOCENE ARCHIDISKODON PLANIFRONS MEASUREMENTS OF RIDGE-PLATES IN MILLIMETERS Falconer (1868) Brown (1922) Max. Min. Max. Min. Third suverior molars, M* Height 123 64 102 88e Length 279 247 254 201 Third inferior molars, M3 Height 115 77 135 53 Length 323 224 313 203 Allowing for individual differences in size due to sex and geologic level, and for probable differences in mode of measurement, especially of the height of the ridge-plates, it appears probable that in both collections we have to do with a series of ascending mutations some of which attain the size and ridge-plate height of Archidiskodon meri- dionalis. It is noteworthy that in the A. planifrons r.M®, the length (221 mm.) and the height of the 7th ridge- plate (7le mm.) agree closely with the 7th ridge-plate height of M? of A. planifrons from the Piltdown gravels of Sussex, England (see Figs. 852, 853, 854). Falconer observes that while the ridge-plates in A. meridionalis increase in number they do not increase correspondingly in height. In two of his figures reproduced in the present Memoir (Fig. 862) the measurements are: PRIMITIVE ARCHIDISKODON MERIDIONALIS [OR A, PLANIFRONS| Length Breadth Height Val d’Arno, r.Mz 10 in. [=255 mm.] 3.4 in. [=87 mm.] 5 in. [=126 mm.] Norwich Crag, r.M,; 11.25,[=287 mm:] 3:8 [=97 mm.] 4.8 [=123 mm.] ‘Or Archidiskodon proplanifrons, an equally primitive species subsequently described by Professor Osborn from Gong-Gong, Vaal River (Osborn, 1934.925, p. 10).—Editor.] THE MAMMONTINA: ARCHIDISKODON AND METARCHIDISKODON 949 UPPER SIWALIKS MUTATIONS TYPICAL ARCHIDISKODON PLANIFRONS: M 3 3#t, ASCENDING TO TYPICAL A. MERIDIONALIS: M 3 4}3=14 TasLe VI. SumMMARY OF MEASUREMENTS OF THIRTY-NINE SPECIMENS REFERRED BY FALCONER, 1868, Tro ELepHas [= ARCHIDISKODON] PLANIFRONS Length Breadth Height SUPERIOR Plate Figure Molar in. mm. in. mm. Index in. mm. Ridge-plates Third Molars XII Footnote, p. 433. Unfig. M3 11 =279 3.7-4.8=95-123 34 8th 4.8=123 Y-12-}5 XI 1 1. M8 11 =279 4.5=114 4 ry, M3 5:4 —=135) 25 5— 63 8+ XIV 8 *r. M3 10) =255) 325— 89 10th? 2.5= 63 8-9-4 r. M? 9.7=247 4 =102 x r.Mé 9.7=247 3.5= 89 4 =102 Second Molars I] 5a Type r. M? 8.7 =221 [7th =. le] 9 XII 5 r. M2 8 =204 3 77 3 8 5a 1. M2 (.0=191 3.2= 42 84 6, 6a r. M2 8 First Molars VI Footnote, p. 427 M! 6.5=165 3.4= 87 52) 4th 3r0— 89 -7-'6 6 M! DL o— 140) 22 7= 169 A9N 4thy oe ene y-7-'6 XII 4, da .M! Dea 2a Oia 55 Us Fourth Deciduous VI 5 Dp? 4 =102 2.4= 61 60 -6-}5 Third Deciduous XII 2 Dp* 3:8= 97 2.8= 59 60 4-6-1 Second Deciduous 1, la Dp? 1 True Premolars VI 4 Noe 1F2— 3028 96 .8= 21 83 with indistinct front and back | ridges INFERIOR Third Molars XII Footnote, p. 433. Unfig. r.M; 11.8=300 3.9=100 3a0) ithe 46115 1444 1.M3 11 =279) 3.3= 84 30 8th 3 = 77 1344 13 1.Ms; 1221, —323) 050 —s9I 28 10th 4.5=114 13-%+ XI 8 M.[M;] 12.1=308 3.8= 97 32 11-4 2 EEMig elie S—300) 03) ana 25 14-13-%4 | 5 r.M; 10.5=267 4.2=105 39 9th 3.5= 89 QF II 5b Paratype 1.M3 OFF XI 7 1.Ms; 10.2=259 4 =102 44 XVIII.A Na 1.M3 10.4=265 4.1=104 2) Ol 8 XI 3 1.M; 10) = 2557 326—" 91 36 if XII 12, 12a ’r.Ms; 10 = 25589 7th 4 =102 ¥-10-% VIII 2, 2a r.M; 8.8=224 3.8= 97 43 9-11 XIV 9 r.M3; 915 =241 ro —1 89 6th 38 = 77 845 XI 10 1.M; B7= OF Sth 3.8= 97 | Second Molars 6 1.Mz. 8 =204 2°8= 71 9-' | 10 1.M> 7 =178 3.7= 95 8% First Molars 6 1.My, G shy 2s 7 6+ XII 10, 10a 1.M, Ge — 1 226 — 566 7 Fourth Deciduous 8, 8a 1.Dps APA i324 aol 4-7-% | Third Deciduous Us UG r.Dps 24 —) 61 14 — 36 6-13 | True Premolars 10, 10a 1. Pidrag:) 1 = 26 1 = 26 1.5= 38 Does not | 8, 8a I) 1 1 — 226s — oll show ridges One of the genotypes of Leith-Adamsia Matsumoto.—Hditor.] “Valuable because found with r.Mz (Pl. xrv, fig. 9). [Pl. x1v, fig. 8, represents one of the genotypes of Leith-Adamsia Matsumoto.—Hditor. | 950 OSBORN: THE PROBOSCIDEA 1, ARCHIDISKODONTS OF SOUTHERN EURASIA From our present knowledge it appears probable that Archidiskodon originated in Africa, because seven stages in the specific evolution of this genus have been discovered on that continent. In India Archidiskodon planifrons suddenly appears in the Upper Pliocene, Pinjor horizon,’ while A. meridionalis appears in the uppermost Pliocene and Lower Pleistocene of western Europe. The subspecific name Elephas [= Archidiskodon]| meridionalis cromeren- sis has been applied by Depéret and Mayet to rather primitive archidiskodont molars found in the Forest Bed of East Anglia. SYSTEMATIC Archidiskodon planifrons Ialconer and Cautley, 1846 [1845] Figures 815, 817, 819, 825, 827-850, 852-856, 865, 871, 876, 914, 1208, 1231, 1239, Pl. xx1 ‘Sewalik Hills’ of Falconer, northern India; Pinjor horizon, Upper Pliocene, Upper Siwaliks (Pilgrim)!; Upper Siwaliks, below Boulder Conglom- erate, Lower Pleistocene (Brown). Syn.: Leith-Adamsia Siwalikiensis Matsumoto, 1927. Sprectric CHARACTERS (FALCONER, 1846, 1863, 1868; OsBorn, 1928).—Species distinguished from Hlephas [=Stegodon] aurore by the more elevated or hypsodont ridge-plates; distinguished from Elephas |= Archidiskodon| meridionalis by the much less elevated ridge-plates; by the flattened forehead to which the specific name planifrons refers in distinction from the concave forehead of A. meridionalis; also by the more primitive ridge-plate formula, M 3 2°, and by the succession of premolars, this being the last species of the Elephantide in which the true premolars, P 3 and P 4, are erupted. Rostrum of jaw elongated and depressed (see Vig. 849). Mareriats.—The above comparison of measurements of thirty-nine specimens in the Falconer and Cautley collection (1830-1840) and of twenty-seven or more specimensinthe American Museum collection of 1922 made by Barnum Brown (see Summary of Measurements, Tables VI and VII, pp. 949 and 954) renders it probable that both collections, totaling sixty-six or more speci- mens, came from the same general locality and represent ascending mutations, from the more primitive typical Archidiskodon plani- frons, M 3 10F | toward the more progressive A. meridionalis stage, M 3\°=**. We have to do, therefore, with ascending mu- tations. Gero Loaic Locauiry AND LrevreLs.—All the chief specimens (numbering 39 in Table VI above) listed, described, or figured by Falconer and Cautley as ‘Elephas planifrons’ are recorded as simply from the Siwalik Hills, since they did not distinguish be- tween ‘upper,’ ‘middle,’ and ‘lower’ Siwaliks. Pilgrim (1913) attributes this species only to the ‘Upper Siwaliks,’ Pinjor horizon, summit of the Pliocene; he writes (1913, p. 294): “There is absolutely no trace of Elephas either in the Middle Siwalik or in the Tatrot zone of the Upper Siwalik. It first appears as the species Hlephas planifrons some 2,000 feet above the base of the Tatrot zone.” DESCRIPTION OF SPECIES In the collection of 1922 made by Barnum Brown, twenty- seven or more specimens are now recorded as, or referred to, Archidiskodon planifrons; this collection (see map, Fig. 826) is recorded by Brown especially from near the towns of Kalka, Charnian, Siswan, Chandigarh, and Mirzapur, as shown in the complete list on page 955 below. None of the twenty-seven or more specimens of the Brown collection was actually found embedded in the rock; all of them were loosely embedded in sand, occurring in gullies or depressions. Consequently the exact geologic age and horizon levels are inde- terminable; yet they may all be safely classed as Upper Pliocene.! History.—Falconer and Cautley first published in the “Fauna Antiqua Sivalensis” the name Hlephas planifrons in the legend of Plate 11, figs. 5a, 5b, of date 1845. In 1846, p. 38, Falconer wrote: “The next serial modification in the disposition of the three dental substances, and in the consequent form of the teeth, is exhibited in fig. 5a of the same plate, which represents a section of the penulti- mate upper molar of another Indian fossil species which we have named Ff. planifrons.”’ As usual in his descriptions the locality is given simply Siwalik Hills, but we know from Pilgrim (1913, p. 294), as quoted above, that this phylum first appears as the species E. planifrons in the Pinjor horizon, Upper Pliocene,! some 2,000 feet above the base of the Tatrot horizon. It also occurs in Austria and Bessarabia. As early as 1863 Falconer remarked (p. 80): affinity, and that a very close one, of the Huropean FL. meridionalis is with the Miocene EH. (Loxod.) planifrons of India.”’ Faleoner thus led the way for our present knowledge, namely, that Archidis- kodon planifrons is directly ancestral to A. meridionalis. E. |Elephas] planifrons Yaleoner and Cautley, 1846 [1845]. “Fauna Antiqua Sivalensis,” 1846, p. 38 [1845, Pl. 11, figs. 5a, 5b]. Lecrorypr.—A second superior molar with nine ridges, the three anterior ridges being in use (Brit. Mus. M.3068). Co- tyPr.—Broken third molar of lower jaw, |.M;, with nine ridges remaining. Brit. Mus. M.2010. Horizon AND Locauity.— Siwalik Hills, India, Upper Siwaliks, Pinjor horizon, Upper Pliocene.! LecroryPre AND Coryrr FigurEs.—Falconer and Cautley, 1846 [1845, Pl. 11, figs. 5a, 5]. cor The nearest ‘Recent field studies in northern India by Dr. Hellmut de Terra and Pére Teilhard de Chardin have offered strong evidence to support the conclusion that the Tatrot and Pinjor horizons are equivalent and are entirely of Lower Pleistocene age. The Boulder Conglomerate ranges up to the Middle Pleistocene (see Chap. XXII below on the Geologic Succession of the Proboscidea for more detailed discussion). This note was prepared by Dr. E. H. Colbert, January, 1937.—Editor.] THE MAMMONTINA: ARCHIDISKODON AND METARCHIDISKODON Derscription.—(Faleoner and Cautley, 1846.1, p. 38): “The next serial modification in the disposition of the three dental substances, and in the consequent form of the teeth, is exhibited in fig. 5a of the same plate, which represents a section of the penultimate upper molar of another Indian fossil species which we have named F. planifrons. This tooth shows nine ridges, the three anterior of which alone have been in use, the two first being worn down to a single dise of ivory. The common nucleus of this substance is of less thickness than in the corresponding tooth of E. insignis, and the divisions which are continued upwards from it into the centre of the ridges are more elongated, with a narrower base, forming irregular-shaped wedges. The layer of enamel is diminished in thickness and is less uniform in outline, and the surface in contact with the cement shows a feathered or ragged edge, indicating superficial inequalities for the firmer cohesion of this latter substance. The enamel is reflected over the ridges of ivory, and down into the hollows zig-zag wise, exactly as in fig. 6a, the principal difference being that the ridges are narrower, with 951 between molars of the upper and lower jaws, and of different ages in the same species.”’ FALCONER’S NOTES OF 1868 ON TYPE AND REFERRED SPECI- MENS OF ELEPHAS PLANIFRONS Falconer, “Paleontological Memoirs,” Vol. I, 1868, pp. 423-442, Plates Il, VI, VII, XI, X11, and xtv of the “Fauna Antiqua Sivalensis.”’ Falconer’s observations on this species are to be found in his successive publications of 1845 (Plates), 1846 (Letterpress), 1857, 1863, 1865, and 1868. In the latter year (‘‘Paleontological Memoirs” of 1868) a summary of all his observations was pub- lished by Murchison, covering in detail thirty-nine or more speci- mens referred to Hlephas planifrons, all presumably collected from the Pinjor horizon, Upper Pliocene.! ELEPHAS PLANIFRONS: [LecroryPE (Brit. Mus. M.3068) AND CoryPr (Brit. Mus. M.2010)|.—Plate n, fig. 5a, M? with 9 ridges, cement filling valleys, enamel folds thick, length 8.7 in. = 221 mm.; fig. 5b, Ms, vertical section, with 9 ridges. LecrotyPE (R.M?) 4np Coryre (L.M3) or ARCHIDISKODON PLANIFRONS Fig. 825. Lectotype and cotype of Elephas planifrons Valconer and Cautley, 1846 [1845, Pl. u, figs. 5a, 56). From the Siwalik Hills, India, Upper Siwaliks, Pinjor horizon, Upper Pliocene.! One-third natural size. See ‘Description of the Plates in the Fauna Antiqua Sivalensis” (Malconer, 1867, p. 3; also Falconer [Murchison], 1868, Vol. I, p. 423): “Pig. 5a.—. .. Vertical section of penultimate upper molar [r.M?’|, with nine ridges, the three anterior of which alone have been in use, the two first being worn down to a single disc of ivory.” Brit. Mus. M.3068. Molar inverted to show natural position. “Fig. 5b.—‘{Cotype.] Vertical section of portion of last molar of lower jaw [l.Ms3], with nine ridges.” Brit. Mus. M.2010. Osborn (1924): The cotype M3 is a broken tooth which typically presents ten ridges, sometimes eleven. Observe in comparing the original figure (5a) with the new figure (Fig. 827) that Falconer omitted the seven ridge-plated r.M!. See also figure 828 after Lydekker. M’, ridge-plates pre-concaye, post-convex. a greater vertical height. The cement substance attains its maxi- mum of development in this species, completely filling up the wide interspaces of the ridges, over which it is continued in a thick mass. This tooth measures 8.7 inches in length.” “Hig. 5b, represents a corresponding section of a portion of the last molar of the lower jaw of the same species, comprising nine ridges. This tooth had been longer in use than that of the upper jaw, and all the ridges are more or less worn except the two last. It presents the same general characters exhibited by fig. 5a, in the elongated cuneiform ivory ridges, unequal enamel, and abundant cement, the differences being merely such as constantly hold See footnote on previous page.—Hditor. | Ms, ridge-plates pre-convex, post-concave. Ripce FormMuLa& IN Upper AND Lower GRINDING TEETH Upper Jaws.—Plate vi, figs. 4, 5, 6. Skull with premolar, ?Pm, Dp*, M!. Left Pm with 3 ridges and 2 half-ridges. Dp‘ with 6 ridges also a heel and front ridge, enamel thick. (Footnote, p. 427): Palate with M!, 7 ridges and 2 half-ridges; length 6.5 in. =165 mm., width 3.4 in.=87 mm., height at 4th ridge 3.5 in. =89 mm., fifth and sixth ridges with 6 conelets each. Plate x1, fig. 1, aged palate with r. and 1.M$, 10 ridges including posterior half-ridge, enamel very thick; fig. 4, M*, right side, remarkable fragment, 6% ridge-plates, length of fragment 5.4 in.=135 mm., width 2.5 in.=63 mm. Plate xu, figs. 1, la, Dp? with 4 ridges; 952 fig. 2, section of r.Dp*, with 6 main ridges, a back talon and front heel, total 6 and 2 half-ridges, length 3.8 in.=97 mm., width 2.3 in.=59 mm.; figs. 4, 4a, M' with 5 ridges and heel remaining, two gone, 7 probable total, length 5.2 in.=132 mm., width 2.8 in.=71 mm.; fig. 5, M' [same as fig. 4, this plate], M?, the latter with 8 ridges and front heel, length of M? 8 in.=204 mm., width 3 in.=77 mm.; fig. 5a, palate with r. and 1.M?, 8 distinct ridges and front and back heel, total of 8 ridges and 2-half-ridges, conelets few | gAMRITSAR we / pe Fig. 826. Favorable exposures, southwest of Simla, of the Archidiskodon planifrons life zone, Upper Siwaliks, India, chiefly Pinjor horizon, from which were collected by Barnum Brown in 1922 twenty-seven or more specimens referable to this species, especially from Kalka, Charnian, Siswan, Chandigarh, and Mirzapur, also transitions to A. meridionalis. Compare figures 820, and 029, PlixXxv. and large, enamel thick, length of 1.M? 7.5 in.=191 mm., width 3.2 in.=81 mm., fig. 6, M? with 8 main ridges. Plate xtv, fig. 8, r.M’, showing 8 or 9 ridges and a heel, enamel thick, length 10 in. =255 mm., width 3.5 in. =89 mm. Lower JAws.—Plate vim, figs. 2, 2a, perfect lower jaw, Ms; with a total of 9-11 ridges, enamel very thick, length of M; 8.8 in. = 224 mm., width 3.8 in.=97 mm. Plate x1, fig. 2, superb lower jaw, two mental foramina, r. and 1.M; with 13 ridges and front and back half-ridges, enamel very thick, length of r.M; 11.8 in. =300 mm., width 3 in.=77 mm.; fig. 3, superb left half jaw, only 7 remaining ridges in M;, enamel thick, cement abundant, ‘‘Proved to be EF. planifrons by the distance between the plates, the very low crown, thick enamel, and two mental foramina,” extreme length of jaw 24.2 in.=614 mm., length of M; 10 in.=255 mm., width of M,; 3.6 in.=91 mm.; fig. 5, r.M3, enormous tooth frag- ment, very thick enamel, low ridges, and mesial expansion, 9 ridges remaining, length of M; 10.5 in.=267 mm., width 4.2 in.=105 mm., height of 9th ridge 3.5 in.=89 mm.; fig. 6, left jaw with 1.M,, ridge-plates 6+-, Moe, ridge-plates 9 and small heel, 5 conelets OSBORN: THE PROBOSCIDEA on 4th ridge, length of M, 6 in.=153 mm., width 2.8 in.=71 mm., length of M2 8 in.=204 mm., width 2.8 in.=71 mm.; fig. 7, left lower jaw, Ms; very aged, length of molar 10.2 in.=259 mm., width 4 in.=102 mm., three mental foramina; fig. 8, fine lower jaw fragment, M2 [M;3, Osborn] complete, with 111 ridges, enamel very thick, plates wide apart, much cement, few conelets, three Type ; Brit. Mus. M3068 S LE > e, Right Second 4a LZ2e S41 Superior Molar New Lecroryrr Figure or ARCHIDISKODON PLANIFRONS, R.M? Fig. 827. New lectotype figure of Elephas planifrons Falconer (Brit. Mus. M.3068), 9+ ridge-plated second superior molar, of the right side, r.M?, with worn crown of a 27} ridge-plated first superior molar of the right side, r.M'. Redrawn by Miss G. M. Woodward for this Memoir; one-third natural size. Compare figure 825. Observe in comparing this figure with figure 854 of the same molar, that the ridge-plates agree in size with those of Piltdown, England (Tig. 853). mental foramina, length of Ms [M3, Osborn] 12.1 in.=308 mm., width 3.8 in.=97 mm.; fig. 9, fragment of 1.M3; much worn; fig. 10, lower jaw, Mz with 84 ridges, cement abundant, enamel thick, no crimping, and no mesial expansion, length of Ms 7 in.=178 mm., width 3.7 in.= 95 mm., 1.Mz; height of 5th unworn ridge 3.8 in.=97 mm. Plate xu, fig. 7, right lower jaw, with Dp, with 6 main ridges and small half-ridge (same formula as in Loxodonta africana); fig. 8, left lower jaw, with small vertically succeeding premolar (c), above it Dp;, also Dp, with 7 main ridges, double front heel and small half-ridge behind; figs. 10, 10a, first left true molar, 1.My, with 7 main ridges, a small ridge in front, no heel behind, length of M, 6.7 in.=171 mm., width 2.3 to 2.6 in. =59 to 66 mm.; figs. 12, 12a, ?r.M3, “apparently of a small sized individual,” length 10 in. =255 mm., width in front 3.5 in. =89 mm., height of crown at 7th plate 4 in. =102 mm., ten main plates and a front plate and heel; fig. 13, left lower jaw, 1.Ms; entire, with about 13 ridges and a heel, or possibly 14, length of Ms; 12.7 in. =323 mm., width 3.6 in. =91 mm., height at 10th ridge 4.5 in.=114 mm. Plate xrvy, fig. 9, last lower molar, right side, r.Ms, length 9.5 in.=241 mm., width 3.5 in.=89 mm., height at 6th ridge 3 in.=77 mm., shows 8 ridges and a heel. Plate xvut.a, figs. 1, la, an 1.M;, an enormous specimen; 8 plates, length 10.4 in. = 265 mm., width 4.1 in. =104 mm., height 3.2 in.=81 mm. THE MAMMONTINA:: ARCHIDISKODON AND METARCHIDISKODON 953 CotuectiveE Ripae Formuna (FALconrer, 1868).—(1) The permanent fourth premolar, P 4, persists, with 3 ridges. (2) From the specimens of Hlephas planifrons, as described by Falconer (39 of which appear in Table VI “Summary of Measurements” above), from the Upper Siwaliks, we deduce the collective ridge formula below; (3) the minimum numbers represent partly worn or partly developed teeth, while the maximum numbers represent fully worn and fully developed teeth; (4) half-ridges develop both in front Elephas planifrons.—Verticai and longitudinal section of the second upper true molar; from the Pliocene of the Siwalik Hills. 3. Fig. 828. Lectotype nine and a third ridged upper molar, r.M?, of Elephas [Archidiskodon| planifrons. Reproduced from a wood engraving by Lydekker (1886.2, p.102, fig. 24). Brit. Mus. M.3068. One-third natural size. Inverted to show natural position of molar. a, cement; b, enamel; c, dentine. We observe that the cement (a) completely fills the valleys. The enamel (b) is extremely thick, since the sides of the enamel ridge-plates diverge to- wards the apex and converge towards the base; the cement interspaces between the ridge-plates become constantly narrower as the crown wears down, while the dentine within the enamel becomes constantly broader, as shown in the figure opposite. and behind the main ridges; (5) the collective formula below indicates the half-ridges as well as the main ridges. Maximum and minimum collective ridge formula [of ascending mutations] of Hlephas planifrons: Dp 2* Dp 3 “Gy (P4***) Dp 4 ers M 1 ee M 2 Saco f 8 -9-Y--1 2-4 M3 8-9-16-1 0-}6-}6-1 2-4-1 4-46 (6) It is interesting to observe the half-ridges arising both in front and behind the main ridges, which seems to be a char- acteristic feature of this species, as compared with Hlephas hysudri- cus. (7) A primitive or simplified formula from Falconer of the minimum ridges in #. planifrons from the Upper Siwaliks of India would be: Dp 24 Dp 3 ¢& (P 47°) Dp 4 =M12M2$M3 2%. SPECIFIC CHARACTERS OF ARCHIDISKODON PLANIFRONS (LYDEKKER, 1886; OSBORN, 1924) The next review of Archidiskodon planifrons is that of Lydek- ker (1886.2, pp. 98-107) in which the cranium (Brit. Mus. M. 3060 —our Fig. 830) was selected as the type; in the present Memoir the specimen first described and figured by Falconer and Cautley is regarded as the type [lectotype] (see Fig. 825). From Lydekker’s descriptions the following characters may be summarized: (1) Both the third and fourth milk molars were vertically succeeded by premolars, thus the dental formula is: Dp?* Dpo-s, Ps P3-,, M3 My-3. This succession is an important character which must be looked for in the ancestors of this species; it does not occur in any other species of the Mlephantide thus far known; nor is it found in Stegodon bombifrons. (2) The correct collective Fig. 25. Elephas planifrons.—The hinder half of the third right upper true molar from the Pliocene of the Siwalik Hills. 2. The lower border of the figure is the inner border of the specimen. Fig. 829. Referred right M® of Hlephas [=Archidiskodon] planifrons. After Lydekker, 1886.2, p. 102, fig. 25. Brit. Mus. M.3070. Two-thirds natural size. Figured by Falconer and Cautley, 1846 [1845, Pl. x1, fig. 4], and made one of the genotypes of Leith-Adamsia Matsumoto, 1927, but regarded by the present author as a synonym of Archidiskodon planifrons. (See Fig. 847 below.) As compared with figure 828 this is a crown view of a much worn third superior molar in which the enamel loops in the middle of the crown are contiguous, the dental areas are expanded, the cement areas are contracted. The cement extends beyond the outer borders of the plates, a highly char- acteristic feature of Archidiskodon. ridge formula is that deduced from Falconer as his maximum- minimum or typical, as given above. The ridge formula of Lydekker (1886.2, p. 99) is incorrect. (3) Crowns of molars broad; cement in the interspaces frequently very great; enamel usually thick, frequently devoid of plication in the middle and near the root of the crown; plication near the summit of the crown relative- ly coarse. (4) Ridge-plates subellipsoidal, frequently with a mid- Fig. 8380. Skull of Hlephas [=Archidiskodon] planifrons from the Siwalik Hills, India, reproduced from Gaudry, 1878, p. 185, fig. 246, after Fal- coner 1846 [1845, Pl. x]. One-sixteenth natural size. This skull (Brit. Mus. M. 3060) was erroneously selected by Lydekker as the type of Hlephas planifrons, and is figured in his “Catalogue of the Fossil Mammalia in the British Museum (Natural History)” of 1886 (1886.2, woodcut fig. 23, p. 100). Compare figures 848 and 817 of the same cranium. Depéret and Mayet regard this cranium as that of a female, which, if true, partly accounts for its small size (see p. 962 of the present Memoir). 954 expansion, sometimes imperfectly lozenge shaped, in slightly worn teeth the middle portion of each ridge frequently forms an isolated disk. (5) Cranium characterized by flatness of the fronto- parietal region, by a small incision of the temporal fosse on the frontals, by a comparatively slight elevation of the vertex (hypsi- OSBORN: THE PROBOSCIDEA cephaly), by smallness of the nasal aperture, by wide divergence of the incisive alveoli, as in A. meridionalis; vertex of skull flattened almost at right angles to occiput, oeciput pitching forward 55° when the grinding teeth are horizontal; tusks very stout and incurved. REVISION OF ARCHIDISKODON PLANIFRONS BY OSBORN (1927-1928), BASED ON THE SIWALIK COLLECTION MADE BY BARNUM BROWN IN 1922 The twenty-seven specimens referable to Archidiskodon planifrons and its ascending mutations were found in the ‘Upper Siwaliks,’ below the Boulder Conglomerate formation, and recorded as follows: TABLE VII. MEASUREMENTS OF TWENTY-SEVEN SPECIMENS COLLECTED BY BROWN AND REFERRED BY OSBORN TO ARCHIDISKODON PLANIFRONS, ASCENDING TO A, MERIDIONALIS All recorded by Barnum Brown from near Kalka, Charnian, Siswan, Chandigarh and Mirzapur (see Figs. 820, 826) SUPERIOR Figure Amer. Molar Length Breadth Index Height Ridge-plates Ridge-plates Mus, mim. mm. est. mm. act. or est. in 10 em. Third Molars 19882 r.M*® 254 96 38 9th =102 12+ 5%-6 834 19950 1.M*® 219 100 46 7th = 88e 9-4-4 = 12 4+ 19880 1.M* 201 88 44 9th = 95 ils 5% First or Second Molars a 19821 1.M2 217 94 43 7th = 66e j#$8-4%%=10 3K-4 ace 199555 eri iz 76 44 6th = 70 ¥-7-4= 9 4-4}, (19955 1. MY 162 80 49 5th = 65 ¥-6-4= 8+ 4-41 831 19881 1.M! 165 90 54 Bol = Gy ¥-6 = 7+ 3-4 19916 ?r.M! 138 80 58 5th =107 Y-6 3-4 INFERIOR 842) Third Molars =| 19951 r.M; 313 101 32 thy 124 11 =12 3K-4' 841 19861 1.M; 250 79 32 9th = 88e 11% =12 5 846 19879 1.M; 190+ 105+ 55 8th =135 10+ 5-54 844 19819 1.M; 292+ 95 32 7th = 88e 10% =11+ 4 855 19798 r.M; 203 80 39 fon, = 5S} 11-- 64% 19870 1.Ms3 2d 85 843 19968 1.M; 213e 95 45 7th = 66e 9+ 5 840 19965 r.M; _ 180 (inc.) 86 48 7th = 55e 9 4-4", 839 19864 r.M; 194+ 78 40 3d = 45 ¥-8-43= 10+ 5 835 19952 1.M; 238e 109 47 6th =114 8ke= 9+ 3h-4 Second Molars 836 19778 r.Mes 140+ (ine.) 90+ 64 yo) 74 ¥-5+= 6+ 19871 1.M. 167+ 87 52 8th = 99 8+ 19870 1.Me 10th = 82 First Molars 838 19862 ?1.M, 160e 80 50e 38rd = 45 ¥-8 = 9+ 5 19917 r.M, 190+ 78 41 5th = 91+ +7% 5 | 19961 ?1.M, 189 90 48 7th = 73 7 4 19871 1.M, 127 85 67 4-6-4 836 19778 r.M, 103 70 68 3+ Fourth Deciduous 837 19873 r.Dp, 121 63 52 7th = 44 ¥-T*%= 9 7 THE MAMMONTINA: ARCHIDISKODON AND METARCHIDISKODON 955 As remarked above, the large American Museum collection of Archidiskodon planifrons, including examples of the entire den- tition, but lacking a skull, renders it highly probable that the Falconer and Cautley collection of A. planifrons, figured and listed in detail, as above described, came from the same geologic Near Kalka 9 miles west Amer. Mus. 19778 6 miles west Amer. Mus. 19798 Near Charnian 2 miles south Amer. Mus. 19819 Near Siswan Amer. Mus. 19821 Amer. Mus. 19870 Amer. Mus. 19879 Amer. Mus. 19880 Amer. Mus. 19881 Amer. Mus. 19882 3 miles north Amer. Mus. 19871 3 miles north Amer. Mus. 19873 3 miles north Amer. Mus. 19968 Amer. Mus. 19965 Amer. Mus. 19862 Near Chandigarh 3 miles west Amer. Mus. 19861 3 miles west Amer. Mus. 19916 3 miles west Amer. Mus. 19950 1 mile west Amer. Mus. 19951 1 mile west Amer. Mus. 19952 3 miles west Amer. Mus. 19955 1 mile west Amer. Mus. 19961 3 miles west Amer. Mus. 19917 Probably near Chandi- garh, record incomplete Amer. Mus. 19967 Near Mirzapur 3 miles north- Amer. Mus. 19864 east localities of the Pinjor horizon level, namely, from deposits around Kalka (see Figs. 826, 820). The following 29 specimens, belonging to 24 individuals, are given with measurements above (Table VII); they are also illustrated in great detail in the accompanying figures (Figs. 831-844). Fragment of right jaw with M, and Mz zn situ (Fig. 836 for M,). Fragment of jaw with r.Ms; in situ (Fig. 855 FOR r.Ms). Fragment of left jaw with 1.M, in sitw (Fig. 844). Fragment of maxilla with 1.M? in situ (Fig. 833). Fragment of jaw with 1.M.» and 1.M3 in situ, Me incomplete, M3; partly erupted (not figured in present Memoir). Fragment of jaw with 1.M3 zn situ (Fig. 846). Left M* (not figured in present Memoir). Left M! (Fig. 831). Fragment of maxilla with r.M* (not figured in present Memoir). Fragment of jaw with 1.M, and My zn stu (not figured in present Memoir). Fragment of jaw with r.Dp, in situ (Fig. 837). Fragment of jaw with 1.M3; in situ (Fig. 843). Fragment of jaw with r.M; 7n sztu (Fig. 840). Fragment of jaw with 1.Dp, and 1.M, (Fig. 838). Fragment of jaw with 1.M3; 7n situ (Fig. 841). ?Right M! (not figured in present Memoir). Fragment of maxilla with 1.M? in situ (Fig. 834). Right M; (Fig. 842). Left Mz; (Fig. 835). Right M! and left M? (Fig. 832). ?Left My, (not figured in present Memoir). Right M, (not figured in present Memoir). Lower jaw, left ramus. Fragment of jaw with r.M; zn situ (Fig. 839). SUMMARY OF DENTAL MEASUREMENTS OF THE AMERICAN MUSEUM COLLECTION MADE BY BARNUM BROWN IN 1922 These measurements accord in general with those of the Falconer and Cautley Collection in Table VI above and exhibit variations in size, length, breadth, and proportions, partly attrib- utable to male or female sex, partly to progressive ascending muta- tions. The increase in number of the ridge-plates and half ridge- plates is probably attributable to progressive evolution or to ascending mutations ranging into higher geologic levels. In general, the smaller animals, with fewer ridge-plates, probably occurred in lower geologic levels than the larger animals, with more numerous ridge-plates and other progressive characters. TxHirp Superior Mouar, M$ Minimum Maximum Length 201 254 Breadth 88 109 Breadth-length index 38 47 Height 66e 102 Ridge-plates 9+ 12 SECOND SupERIOR Morar, M? (Only specimen available in Amer. Mus. Coll., No.19821) Length 217 Breadth 94 Breadth-length index 43 Height 66e Ridge-plates 10 First Superior Mouar, M! Minimum Maximum Length 138 172 Breadth 76 90 Breadth-length index 44 58 Height 65 107 Ridge-plates a 9 Tuirp Inrertor Morar,M; Length 190+ 313 Breadth 7 i 105 Breadth-length index 32 55 Height 45 135 Ridge-plates 9+ 12 Sreconp Inrertor Morar, Me Length 140+ 167+ Breadth 87 90+ Breadth-length index 52 64 Height 72 99 Ridge-plates 8+ 8+ First Inrertor Morar M, Length 127 190+ Breadth 85 78 Breadth-length index 67 41 Height ies 91-- Ridge-plates 6-14 +7% 1 ARCHIDISKODON PLANIFRONS Re. Amer (Jus. /982/ ' ' ! ! 1 i] 1 ARCHIDISKODON PLANIFRONS Ref. Amer: (us. (9955 Pig. 831 Fig. 832 Fig. 833 ARCHIDISKODON PLANIFRONS Rg. AMERICAN MusrumM COLLECTION Upper PLiocENE ARCHIDISKODON PLANI- FRONS OF THE Pryzor Horizon (SEE Fia. 826) Figs. 831-835. Archidiskodon planifrons, referred superior and inferior grinding teeth of the Barnum Brown Siwalik Collection, listed with measurements in Table VII. All figures drawn to the same one-fourth scale. Cement (dotted), dentine (horizontal lining). Com- pare Falconer’s measurements in Table VI. Figures: (831) First left superior molar, 1.M!. (832) First left and right superior molars, 1.M?, r.M?. (833) Second left superior molar, 1.M2; a primitive stage as shown in midsection (Fig. 855); this M® agrees closely with that of Piltdown, Sussex (Fig. 853). (834) Third left superior molar, 1.M, 12 plated. (835) Third left inferior molar, 1.Ms3. ' ARCHIDISKODON PLANIFRONS Ref. { 1 i L.M3Z Amer. \hes. 79952 U Pee AU \0 = = = = = == = - N Tig. 835 956 Yr ARCHIDISKODON PLANIFRONS [iJ. ' A. PLANIFRONS mer. (us. 19778 ' A./7. (9862 Fig. 837 ARCHIDISKODON PLANIFRONS Ref R.M3 ARCHIDISKODON PLANIFRONS Ref. Amer. us. 19965" ' | 1 1 i Amer: (Mus. 1/9864. % Nav. seize Fig. 839 Fig. 840 AMERICAN Museum. Upper PLIOCENE ARCHIDISKODON PLANIFRONS OF THE PINJOR HORIZON (SEE Fra. 826) Figs. 836-840. Inferior grinders of Archidiskodon planifrons (Figs. 836-840), in the Barnum Brown Siwalik Collection, fully listed with measurements in Table VII. All figures one-fourth natural size, excepting 837 which is one-third natural size. Cement (dotted), dentine (horizontal lining). Compare Ialconer’s measurements, Table VI. Figures: (836) Portion of first and second right molars, r.Mj, r.My. (837) Fourth deciduous premolar, Dps, with 9 ridge-plates. (838) Worn crown of left fourth premolar, 1.Dp4, and first molar, 1.M;. (889) Portion of lower jaw with third right molar, r.M3. (840) Portion of lower jaw with third right molar, r.Ms3. 957 ARCHIDISKODON PLANIFRONS Re. f Am er. /4us.7986/ ? Amer. Mus. 19951 ARCHIDISKODON PLANIFRONS Ref. Wig. 842 | Amer (Tus. 198/9 | ARCHIDISKODON PLANIFRONS Ref. ' i 4 Nat- suze 1 ARCHIDISKODON PLANIFRONS Ref | 1 ' Amer. Mus. 19968 Fig. 843 Fig. 844 AMERICAN MUSEUM ARCHIDISKODON PLANIFRONS OF THE PinJoR Horizon (sEE Fa. 826). Inrertor Mouars Figs. 841-844. Archidiskodon planifrons referred inferior grinding teeth and jaws in the Siwalik Collection of the American Museum made by Barnum Brown, as fully listed with measurements in Table VII. All figures one-fourth natural size. Cement (dotted), dentine (horizontal lining). Compare Falconer’s measurements, Table VI. Figures: (841) 12 plated left third molar, 1.M3. (842) 12 plated right third molar, r.Mg, of a slightly older individual; a progressive stage as shown in midsection (Tig. 855B). (843) 94 plated left third molar, 1.M3, imperfect anteriorly. (844) 11 plated left third molar, 1.Ms. 958 THE MAMMONTINA:: ARCHIDISKODON AND METARCHIDISKODON ae ae —— 2 nee OEE Amer. (us. /9951 ARCHIDISKODON PLANIFRONS Rg. Upper PLIOCENE ARCHIDISKODON PLANIFRONS OF THE Pinson Horizon (SEE Fas. 826, 820) Fig. 845. Archidiskodon planifrons ref., third right inferior molar, r.M3 (Amer. Mus. 19951), in the Barnum Brown Siwalik Collection, one-third natural size. Compared with the collective figures above, this specimen is very close to the typical ridge formula of Archidiskodon planifrons, namely, M 3714 ridge-plates. Length 313 mm., width 101 mm., height of seventh ridge-plate 124 mm. (see Table VII, p. 954). ASTAIIEBIS 4s Nat- Szze Upper PLIOCENE ARCHIDISKODON PLANJFRONS OF THE PrnJor Horizon (sEE Fia. 826) Archidiskodon planifrons ref., a third inferior molar of the left side, 1.M3 (Amer. Mus. 19879) in the Barnum Brown Siwalik Collection, listed with measurements in Table VII; portions of ten ridge-plates are preserved, as displayed in the transverse vertical section of the same tooth (left). Fig. 846 (right). All figures one-third natural size. 959 LEITH-ADAMSIA SIWALIKIENSIS Figure 847 LEITH-ADAMSIA SIWALIKIENSIS! Marsumoro, 1927.—‘On Leith-Adamsia siwalikiensis, a New Generic and Specific Name of Archetypal Elephants.” Japanese Journ. Geol. and Geog., V, No. 4, Art. 12, 1 page. Typr.—Two superior molars, both of the right side, r.M%, in the collections of the British Museum (Brit. Mus. M.3070 and 36695). From India. TYPE Ficurr.—Falconer, “Fauna Antiqua Sivalensis,”’ Pls. x1, fig. 4, and xv, fig. 8. Type Description.—(Matsumoto, 1927.2): “Tn the course of a study of fossil Elephants, the writer has come to be faced with the serious need for a proper name for a certain type of archetypal Elephants of India. He means the small and narrow-molared form recorded under the name of Elephas plani- frons. This form appears, in all likelihood, to stand at the starting point of the entire phylum of the Loxodontine Elephants. As it would appear to occupy too important a position among the Hlephantide to be left unnamed, the writer proposes here to call Compare Falconer’s measurements Table VI, p. 949. Regarded by Professor Osborn as a synonym of Elephas [Archidiskodon] planifrons.—Kditor.] 960 it by a generic name as follows, in honour of the late Professor Doctor Leith Adams, the eminent palzeontologist and specialist on ” fossil Elephants. “Teith-Adamsia, gen. nov. A genus of archetypal Elephants. Cheek-teeth subhypsodont, narrow-crowned, with a low ridge- formula; loxodont sinus present, and of an obtuse type; disks of well-worn ridges may be more or less lozenge-shaped.” REFERRED ARCHIDISKODON PLANIFRONS Fig. 847. Type molars of Leith-Adamsia siwalikiensis Matsumoto, 1927. After Falconer and Cautley, 1846 [1847, Pl. x1, fig. 4, and Pl. xrv, fig. 8], one-third natural size, reduced to one-fourth natural size. (Pl. x1, fig. 4) Brit. Mus. M.3070. An r.M%, posterior half of crown, exhibiting 64 ridge-plates (same molar as in Fig. 829, above). (PI. xrv, fig. 8) Brit. Mus. 36695. An r.M®%, with 6} posterior ridge-plates preserved of a probable total of 10 ridge-plates. CRANIAL CHARACTERS OF ARCHIDISKODON PLANIFRONS The single cranium known in the British Museum collection (Brit. Mus. M.3060—see Figs. 830, 848, and 817) is extremely primitive, somewhat resembling that of certain species of Stegodon, for example, Stegodon pinjorensis type (Amer. Mus. 19772—see Figs. 711, 765, and 817C), except that Archidiskodon planifrons is much less elevated (acrocephalic) than S. pinjorensis, as clearly shown in the comparative profiles (Fig. 817). This is in adaptation to the relatively abbreviate 10+ ridge-plated crown of M® in A. planifrons, as compared with the extremely elongate 15 ridge- Also compare the flattened planifrons with the abbreviated and extremely plated crown of M* in S. pinjorensis. forehead of A. elevated forehead of S. pinjorensis. This flat-faced condition, to which Falconer assigned the specific name planifrons, develops into the concave-forehead OSBORN: THE PROBOSCIDEA profile of A. meridionalis (Fig. 817). Meanwhile the cranium heightens (hypsicephaly) and deepens (bathycephaly), in adapta- tion to the enlarging and deepening of the third superior molars, also, as fully explained in Chapter XV, p. 915, on the mechanies of the proboscidean cranium, the heightening of the occipital crest (acrocephaly) is in adaptation to the elongating tusks and the strengthening of the cervical ligaments and muscles which sway the great tusks and proboscis. Nothing is known of the limb skeleton of A. planifrons, but it is inferred that the animal was greatly inferior in shoulder height, in length, and in proboscis development to its giant suecessors A. meridionalis and A. impe- rator. The completely preserved tusks in members of this species discovered in southern France (Fig. 850) lack the strong outward curvature and meridionalis (lyrodon), see figure 864, reaching a supreme stage in A. imperator. incurvature characteristic of > M 235 M3ita 9-11 1Li-14° This formula of 1877-1881, together with Leith Adams’ sum- mary of the points distinguishing the molars of Elephas meridionalis from those of the narrow-toothed Elephas antiquus (p. 232), are to be found in his important Memoir (p. 208). He assigns the follow- ing characters to Hlephas meridionalis. Cuaracters.—(Cf. op. cit., p. 30): M?® in EH. meridionalis, ridges nearly as broad as they are long, thick plates, grosser masses of intervening cement, macherides uncrimped, X 10 X. (Cf. p. 44): M® massive, enamel and plates very thick, macherides scarcely plaited, great breadth and low ridge formula, which rarely if ever exceeds that of M? of #. antiquus. (Cf. p. 48): In M?* the highest ridge formula of £. meridionalis equals lowest of E. anti- quus; so that in number of ridges we find EL. primigenius, E. antiquus, E. meridionalis, and E. namadicus meeting at their extremes. Leith Adams (cf. pp. 129 to 144) compares the cranium of E. meridionalis in great detail with that of EL. planifrons, E. hysudricus, E. bombifrons, E. africanus, ete., concluding that the skull and dentition of #. planifrons make the nearest approach to E. meridionalis (see pp. 186, 208-210, 239, 244). Error.—Originally Leith Adams also made the error of including the low ridge-plate formula (M 3 +2) of E. planifrons with the high ridge-plate formula (M 3 }4+4) of EZ. meridionalis, in describing the range of evolution and variation in the ridge formula of the British specimens of #. meridionalis. Werrnorer, 1890, LypeKknr, 1886 CoutuectivE Ripce Formuta.—The collective ridge-plate formula [of A. planifrons and A. meridionalis| may also be cited from Weithofer (1890, p. 172): “Als Gesammtformel fiir die Ziihne des El. meridionalis, soweit sie hauptsichlich aus dem Material des Museums zu Florenz resultirt, ergiibe sich demnach’’: WeirHorer, 1890, E. meripionatis: Dp 253°; Dp 3 Dp 475 Mle M 222° M 3 i233 9-11 11-12° 5-6 3 5-6 PROBOSCIDEA Weithofer in his collective formula, like Faleoner (1868), un- doubtedly erred by including specimens having the typical Archi- diskodon planifrons ridge formula with those of A. meridionalis; thus this collective formula ranges from the formule of Archidis- kodon planifrons, e.g., M 3 +4, to A. meridionalis, e.g., M 3 +3. LyprkKker, 1886.—Lydekker in his ‘‘Catalogue of the Fossil Mammalia in the British Museum (Natural History),” 1886.2, p. 107, adopts the Leith Adams formula of 1877-1881 as follows: (1) The molars of #. meridionalis so closely resemble those of F. planifrons, that if they both occurred in the same area it is more than doubtful if they could be specifically distinguished; (2) both frequently exhibit partial denudation of the enamel ridges; (3) there is variation in the thickness of the enamel and in the breadth of the ridges, in some molars the enamel being relatively thin and considerably plicated; (4) the cranium is characterized by large, slightly curved tusks and widely diverging alveoli; (5) in general contour it is intermediate between the cranium of FH. planifrons and #. hysudricus, although nearer the latter; (6) it agrees with the cranium of H. planifrons in the relative distance between the nasals and the vertex, but has the vertex more vaulted, the frontal profile concave, the temporal fosse intruding largely on the frontal aspect; (7) the species attained an enormous size, the height of some individuals being estimated at upwards of fifteen feet. ARCHIDISKODON MERIDIONALIS REF. Tig. 863. Referred Elephas meridionalis of Chagny (Céte-d’Or), one- fourth natural size, after Gaudry, 1878, p. 178, fig. 237. This fourteen ridged third superior molar, M®, of Chagny, is to be compared with the thirteen ridged molar stage of Elephas meridionalis of the Val d’Arno, i.e., M3 ygrr5) 88 deduced by Falconer (see Falconer, 1868, Vol. II, p. 118). Observe superior ridge-plates pre-concave, post-convex (cf. Fig. 827, A. planifrons). CRANIAL AND DrentaL CHaracters.—Weithofer (1890, pp. 136, 137) refigures Nesti’s type skull (Fig. 861 of the present Memoir) showing the right side (Taf. 1, fig. 1): “Hlephas meri- Nesti; Cranium ©; oberes Arnothal; von rechts.” Faleoner also refigures this skull (1846 [1847, figs. xix of Pls. XL and xuiv]), as well as Depéret and Mayet (1923, p. 128, fig. 16). The three skulls in the Florence Museum exhibit the follow- ing characters: (1) Pointed nasals, (2) concave frontals, (8) high occipital erest flattened anteroposteriorly, (4) extreme brachy- cephalie cranium shortened anteroposteriorly, (5) broad narial dionalis THE MAMMONTINA: openings, (6) parallel sides of the premaxillary sockets of the tusks, as contrasted with the broadly flaring sides of the sockets similar to those in the contemporary Hesperoloxodon antiquus. All these characters point to the cranial relationship of the Italian Archidiskodon meridionalis and its ancestor A. planifrons. WertTHorer’s TypE SKULL oF ELEPHAS LYRODON = ARCHIDISKODON MERIDIONALIS FEMALE Fig. 864. Types of two individuals of Hlephas lyrodon Weithofer, Florence Museum, upper Val d’Arno deposits (Weithofer, 1889, pp. 79 and 80): ‘Hier sei beziiglich der zum erstenmal genannten Species Hlephas lyrodon nov. sp. nur bemerkt, dass sie auf zwei vollstindige Schidel sammt Stoss- zihnen, sowie Schidelfragmenten mit Stosszihnen und mehreren Unterkiefern und isolirten Stoss- und Backenzihnen des Museums von Florenz basirt ist.’ After Weithofer, 1890, Taf. 11, fig. 2 (right); rv, fig. 2, Schadel a (left); v, fig. 1, Schidel a (middle). All one-twentieth natural size. Compare figure 865 (11, 11a). MS. ELepHas GIGANTEUS AYMARD (IN Fatconer, 1857, p. 321).—(Lucien Mayet, letter, December 11, 1922): ‘‘Hlephas giganteus Aymard n’est qu’une désignation portée sur des étiquettes de sa collection par ce paléontologiste. I] y aurait lieu de retrouver les piéces et de les déterminer; ce serait peut-étre 2. meridional- is(?) ou EL. trogontherii(?) ou E. antiquus(?). Ils’agit trés probable- ment lA—comme pour beaucoup des ossements recueillis par Aymard dans le Pliocéne supérieur et le Pleistocéne du bassin du Puy—de désignation inscrite sur des étiquettes, sans que les piéces correspondantes aient jamais été déterminées ou aient fait Vobjet d’une révision ultérieure. C’est un nom—celui d’Hlephas giganteus—a faire disparaitre purement et simplement.” ARCHIDISKODON AND METARCHIDISKODON 975 SKULLS AND SKELETONS OF ARCHIDISKODON MERIDIONALIS In figure 865 are displayed outlines of the lectotype (Nos. 7,18, 14), cotype (Nos. 8, 9,11, 11a, 15), and other skulls of Archi- diskodon meridionalis, male and female, of which we have been able to find figures in the literature. They include the following: 1) Male skull figured in error by Falconer and Cautley in 1847 as Hlephas antiquus: front view, Pl. xu, fig. x1x; side view, Pl. xurv, fig. xrx. Cranium C of Nesti [lectotype of Elephas meridionalis Nesti, 1825, Tav. 1, figs. 1, 2]. 2) Cranium of EHlephas meridionalis, side view, after Weit- hofer. Cranium C of Nesti. 3) Cranium A of Nesti [cotype], palatal view after Nesti, 1825, Tav. 1, fig. 3; front and side views after Weithofer, 1890, Taf. 1, figs. 1, 2. 4) Front and side views of female cranium, type of Hlephas lyrodon Weithofer, 1890, Taf. v, fig. 1, Taf. rv, fig. 2. CHARACTERS (OsBoRN, 1924).—These crania agree in the following points: (1) Extremely broad and vertically shallow anterior narial openings, a feature shared by Archidiskodon im- perator (Fig. 896); (2) extreme cranial abbreviation (hyper- brachycephaly) and depth (bathycephaly) resulting in hypsicephaly and acrocephaly; (3) concave forehead or frontoparietal profile; (4) parieto-occipital crest rising high in profile (ef. Weithofer, 1890, Taf. 1, fig. 2, 2. meridionalis cotype, with EH. 7mperator ref., Los Angeles, Calif.); (5) through hypsicephaly, orbits and occipi- tal condyles approximated, vertical diameters greatly exceeding anteroposterior diameters; there seems to be little doubt of the phylogenetic kinship of the Archidiskodon meridionalis with the A. umperator crania; (6) cranial profiles of A. meridionalis and A. imperator analogous to the hypsicephalie and acrocephalie profile of Mammonteus primigenius; (7) in comparison with the more primitive, more platycephalic A. planifrons, with smaller narial openings (Fig. 848), the known crania of A. meridionalis are much larger and more progressive than the known crania of A. planifrons. CRANIA IN THE FLORENCE MUSEUM (W. D. Marruew, September, 1920).—Here are found be- sides the type skull of Hlephas meridionalis Nesti [Cranium C] several more or less incomplete skulls, many jaws and teeth, in- cluding those from the Upper Pliocene near Magello. . ARCHIDISKODON IMPERATOR IN THE U. 8S. NATIONAL MUSEUM FLORIDA COLLECTION Specimens collected for the U.S. National Museum by James W. Gidley in the locality of Melbourne, Florida, and presumably all from the “No. 2”’ bed of Sellards, are as follows: Nat. Mus. 11805. A third inferior molar, M3, with 15-18 ridge-plates; laminar frequency 5 ridge-plates in 10 cm. Nat. Mus. 11814. A much worn third inferior molar, M3; laminar frequency 5 ridge-plates in 10 em. Total ridge-plates 12+ (tooth worn to base). Nat. Mus. 11620. The Venice mammoth from near Mel- bourne, Florida. Aged individual, with third superi- or and inferior molars much worn, also jaw, por- tions of skull, right tusk, and hind foot. Third superior molar, 1.M%, total ridge-plates estimated at 184-19, 3 to 4 anterior worn off, length 200 mm., breadth 106 mm., 5}5 ridge-plates in 10 em.; maxi- mum breadth 106 mm. as compared with 125 mm. in Leidy’s Nebraska type (Fig. 884). Third inferior molar, r.M;, estimated ridge-plates 16-17, 3 to 4 Amer. Mus. 2568 United States Amer. Mus. 11871 See figures 884, 886, 887. Mexico See figures 886 Al, 888. Amer. Mus. 14476 Texas County, Texas. Amer. Mus. 14558 Imperfect lower jaw, right and left third inferior molars. Kansas From Ness County, Kansas. Amer. Mus. 10598 Texas Amer. Mus. Cope Coll. 14475 Texas Tule Canon, Briscoe County, Texas. Lower jaw with partly worn M; (see Figs. 892, 898), also right forelimb (see Figs. 906, 907, 908, 912). 1005 anterior worn off, length 228 mm., breadth 97 mm. Gidley (letter, Oct. 27, 1928) adds that the specimen from Venice has an extremely deep jaw but otherwise is like A. impera- tor; that both A. [=Parelephas] columbi and A. imperator occur at Melbourne and at Vero. “I think, therefore, there can be no doubt that these two species were contemporaneous in Florida, and perhaps also in other localities of the southern and south- western United States.” Osborn, 1922-1928: This is a very important case of the ap- parent association of Archidiskodon imperator ref. and of Parelephas columbi ref. in the same geologic locality, if not actually from the same level. ARCHIDISKODON IMPERATOR IN THE AMERICAN MUSEUM COLLECTION See figures 885, 886, 888, 889, 891, 892, 896-898, 900, 906-908, 912, 920 of the present Memoir. The type cast, the neotype, and the finest referred specimens referable to Archidiskodon in the American Museum collection are listed herewith (bottom of this page). These superb materials are illustrated in various figures of the present Memoir and add greatly to our knowledge of this imperial species of southern mam- moth. They display clearly the specific characters enumerated above. COMPARISON OF THE JAW AND DENTITION WITH OTHER PROBOSCIDEANS A great deal of study has been given to comparison of the jaws of various types of the Proboscidea, especially in correlation with: (1) Abbreviation (brachycephaly) and elevation (hypsicephaly) of the cranium, also with (2) insertion of the third inferior molar tooth which naturally is the chief functional organ of the body. It is seen in the accompanying diagram (Figs. 892, 893) that the progressive jaws of Archidiskodon imperator (Fig. 892 A, B) are profoundly different from the relatively elongate jaws represented in figures 898, A, B, C (Archidiskodon hayi, Parelephas washing- tonii, Loxodonta africana). The jaws of Archidiskodon imperator (Fig. 892 A, B) are also readily distinguished both from those of Parelephas jeffersonii (Fig. 892 C, D) and of Elephas indicus (Fig. 893, D, E). It is observed that in the Texas jaw (Amer. Mus. 10598—see Fig. 892 B) Ms; exhibits sixteen ridge-plates, while in the Kansas jaw (Amer. Mus. 14558—see Fig. 892A) M; exhibits five accessory ridge-plates, namely, 16, 17, 18, 19, 20; this difference cannot be due Type r.M3 of Elephas imperator, cast (Amer. Mus. 2568) after original in U.S. National Museum (Nat. Mus. 185). Neotype (Osborn) of Elephas imperator, right third superior molar, r.M%, from Guadalajara, Jalisco, Mexico. Lower portion of cranium with tusks, here represented in figures 889B, 896, 897, 906. From Victoria, Victoria Crown view (Fig. 889A); side view (Figs. 892A, A1). Total of 19-20 ridge-plates, of which the 19th and 20th are extremely short and rudimentary (see Fig. 892). From near Referred young male skull of Archidiskodon imperator, with tusks and |.M?, discovered near Dallas, Texas, and described by Cope in 1889 (1889.2) as “Elephas primigenius columbi Fale.” (see Fig. 891), . 7 ae: Elephas jeffersonii Amer. Mus. 13225 12 / D 42 D Wyn \ oF i EZ BA AMM: % ZN Rat / LZ 3456789 uM \ K \ 18 et ye ~ Swen” -— U \ Elephas jeffersonii Osb Amer. Mus. 9950 Type Ea 17 D> 12 15 218 \Z Yi pa 2, gone es VAAL D Sas COLCLIU) 2,18 P19 Cc us 19 one x AG { N Ty ‘ 4 SK SA ae ee E. imperator Amer. Mus. 10598 Ref. se E. imperator Amer. Mus, 14558 Ref. All 1/8 nat. size E. indicus E, indicus Amer. Mus. Dept. Mam. 54261 Loxodonta africana \ Amer. Mus. Dept. Mam. 39083 E. washingtonii Amer. Mus. 8681A Type at 456 wa) Fs OUTED Nr s/o a SROBQRETELALEie A ae ale E. hayi Neb. Mus, 23-6-14 Type JAws OF ARCHIDISKODON IMPERATOR IN COMPARISON WITH PARELEPHAS JEFFERSONI, Vey WASHINGTONII, LoxopoNnTA AFRICANA, AND EL»PHAS INDICUS Fig. 892. Internal aspect of the lower jaws and Msg of Elephas [ = Parele- phas) jeffersoni (paratype and type) and EHlephas { = Archidiskodon| imperator, also sections D1, Cl, Bl, Al, cut at point indicated by the dotted line S. One-eighth natural size. D, Elephas | =Parelephas) jeffersonii paratype (Amer. Mus. 13225); M3 in situ with fifteen worn plates and eight unworn plates, a total of twenty-three plates. [See Fig. 960 for final count of twenty-four ridge-plates in this para- type specimen. | C, Type of Elephas [= Parelephas| jeffersonit (Amer. Mus. 9950), a very aged individual, Mg with seventeen worn plates and three additional unworn plates im situ, a total of twenty plates. This, as explained elsewhere, is a very aged individual and it is probable that one or two of the extreme an- terior plates have been worn off. B, Jaw of Elephas [= Archidiskodon| imperator ref. (Amer. Mus. 10598), Mz with fourteen worn plates and a total of sixteen plates in situ; a very robust individual. From Tule Cafon, Texas. A, Elephas |=Archidiskodon| imperator ref. (Amer. Mus. 14558), from Ness County, Kansas, M3 with fourteen to fifteen (est.) worn plates and five additional, a total of twenty plates. See also figure 889A, crown view of same tooth. Fig. 893. Internal views of jaws of Elephas indicus, Loxodonta africana, Elephas {= Parelephas| washingtonii, and HE. [=Archidiskodon| hayi, for com- parison with figure 892. One-eighth natural size. E, Elephas indicus, a fully developed Mg of the right side with twenty- seven plates. Drawn after de Blainville, 1839-1864, Pl. rx, fig. 6° (reversed). D, Elephas indicus, right jaw (Amer. Mus. Dept. Mam. 54261); Mg with nineteen plates fully formed and five imperfect plates in the jaw, a total of twenty-four plates. C, Loxodonta africana jaw (Amer. Mus. Dept. Mam. 39088, Akeley Coll.); ?Mp with twelve plates developed. B, Type jaw of Elephas [= Parelephas| washingtonii Osborn (Amer. Mus. 8681A), with twenty-one plates developed in M3. Compare figure 969, type of Parelephas progressus. A, Type jaw of Hlephas [=Archidiskodon| hayi Barbour (Neb. Mus. 23-6-14). 1006 All] B nat size THE MAMMONTINA: ARCHIDISKODON AND METARCHIDISKODON to age or to sex; it is more probably due to the progressive addi- tion of ridge-plates in the Kansas specimen. Majestic as are these imperial mammoths from Texas the largest pair of tusks known is from Mexico and is described below, with change of name, by Osborn (Fig. 894). APPOSITION OF THE GRINDERS.—In the American Museum collection there is no case of skull and jaws being found together, so that we cannot positively determine the apposition of the grinders; it is almost certain that an equal number of ridges are in use at the same time in animals of the same age. Consequently the superior and inferior molars represented in figure 889, in each of which fifteen ridge-plates are in use above and below, probably represent animals of the same age, although they may not be ex- actly in the same stage of dental progression. The teeth from Kansas (Amer. Mus. 14558—Fig. 889A, 892A) have 14-15 ridge- plates in use, also the maxillary teeth from Texas (Amer.Mus. 14476—see Fig. 889B) have fifteen ridge-plates in use. These grinders (Fig. 889) are examples of typical A. imperator molars exhibiting the following characters: (1) Inferior ridge-plates con- eave anteriorly, partly worn summits of ridge-plates exhibiting Fig. 894. A right superior tusk of record size (16+ ft.) of Archidiskodon imperator (Amer. Mus. 22481), from Post, Texas, one-twentieth natural size, in comparison with the superb left tusk (13 ft. 9% in.) formerly in the National Museum, Mexico, after photograph of original taken by Barnum Brown in 1910, A letter from Sr. Francesco Con- treras of the Universidad Nacional de Mexico, May, 1935, states that the orig- inal tusk was unfortunately destroyed. 7-8 conelets; (2) superior molars, 1.M*, ap. 290 mm. (restored), tr. 111 mm., index 38; r.M?’, ap. 295 mm. (restored), tr. 120 mm., index 41; (3) M3, ridge-plates concave posteriorly, thus following the mechanical principle of reversal; (4) breadth-length index, M°=41. 1'This statement was true at the time it was written about eight or ten years ago. the tar pools of Rancho La Brea, California. (skull), 3801-1 (jaws). 1007 Incistve TusKs.—The characters of the tusks are superbly displayed in the cranium (Amer. Mus. 14476) as shown in figure 896; these tusks curve downwards, outwards, upwards, inwards, until finally they cross each other on the median line, forming a complete superior arch, as in the aged type specimen of Hlephas [=Parelephas] jeffersonii. The tusks are parallel but not closely opposed to each other where they issue from the alveoli, as in the specimens referred to this species in the Nebraska State Museum (Fig. 899) found at Hay Springs, Sheridan County, Nebraska. This comparative study of the superior and inferior teeth and jaws enables us to define the species A. closely than before. imperator much more Recorp ProposcipEAN Tusk A record proboscidean tusk (Amer. Mus, 22481) was presented to the American Museum in 1934 by Mr. George D. Doughty of Post, Texas. This tusk was found in the vicinity of Mr. Doughty’s home, namely, Post, Garza County, Texas, in which region the Imperial Mammoth (Archidiskodon imperator) seems to have flourished in Pleistocene time. This gift was first noted in the January, 1935, number of Natural History, page 84, followed in the April number, page 357, by a brief description with compara- tive estimated measurements. Since that time the tusk has been restored conservatively to conform to the measurements given by Mr. Doughty, who was unable to save the anterior end; in life it Fig. 895. Referred young male(?) skull of Archidiskodon imperator from Los Angeles Museum 3800-1 Although partly restored at the summit of the occiput, this is one of the most highly characteristic skulls ever found, in the sharp concavity of the forehead and the peaked cranium, resembling closely the skull of Mammonteus. In 1934, however, Mr. George D. Doughty of Post, Texas, presented the American Museum with a giant tusk of Archidiskodon imperator (Amer. Mus. 22481), the size and weight of which establishes the record for this species, or, as a matter of fact, for any proboscidean, namely, a length of 16+ feet. This gift was noted in 1935 in the January number of ‘Natural History,” page 84, followed in the April number (p. 357) by a brief description with measurements (see Fig. 894).—Editor.] £ lephas impera Zor A.M [4476 Fig. 896. Skull of Archidiskodon imperator (Amer. Mus. 14476), found at Victoria, Texas. The upper portion of this skull is entire- ly restored. The premaxillaries, the palate, and the superior grinding teeth are perfectly preserved. The palate and grinding teeth of this specimen are also represented in figures 897 and 889. The tusks are complete and natural, without restoration; they measure 4m. 21.5 em. or 13 ft. 10 in., as compared with 4 m. 20 cm., est., or 13 ft. 94 in., the measurement of the tusks of A. imperator in the Geologi- cal Institute of the City of Mexico (Fig. 894), as remeasured by Barnum Brown. [See footnote on preceding page which gives the length of the record tusk of this species (Amer. Mus. 22481) as 16+ ft.—Editor.] 1008 THE MAMMONTINA: ARCHIDISKODON AND METARCHIDISKODON 1009 was evidently longer than now restored, namely, 16+ feet. Associated with this giant tusk were two superior molars (M’), probably belonging to the same individual. CRANIAL CHARACTERS OF ARCHIDISKODON Compare Chapter XV, pp. 915-926, also figures 865, 891, 895, 896, 897, 902, and 906 of the present Memoir. In the progress from youth to maturity, the occipital summit and profile of the Archidiskodon cranium changes profoundly with growth, with sex, and with the development of the enormous’ superior tusks; these progressive stages are displayed in a comparison of the following eight more or less com- plete crania in the Los Angeles, American, and Nebraska State museums, and the Geological In- stitute of Mexico: Fig. 895. California. Primitive stage. Young male(?) skull from Elephant Pit No. 9, Rancho La Brea tar pool (Los Angeles Mus. Nos. 3800-1 (skull) and 3801-1 (jaws). [= Archidiskodon imperator. Fig. 891. Texas. Young male stage. Skull from near Dallas (Amer. Mus. 14475). [= Archidiskodon imperator. Fig. 902. Mexico. Adult male stage. Skull from Tepexpan (Geol. Inst. 212). [= Archidiskodon imperator. Fig. 897. Texas. Aged stage. Skull (summit restored) from Victoria, Victoria County (Amer. Mus. 14476). : [= RNASE imperator. | Elephas Fig. 918. Nebraska. ‘Lincoln County Mam- moth.’ Type skull, mandible, and tusks from near Curtis, Lincoln County (Neb. Mus. 5-9-22). [= Archidiskodon imperator maibeni.] Nebraska. “Adams County Mam- moth.’ Not figured in present Memoir. Skull, mandible, and teeth from sandpit 6 miles due south of Hastings (Neb. Mus. 11-3-18). [= Archidiskodon imperator.| Nebraska. ‘Howard County Mam- moth.’ Skull, mandible, teeth, and tusks, from near Dannebrog (Neb. Mus. 2—7-17B). [= Archidiskodon imperator.| Nebraska. ‘Custer County Mammoth.’ Skull, tusk, and teeth from Callaway (Neb. Mus. 16-6-16). [= Archidiskodon imperator.| Texas, VicroriA, SKULL (Amer. Mus. 14476—F ics. 896, 897, 906).—Only the lower portion of the cranium, including the palate, condyles, and paired grinding teeth, a is preserved, the upper portion, as shown : A.M. (4476 in figure 896, being entirely restored above the white dotted line; as the summit of the Victoria, Texas, skull (Amer. Mus. Fig. 897. Skull of Archidiskodon imperator from Texas, as largely restored and mounted in the American Museum. 14476) is restored, we have not ventured to ss 0 elt = Oe ate : : J “ Nas a is e not ventuned i Skull (Amer. Mus. 14476) found near Victoria, Victoria County, Texas; upper part of cranium give this hypsicephalic character full expres- entirely restored. Side view one twenty-fourth natural size; palatal view one-sixth natural size. sion; it characterizes all the young speci- See also figure 889 for view of palate; figure 896 for front view of same skull. tmperareor 1010 mens of the species A. imperator; but an aged Mexican skull (Fig. 902) has a broad, massive occiput. The hypsicephaly or peaked contour of the superior crest is based upon a beautiful cranium of a young male (?) in the Los Angeles Museum, from Rancho La Brea (Figs. 865, No. 10, and 895); we observe a remarkable similarity between the profile of this Los Angeles cranium and that of the true Mammonteus primigenius, namely: (1) Forehead concave; (2) parieto-occipital union acute, hypsicephalic; (3) occipital condyles and orbits approximate, i.e., brachycranial; (4) depth extreme from peak of cranium to base of lower jaw. Reverting to figure 865, these distinctive cranial characters of Archidiskodon imperator are E. imperator Amer. Mus, 10598 Ref. E. imperator Amer. Mus. 10598 Ref. 1/B nat. size Fig. 898. Jaw of Archidiskodon imperator ref. (Amer. Mus. 10598), found at Tule Cafion, Texas, by the American Museum Expedition of 1899. One- eighth natural size. Section and inside view of the same jaw, completely exposing M3, shown in figure 892B, B1. A, Right ramus, outer aspect; jaw uptilted. Al, Top view; tooth greatly shortened by perspective. A2, Same tooth in perpendicular view of crown, showing sixteen ridge- plates, fourteen of which are worn, indicating that this was a fully adult animal. See figure 907 for forelimb of same individual. OSBORN: THE PROBOSCIDEA closely analogous if not genetically related to those of the true mammoth (Mammonteus primigenius), while they are widely distinct from those of the trogontherian elephant (Parelephas trogontheriz) or from the typical elephant (Elephas indicus). Comparing the profile and palate of this cranium with that of Archidiskodon planifrons and of A. meridionalis, we observe the close similarity in fore-and-aft compression (cyrtocephaly) and corresponding vertical elevation (hypsicephaly, acrocephaly). FLATTENED CRANIA OF ARCHIDISKODON IMPERATOR FROM HAY SPRINGS, NEBRASKA, IN THE NEBRASKA AND AMERICAN MUSEUMS. AFTONIAN AGE. As provisionally determined by Matthew (1902.1, pp. 317, 318, and 1918.1, pp. 226, 227) from the American Museum col- lections of 1893 and 1897 (modified by Hay and by Osborn), also by Frick (1929.1, p. 107, and 1930.1, p. 79), [and finally by Barbour and Schultz who have published a preliminary list of the Hay Springs fauna (1937.1, pp. 3-6)], the following includes the species so far recorded by the above-mentioned authors: EDENTATA Mylodon garmani Allen Mylodon nebrascensis (Brown) Megalonyx leidyt Lindahl RopENTIA Cynomys niobrarius Hay Geomys sp. Thomomys sp. Castoroides ohioensis nebraskensis Barbour Castor sp. Ondatra nebrascensis (Hollister) Microtus? CARNIVORA Canis latrans? Say Canis (Aenocyon) dirus nebrascensis Frick Arctodus simus nebrascensis Frick Mustela vison? Schreber Smilodon nebrascensis Matthew PROBOSCIDEA Archidiskodon imperator (Leidy) PERISSODACTYLA Equus excelsus Leidy Equus excelsus niobrarensis Hay Equus calabatus nebrascensis Frick ARTIODACTYLA Platygonus vetus Leidy Camelops kansanus Leidy Camelops vitakerianus? (Cope) Tanupolama americanus (Wortman) Odocoileus sheridanus Frick Capromeryx furcifer Matthew Tetrameryx (Hayoceros) falkenbachi Vrick Bovid It is a striking fact that two extremely flattened crania have been discovered in the deposits of Hay Springs, Sheridan County, THE MAMMONTINA: ARCHIDISKODON AND METARCHIDISKODON Nebraska, which are considered of Lower Pleistocene (Aftonian) age. The first is the flattened cranium (Fig. 899) in the Nebraska Museum; the second is the flat cranium (Fig. 900) in the Ameri- can Museum of Natural History. The flattening of these crania is a proof of the heavy geologic pressure to which these beds were formerly subjected. Nersraska Musrum.—In figure 899 is represented the palate of a flat skull referred by Prof. E. H. Barbour to Elephas imperator (Neb. Mus. 1-11-8-17E); it was found in 1917 at Hay Springs, Sheridan County, Nebraska, in a bed of diatomaceous earth, flattened to four inches in thickness. The broad-plated grinding teeth enable us to confirm this reference, the molars presenting marked resemblance to those of the typical Archidiskodon im- perator. The geologic age of this specimen is very important, since the Hay Springs fauna is now regarded as of Lower Pleistocene (Aftonian) age.' In describing this skull, Barbour observed ten Fig. 899. Archidiskodon imperator cranium of aged male (Neb. Mus. 1-11-8-17E) found in a bed of diatomaceous earth, flattened to four inches in thickness; discovered in 1917 at Hay Springs, Sheridan County, Nebraska. [Middle Pleistocene (see Barbour and Schultz, 1937.1, p. 3).—Editor.] 1011 exposed ridge-plates, also three on the deeply worn anterior portion of the tooth, thirteen worn ridge-plates in all; the total number of ridge-plates is unknown. The tusks are very massive in transverse section and closely approximated where they issue from the alveoli; when found the left tusk was 9 feet long, but it was partly destroyed by careless collecting. AMERICAN MuseUuM CRUSHED 900).—This SKULL (Fic. Sls A.M.17355 Fig. 900. 17355), extremely flattened, exhibiting small posterior nares and cranial foramina. Discovered in 1916 at Hay Springs, Sheridan County, Nebraska. Archidiskodon imperator cranium of adult male (Amer. Mus. 1012 crushed cranium (Amer. Mus. 17355) closely resembles in size and character that in the Nebraska Museum (Neb. Mus. 1-11-8-17E). It was discovered in 1916 by Albert Thomson in the famous Hay Springs quarry, Sheridan County, Nebraska. The cranium belongs to a fully adult male, somewhat less aged than the Nebraska Museum cranium (Fig. 899), because the third superior molars, r.M3, |.M%, are less worn, displaying anterior ridge-plates. Besides the anomalous crushing, this specimen finely displays the char- acters of the palate, especially the ‘backward’ posterior nares, which contrasts with the very large ‘forward’ posterior nares of the superb Parelephas jeffersonii cranium (the Franklin County Mammoth) in the Nebraska Museum (Neb. Mus. 1-4-15—see Fig. 963), which for a time was erroneously referred to E. [= Parelephas] columbi. This palate of Archidiskodon imperator (Amer. Mus. 17355—Fig. 900) should be compared with that of Parelephas jeffersonii, which more nearly resembles that of Hlephas indicus bengalensis (Fig. 800). The backwardly placed opening of the posterior nares appears to be a very distinctive character of Archidiskodon. Hay Sprincs, Nepraska, Frick Cottection.—The follow- ing specimens, collected under the direction of Mr. Childs Frick, serve to confirm Archidiskodon imperator as the characteristic proboscidean of the Hay Springs horizon. Amer. Mus. numbers: 25501A, fragment of superior molar, ridge-plate maximum height 244 mm.; 25501, posterior portion of inferior molar, ridge-plate maximum height 164 mm.; 25501B, anterior portion of extremely worn superior molar, maximum width 111 mm.; 25500A, middle portion of an aged inferior molar, width 97 mm., laminar frequency 6% in 10 em.; 25500, portion of half-worn superior molar, width 107 mm., laminar frequency 6 in 10 em.; 25505 A—D, portions of inferior milk molars; 25506, por- tion of first milk molar. CRANIAL MATERIAL IN THE NEBRASKA STATE MUSEUM The finest Archidiskodon collection is in the Nebraska State Museum as recently listed by Director Barbour (1925.3, pp. 117, 118) in connection with his description of the “Columbian Mam- moth Elephas Maibeni.”’ This list is entitled ‘“Columbi Material in the State Museum”; as revised by Barbour to July, 1925, it is as follows: SKULLS 11-3-13 Skull, mandible, and teeth from sandpit 6 miles due south of Hastings, Adams Co., Nebraska. Apams County MaAmMortu. [= Archidiskodon imperator.| Skull, mandible, and tusks complete, of extraor- dinary size. Campbell, Franklin Co., Nebraska. FRANKLIN County Mammotu (compare T['igs. 963 and 964 of present Memoir). [= Parelephas jeffersoni.| Skull, tusk, teeth, and femur. Callaway, Custer Co., Nebraska. Custer County MAammMoru. [= Archidiskodon imperator.| Skull, mandible, teeth, tusks, two tibizw, scapula, and ribs. Found 7 miles south of Farwell, near Dannebrog, Howard Co., Nebraska. Howarp County Mammortu. [= Archidiskodon imperator.] 1-4-15 16-6-16 2-7-17B OSBORN: THE PROBOSCIDEA 5-9-22 Skull, mandible, tusk, forelimbs, six cervicals, several dorsals, four lumbars, sacrum, femur, part of pelvis, ribs, ete. Lincoln Co., Nebraska. Described as Elephas (Archidiskodon) maibeni. Lincotn County MAmMorTH. [=Archidiskodon imperator maibeni.] 1-4-26 Skull, teeth, and tusks of a form apparently identi- cal with A. maibeni; the same type of teeth and the same size and curvature of tusk (see Barbour, 1926.1, p. 122). From Lingle, Wyoming. [= Archidiskodon imperator maibeni.| {1-11-8-17E Palate of a flat skull referred by Barbour to Elephas imperator, Hay Springs, Sheridan Coun- ty, Nebraska. SHpripAN County Mammorn. [= Archidiskodon imperator.| MANDIBLES 19-9-17 Mandible, left half, with teeth. Powell, Jefferson Co., Nebraska. [= Archidiskodon imperator.| 29-25-11-18 Mandible and teeth. Inland, Clay Co., Nebraska. [=Archidiskodon imperator.| 2's-3-8-19 Mandible and teeth. Republican City, Harlan Co., Nebraska. [= Archidiskodon imperator.] 8-7-08 Mandible and teeth. Young. Benkleman, Dundy Co., Nebraska. [= Archidiskodon imperator.| 18-2-22 Mandible with teeth. In Aftonian gravels, Staple- hurst, Seward Co., Nebraska. Type of Hlephas scott? Barbour, 1925. [=Archidiskodon im- perator scotti, or juvenile A. imperator.| 23-6-14 Mandible with teeth. From Crete, Saline Co., Nebraska. Type of Hlephas hayi Barbour, 1915. [=Archidiskodon hayi.] GRINDING TEETH Largely as plotted by Hay (1924) under Elephas [= Archi- diskodon] imperator, thirty-two or more upper and lower grinding teeth, usually associated, from the following counties in Nebraska, Kansas, Indiana, and Wyoming: Dawes, Furnas, Jefferson, Platte, Richardson, Gage, Cass, Thayer, Howard, Adams, Fillmore, Buffalo, Cheyenne, Valley, Harrison, Clay, Butler—of Nebraska; also Herndon—of Kansas, Tipton—of Indiana, and Goshen— of Wyoming. Neb. Mus. 1-4-26 Archidiskodon imperator maibeni ref. of Lingle, Goshen County, Wyoming. Walls of cranium very sloping; maxi- mum transverse measurement of pal- ate 30 in., breadth of nasal opening 20 in.; circumference of tusk 25 in.; ridge-plates as thick as those in A. imperator. 5-11-20 Archidiskodon imperator ref., from Bell- wood, Butler County, Nebraska (cast Amer. Mus. 20069). Posterior half of a third right inferior molar, r.Ms, with eight broad ridge-plates; six arched ridge-plates in 20 em. as com- THE MAMMONTINA!: ARCHIDISKODON AND METARCHIDISKODON pared with nine arched ridge-plates in 20 em. in the corresponding tooth of Parelephas columbi (Amer. Mus. 13707, 1.Ms3); breadth 445 in.=112 mm. This is the most broadly ridge-plated molar on record. 4-12-13 Portions of inferior molars and femur. Arched lamine of inferior molar, four in 10 cm., as compared with three arched lamine in 10 cm. (Neb. Mus. 5-11-20). Femur 56 in. long or 1422 mm.; same measurement as that of Neb. Mus. 13-24-10-14 (see Fig. 908). Fig. 901. Referred superior molar, r.M°, of Archidiskodon imperator in the Geological After photograph kindly furnished by Seforita Compare Reyes, 1923, fig. 3. This tooth was described by Senorita Reyes (1923, p. 229) as follows: “Fig. 3. Escala: }4. Ejemplar nim. 207 del I. Institute of the City of Mexico (No. 207). Reyes. One-half natural size. ’ Geolédgico. Elephas imperator, Leidy. Molar derecho.’ felicis. Length of worn surface 220 mm., breadth 82 mm. ARCHIDISKODON IMPERATOR IN THE MUSEUMS OF THE CITY OF MEXICO See Parelephas columbi below (Chap. XVII) We owe to Osborn (1905.270), to Freudenberg (1922), and finally and most fully to Senorita Reyes (1923) descriptions of the superb materials referable to Archidiskodon imperator and to Parelephas columbi in the museums of the City of Mexico. PROGRESSIVE StTaGes (Mexico).—As described by Osborn (1905.270, p. 931): “The elephant remains in the National Mu- seum have usually been ascribed to Elephas columbi; but they in- clude molar teeth not only of this species, but of the much larger Locality: Zumpango, Mexico. Observe that there are 5! ridge-plates in 10 em. as compared with 9! in P. columbi 1013 form, Elephas imperator. In the collection of the Geological Survey of Mexico in the new survey building are the skull and tusks of an 7. umperator of magnificent proportions, the tusks measuring 5m. 10 cm., or 16 feet 10 inches [4 m. 20 cm. est., or 13 ft. 9% in., as re- measured by Barnum Brown] in length; this specimen was secured during the excavations for the great drainage canal of the Mexican Valley.” In the collection of the Geological Institute of Mexico, under the direction of Dr. José G. Aguilera, there are also several single teeth of Archidiskodon imperator, molars of the Parelephas columbi type from the village of Zacapti in Michoacan and of A. imperator from the valley of Puebla (ef. Osborn, op. cit., p. 931). Reyes, 1923.—The specimens described and figured (1923) by Senorita Reyes, are as follows: Elephas imperator Escuela de Ingenieros No. 1. M 3, with 6 ridge-plates exposed. quiac. Figs. 1 and 2, p. 228. Geol. Inst. 207. Palate with r.M*, 13 ridge- plates exposed; length 220 mm., breadth 82 mm. From Zumpango. Fig. 3, p. 229 (see Vig. 901 of the present Memoir). Jaw containing From Tequix- Geol. Inst. 210. R.M?*, length 300 mm., breadth 115 mm., 12s ridge-plates in 25 em. I’rom Tepexpan. Fig. 4, p. 230. Geol. Inst. 212. Cranium of young [adult] individual, a beautiful specimen well conserved. It is without tusks. Enormous prominence of occipital crest. Associated lower jaw. Dimensions of second superior molar, M?: length of worn area 168 mm., breadth 108 mm. From Tepexpan. Tig. 7, p. 233, Fig. 8, p. 234, and Fig. 9, p. 235 (see Fig. 902 of the present Memoir). Posterior view (Fig. 7). Anterior view (Fig. 8), forehead concave. Summit of occipital crest broadly rugose. Crown view of lower jaw (Fig. 9). Geol. Inst. 211. Left ramus of jaw! with 1.Ms, 12's ridge-plates in 25 em., length 220 mm., breadth 98 mm.; length of femur 1360 mm. From Tepex- pan. Fig. 5, p. 231, Fig. 6, p. 232 (see Fig. 903 of the present Memoir). ARCHIDISKODON (?)HAYI Rer. iv Mexico.—A mandible! (Fig. 903) from Mexico (Geol. Inst. v-211), referred by Senorita Reyes to Elephas hayi, supports the evidence afforded by Barbour’s type of EF. hayi that an Upper Pliocene stage of Archidiskodon, apparent- ly similar in progression to the A. planifrons of Asia and of southern France, entered North America. Osborn, 1924: This mandible from Mexico is intermediate between the #. hayi type of Barbour and the HL. imperator type of Leidy; it shows a prolonged rostrum (see Fig. 903); it appears to us like a progressively modified rostrum derived from an ancestor with a jaw like that of A. planifrons. While this mandible has been referred to Elephas hayi by Senorita Reyes, it seems to Osborn to 1Osborn, 1929: A similar mandible is described below (p. 1033) as Archidiskodon sonoriensis, to which species this specimen may also be referable. 1014 OSBORN: THE PROBOSCIDEA CranruM oF AN ApuLT MALE OF ARCHIDISKODON IMPERATOR IN THE GEOLOGICAL Musrum or Mexico Fig. 902. Referred mature cranium of Archidiskodon imperator in the Geological Institute of the City of Mexico (Geol. Inst. 212), reduced to one- twelfth natural size. After photograph kindly furnished the present author by Sefiorita Reyes. Locality: Tepexpan, Mexico. Compare Reyes, 1923, figs. 7, 8, and 9. This robust male cranium lacks the sharply peaked, acrocephalic, occipital crest structure seen in the young male(?) cranium from Rancho La Brea (Fig. 895). Judging by the elongated and massive alveoli and rugose exoccipital muscular attachments this cranium supported an enormous pair of tusks. The adult male tusks are estimated at 13-+—16-++ feet in length (see Fig. 894). represent rather an intermediate and distinct species in which there is a much larger number of ridge-plates than in the type of 2. hayz. It is another proof of how much we have still to learn regarding the characters and migrations of Archidiskodon in North America. Doubtless future excavation will reveal additional material of great value. We turn to Freudenberg (1922) for his views regarding the relationship of the true Hlephas columbi Falconer to the true E. imperator of Leidy (see p. 1017 below); also for his remarks on the relationship of A. 7mperator to the A. meridionalis of Europe and to the A. planifrons of Asia and southern France. ANCESTRY OF ELEPHAS IMPERATOR (I'REUDENBERG, 1922, SoerGEL, 1915').—(Freudenberg, 1922, p. 171): ‘Die Mastodonten wanderten friiher nach Amerika iiber, die Mlefanten spiter. Die innerasiatischen Hochlinder, die den meisten Saéugetierstammen den Ursprung gaben, sind auch hier in diesem [all als Heimat der mexikanischen Arten anzusehen. [Footnote: ‘Vgl. W. D. Matthew, Climate and Evolution. Annals of the New York Acad. of Science. Vol. 24. pag. 171-318.’]. Fir die Elefanten gilt das mit ziemlicher Sicherheit. Hine Ableitung des Hl. imperator von El. meridionalis Europas, wie Soergel [Footnote: ‘Ww. Soergel, Die Stammesge- Fig. 903. Mandible of Blephas [= Archidiskodon] hayt(?) ref. in the eollems schichte der Elefanten. IV. Die amerikanischen [lefanten. tion of the Geological Institute of the City of Mexico (v-211). Photograph Centralbl. f. Mineralog., Geolog. u. Paliiontolog. 1915. No. 9. through the courtesy of Senorita Reyes (compare Reyes, 1923, figs. 5 and 6). pag. 278-283.’| das will, ist gesucht. Hl. planifrons ist weit eher About one-seventh natural size. The prolongation of the symphyseal rostrum der Stammvater aller spiiterer Elefantiden als der LL. meridionalis, SURE SIe ae SES CINS NED bie type of archidishodon hays Barboms (Pig. 913 of the present Memoir); this feature, however, is more character- 'See citation below, page 1015. istic of A. sonoriensis. [See footnote on p. 1013.—Editor.] THE MAMMONTINA: ARCHIDISKODON AND METARCHIDISKODON der auch sicher mit Unrecht als der Ahne des El. antiquus von genanntem Autor angesprochen wird.”’ ANCESTRY OF ELEPHAS IMPERATOR, SOERGEL, 1915, p. 281.— “Wir miissen annehmen, dass die nach Amerika tiberwandernden Formen der Meridionalis-Trogontherii-Reihe schon beim Uber- wandern resp. kurz vorher besondere Charaktere gegeniiber dem europaischen El. trogontherii meridionalis ausgebildet hatten, Charaktere, die in der weiteren Entwicklung sich zum ‘Imperator- Typus’ steigerten. Es war also im dltesten Diluvium im Kreis der kontinentalen Elephantenformen eine Variationsbreite mit 2 Polen, El. trogontherii meridionalis im Westen und dem direkten Vor- fahren des El. imperator im Osten vorhanden. Es ist klar, dass als direkter Vorfahr beider Pole die nachstiltere Mutation in der kontinentalen Reihe zu gelten hat, das ist El. meridionalis des altesten Diluviums, vielleicht auch des Oberpliociin. Als Vorfahr des El. imperator Leidy hat also jedenfalls El. meridionalis Nesti zu gelten.” In our judgment Freudenberg was less fortunate in his treat- ment of Hlephas imperator as a subspecies of HF. columbi. For reasons originally stated by Osborn (1922.555) the type of Hlephas columbi Fale. is readily distinguishable from the type of Hlephas imperator Leidy; consequently the treatment of H. imperator as a subspecies of #. columbi is invalid; it should stand FL. [= Archi- diskodon| imperator. SYNONYMS OF ARCHIDISKODON IMPERATOR.—The four sub- species of Elephas columbi proposed by Freudenberg (1922) in this manner are redetermined by Osborn as follows: El. Colwmbi var. Felicis Freudenberg, 1922, p. 147, Taf. xv1 (vit), fig. 4, Tecamachaleo, Puebla, Mexico = Parelephas columbi felicis. El. Columbi var. silvestris Freudenberg, 1922, p. 152, fig. 19, Ejutla, Oaxaca, Mexico = Archidiskodon imperator silvestris. El. Columbi var. Falconer Freudenberg, 1922, p. 153, fig. 21, Tequixquiac, Mexico = Archidiskodon imperator falconert. El. Columbi var. imperator Freudenberg, 1922, pp. 160-171 = Archidiskodon imperator. The subspecifie value of Archidiskodon imperator silvestris and of A. imperator falconeri remains to be determined. We may here quote in full Freudenberg’s description of these subspecies and reproduce his type figures. Archidiskodon imperator silvestris Freudenberg, 1922 Figure 904 Kjutla, State of Oaxaca, Mexico. Probably Lower(?) Pleistocene. The type of this subspecies, related by Freudenberg to the species Hlephas [= Parelephas] columbi, is obviously referable as a synonym or subspecies of Archidiskodon imperator. The type (Fig. 904), with at least 16 ridge-plates, is too elevated to be re- garded as an M?; it appears rather to be a left M*; the six ridge- plates exposed (Fig. 904 B) resemble those of A. 7mperator rather than of P. columbi; this Ejutla type has the very broad outer coating of cement characteristic of A. imperator and lacking in P. columbi. El. Columbi var. silvestris Freudenberg, 1922. ‘Die Situgetier- fauna des Pliocdins und Postpliocins von Mexiko.”’ Geol. und Paleont. Abhand., N. F., Band XIV, Heft 3, pp. 152, 153. Type. 1015 Second [third] superior molar of the left side, M2 {I1.M*]. Pal. Coll. Univ. Leipzig 4402. Horizon AND Locauiry.—Hjutla, Estado de Oaxaca, Mexico. Tyrer Figurn.—(Op. cit., p. 146, fig. 19): “Fig. 19. a. El. Columbi var. silvestris. WKronenansicht des zweiten oberen Molaren, in *% nat. Gr. b. Seitenansicht in etwa /s nat. Grésse. Original in Leipzig, von Ejutla, Estado de Oaxaca in Mexiko. Coll. Felix, wohl aus jiingerem Diluvium. Subtropische Waldform.”’ I’REUDENBERG’S TypE OF ARCHIDISKODON IMPERATOR SILVESTRIS Fig. 904. Type, left M°, of Elephas Columbi var. silvestris Freudenberg, 1922, p. 146, fig. 19, reduced to one-third natural size. From Ejutla, Estado de Oaxaca, Mexico. After photographs sent by Doctor Freudenberg. Original in Leipzig (Pal. Coll. Univ. Leipzig, 4402), Coll. Felix. B, Crown view; A, side view. Type Descrietion.—(Op. cit., p. 152): “3. Der Zahn von Kjutla (El. Columbi var. silvestris Freudenberg). Wenden wir uns genauer dem oberen Molaren von Hjutla (Mstado de Oaxaca) zu, welchen Felix als ‘Zl. Columbi’ etikettiert und nur mit Vorbe- halt wegen der relativ enggestellten Lamellen unter El. primi- genius Blumenbach var. angefiihrt hat. Dieser Zahn trigt jetzt die No. 4402 der paliontologischen Sammlung der Universitit Leipzig. Es ist ein zweiter oberer Molar der linken Seite. Ich bilde ihn ab von der Seite als Textfig. 19b und a und von der Kau- fliche aus. Das Hinterende weist eine merkliche Einwaértsdrehung der Lamellen auf und zugleich eine Vertiefung in 5 em Nntfernung 1016 unter der letzten angekauten Lamelle. Diese Grube halte ich trotz ihrer Rauhigkeit fiir eine Pressionsmarke, hervorgebracht vom nachdriingenden M*. Die vordersten 4 Lamellen sind abge- brochen; einige davon diirften abgekaut sein. Dadurch ist die Lamellenzahl im Minimum=16.” The author rightly contrasts this species as identical in the ridge formula of M‘ with that of the subspecies Parelephas columbi felicis. He remarks (op. cit., p. 153): “Tn der Art der Abkauung gleicht dieser Zahn sehr dem von Pohlig—Nova Acta Acad. C. L. C. G. Nat. Cur. Vol. 57 abgebil- deten M2 des El. Americae Pohlig aus Chihuahua, Mexiko. Es wird auch ein M’ von dort an gleicher Stelle abgebildet mit dersel- ben schiefen Stellung der Lamellen zur Lingsachse des Zahns wie an den Molaren von Hjutla und ganz verschieden von den Hoch- plateauformen der Mesa central von Mexiko oder Puebla. Es empfiehlt sich aus oben genannten Griinden nicht, Pohligs Namen El. Americae {= Elephas americanus De Kay, 1842] fiir die Wald- form des El. Columbi anzuwenden, dessen Verbreitungsgebiet die siidwestlichen Randgebirge des mexikanischen Hochplateaus waren. Ihnen gegeniiber stehen die als Hl. Columbi var. Felicis aus der Mesa central bezeichneten Steppenformen. Archidiskodon imperator falconeri Ireudenberg, 1922 Figure 905 Tequixquiac, Valley of Mexico; probably Lower(?) Pleistocene. This Tequixquiac jaw (Fig. 905) represents the cotype and is undoubtedly related to Archidiskodon imperator rather than to Parelephas columbi. From the figure it is difficult to give the characters of this jaw, but it appears to be somewhat longer and more primitive than the type of A. imperator. The lower grinding teeth apparently present the following formula: M 3,947. In this jaw the 16+ ridge-plates exposed in the 1.M; are too widely separated to be related to P. columbi; the 1.M; measures ap. 175+ mm., tr. 95 mm., the dimensions apparently equal those of P. columbi, namely, neotype 1.M; ap. 298 mm., tr. 91 mm. The locality of Tequixquiac yields both A. imperator and P. columbi, according to Reyes. El. Columbi var. Falconer? Freudenberg, 1922. “Die Siugetier- fauna des Pliociins und Postpliocins von Mexiko.” Geol. und Pale- ont. Abhand., N. F., Band XIV, Heft 3, pp. 153-160. Lxrcto- typE.—lirst left lower molar, 1.M;, Mus. Royal Coll. Surg. 741a.! Coryrr.—Lower jaw with both third molars Ms; in situ. Horizon anp Locatity.—Tequixquiac, Mexico. Lxc- tTotyPe Ficure.—After Falconer, 1863.1, Pl. n, fig. 1. Corypr Ficurn.—Freudenberg, 1922, p. 154, fig. 21. DescripTion.—(Op. cit., p. 153): “Als Typus der Form gilt mir der von Falconer abgebildete M; von Mexiko (London) und ein Ms;, den ich auf S. 54 [154], Fig. 21 abbilde. Dieser M* [M3] gehort wohl der linken Seite an. Er stammt vermutlich aus dem Valle de Mexico. Die Hinterseite des Zahnes ist abgebrochen. Die Zahl der fehlenden Disken lisst sich nicht mehr bestimmen. lbensowenig weiss man, wie viele Lamellen vorn durch Abkauung verschwunden sind. Die Kaufliche ist intakt und wurde in natiir- licher Grosse abgebildet. Die Linge der Kauflaiche misst 175 mm, die Breite in der Mitte, einschliesslich des Zements, misst 95 mm. Die Kaufliche ist also etwa doppelt so lang wie breit. Der Ver- schmelzungstypus der eben erst angekauten Disken (im Sinne der OSBORN: THE PROBOSCIDEA Soergelschen Nomenklatur) ist median lamellar, lateral anular. Besonders an den drei ersten Jochen ist dieser Bau der Schmelz- pfeiler zu beobachten. Ein mir vorliegender M* von El. trogon- therii Pohlig [Footnote: ‘1) Abgebildet in meiner Arbeit: Die Sdugetiere des dlteren Quartirs von Mitteleuropa ete. Geolog. u. palaont. Abh. N. F. Bd. 12. Heft 4/5. Jena 1914. t.3.f.5. Siehe auch unsere Taf. rx [xvu].’] von Jockgrim in der Pfalz hat eine Kaufliche von 195 mm bei einer maximalen Breite von 82 mm. Das Verhaltnis von Lange zu Breite ist also iiber 2:1 (Breite=1). Wahrend bei El. Columbi die Verhialtniszahl fiir die Linge (Breite = 1) unter 2 ist.” CuHaractrers.—(Op. cit., p. 154): “Als altquartiir haben sich jene Formen herausgestellt, welche wir nach Falconer als Elephas Columbi var. Falconeri bezeichnen miissen. Ihre Lamellen sind kiirzer und stehen isolierter (wie z. B. an dem Taf. vir [xvi], Fig. 1 abgebildeten Molaren) verglichen mit der var. silvestris und erst recht mit der var. Felicis. Bei der typischen (altquartiiren) Marctbula interior de/ ELEPHAS PRIMIGENIUS, BLUM (06lTajo de Teguixguiad Coryrr oF ARCHIDISKODON IMPERATOR FALCONERI I’ REUDENBERG Vig. 905. Cotype jaw (one-sixth natural size) of Hlephas Columbi var. Falconeri Freudenberg, 1922, p. 154, fig. 21; crown view one-half natural size. From Tequixquiac, Mexico. Originally figured by Villada, 1903, Lam. vurr. ‘By inference this is Professor Osborn’s lectotype. It is not figured, however, in the present Memoir.—Hditor.] THE MAMMONTINA:: ARCHIDISKODON AND METARCHIDISKODON Columbi-Form, der gerade auch das Original Pohligs aus Mexiko zugehért, kommen am oberen M* vier Lamellen auf 53 mm, wiihrend deren 6 sich auf die gleiche Strecke verteilen bei dem jung- diluvialen Zahn von Mexiko Taf. vim [xvi], Fig. 4. Es mag nun dieser weniger Waldform als Steppenform gewesen sein.” El. Columbi var. imperator Leidy [=Archidiskodon imperator] Freudenberg, 1922 Spokam Bar near Helena, Montana. Under the true specific name Archidiskodon imperator should apparently be included the El. Columbi var. imperator of Freuden- berg, 1922, in the Geological Museum of Bonn. El. Columbi var. imperator Leidy Freudenberg, 1922. ‘Die Siiugetierfauna des Pliociins und Postpliociins von Mexiko.”’ Geol. und Palewont. Abhand., N. F., Band XIV, Heft 3, pp. 160-171. Freudenberg frequently introduces the subspecific name Elephas columbi var. imperator Leidy (op. cit., pp. 155, 156) and finally definitely employs it on page 160. Tyrer Descriprion.— (Op. cit., p. 160): ‘5. El. Colwmbi var. imperator Leidy. Wegen der vielfachen Verwechslungen und wegen der mutmasslichen Ab- stammung des El. Columbi von El. imperator ist es nétig, niher auf diese Form einzugehen. Es ist das um so notwendiger, als H. F. Osborn [Footnote: ‘Recent Vertebrate Palaeontology. Fossil mammals of Mexico. Science. Vol. 21. 1908 [1905]’.] diese Art aus Mexiko anfihrt. Osborn griindete seine Bestimmung wohl in erster Linie auf den gewaltig langen Stosszahn, der 16 Fuss misst und somit der liingste Stosszahn ist, von dem eine Kunde vorliegt. Aber warum sollte nicht ein Hl. Columb: gelegentlich riesige Dimen- sionen erreichen, zumal da diese gute Art in eine ganze Anzahl von Unterarten einmal zerlegt werden muss, sowie Matschie 4 Arten des afrikanischen Elefanten heute unterscheidet.”’ CHARACTERS.—After quoting Leidy, Osborn, Lull, Lueas, Holmes, Pohlig, Soergel, and Cope (op. cit., pp. 160-169), Freuden- berg concludes (p. 170): ‘‘‘Hl. imperator’ gewinnt mit seinem Aufsteigen ins jiingere Diluvium immer mehr Aehnlichkeit mit den englamelligen, schmalkronigen, eigentlichen ‘Colwmbi-Rassen.’ Vielleicht lésen sich die letzten Herde des waldlebenden Kaiser- elefanten auf im Colwmbi-Hauptstamm als sogenannte var. silvestris. Das hingt doch wohl mit dem immer weiter ziiruck- weichenden Seen- und Waldklima zusammen, das offenbar der Existenz der Imperator-Rasse in der Sonorischen Faunenprovinz giinstig war. Die zunehmende Austrocknung der diluvialen Seen- gebiete forderte die Entstehung von mammutihnlichen Colwmbi- Rassen, die wohl vorwiegend auf die spiirliche Koniferenkost und auf harte Steppenpflanzen angewiesen waren. Die im tropischen Laubwald isenden Jmperator-Rassen, die, wie El. indicus auf Ceylon gute Bergsteiger gewesen sein mochten, haben bei Afton wie im Becken von Puebla, Chihuahua und Mexiko als deutlichen Hinweis auf einstige Holziisung tief ausgehohlte Zementintervalle zwischen den dusserst kriftig gebauten Dentinpfeilern und ihrer oft stark gefalteten Schmelzhiille.”’ SKELETAL CHARACTERS OF ARCHIDISKODON Much remains to be done in establishing the skeletal characters of Archidiskodon imperator. In the fluviatile sand and gravel deposits of early Pleistocene time, in which these imperial mam- moths occur, skulls, jaws, and skeletal parts are widely scattered. 1017 The principal partly or fully associated materials hitherto described are the following: Figs. 907 and 906. Forelimb from Tule Canon, Briscoe County, Texas (Amer. Mus. 10598). Fig. 908. Femur from near Reynolds, Jefferson County, Nebraska (Neb. Mus. 13-24— 10-14). Fig. 912. Forelimbs associated with numerous ver- tebre, skull, jaw, one tusk, and parts of hindlimbs—the finest material thus far discovered [=Archidiskodon im- perator maibeni| from near Curtis, Lincoln County, Nebraska (Neb. Mus. 5-9-22). Za Z SA Zz Ze \\ ARCHIDISKODON IMPERATOR. AGED MALE Fig. 906. Outline reconstruction from right forelimb in the American Museum (Amer. Mus. 10598—Fig. 907) and from skull (Amer. Mus. 14476— Vig. 896), one-sixtieth natural size. The forelimb (Amer. Mus. 10598) measures 348 2mm. or 11 ft. 5 in., as shown in figure 912. This gives a shoulder height in the flesh of 3702 mm., which may be taken as a conservative estimate of the shoulder height in the flesh of a nine-tenths grown Archidiskodon imperator. Compare figure 912, also figure 907, lateral and front views of the same forelimb, one twenty-fourth natural size. The diagram of the forelimb, found in 1899 near Tule Canon, Texas, is taken from the mounted forelimb in the American Museum (Amer. Mus. 10598). The diagram of the skull is taken from Archidiskodon imperator (Amer. Mus. 14476), found at Victoria, Texas. Forr- AND HINpDLIMBs OF A. IMPERATOR (FIG. 907).—For- tunately the complete right forelimb (Amer. Mus. 10598), dis- covered in 1899 near Tule Cafion, Briscoe County, Texas, along with other parts of the skeleton belonging to a single individual, gives us the means of estimating the height of Archidiskodon imperator, namely, 12 ft. 1% in. =3702 mm. at the withers, 13 ft.= 3960 mm. at the top of the head when elevated. This elevation is shown diagrammatically in figure 906 which combines the skull (Amer. Mus. 14476) from Victoria, Victoria County, Texas, with the forelimb (Amer. Mus. 10598) from Tule Cafion, Briscoe County, Texas. 1018 OSBORN: THE PROBOSCIDEA In the rich collections made by Professor Barbour for the State Museum, University of Nebraska, are a femur (Fig. 908) and a humerus which may be compared with corresponding limb ele- ments in the skeleton of “Jumbo” (Loxodonta africana oxyotis) a2 ees \ | \ i in the American Museum. The specific reference is somewhat \ \ ; uncertain. The femur probaby does not represent a full-grown \ | i Archidiskodon imperator, which would greatly exceed the specimen ' here represented in the photograph (Fig. 908) kindly furnished by Professor Barbour. i FORELIMB OE ARCHIDISKODON IMPERATOR. AMERICAN MUSEUM Compare Figure 912 Fig. 907 tight forelimb referred to Archidiskodon imperator, as mounted in the American Museum (Amer. Mus. 10598). one twenty-fourth natural size, found by Alban Stewart in 1899 near Tule Cajon, Briscoe County, Texas, along with other parts of the skeleton belonging to a single individual. The same forelimb is re presented, vertically extended, in figure 912. The vertical measurements of the separate , radius 890 mm height 3482 mm. or I1 ft cartilages, segments are: Scapula 1017 mm., humerus 1095 mm , manus 480 mm., sum of total vertical 5 in. As the limb is always somewhat flexed, the foot pads, and the flesh and skin above the shoulder give this animal a total height of 3702 mm., or 12 ft. 1°4 in , a conservative estimate of the height of this animal E rig. 908. Limp Bones OF ARCHIDISKODON IMPERATOR (YOUNG ADULT) AND LOXODONTA AFRICANA OXYOTIS COMPARED (Lower) Three views (E) of femur of referred Archidiskodon imperator (Neb. Mus. 13-24-10-14) from Reynolds, Jefferson County, Nebraska, one- tenth natural size. Height of femur 1422 mm. =56 in. (Barbour, 1925.3, p. 116), same measurement as the femur (Neb. Mus. 4-12-13), namely, 4.8 feet. (Upper) The same femur (Neb. Mus. 13-24-10-14), one-twentieth natural size, compared with (A, B) femur and humerus of Loxodonta afri- cana oxyotis (skeleton of “Jumbo’’) in the American Museum; also with (C, D) humerus and femur (latter computed) referred to Archidiskodon im- perator (Amer. Mus. 10598). Compare figure 907 opposite. THE MAMMONTINA: ARCHIDISKODON AND METARCHIDISKODON BARBOUR’S DESCRIPTION (1925) OF THE SKELETON OF ARCHIDISKODON MAIBENI (See type description of skull, Jaws, and dentition below.) As above noted, this is the most nearly complete skeleton of Archidiskodon thus far discovered. The following is a free rather than a literal citation from Professor Barbour’s valuable paper of August, 1925 (Barbour, 1925.3), entitled, “Skeletal Parts of the Columbian Mammoth, Elephas maibeni, sp. nov.” For reasons shown in the systematic treatment of H. (Archidiskodon) maibeni below, there appears to be no doubt that this skeleton is properly 1019 flocks of poultry on the Karriger farm, before it was realized that they were out of the ordinary, after which the remaining parts were dug out and tared for with unusual appreciation and discernment. .. . The bones of Elephas maibeni were found pro- jecting from a loessial wall at the bottom of a small canyon. The general thickness of the loess at this point is about 100 feet. In an attempt to find additional material the writer, aided by Mr. William Hall and Mr. H.S. Karriger, blasted out many cubie yards of the loessial wall.” “The skeletal parts preserved are the skull, mandible, one Vig. 909. natural size. referable to Archidiskodon rather than to Elephas; also that it is closely related to Archidiskodon imperator rather than to Parelephas columbi. The narrative of discovery is as follows: “The last and most remarkable specimen of the columbi [imperator] type was found in Lincoln County, about 16 miles It was discovered by Mr. and Mrs. H.S. Karriger, and was dug out and preserved by them. Later it was procured of them for the palaeontological collections of Mr. Hector Maiben, who, next to Mr. Charles H. Morrill has been the most generous contributor of funds for the purchase and pres- ervation of choice Nebraska specimens. . . . It should be recorded in connection with this specimen that an unknown number of bones and parts of bones were pounded up to furnish lime for the large north of Curtis, on the Karriger farm. Imperial Mammoth (Archidiskodon imperator) of Nebraska and Texas. About one-fiftieth After restoration by Osborn and Knight, 1908. tusk, the atlas, axis, and four other cervicals, several thoracics, lumbars, and the sacrum, ribs, and double ribs, both fore limbs and parts of the hind limbs. Both fore limbs are practically complete and are essentially perfect save that but one foot bone was found, hence the feet must be supplied. The hind quarters are represented by parts of the pelvis, the shaft of a femur, and the major portion of a fibula. The dentition is perfect. The molars have 14 ridges bonded together by an uncommon thickness of cement, which is a character of columbian elephants. Even the great columbian elephant from Franklin County [referred to Parelephas jeffersonii in the present Memoir] seems surpassed in size. Heretofore, the tusks of the Franklin County elephant [Neb. Mus. 14-15] have been considered the largest reported, namely about 13! feet long OSBORN: with a maximum circumference, near the incisive sheath, of 29 inches. The skull and mandible of Elephas maibeni is noticeably larger. The incisive sheath shows that the tusk had a diameter of 104 inches and the incredible circumference of 33 inches.” “The outstanding characters of Hlephas [= Archidiskodon| maibeni are size, extreme curvature and divergence of tusks and incisive sheaths, unusual shortness of centra coupled with great width. The tusk must have lain in a plane or nearly so and must have described a circle, the radius being 28 inches (711 mm.). The diameter of the tusk is 6% inches (165 mm.) at the tip, 7}: inches (190 mm.) four feet back of the tip, and 10 inches (297 mm. [267]) at the incisive sheath. Originally it was a magnificent piece of ivory.” “The fore quarters have bones unexpectedly large and mas- sive, especially the humerus. The humerus is huge beyond the visualization of those who have not seen it, hence must judge of it from figures and measurements. In the hind quarters the bones are, if anything, less massive than might be expected. They seem in contrast to those of the fore limbs. Judging from the very short vertebra the body must have been unduly foreshortened.” “The centra of the vertebrae are very short, compared with their width, consequently certain ribs, presumably the fifth and sixth pairs, came in contact and became completely fused into one shaft with a double head and double tubercle, making huge and peculiar mammal ribs. A slight longitudinal depression is a vestige of the original boundary between the two shafts. It is interesting THE PROBOSCIDEA to note that the cancellous portion is uniform and continuous and is without partitions or vestiges of the dual origin. From this it may be inferred that they had been in coalescence for ages and that the character may have become fixed. At any rate it is not a case of pathology. The great fore limbs, several vertebrae, and their corresponding ribs, have been mounted as an arch, a palaeo- zoologic arch [Fig. 910 of the present Memoir], through which all students and visitors must pass on entering the main floor. It is but a temporary mount which must be dismantled and moved into the new museum sometime in 1926, where the complete skeleton will be carefully articulated and properly installed. So many skeletal parts are at hand that this huge elephant when ready for exhibition will seem complete. Its proper installation demands a ceiling 18 feet high.”’ [See Fig. 911 for present mount.—lHditor.] Vig. 910. Forelimbs of type skeleton of Archidiskodon imperator maibeni as mounted in the Nebraska State Museum (Neb. Mus. 59-22) in the year 1925. Professor Barbour is standing in the background. The dimen- sions of this superb skeleton are shown (I'ig. 912) in direct comparison with the forelimbs of A. imperator from Texas, of Lozxodonta africana, and of Elephas indicus. The cranium and tusks in the back- ground to the left, from Franklin County, Nebraska, belong to Parelephas jeffersoni. Barbour notes (1925.3, legend to Fig. 58): “The great skull and tusks in the background are those of Elephas columbi from Jefferson [Franklin] County.” {for present mount, see figure 911 on following page.—Editor. | “Mammoth scapulae are large, heavy,and very thin in portions, so it seems the more remarkable that the two huge shoulder blades should have been preserved practically without blemish. The right humerus is likewise perfectly preserved; in the left the head is wanting but has been modeled on from the right humerus. The right ulna is perfect save that the distal epiphysis is missing. This has been modeled after the left ulna in which the epiphysis is present but the shaft missing... . The fore limbs, four vertebrae, two pairs of single ribs, and one pair of double ribs of this ex- ceptional mammoth are mounted in approximate position and make an impressive arch, the height of which is 13 feet from the tip of the toe to the top of the spine, see fig. 58 [Fig. 910 of the present THE MAMMONTINA: Memoir; see also Fig. 912]’’. “Tn the flesh the height of Elephas [= Archidiskodon| maibeni at the shoulder must have been about 13 feet, and the top of the head of this magnificent beast must have been about 14 feet above the ground.” “From the tip of the toes to the top of the scapula is 11 feet, 6 inches.” “This specimen is believed to hold the record for size amongst the Columbian group of mammoths.” “In point of size, Elephas [= Archidiskodon| maibeni was a rival of the Imperial elephant itself, which stood 13} feet high. ARCHIDISKODON AND METARCHIDISKODON 1021 The tallest living African elephant stands 11 feet high and the average elephant of the menagerie and circus 8 to 9 feet.”’ Referring to figure 912 of the present Memoir, we observe that A 1 (right) is foreshortened, while A, B, C, D represent ortho- gonal full length projections of each limb segment, with the actual measurement of each segment in millimeters. This affords an absolutely reliable comparison of the ascending height of these four animals. The following entirely consistent comparative measurements appear to demonstrate that Archidiskodon imperator and the more primitive giant species A. maibeni towered in height far above the largest existing elephants: Manus Forearm Humerus Scapula Fig. 912, A, Archidiskodon imperator maibeni 297 (as mounted) 1085 1251 1066+ 424e (fully extended) B, Archidiskodon imperator 480 (fully extended) 890 1095 1017 C, Loxodonta africana oxyotis (‘Jumbo’) 361 (fully extended) 830 1078 925 D, Elephas indicus 391 (fully extended) 744 1008 864 Fig. 1083, Hesperoloxodon antiquus (Upnor) 1290 1170 Fig. 911. ty of Nebraska (Neb. Mus. 5-9-22): for dimensions of the forelimbs of this superb specimen, see figure 912, which was based on figure 910 showing the forelimbs as first mounted in 1925. a juvenile Hlephas indicus. Barbour. Present mount of the type skeleton of Archidiskodon imperator maibeni in Morrill Hall of the Universi- In the left foreground is a skeleton of After original photograph kindly furnished the present author by Professor Erwin H. Est. hewght — 4068 eooo Imperial Mammoth (maivent) Imperial Mammoth Esv. height African Elephant Max.height Tee 3482 \ 3826 Indian £ lephant Max. height —3G200 £. iNDICUS L. AFRICANA A, IMPERATOR A. MAIBEN! "Tumtzo" A.M. /0598 Neu. Nuss. §-9-22 97107%" 10-574" 11-5 12-67%" D Cc B A Al Fig. 912. SHouiper Hereuts or Livina AnD Extincr ELEPHANTS. Compare Fiaure 1194 [Professor Osborn’s method, used in the present Memoir, of estimating the height in the flesh is to add six and a third per cent. to the skeletal height.— Editor.} SKELETON LESH Elephas indicus: Standard skeletal height of full-grown male 3007 mm., 9 ft. 10° in. 3200 mm., 10 ft. 6 in. est.! Lozodonta africana oxyotis (“Jumbo’’): ‘ Adult male, extended skeletal height at shoulder 3194 mm., 10 ft. 5° in. 3396 mm., 11 ft. 1% in. est. Archidiskodon imperator (Amer. Mus. 10598); sex unknown: Extended skeletal height 3482 mm., 11 ft. 5 in. Estimated height in flesh 3702 mm., 12 ft. 14 in. Archidiskodon imperator maibeni (Neb. Mus. 5-922): (Right) Perspective drawing after mount in the Nebraska Museum. (Left) Right forelimb, also bony projection of each segment. Skeletal height: Fully extended forelimb 3826 mm., 12 ft. 6% in. Estimated, at shoulder, in flesh 4068 mm., 13 ft. 44 in. "Rowland Ward’s “Records of Big Game,” 1922, p. 468. A subsequent record (Ward, edition of 1928, p. 451) gives a height of 10 ft. 8 in.—Editor. | The record skeletal height of the African elephant (Lozodonta africana) is 3290 mm. =10 ft. 9) in. (“set up forelimb in Quex Museum,”’ Birchington, England—letter of January 6, 1931, from Major P. H. G. Powell-Cotton to Mr. J. W. Walker of Michigan) or 3520 mm.=11 ft. 6% in., in the flesh, as measured on the same specimen in the field by Major Powell-Cotton (op. cit., Ward, 1928, p. 456).- Editor.] 1022 THE MAMMONTINA:: ARCHIDISKODON AND METARCHIDISKODON The principal measurements of the type skull and skeleton of Archidiskodon imperator maibeni (see Barbour, 1925.1, pp. 114— 117) are as follows: SKULL Occipital condyle, greatest diameter Molar (r.M°), breadth Breadth-length index MANDIBLE Molar (M3), length of grinding surface Molar (M3), greatest breadth Breadth-length index Transverse ridges 14 ScAPULA Total height Extreme width suprascapular border Length of glenoid border HuMERUS Total length Transverse diameter of head ULNA Total length Fore Limp Total height In the flesh this huge creature must have stood not less than 13 feet high at the shoulders [13 ft. 444 in. est.] Tusk Diameter 8 inches from tip Diameter at incisive sheath ATLAS Total width Length AxIs Height CERVICAL 5 Posterior centrum, vertical diameter Posterior centrum, transverse diameter CERVICAL 6 Centrum, height Centrum, length measured at center CERVICAL 7 Posterior centrum, vertical diameter Posterior centrum, transverse diameter Dorsat 9 Centrum, vertical diameter, anterior Centrum, transverse diameter, anterior Dorsat 10 Posterior centrum, vertical diameter Posterior centrum, transverse diameter LUMBAR Centrum, length Centrum, height Average length of 4 lumbars Molar (r.M®), length of grinding surface Molar (r.M®), greatest thickness of cement Mm. Inches 125 5 245 We 110 4\4 45 13 My 240 9s 95 34 40 1,257! 49}, 889 35 556 22 1,226} 48 330 13 1,080! 42', 3,505! 11 ft. 6in.! 165 64 267 1015 440 1735 140 515 336 13% 186 1% 192 74 175 ys 39 1 185 74 192 746 167 635 144 5% 175 636 156 616 80 3}s 131 5 83 3h 1023 SACRUM Length 325 125 Greatest breadth 296 11% Centrum, anterior, transverse diameter 205 8 Femur Circumference at slimmest point, about middle 432 7 Transverse diameter at narrowest point of shaft 157 6.4 Anteroposterior diameter at narrowest point of shaft 115 4\ Archidiskodon hayi Barbour, 1915 Figures 893A, 903, 918, 915 Crete, Saline County, Nebraska. Lower Pleistocene. This appears to be the most primitive species of proboscidean thus far discovered in America, distinguished by its shallow jaw and elongated rostrum. It obviously belongs to the broad-plated Archidiskodon phylum, but with 10-11 ridge-plates like A. plani- frons. The relatively long, shallow jaw, the prominent rostrum, the low coronoid process are totally different from the adult Archidiskodon imperator, also from the adult A. meridionalis, and there is certainly a very strong resemblance to the jaws of A. plani- frons as figured by Mayet and Roman (Figs. 894 above and 914). A juvenile jaw of A. 7mperator would have this shallow character but Professor Barbour, who discovered this important specimen, remarks: (Letter of January 17, 1923): “... we both examined it [the type] as well as we know how, and count it an old not a young individual. I feel quite sure of this. [H#.] hay? is very distinctive.” Elephas hayi Barbour, 1915. ‘“‘A New Nebraska Mammoth, Elephas hayi.”” Amer. Journ. Sci., (4), XL, No. 236, pp. 129- 134 (1915.2). Typr.—Mandible and teeth. Associated with this jaw were fragments of a large tusk. (Barbour, op. cit., 1915, p. 132): ‘The distinguishing character on which this new mammoth must depend is derived, first of all, from the teeth. Especial care was exercised to determine whether the teeth in the jaw of H. hayi are penultimate or ultimate molars. If penultimate, a successor should be in evidence in each ramus, but not a fragment of a tooth or plate could be found in the cavities, which were filled with com- pact sand and gravel; nor could any such fragments be found in the surrounding gravels when screened. Undoubtedly the two teeth are the sixth molars, a point of consequence in this connection.” Collections of Hon. Charles H. Morrill, the Nebraska State Mu- seum (Neb. Mus. 23-6-14). Typr Locatiry AND Horizon. (Op. cit., p. 129): ‘Hurlbert sand pit at Crete, Nebraska, eight blocks east and three blocks north of the center of the town... . Aftonian. . . . 11 feet below the surface.” Type Ficgure.— Barbour, op. cit., 1915, p. 180, fig. 1, p. 133, fig. 3, p. 134, fig. 5d. Type DrscripTion.—(Barbour, op. cit., p. 129): ‘The chief distinguishing characters of Hlephas hayi are: unusual length of mandible; the last molar small, narrow, and anterior to the coro- noid; transverse ridges 10 to 11; angle distinet and sharp pos- teriorly; coronoids uncommonly prominent, deeply pitted, and set very obliquely. ... The mandible... measures 29! inches (750) from the tip of the symphysis to the angle, .... The depth of the jaw at the coronoids is 9% inches (241 ™"), .... The coronoid ‘etter of E. H. Barbour, May 8, 1929, gives corrected measurements of Barbour, Colbert, and Shanafelt, namely, scapula 1066-++ mm. =42 + in.; humerus 1251 mm. =49% in.; ulna 1085 mm. =42* in.; forelimb 3826 mm. =12 ft. 6% in.; length of grinding surface of left lower molar 238 mm. 1024 OSBORN: process is conspicuously robust, being 2% inches (70"™") through near its base, and an inch (25"™") near the summit. It stands 4 inches (102™") above the superior mandibular border, and 2 inches (51 ™") above the crown of the teeth. It is set more obliquely than in other mammoths. Its inner surface is deeply pitted, and extends from the outer to the inner alveolar border. . . . The teeth are those of a mature individual, with the crowns well Barpour’s Type JAW OF ARCHIDISKODON HAYI Fig. 913. 23-6-14), one-eighth natural size. restored. Compare jaw of Archidiskodon planifrons of Chagny, France (I’ig. 914). Professor Barbour (letter of January 17, 1923) remarks: “. . . we both examined it [the type] as well as we know how, and count it an old not a young individual. [E.) hayz is very distinctive.” Type of Elephas hayi Barbour, 1915, p. 130, fig. 1 (Neb. Mus. Ramus perfect to summit, condyle only I feel quite sure of this. worn. Though well cemented and strong, the teeth of 2. hayi are noticeably small. The postero-anterior diameter is but 9 inches (229 ™™"), and the greatest transverse diameter 3 inches (76 ™™). The dimensions of these teeth agree more closely with those of our earlier Nebraska mastodons than with those of our mam- moths. The number of transverse plates is noticeably reduced, for THE PROBOSCIDEA there are but 10 in one tooth, and 11 in the other, with no plates missing. .. . In #. hayi, there are 4 and a fraction transverse enamel ridges to the decimeter. The valleys are deep and bordered by highly crenulated enamel ridges. The great anterior prong branches widely and carries 3 plates. The teeth lack the sym- metrical development common to mammoths. They are notice- ably constricted back of the anterior prong, and taper posteriorly to 143 inches (88™™) .. . there are but 11 transverse ridges at most [to each molar tooth], the last being small, perhaps a heel. This form seems to be an earlier and more primitive type of mammoth than any other known to the State [Nebraska]. The inferior dental foramen is small, and has a circular border, while in H. imperator it is very large and deeply notched, as shown in the accompanying All to YB Wat. 57je ‘= J VY Li! From Seneze WY Fig. 914. Primitive mandibles of Hlephas |Archidiskodon| planifrons of the Siwaliks, India, Chagny-Bellecroix and Senéze of France, after Mayet and Roman, 1923, p. 81, fig. 13, inserted for comparison with type figure of EB. [Archidiskodon| hayi (Fig. 913 opposite). See caption to figure 849, p. 962 above. One-eighth natural size. Observe similarity to the Chagny mandible. ARCHIDISKODON = PLANIFRONS THE MAMMONTINA: ARCHIDISKODON AND METARCHIDISKODON figures. Although inferior dental foramina differ in individuals, and even between opposite sides of the jaw, the differences shown by the cuts are significant. The ascending rami of our probo- scideans also vary between wide limits.” Spreciric CHaractTers.—(1) Mandibular ramus elongate, de- pressed, coronoid process low. (2) Third inferior molars broad in center, with ten to eleven ridge-plates set very far apart; laminar frequency 443in 10 em.; molars narrowing posteriorly. (3) Ramus of jaw shallow, moderately expanded; rostrum relatively promi- nent. (4) Agreeing with Archidiskodon planifrons in ridge formula, differing in greater width of M;. (5) Dimensions of M3, length 229 mm., maximum breadth 76 mm., index 33. COMPARISON WITH ARCHIDISKODON PLANIFRONS (OSBORN, 1924).—It is remarkable that the type of Archidiskodon hayi re- sembles closely in the profile of the jaw the Archidiskodon plani- frons of India and of southern France, apparently justifying Bar- bour’s statement that A. hayi seems to be an earlier and more primitive type of mammoth than any other known to the state [Nebraska], and suggesting the possibility that we have to do with Fig. 915. 1025 Archidiskodon imperator scotti Barbour, 1925 [=A. imperator ?juvenile] Vigures 916, 917, Pl. xx1 Five miles south of Staplehurst, Seward County, Nebraska. Lower Pleistocene, Aftonian gravels. Awaiting further evidence, Osborn is inclined to regard the type of ‘Elephas scott:’ as representing a young individual of Archidiskodon imperator. This Archidiskodont, discovered in 1922 in the Aftonian gravels, 5 miles south of Staplehurst, Seward County, Nebraska, and 20 miles south of Crete, Saline County, where the type of Barsour’s Type oF ARCHIDISKODON HAYI COMPARED WITH A. IMPERATOR REF. (Left lower) Type right mandible of Archidiskodon hayi, sectioned at three points, grinding teeth with 10-11 ridge-plates. (Left upper) Archidiskodon imperator ref. Same aspect of right mandibular ramus, grinding teeth with 18 ridge-plates. (Right upper and lower) Archidiskodon imperator with 18 ridge-plates, lateral and superior aspects of mandible. Both specimens in the Nebraska State Museum, Morrill Collection. After Barbour, 1915.2, figs. 2, 3, and 4. the arrival in North America in the late Pliocene or early Pleisto- cene of a primitive proboscidean, displaying some of the chief A. planifrons characters, namely: (1) Jaw long and shallow; (2) rostrum prominent; (3) coronoid relatively depressed; (4) grind- ing teeth broad with 10-11 transverse ridge-plates. Supplementary figures and sketches of the jaw and teeth of A. hayi (jaw, Fig. 893 A) serve to emphasize the wide contrast between this type jaw and the adult jaw of A. imperator (Fig. 892 B, A). Elephas |= Archidiskodon| hayi was found (1914), is regarded by Barbour (1925.1, p. 22) as a mature individual, as primitive as A. hayi, if not more so. This statement rests upon the identifi- cation of the small 8 ridge-plated molars as representing third inferior molars, M;, which appears to Osborn very doubtful, especially as the jaws of the type of ‘Hlephas scotti’ (Fig. 916) are very robust in section with greatly abbreviated symphysis, alto- gether different from the relatively slender, elongated mandibular 1026 rami of A. hayi. Consequently it seems probable that they are second inferior molars, Me. Elephas scotti Barbour, 1925. “‘Elephas scotti, A New Primitive Mammoth from Nebraska.’”’ The Nebraska State Museum, Bull. 2, Vol. I, April, 1925, pp. 21-24 (Barbour, 1925.1). Typr.— Mandible with last [second?] lower molar of each side (Neb. Mus. 18-2-22, the Maiben Paleontological Collections. Cast of type right Ms; [r.M2], Amer. Mus. 14610). Horizon AND Locatiry.—Discovered on farm of Mr. E. J. Hartman, five miles south of Staplehurst, Seward County, Nebraska. TYPE Vig. 916. 18-2-22), from Seward County, I. H. Barbour who in 1925 made this the type of Hlephas scottt. Juvenile jaw of Archidiskodon imperator ref. (Neb. Mus. Nebraska. Prof. Compare After photograph by figure 917. Scale about one-sixth natural size. These 8 ridge-plated grinding teeth, described as third inferior molars, Ms, are regarded by Osborn as second inferior molars, Me (cast of r.M», Amer. Mus. 14610). OSBORN: THE PROBOSCIDEA Figure.—Op. cit., text figures 7-10. Tyre Derscriptrion.—(Barbour, 1925.1, pp. 21-24): “On February 18, 1922, the mandible of an unusually primitive mam- moth was secured for the palaeontological collections of Mr. Hector Maiben by Mr. E. T. Engle. . . . The peculiarities of this mam- moth seem to entitle it to a position as a distinct species, for which we are proposing the name, Elephas scotti, named for Professor William D. [B.] Scott. It cannot be compared with the later and more advanced mammoths, such as imperator, jeffersoni (columbi), or primigenius. It is comparable instead, with the earlier and more conservative mammoth, Elephas hayi. . . . The new mammoth is as primitive as Elephas hayi, if not more so. Like hayi, it is un- doubtedly a mature individual. Its teeth are taken to be last [second] molars. The enamel plates, which are highly crenulated, incline noticeably backward, and are worn with extreme obliquity. At the same time the valleys, or dental spaces, are so deeply in- dented, as to still further heighten and exaggerate the effect. There are but five pronounced ridges, and in all, but eight and a cone. Two of the anterior ridges are so confluent that the count is ren- dered somewhat uncertain, as shown in figure 10 [our figure 917A, Al, A2]. The transverse ridges in E. hayi are eleven. The molars of scotti measure 219 mm. (8% in.) in length, by 117 mm. (4%, in.) in extreme width. They are short and abruptly expanded in the middle. In the mammoths the number of enamel ridges to the decimeter serves, in a general way, in the recognition of species. In E. seotti there are three and a fraction, transverse, grinding ridges to the decimeter; in FE. hayi four and a fraction; in E. imperator five to six; in E. jeffersoni (columbi) six to eight; and in E. primigenius nine to ten. It is a noteworthy feature that the robust jaws of Elephas scotti come within three-fourths of an inch of meeting on the middle line, as is plainly shown in the figures [see Figs. 916 and 917A of the present Memoir]. This is not due to crushing, as far as can be learned, for the specimen in hand is essentially perfect. The coronoids of the earlier and the later mammoths differ widely and are worthy of notice. Those of scotti and hayi are much more robust, thick, and heavy, and flare out- wardly, and are posterior to the molars. The inner wall is broader and more heavily roughened and pitted for ligamentous attach- ment. Each ramus, measured back of the molar, has a width of 185 mm., (7%.in.) and a depth of 180 mm., (7's in.). On the middle line the jaws are but 19 mm., (*4 in.) apart.”’ Osborn, 1928: The Seward County jaw (Neb. Mus. 18-2-22 —see Fig. 916, and as seen from above, Fig. 917A) in Osborn’s opinion resembles a juvenile jaw of A. imperator rather than the type jaw of #. hayi Barbour. In superior view (lig. 916) the jaw appears fairly robust, less swollen than that of A. imperator, with highly characteristic outwardly flaring coronoid processes. In lateral view (lig. 916) it is relatively short and deep, the rostrum is short and depressed, in wide contrast to the long, shallow jaw of the type of A. hayi. The single grinding tooth, in which 8-9 ridge- plates appear, may represent My» of a young A. imperator; the tooth is obliquely worn and consequently the dental space between the broad enamel ridges appears to be much greater than it actually is; the disparity in the actual distance between the ridge-plates (ig. 917 A2) and the apparent distance due to obliquity of wear (Fig. 917 Al) are clearly shown in this diagrammatic represen- tation. THE MAMMONTINAS: ARCHIDISKODON AND METARCHIDISKODON Archidiskodon imperator maibeni Barbour, 1925 (See full description above, page 1019, of the skeleton of A. mazibenz) Figures 815, 824, 910-912, 917, 918, 1239, Pl. xx1 Lincoln County, about sixteen miles north of Curtis, Nebraska. Aftonian,! loess 100 feet in general thickness. Upper Pleistocene (see Fig. 1239, also Pl. vin, Vol. 1). The type of this species is an unusually complete skeleton, entitled the ‘Lincoln County Mammoth,’ in the Nebraska State Museum, described by Barbour in 1925 (1925.3) as the “Columbian Mammoth Elephas maibeni, sp. nov.” and subsequently (June, 1926) transferred by him to Archidiskodon maibeni, now mounted in the new Museum, Morrill Hall, University of Nebraska. To Neb, Mus, 5-9-22 1/B nat. size Fig. 917. third inferior molars, M3, in place, exhibiting 15+ ridge-plates, 14 exposed; and (A) type jaw of Elephas scotti [= Archidiskodon imperator scolti or ?juvenile A, imperator| (Neb. Mus. 18-2-22), containing second inferior molars, Mo, exhibiting 8 ridge-plates, and measuring ap. 219 mm., tr. 117 mm., i.e., short and abruptly expanded in the middle; Al, A2, enlarged views of right second inferior molar of the type of ‘#. scotti’ (cast Amer. Mus. 14610). the above full description of the skeleton (p. 1019) by Barbour (1925.3) may now be added the type description and characters of the skull and dentition (Barbour, 1926.1), as follows: Archidiskodon maibeni Barbour, 1926. Professor Barbour in his supplementary description (‘“‘Archidiskodon maibeni,’”’ Nebras- ka State Museum, Bull. 11, Vol. I, June, 1926, pp. 119-122) states: “‘Archidiskodon maibeni was first described in Bulletin 10 of the Nebraska State Museum under the title ‘Skeletal Parts of the Columbian Mammoth, Elephas Maibeni, sp. nov.’”’? Under this designation Barbour gives additional measurements and comments on the skeleton. He also mentions the discovery of another skull (Neb. Mus. 1-4-26) referable to A. mazbeni. Elephas maibeni Barbour, 1925. ‘Skeletal Parts of the Columbian Mammoth, Elephas Maibeni, sp. nov.,’’ Nebraska State Museum, Bull. 10, Vol. I, August, pp. 95-118. TyPr.— (Op. cit., p. 98): “The skeletal parts preserved are the skull, mandible, one tusk, the atlas, axis, and four other cervicals, several 1027 thoracies, lumbars, and the sacrum, ribs, and double ribs, both fore limbs and parts of the hind limbs. . . . The hind quarters are represented by parts of the pelvis, the shaft of a femur, and the major portion of a fibula. The dentition is perfect.’”’ Neb. Mus. 5-9-22. Horizon AND Locatiry.—Discovered by Mr. and Mrs. H. 8. Karriger about sixteen miles north of Curtis, Lincoln County, Nebraska, on the Karriger farm. Tyrer Figure.— Op. cit., Figs. 58-60, 63-70, 72, 74, 76-87. Typr Descriprion.—(Barbour, 1925.3, pp. 97-111): “Con- fusion has long surrounded the columbian and_ jeffersonian mammoths. But the one under consideration is undoubtedly of the true columbian type. The bones of Elephas maibeni were Neb, Mus. 18-2-22 V4 nat size Comparison of: (B) Mandible of the type of Elephas (Archidiskodon) maibeni [= Archidiskodon imperator maibeni] (Neb. Mus. 5-9-22), with found projecting from a loessial wall at the bottom of a small canyon. The general thickness of the loess at this point is about 100 feet. . . . The outstanding characters of Elephas maibeni are size, extreme curvature and divergence of tusks and incisive sheaths, unusual shortness of centra coupled with great width. The tusk must have lain in a plane or nearly so and must have described a circle, the radius being 28 inches (711 mm.) The diameter of the tusk is 6}5 inches (165 mm.) at the tip, 7's inches (190 mm.) four feet back of the tip, and 105 inches (297 [267] mm.) at the incisive sheath. Originally it was a magnificent piece of ivory. ... The humerus is huge beyond the visualization of those who have not seen it, hence must judge of it from figures and measurements. In the hind quarters the bones are, if anything, less massive than might be expected. ... Judging from the very short vertebrae the body must have been unduly foreshortened. ... The skull is broken into several large, and numerous small pieces, which have not been set permanently in place. ... The ‘{Lugn and Schultz (1934.1, p. 376, also Table A) regard it as of Iowan, late Pleistocene age.—Editor.| 1028 OSBORN: THE PROBOSCIDEA accompanying free hand sketches were made with the pieces set apart... . The anterior borders of the coronoids are excessively approximately in position, and give our impressions of the skull. roughened; they are even nodular, and we find no parallel... . ... The head must have presented a bull-dog effect or aspect. The lachrymal process, which is uncommonly large, prominent, It might be called the bull-dog mammoth. . .. The mandible is and acorn-shaped, is dissimilar to all others available for study and well preserved, and in point of size passes the largest in our col- comparison. . . . The fore limbs, four vertebrae, two pairs of single lections. The rami are widely divergent and the condyles far ribs, and one pair of double ribs of this exceptional mammoth are mounted in approximate position and make an impressive arch, the height of which is 13 feet from the tip of the toe to the top of the spine, see fig. 58 [Fig. 910 of the present Memoir]. In the flesh the height of Elephas maibeni at the shoulder must have been about 13 feet, and the top of the head of this magnificent beast Mo Barsour’s Type or ARCHIDISKODON IMPERATOR MAIBENI Vig. 918. Archidiskodon imperator maibeni, the ‘Lincoln County Mammoth,’ superior and inferior dentition of the type skeleton (Neb. Mus. 5-9-22). After photographs kindly furnished the present author by Prof. E. H. Barbour (compare Barbour, 1925.3, figs. 64, 65, 67, 68). A, Mandible of A. imperator maibeni from above, about one-cighth natural size. Right and left inferior molars, r.M3, 1.M3, with 15+ ridge-plates, 14 exposed, more or less worn, broad cement, somewhat sinuous, but. slightly concave posteriorly, crown externally plane, internally convex. Al, The same in lateral view. One-eighth natural size. B, Palate containing right and left third superior molars, r.M*, 1.M*. One-cighth natural size. B1, Right third superior molar, r.M*. One-fourth natural size. C, Extremity of tusk 4 feet in length measured on outer curve. About one-tenth natural size. Observe in B, B1, 16+ ridge-plates, of which 1-13 show signs of wear, quite strongly concave posteriorly with heavy border of cement; crown externally convex, internally slightly concave. Very similar in contour and ridge formula to A. imperator (Amer. Mus. 14476—I'ig. 889B), from Victoria, Texas. Observe in A, Al, 15+ ridge-plates, 14 exposed. THE MAMMONTINA:: ARCHIDISKODON AND METARCHIDISKODON must have been about 14 feet above the ground. From the tip of the toes to the top of the scapula is 11 feet, 6inches. This specimen is believed to hold the record for size amongst the Columbian group of mammoths. In point of size, Elephas maibeni was a rival of the Imperial elephant itself, which stood 13}5 feet high. The tallest living African elephant stands 11 feet high and the average elephant of the menagerie and circus 8 to 9 feet.” Type or ARCHIDISKODON HAROLDCOOKI Fig. 919. in situ of Elephas haroldcooki Hay, 1928. fig. 1. One-fifth natural size. Type mandible and third inferior molar of the right side After Hay and Cook, 1930, PI. 1m, Archidiskodon haroldcooki Hay, 1928 Tigure 919 Found in Holloman’s gravel quarry, Frederick, Oklahoma. Aftonian? gravels. Elephas haroldcooki Hay, 1928. “Preliminary Descriptions of Fossil Mammals Recently Discovered in Oklahoma, Texas and New Mexico.” Proc. Colo. Mus. Nat. Hist., Vol. VIII, No. 2, Pt. 1, February 2, 1928, p. 33. Typr.—Nearly complete lower jaw containing last right and left molars in situ (Colo. Mus. 1057). Horizon AND Locaniry.—Holloman’s gravel quarry, 1029 at Frederick, Oklahoma. Type Fiagure.—Hay and Cook, 1930.1, Pls. 11, fig. 1, v, fig. 1, x1 and xtv. Type Descriprion.—(Op. cit., 1928, p. 33): ‘“Hlephas harold- cooki Hay. Based on a nearly complete lower jaw containing the right and left last molars. Anterior fang and 2 ridge-plates absent through wear. 12 ridge-plates and the rear talon present. 4.4 ridge-plates in 100 mm. Enamel thick, moderately folded. A loxodont expansion present in some of the ridge-plates. The crown very high. Found in Holloman’s gravel quarry at lrederick, Oklahoma. No. 1057, Colorado Museum of Natural History, Denver.” SUPPLEMENTARY DescripTION (Hay Ann Cook, 1930.1, PP. 32, 33).—“This jaw was embedded in the cemented gravel very close to the Permian red clay .. . It will be seen that nearly all of the jaw behind the teeth is lacking. These teeth are in fine con- dition and show that the animal was somewhat beyond middle age. The height of the jaw where the ascending ramus arises is a little more than 200 mm. Its thickness, where greatest, at the middle of the tooth is 155 mm.; the width of the bone, taken at the rise of the ascending rami, is about 390 mm.” “As will be observed from the views of the jaw, it is wholly without a beak in front; and does not turn downward, also the symphysis is short, about 60 mm. in length. The teeth present are parts of the hindmost molars. This is shown by the arrangement of the rear ridge-plates, only partly shown in the figures. In front of each tooth (plate m1, figure 1 [=Fig. 919 of present Memoir]), especially of the one of the right side, is seen a cavity in which was lodged the anterior fang. The tooth had been worn down to the bar of bone separating this fang from the part behind it and the fang had fallen out. With it went three worn-out ridge-plates. Behind the bar of bone may be counted on the grinding face 11 ridge-plates, the tenth being represented by a small circle of enamel, the eleventh by a dot of enamel. The length of the abraded surface of the molar (plate v) is 182 mm.; its width, with the cement, is 85 mm.; without it, 80 mm.” “Behind this eleventh ridge-plate is a mass which may be regarded as a talon, a ridge-plate as yet undeveloped. Adding now to these eleven ridge-plates the two supported by the anterior fang and the one by the bar of bone behind it, we have 14 ridge- plates for the hindmost molar of the species. From plate x1v it is seen that the ridge-plates are very thick. Measured at one-half their height three are spanned by a line just a little less than 75 mm. Measured at one-half their height 4.44 plates are crossed by a 100 mm. line. The hind end of the crown is almost 6 inches high, nearly as high as the grinding surface was long.” “The view (plate x1v) of the inner face of the tooth shows that the ridge-plates are at first directed strongly forward, then are turned abruptly upward and (in relation to the grinding face) somewhat backward. It is due to this obliquity that the front enamel plate of the hinder ridge-plates is more exposed to view than they are farther forward. It will be seen that the outer ends of the loops of enamel are rather strongly turned forward. The enamel, as usual in primitive elephants, is thick, here about 3 mm. It is moderately crimped. A feature of interest is the presence of a lozenge-shaped expansion at the middle of each ridge-plate. This is a characteristic of the earlier elephants, as E. planifrons, E. meridionalis, and E. imperator.” Wo Natural Sz2ze ARCHIDISKODON EXILIS ARCHIDISKODON IMPERATOR Ref —— Dwarr INsuvar Species or ARCHIDISKODON COMPARED WITH A. IMPERATOR Jaws of Archidiskodon exilis to same one-sixth scale as Jaws of A. imperator Fig. 920. Comparison of Plephas [Archidiskodon] exilis (Calif. Inst. Tech. Coll. Vert. Pal. 14) with Archidiskodon tmperator (Amer. Mus. 10598) from near Tule Cation, Texas, to the same one-sixth scale, with the exception of r.M? and 1.M3, crown views, which are one-half natural size, reveals the marked diminution in size of the species from Santa Rosa Island, California, described in 1928 by Stock and Furlong. After photographs kindly forwarded to the present author by Dr. Chester Stock of the California Institute of Technology. 1030 THE MAMMONTINA:: ARCHIDISKODON AND METARCHIDISKODON Archidiskodon exilis Stock and Furlong, 1928 Figures 815, 920, 921, Pl. xx1 Santa Rosa Island, California. Pleistocene. This dwarfed insular form of imperial mammoth, estimated at 6-8 feet in height measured at the shoulder, as compared with 12 ft. 1%in., distinctive height of Archidiskodon imperator, is of the greatest interest. There is little doubt from the drawings received (Fig. 920) that this is a diminutive insular form related either to A. imperator or to Parelephas columbr. Elephas exilis Stock and Furlong, 1928. ‘‘The Pleistocene Elephants of Santa Rosa Island, California,” Science, Vol. LX VIII, No. 1754, p. 140. Typr.—‘A skull and mandible including four cheek-teeth and two tusks” (Calif. Inst. Tech. Coll. Vert. Pal. 14). Horizon AND Locauity.—Santa Rosa Island, California, the second largest of four islands separated from the mainland by the Santa Barbara Channel. Pleistocene. TYPE 1031 recently summarized the available information and recognizes the presence of Elephas imperator and of an undetermined species. During the past year Dr. Spencer Atkinson and Mr. J. A. Barbieri, of Pasadena, secured a fragmentary elephant skull on Santa Rosa Island and presented the specimen to the California Institute of Technology. Through the courtesy of the Vail Company of Los Angeles, owners of the island, the California Institute, with the cooperation of the Carnegie Institution of Washington, have been given the opportunity to investigate the occurrence, and facilities were kindly made available to collect further remains. . . . The collections secured by the California Institute include a number of teeth, parts of skulls and skeletal material. Occasionally several skeletal elements and teeth are found associated in the deposits. Usually the remains are scattered. One curious feature of the occurrence is the apparent total absence of associated mammalian types. The proboscidean remains are referable to the genus RESTORATION OF ARCHIDISKODON EXILIS OF SANTA Rosa ISLAND Pig. 921. Quaternary deposits, Santa Rosa Island, California. Stock. Ficgurr.—Stock, 1935.1, p. 210, fig. 6. SUPPLEMENTARY DE- scriIpTION.—Stock, “Exiled Elephants of the Channel Islands, California,’ Scientific Monthly, 1935, XLI, September, pp. 205 214, text figs. 1-10. Tyrer Description.—(Stock and Furlong, 1928.1, pp. 140, 141): “W. G. Blunt’s discovery of fossil teeth of an elephant on Santa Rosa Island, one of the Channel Islands off the coast of southern California, was recorded by Stearns [Footnote: ‘Stearns, R. E. C., Proc. Calif. Acad. Sci., Vol. 5, p. 152, 1873.’] in 1873. Since that time this interesting and significant occurrence has been referred to by several authors. Hay [Footnote: ‘Hay, O. P., Carnegie Inst. Wash. Pub. 322B, pp. 42, 48 and 51, 1927.’] has Facial portion of the skull, with tusks and lower jaw, of Hlephas exilis Stock and Furlong, 1928. After unpublished photograph kindly furnished by Dr. Chester From Elephas. The individuals exhibit considerable variation in size, and this is undoubtedly to be ascribed in part to differences in age. A survey of the collection as a whole yields the impression rather strongly that the elephant types were of relatively small size. Some of the forms may have a height of six to eight feet as measur- ed at the shoulder. The larger individuals are perhaps comparable in size to the American mastodon and are certainly smaller, possibly considerably smaller, than the Pleistocene mammoths of the southwestern United States. While the Santa Rosa Island elephant has been determined as representing the species Hlephas primigenius Blumenbach and FE. (Archidiskodon) imperator Leidy, the difference in size, coupled with differences noted in the skull and 1032 OSBORN: THE PROBOSCIDEA dentition, seem quite clearly to distinguish the island form as a distinct species for which the name Elephas exilis is here pro- posed.” SuPpPLEMENTARY Description (Stock, 1935.1, pp. 206, 207, 212-214).—“‘Remains of extinct elephants are now known to occur on three of the Channel Islands, namely, on San Miguel, Santa Rosa and Santa Cruz (see Fig. 1 [=Fig. 922 of present Memoir]). The first material was found on Santa Rosa more than sixty years ago, and this island has furnished by far the largest collection of fossil specimens representing these types. Similar material has been brought to light on San Miguel. In contrast to the rather numerous finds of elephant remains in Quaternary de- posits of Santa Rosa, the presence of elephants on Santa Cruz is known thus far by only two fragmentary enamel plates of a cheek- tooth.” “San Miguel: Although this island is wind-swept and shifting sand dunes mantle much of the area underlain by sediments of Tertiary and Quaternary age, the incision of the present land sur- face by ravines and gullies and the constant though gradual re- Conclusions (pp. 212—214).—‘‘Whether or not more than one species of elephant is present among the island forms remains to be definitely determined. In this connection, it should be recognized that an interesting and perhaps significant difference may exist between those forms on Santa Rosa and the types of San Miguel.” “Numerous cheek-teeth and tusks, fragmentary jaws and skeletal elements comprise the bulk of the collections obtained on Santa Rosa. Individuals of all ages are preserved, from an unborn type to fully grown adults. The youngest specimen, evidently belonging to a foetus, is represented by a lower jaw (Fig. 6 [not figured in present Memoir]) in which the enamel plates had not firm- ly consolidated to form the lower cheek-teeth and had not erupted through the gums. One fairly complete skull represents an adult in- dividual and furnishes valuable information as to the specific char- acters of the island elephants. When found in Quaternary strata, exposed in the sea-cliff near the mouth of the Canada Corral, only the weathered cranial portion was visible. ... Excavation revealed the rest of the skull and upper tusks . . . with the lower jaw in position below the palate. [This is the type.] Illustrations of this specimen and of a young adult skull of the im- perial mammoth (Archidiskodon imperator), drawn Scale 20 10 30 Submarine depths in fathoms we», Santa Barbara Carpinteria — Ventura anacra SAD . to the same scale, are shown in Figure 9 [not fig- Bae ured in present Memoir].” “Comparison of fossil remains of elephants found on Santa Rosa with comparable materials occurring on the mainland establishes clearly the fact that the island forms were smaller in stature than their relatives of the mainland. Consider- able variation in size exists among the island types, but the difference in stature between island and mainland forms remains a notable feature. . . . While the elephants of the mainland ranged in height from approximately 10! feet to 131% feet as measured at the shoulders, those of the islands presumably never exceeded 8 or 9 feet in height and the smaller individuals were probably no taller than 6 feet. Thus, the smaller size of these elephants presents a character wherein they re- semble the fossil or subfossil, dwarfed elephants described from the Maltese Islands of the Medi- Vig. 922. ‘Map of coastal province of southern California in vicinity of Location of some occurrences of fossil elephants on Channel Islands shown line indicates hypothetical border of land during Pleistocene time, after Chaney and Mason.” “As mentioned before, the elephants of San Reproduced from Stock, 1935.1, p. 206, fig. 1. cession of the sea-cliffs develop exposures on which occasionally the weathered-out materials of fossil mammals have been discovered. Several tusks and cheek-teeth of elephants were found in a thin series of Quaternary alluvial deposits lying beneath a table-like surface and exposed in the sides of gullies near the northwest end of San Miguel. Scattered proboscidean teeth have been found from time to time elsewhere on this island. Among the fossil materials are specimens which clearly point to the fact that the San Miguel elephants are among the largest types to be obtained in the insular region.” = terranean. The diminution in size, however, has < 1 © ] V « SantasBarharal | 2008 been carried so far in the Channel Island by x. Dotted elephants as in the Maltese species.” Miguel are among the largest types to be recorded from the island region. Tusks of these forms have been found which measure 5 feet in length and 6 inches in diameter at the base. While some of the fossil materials on Santa Rosa likewise indicate the former presence of relatively large individuals, it is possible that the average size of the San Miguel elephants was larger than that of the Santa Rosa types. Were this ultimately established to be the case, on the basis of a comparison with more extensive collections than are now avail- able from San Miguel, it is interesting to speculate whether the difference may not have been the result of an earlier extinction of elephants on the smaller of the two islands.” THE MAMMONTINA: ARCHIDISKODON AND METARCHIDISKODON Archidiskodon sonoriensis Osborn, 1929 Figure 923 One mile east of Arizpe, northern Sonora, Mexico. Lower Pleistocene. “c Archidiskodon sonoriensis Osborn, 1929. ‘‘New Eurasiatie and American Proboscideans,’’ Amer. Mus. Novitates, No. 393, Dec. 24, 1929, p. 18. Typr.—‘Nearly complete skeleton,! of which the palate with third superior molar, M*, of both sides, right lower jaw (lacking ascending ramus), with third inferior molar, r.Ms, zn situ, also symphysis, are in the American Museum.”’ Amer. Mus. 22637 (Osborn, 1929.797). HorIzoN AND Locauiry.—‘‘One mile east of Arizpe, northern Sonora, Mexico, on the Sonora River, 60 miles southeast of Cananea and approximate- ly 100 miles north of La Prietas and San José de Pimas. .. . The “AM.22637 > TYPE 1033 Archidiskodon meridionalis nebrascensis Osborn, 1932 Tigures 815, 924, 927, 928, 1239, Pl. xx1 One mile northwest of Angus, Nuckolls County, Nebraska. Lower to Middle Pleistocene. Archidiskodon meridionalis nebrascensis Osborn, 1932. “The ‘Elephas meridionalis’ Stage Arrives in America,’ Proc. Colo. Mus. Nat. Hist., XI, No. 1, Sept. 7, 1932, pp. 1-3 (Osborn, 1932.893). Typr.—skeleton, lacking cranium also tusks, ex- cepting mid-portion of left tusk, with lower jaw in complete state of preservation. Colo. Mus. 1359. Horizon anp Locatiry.— Found ‘‘one mile northwest of Angus, Nuckolls County, Nebras- ka, some fourteen or fifteen years ago.”” Lugn and Schultz (1934.1, Table A) regard this species as of Yarmouth (Upland) age, equiv- Fig. 923. Archidiskodon sonoriensis, anterior portion of type mandible and maxilla showing r. M3, r.M? (Amer. Mus. 22637), one-sixth natural size. Arizpe horizon is regarded by Barnum Brown as Lower Pleistocene (lake deposit).”’ Typr Ficurr.—Op. cit., p. 18, fig. 18. Specific CuHaracrers.—‘‘Mandibular prolonged obliquely downwards, with downturned beak, as seen both in front and side views; length from symphysial groove to tip of rostrum 230 mm., exposed length of M* 246 mm., of M; 346 mm.; depth from third unbroken plate to bottom of jaw 244 mm. A total of 11+2(?) exposed ridge-plates in M?, of 2(?)+11+3 in M3.” rostrum Remainder of skeleton unintentionally destroyed by discoverer.—Editor.] Compare Osborn, 1929.797, p. 18, fig. 18. alent to Lower to Middle Pleistocene. Type FigurE.— Osborn, 1932.893, figs. 1 and 2. CuaractTEers.—‘‘The inferior grinding teeth similar in charac- ter and in ridge formula to the ‘Hlephas meridionalis’ of Durfort, but somewhat broader with much thicker surrounding layer of cement. ... Mandible: (1) A very prominent rostrum. (2) A rel- atively elongate and shallow ramus. (3) Measurements as follows: Length mandibular condyle to symphysis. ... . 943 mm. Depth below M3 of mandibular ramus........ 220 mm. Fig. 924. Type ManpisLe or ARCHIDISKODON MERIDIONALIS NEBRASCENSIS (Coxo. Mus. 1359). Boru FIGURES ONE-FIFTH NATURAL SIZE (Upper) Crown view of mandible with right and left third molars in situ. After Osborn, 1932.898, fig. 1. (Lower) Lateral view of mandible and rostrum, After Osborn, 1932.893, fig. 2. 1034 THE MAMMONTIN: ARCHIDISKODON AND METARCHIDISKODON 1035 Drawn from cast Amer (us 2/895 Length of r.M3, Third Inferior Molar. ....... 289 mm. Width of r.M3, Third Inferior Molar........ 84 mm. ReNSEvoullerid se=-crestsenein serene +134 imMisncovleridge-crestseaeneeen eerie 13 3 Inky anacus 1m 10) GI. 6 heseecaccuungccobed 5% Widestirid re=crestz ncn ce occa ner sek 73 mm. The above ridge-crest formula, together with the dimensions of the third grinders, agree closely with those which prevail among most of the specimens referred to ‘EHlephas meridionalis’ in the British Museum as described and figured by Falconer.” ARCHIDISKODON MERIDIONALIS Ref of Durfort France Y Natural size “Unfortunately, the cranium was the first part of this animal to be exposed and was completely weathered out. Only the ex- tremity of one of the superior incisive tusks remained; this was lost; a mid-portion of the left incisive tusk remains. Fortunately, every other part of the skeleton was preserved in absolutely com- plete condition on one side of the animal or on both sides so that the skeleton is now superbly mounted and becomes a classic in Piscine, ER SOB all its dimensions as follows: Drawn from cast Vig. 925. Referred superior and inferior molars found associated with Vertebral column: Durfort skeleton (Archidiskodon meridionalis) in the Muséum d’ Histoire Naturelle, Paris, after casts kindly furnished the present author by Dr. 7 cervical vertebrae measuring. ......... 543 mm. Marcellin Boule in January, 1930. Both figures one-third natural size. 19 dorsal vertebrae measuring.......... 1640 mm. (Upper) Right second superior molar, r.M? (cast Amer. Mus. 21895), 4 lumbar vertebrae measuring. ......... 400 mm. with +8 worn ridge-plates; 5} ridge-plates in 10 em.; coronal surface length Sacrals not preserved. 162 mm., maximum breadth 81 mm. = (Lower) Summit of crown of left third inferior molar, 1.M3 (cast Amer. Y caudals only preserved. Mus. 21894), with 14% ridge-plates, 6 partly worn; 5 anterior ridge-plates Height dorsal spine to ground (as in 10cm.; maximum length 276 mm., maximum breadth 83 mm. TNOUNGCC) sce sesies ae enaceaes ies eee orev 3695 mm. ARCHIDISKODON MERIDIONALIS Pex of Duryort France Drawn 7rom cast Amer (us 2/8H ARCHIDISKODON p p - MERIDIONALIS Ref oF Duryort France ' 4y Natural size Drawn From cast Amer Mus. 2/89/ = ) 4 Natural seje Fig. 926. Superior and inferior molars of Durfort skeleton (Archidiskodon meridionalis) in the Muséum d’Histoire Naturelle, Paris, after casts kindly furnished the present author by Dr. Marcellin Boule in April, 1930. All figures one-third natural size. Compare with figure 924, showing type mandible of A. meridionalis nebrascensis of Nebraska, with third inferior molars in situ. é (Left) Left and right second and third superior molars (M**), after cast (Amer. Mus. 21891). Ae al Right second and third inferior molars (r.Mo.3), same as opposite figure, and left second and third inferior molars (1.Mo-g), after cast (Amer. Mus. 1). 1036 OSBORN: THE PROBOSCIDEA Fore and Hind Limbs: Height fore-limb scapula to ground....... 3454 mm. Pelvis, length of os innominatum........ 1350 mm. Scapulaenpthvotee.-me cs ere 1020 mm. Pelvis, width of os innominatum........ 1754 mm. FUumendsy lenge unyOlerer ect rer 1220 mm. Hemurylengthvolasreeeeae eee 1390 mm. Winawlengthioterer neva c.cct- oeatciar ent: 910 mm. Tibia; articularnilengthiofere eee ete 840 mm. J RGVo bP) eyelid SW) Coane, Oheaeee Glee omen rooms 980 mm. Pes: astragalus to tip of Mts. III........ 475 mm. Median metacarpus III................ 200 mm. Res: depthiof Mts Meer ere 150 E.” Fig. 927. Restoration of Archidiskodon meridionalis nebrascensis Osborn, 1932, from the complete skeleton and mandible with lower portion of the tusks, lacking only the cranium. One-fiftieth natural size. The grinding teeth of this priceless specimen are in the same stage of evolution as those of the famous ‘Hlephas’ meridionalis of Durfort, France, as established by casts of the Durfort grinding teeth kindly sent to the American Museum by Director Marcellin Boule (see Figs. 925, 926). THE MAMMONTINA: ARCHIDISKODON AND METARCHIDISKODON TABLE XI. MANDIBLE AND THIRD Morars or A. MERIDIONALIS NEBRASCENSIS) Length Cast Amer. Mus. 21891 L. M2 152 R. M2 157 L. M3 R.M? R.Me 140 L.Me 143 R.Ms L.Ms3 Associated molars Cast Amer.Mus. 21894 L. Ms 278 Cast Amer. Mus. 21895 R.M? 157 MEASUREMENTS OF MOLARS OF ARCHIDISKODON MERIDIONALIS OF Durrort (cr. Frias. 925,926, atso Fic. 924, 1037 Breadth Ridge-plates 94 YT 5\4 ridge-plates in 100 mm. 93 Y-7 545 ridge-plates in 100 mm. 85+ 4 partly exposed, heavy cement 82+ 3 exposed 86 13-6 5 ridge-plates in 100 mm. 92 1-6 5 ridge-plates in 100 mm. 90 4 exposed 85 4 only exposed 83 14% 5) ridge-plates in 100 mm. 81 V7 544 ridge-plates in 100 mm. [This species was described by Professor Osborn in 1932, while visiting the Colorado Museum of Natural History at Denver. In his article in the Proceedings of the Colorado Museum (Osborn, 1932.893) he states: “In the years of study which the present writer has devoted to the evolution of the fossil elephants, he became convinced that the ‘EHlephas meridionalis’ of France is the direct ancestor of the ‘Hlephas imperator.’ New and positive evi- dence of the correctness of this theory is now afforded by the dis- covery of the complete skeleton which forms the subject of the present paper. This skeleton with the lower jaw in a complete state of preservation proves to resemble very closely indeed in every detail the ‘Hlephas meridionalis’ of Durfort, France, as fully described by Albert Gaudry.... In August, 1931, it [the skeleton] was brought to the attention of Director Figgins of this Museum [Colorado Museum], who immediately took steps not only to 3721mm.,12/2 %” DURFORT ARCHIDISKODON MERIDIONALIS 1931 S. FRANCE rescue the fragments from careless visitors but to institute the complete excavation by the most skillful modern methods. The result will be most gratifying to palaeontologists all over the world, mainly for two reasons; first, because it is by far the most com- plete and perfectly preserved skeleton of Archidiskodon ever found, lacking only the cranium; second, because it enables us to record the early migration of ‘Hlephas meridionalis’ to North America, and thereby establish a direct ancestral relationship of the Durfort form to the present Nebraska mammoth.” “Fortunately, the writer had recently secured from Director Boule of the Paris Museum a series of casts of the upper and lower grinding teeth of the Durfort specimen [Figs. 925 and 926 above]. Placed side by side with corresponding teeth of the Nebraska specimen, there can be no doubt that the two forms are closely related.”’—Editor.] ( | ARCHIDISKODON MERIDIONALIS NEBRASCENSIS 3672M M12” 1933 S. NEBRASKA Fig. 928. Restoration by Margret Flinsch Buba of Archidiskodon meridionalis of Durfort and A. meridionalis nebrascensis of Nebraska, under the direction of Henry Fairfield Osborn. One-fiftieth natural size. 8801 “HIOWS IL S,NHOdSO uossa4oudg OL ISILay GHL dO NOLLASIYLINO(D) Soouangdo GAHOLSaY “WOTAHG SVHda1dUVgd AHL AO saloddg LSaouv’T aHL ‘HOVASOJY JO IIMAHLNODOUL SVHAATANY F "626 “SI MHL SI DNIMVUC SIBY, “E61 ‘VEO HOSNITY LINDAVY, AA A CHAPTER XVII THE GENUS PARELEPHAS (SUPERFAMILY ELEPHANTOIDEA), OF THE SUBFAMILY MAMMONTIN-, INTERMEDIATE BETWEEN ARCHIDISKODON AND MAMMONTEUS, DISTRIBUTED IN THE NORTH TEMPERATE ZONE OF EURASIA AND NORTH AMERICA PROFOUND CRANIAL AND INCISIVE TUSK RESEMBLANCES TO ARCHIDISKODON AND MAMMONTEUS. CLEAR DIS- TINCTIONS FROM THESE GENERA IN GRINDING TOOTH STRUCTURE. FIRST APPEARANCE IN THE UPPER PLIOCENE! AND LOWER PLEISTOCENE OF EUROPE. MIGRATION THROUGH ASIA MINOR AND ASIA TO THE UNITED STATES— WASHINGTON, NEBRASKA, TEXAS, ILLINOIS, INDIANA, OHIO, SOUTH CAROLINA, GEORGIA, FLORIDA—-AND FRENCH GUIANA, SOUTH AMERICA. EXTINCTION IN UPPER PLEISTOCENE OR POSTGLACIAL(?) TIMES. 1. European north temperate origin. History of separation from 3. North and South American species of Parelephas. other extinct proboscideans. Parelephas jacksoni, Ohio, exact locality unrecorded. Complete separation by Osborn of the phylum Parelephas. Parelephas(?) mississippiensis(?), Indiana. Order of discovery and description of species of Parelephas. Dareeheecatnke s Phylogenetic order of succession of Parelephas. GT EcePIbOS (COUT, COLE: Distinctive cranial characters of Parelephas. Parelephas columbi felicis, Mexico. 2. Systematic description of European and Asiatic species in Parelephas columbi cayennensis, French Guiana, ascending progressive order. South America. Parelephas trogontherioudes, Italy. Parelephas jeffersonii, Indiana. ; Parelephas trogontheriv, Germany. Elephas roosevelti (synonym of P. jeffersonii), Ulinois. Parelephas(?) trogontherit nestii, England. : F 2 Parelephas progressus, Ohio. Parelephas armeniacus, Asia Minor. Deeg ; Parelephas unrecorded in China and Japan. Parelephas washington it, Washington. Parelephas intermedius, France. Parelephas eellsi, Washington. Parelephas wiisti, South Russia. Parelephas floridanus, Florida. 1. EUROPEAN NORTH TEMPERATE ORIGIN. HISTORY OF SEPARATION FROM OTHER EXTINCT PROBOSCIDEANS The generic phylum separated as Parelephas by Osborn in the year 1924 is briefly mentioned in Chapter II of the present Memoir and distinguished especially in its cranial characters in Chapter XV, in which it appears that Parelephas is linked with Archidiskodon and Mammonteus in its cranial resemblances and great incurved incisive tusks, while in its grinding teeth and ridge-plate formule it is so nearly intermediate between these two genera as to have been mistaken for an actual connecting link. In the present chapter it is shown to be an entirely distinct generic phylum which during the more temperate interglacial periods (Fig. 795) occupied the same geo- graphic range as that of the true woolly mammoth (Mammonteus) during the glacial periods. Hisrory.—The sixteen species which are grouped in the genus Parelephas constitute a very ancient and distinct generic phylum for which Osborn’s name (Parelephas) seems appropriate, in reference to the convergence or parallelism of the grinding teeth in this phylum with those of the true Hlephas. For more than half a century, owing to the similarity in appearance of the grinding teeth to those of the true mammoth (Mammonteus), all English and American authors, including Falconer, Leith Adams, and Lydekker, confused the grinding teeth of specimens which Osborn now refers to Parelephas with those of the northern mammoth Elephas [Mammonteus] primigenius, and this accounts for the great discrepancies in the collective ridge formule attributed to the species E. primigenius, i.e., M 3 +®2? as given by Leith Adams (1877-1881, p. 127) and by Lydekker (1886.2, p. 175), and copied by Hay (1914, p. 395). This unfortunate confusion in the ridge formula arose despite the fact that Falconer See footnote 1 on page 1049 below.—Editor.] 1039 1040 OSBORN: THE PROBOSCIDEA as early as 1863 (p. 65) had correctly defined the ridge-plate formula of #. promigenius as M 3 #4, a formula which is fully confirmed by Osborn’s researches for this Memoir (see Chapter XVIII). The lower ridge-plate count (M 3 +) attributed by Adams and Lydekker to Hlephas primigenius really belongs to grinding teeth of primitive Middle Pleistocene species of Parelephas, such as P. trogontherii, which exhibits M 3 73; ridge-plates. As described in detail below, Jourdan (1861) was the first to separate from HE. primigenius one of these species of Parelephas under the name Elephas intermedius, a stage which exhibits M 3 3; ridge-plates; he was followed by Pohlig (1885) who clearly defined the more primitive Elephas trogontherit. RESTORATION OF THE TYPE OF PARELEPHAS JEFFERSONIL One-thirtieth natural size Fig. 930. This painting by Charles R. Knight, in the year 1909, was taken directly from the type skeleton of Parelephas jeffersonii in the American Museum of Natural History. The characters of the typical Parelephas cranium and tusks are particularly well shown, with short concave forehead and prominent convex occipitofrontal crest. The shape of the ears is entirely conjectural. The hairy covering, unlike the hairy and woolly covering of Mammonteus primigenius, is a wholly conjectural character, because no remains of the hair have been discovered, but the presence of this Jeffersonian mammoth in north temperate regions, appearing in post(?)-Wisconsin times, furnishes indirect evidence of a hairy if not of a woolly coat. In the present chapter, chiefly from the researches of Depéret, Mayet, and Osborn, it is shown that the phylum Parelephas constitutes a long line of progressive ascent wholly distinct from that of Mammonteus primigenius. Its first appearance is in Upper Pliocene time in Italy. Its final appearance in IV Guaciau and Postglacial times, principally on or near the 40th parallel of the United States, in the species Parelephas jeffersonii (M 3 2°), is 30 followed by the closing stage Parelephas progressus (M 3 #8). THE MAMMONTINA:: PARELEPHAS 1041 It has also been a long and difficult matter both in Europe and America to clearly separate the members of the generic phylum Parelephas from the newer phylum of the broad-toothed, narrow-plated true mammoth (E. [= Mammonteus] primigenius) on the one hand, and from the older phylum of the broad-toothed, broad-plated southern (Elephas meridionalis) and imperial (#. ¢mperator) mammoths of the genus Archidiskodon on the other. The chief grounds of separation are as follows: (1) The cranium in Elephas intermedius, E. trogontherii, and E. jeffersonii is now known to be readily distinguishable from the crania of either L. [= Mammonteus] primigenius or E. [= Archidiskodon| imperator; (2) the grinding teeth are intermediate in form and in the number of plates, as Sreconp Fyaure or THE TYPE SKELETON OF PARELEPHAS JEFFERSONII One-thirtieth natural size Fig. 931. Second figure of the aged type skeleton of Parelephas jeffersonii Osborn, 1922, p. 11, fig. 10, as mounted in the American Museum (Amer. Mus. 9950). For further information about this type skeleton, see legend of figure 966, also the description below. observed by Jourdan in 1861 in applying the name Elephas intermedius; (3) it is noteworthy that both in France and Germany the grinding teeth of Parelephas have been independently described as intermediate in structure between those of E. [= Archidiskodon] meridionalis and E. [=Mammonteus] primigenius, as shown in the full historic notes below. EHlephas columbi proves to be Parelephas columbi. FRANCE: JOURDAN, 1861.—Despite the early separation by Jourdan of Elephas intermedius from E. primi- genius and description of the grinding teeth as intermediate between HZ. primigenius and E. meridionalis (1861), 1042 OSBORN: THE PROBOSCIDEA Gaudry, who was very conservative in the matter of applying new names to species, continued to describe (1876, p. 40, also Pl. 1x) these animals as “Hlephas primigenius a lames écartées.’’ The steps in the gradual separation of this phylum by paleontologists of France are clearly described by Depéret and Mayet (1923, pp. 176-190). Tig. 932. Chief Lower to Upper Pleistocene localities in which occur species of Archidiskodon, Parelephas, Mammonteus, Loxodonta, and Palzoloxodon (syn. Pilgrimia), after Osborn, 1910.346, p. 391, fig. 176. Fossils attributed to the phylum Parelephas, and of large size, are especially abun- dant in the Middle Pleistocene of Siissenborn (Fig. 932,11), of Mosbach (12), and of Taubach (19). At Mosbach (in 1st? Interglacial times) they occur in the same layers, but with much greater frequency, than those of H. [Hesperoloxodon] antiquus, namely, three to one (Soergel, 1912, p. 32); more recently it is estimated (Schmidtgen, 1926) that in the Mosbach sands HF. [Parelephas| trogontherii is about ten times as abundant as E. |Hesperoloxodon| antiquus; in Mosbach also this elephant attains its greatest size. PLEISTOCENE EUROPE. —1 Forest Bed of Cromer (Norfolk). Sables de 2 St. Prest near Chartres (Eure-et-Loire). 8 Malbattu (Puy-de-Dome). 4 Peyrolles (Bouches- du-Rhone). 5 Solhilac near Puy. Clay deposits of 6 Durfort (Gard). 7% Cajarc (Lot-et-Ga- ronne). 8 Val d'Arno (Tuscany). 9 Leffe near Bergamo (Lombardy). 10 Rizdorf near Pots- dam (Brandenburg). Gravels of 11 Siissenborn near Weimar. Sands of 12 Mosbach in northern Baden. Freshwater deposits of 13 Clacton (Essex). Sands of Mauer near 14 Hei- delberg (western Germany). 15 Chelles on the Marne, near Paris. 16 St. Acheul (Somme). 17 Ilford and Grays Thurrock.(Essex). Lignites of 18 Diirnten and of Utznach, near Ziirich. 19 Taubach near Weimar. 20 Wildkirchli cave on Mont Sdntis (eastern Switzerland). Tufts of 21 the Tiber Valley, near Rome. Caves of 22 Neandertal, near Diisseldorf (western Ger- many), 23 Spy, near Amur (Belgium), 28a Krapina (Croatia), 24 Chapelle-auz-Saints (Cor- réze). Caves and alluvial deposits of 25 Ternifine (or Palikao) near Oran (Algeria), 26 Pointe Pescade, near Algiers (Algeria). 27 Prince's Cave (Monaco). Sandy clays of 28 Véklinshofen (Alsace). 29 Saalfeld (Saxe-Meiningen). Travertines, etc., of 30 Gera, Jena (Saxe-Weimar). 31 Leipzig (Saxony). 32 Solutré, north of Lyons. Loess of 38 Wirzburg (Bavaria). 34 Thiede near Braunschweig (Prussia). Cave of 35 Montmaurin (Haute-Garonne). 36 Chdteauneuf- sur-Charente (Charente). Caves of 37 Schweizersbild near Schaffhausen, and Kesslerloch near Thayngen (northern Switzerland). Remains of lake dwellings at 38 Wauwyl (Lucerne), 39 Ro- benhausen, south of Lake Pfaffikon, 40 Concise on Lake Neuchatel (Switzerland). Peatbogs of 41 Hassleben, near Weimar. Travertines of 42 Langensalza (Erfurt) in central Germany. Caves of the 43 Island of Malta, 44 Island of Crete, 45 Island of Cyprus. [Not on map: 46 San Paolo de Villafranca (Piedmont), Italy. 47 Erzerum, Armenia. 48 Plateau loess, Lyon, France. 49 Tiraspol, S. Russia] Iraty: FaLconrr, 1868.—As early as 1868 (see below under Parelephas armeniacus), Fal- coner entertained a strong suspicion that a form closely related to HL. armeniacus occurred at St. Paolo, Italy, which he identified (1868, II, pp. 249, 250) as resembling HE. armeniacus, but which we now know belonged to Parelephas trogontherioides (see p. 1055). This is a fine ex- ample of Falconer’s unerring sense of form. GERMANY: Pon.iac, 1885.—In 1885 Pohlig, on discovering two grinding teeth of one of these mammoths in the Jnterglacial sands of Siissenborn near Weimar, proposed the specific name EHlephas trogontherti; he maintained that both geologically and zoologically this species was a link, or intermediate, between Elephas primigenius [i.e., Mammonteus|]| and Elephas meridionalis [{i.e., Archidiskodon|. To quote Pohlig’s own language (op. czt., 1885, p. 1027): “Unter der Bezeichnung ‘Elephas trogontherw Pohl.’ fiihre ich in meiner Monographie eine europiische Molarenform auf, welche zwischen denjenigen des EH. primigenius und E. meridi- onalis zoologisch, wie ihrer geologischen Lager- stitte nach, in der Mitte steht.” Pohlig also observed the relationship of 2. trogontheriz to the species which Falconer named Elephas armeniacus in 1857. At the same time Pohlig erred in suggesting the relationship of H. trogontherii to Elephas | Palxoloxodon] namadicus Fale.; he also erred in suggesting that both in craniology and dentition there was a direct phylogenetic or ancestral succession between E. meridionalis, E. trogontherii, and E. primigenius (i.e., ‘“directer Verwandtschaft’’). In 1886, p. 181, Pohlig remarks: ‘Under the name of Elephas trogonthervi, Pohlig, I have described European molars which hold a middle place, both zoologically and geologically, between those of H. primigenius and E. meridionalis, most closely approaching those of #. antiquus in the ridge-formula, but differing more from them than from the other two in the form of the erown. The position of 1. trogontherti with regard to EL. armeniacus, Fale., and F£. namadicus, Fale., still remains to be investigated.” THE MAMMONTINA: PARELEPHAS 1043 FrANcE: DEpg&ReET, 1923.—In 1923, Depéret and Mayet in their invaluable review of “Les Eléphants Pliocénes,”’ Deuxiéme Partie, out of respect for Pohlig’s clear distinction and definition of the species /. trogon- ther, decided not to adopt the term Elephas intermedius of Jourdan, but to name this phylum ‘“‘Rameau de ”) )Elephas trogontherv.”’ They recognized its remote kinship to Hlephas primigenius (see “IIT Groupe des Elephas trogonthervi et LE. primigenius (Mammouths)”’) but established it (p. 176) as a distinct branch of the “groupe des Mammouths.”’ Osborn believes this to be its true phyletic position, resting on two chief characters, namely: (a) The cntermediate character of the grinding teeth, observed by Jourdan, by Gaudry, by Pohlig, and by Depéret and Mayet; (b) the dome-shaped cranium, figured and observed by Pohlig and by Depéret and Mayet. In their analytical treatment of the geologic succession of species, Depéret and Mayet clearly pointed out that the type locality (3d Interglacial plateau loess near Lyons) of Elephas intermedius Jourd. is of more recent geologic age than the type locality (2d Interglacial sands) of Hlephas trogontherii Pohl.; moreover in seeking a Pliocene ancestor of this phylum, they believe they have discovered it in the Upper Pliocene subspecies of Italy, named by Zuffardi in 1913 Elephas antiquus var. trogontherioides. They thereby confirmed Faleoner’s observation of 1868 that an elephant closely similar to HL. armeniacus oceurred in northern Italy (see Parelephas armeniacus descrip- tion, p. 1060 below). Consequently the actual geologic succession of the three types of Parelephas thus far discovered in France and Italy is as follows: Pleistocene (3d Interglacial plateau loess), Lyons, I’'rance Hlephas intermedius Jourdan, type Pleistocene (2d Interglacial sands), Stissenborn near Weimar, Germany Elephas trogontherzi Pohlig, type Upper Pliocene (Villafranchian), Italy [see footnote, p. 1049 below.—Ed.] Elephas antiquus var. trogontherioides Zuffardi, type AMERICA: OSBORN, 1922.—Among the abundant remains of members of this phylum in America, a precisely similar confusion between specimens belonging to Archidiskodon, Parelephas, and Mammonteus arose in the minds of American paleontologists (Cope, Osborn, and Hay). Cope (1889.2, pp. 208, 209) referred the remains of a very fine skull of Archidiskodon (Fig. 891) from Texas to “Elephas primigenius columbi Fale.” Osborn also saw only resemblances to ‘‘Elephas columbi”’ in the fine skeleton found in Indiana (type of Parelephas jeffersonii Osborn, Figs. 961, 931); whereas Hay (1914) referred the same skeleton to Hlephas primigenius. No stronger proof could be given of the truly intermediate character of members of this phylum in America than this alternate reference of the same skeleton by Osborn to Elephas columbi and by Hay to Hlephas primigenius. In Hay’s exhaustive summary (1914) of grinding teeth and other remains from various parts of the United States (Alaska to Florida) which he referred to ‘‘Elephas columbi,” he includes teeth which certainly belong to Parelephas columbi as well as to the truly intermediate form (P. jeffersoniz). Osborn (1922.555) was the first to separate these intermediate animals, previously described by Cope, Hay, and himself as “H. columbi”’ and ‘‘E. primigenius,”’ under the new specific name of Elephas jeffersonii, a species which he subsequently (1924.633, p.4) made the genotype of Parelephas. This separation was based principally upon cranial characters (Osborn, 1922.555, p. 15): “Still more obvious are the differences between the relatively long, broad, and shallow crania of F. jeffersonzi and the relatively short, narrow, and deep crania of EL. primigenius, proportions which are correlated respectively with the corresponding proportions just described and figured in the teeth.” PHYLOGENETIC RELATIONSHIPS (POHLIG, SORGEL).—Inasmuch as the same species of animal had previously been erroneously described by Cope and Osborn under the name of “‘Elephas columbi,” we must credit Soergel (1921, p. 60) with the prior observation that this supposed “Hlephas columbi”’ of America (= Elephas |Parelephas} jeffersonit Osborn) shows phyletic relationships to the Hlephas [= Parelephas| trogontherii of Pohlig. 1044 OSBORN: THE PROBOSCIDEA Schmidtgen and Freudenberg (1926, pp. 62, 68) describe and discuss EHlephas [Parelephas] trogontherii of Mosbach, especially the supposed phylum (“Stammreihe’’) Elephas meridionalis—E. trogontherii—E. primi- genius, which in unbroken succession lived in the region of Wiesbaden. Osborn, 1924-1928: It has remained, however, for Osborn in the present Memoir to institute a close compari- son between all the known characters of the Hlephas trogontherti phylum of Europe and the Hlephas jeffersonii of America and to establish the fact that there is a close phyletic relationship which justifies the linking of the European and American species in the new and distinct genus Parelephas. This affinity is most clearly indicated in the cranium, as shown in the resemblance of several crania, erroneously figured as “Elephas primigenius”’ by Falconer, to those of EF. trogonthervi as figured by Pohlig, and to the cranium of E. jeffersonii as figured in the present Memoir; there can be no doubt that this relatively broad, elongate, and rounded cranium is entirely distinct from the extremely short, compressed, and peaked cranium of the true mammoth Elephas primigenius. HABITAT OF PARELEPHAS TROGONTHERII (SOERGEL, 1912).—In his masterly monograph of 1912, entitled “Elephas trogontherii Pohl. und Elephas antiquus Falc.,’”’ Soergel observes (pp. 105, 106) that the Steppe [plains] and Wald |forest] faunas lived contemporaneously in central Europe; as observed in the /st [nterglacial period, they are found in the lower sands of Mosbach, the ‘“‘kiese”’ [gravels] of Mauer, the sands of Petersdorf bei Gleiwitz in Schlesien, the ‘“Tone”’ [clays] and “‘grauen Rheinsande”’ [gray sands of the Rhine] near Jockgrim in Pfalz, and the gravels of Siissenborn (which persist into II GLacra time).’ He divides the plains or ‘‘Steppe” and forest or “Wald” faunas of the /st Interglacial period as follows: CHIEFLY INLAND STEPPES OR PLAINS CurerLty OcrAN Borpurs AND Forusts E. (Parelephas] trogontherti Pohl. E. |Hesperoloxodon| antiquus Fale. Very abundant in Mosbach; absent in Mauer. Very abundant in Mauer. Dicerorhinus etruscus Fale. Dicerorhinus etruscus Fale. Equus stenonis Cocchi, E. siissenbornensis Wiist, Equus stenonis Cocchi, EL. mosbachensis v. Reich. E. mosbachensis v. Reich. Leptobos etruscus Fale. Bison priscus Bo}. Bison priscus Bo}. Cervus elaphus Linn., C. elaphus trogontherti Pohl., Cervus elaphus antiqui Pohl., C. capreolus Linn., C. capreolus Linn., Alces latifrons Johns. Alces latifrons Johns. Ursus arvernensis Croiz. and Job., U. Deningeri v. Ursus arvernensis Croiz. and Job., U. Deningeri v. Reich. Reich. Felis leo fossilis Goldf. Felis leo fossilis Goldf. Hyena arvernensis Croiz. and Job. Canis neschersensis Croiz. and Job. Canis neschersensis Croiz. and Job. [Homo heidelbergensis Schoet. (lower sands of Mauer)]. Certainly Hesperoloxodon antiquus, a forest loving form with coarser teeth, occurred nearer the ocean borders, whereas Parelephas trogontherii, with finer teeth, frequented both the steppes (plains) and forests. Since animals of both habitats are buried together in river sands, gravels, and clays, there is an intermingling of plains and forest faunas, as shown in Mosbach and Mauer (Osborn). Of the habitat and geographic distribution, Soergel (op. cit., p. 110) observes that whereas Hesperoloxodon antiquus preferred a warmer ocean bordering climate, not under the direct influence of the northern inland ice masses, Parelephas trogontherti sought the cooler northern-northeastern continental localities. Accordingly in Italy, Spain, and Greece it is almost entirely wanting; in France P. trogontheri? is less abundant than H. antiquus, at least, it appears in less characteristic forms. The homeland of the P. trogontherii type appears to be confined to England and Germany, perhaps also to Russia. During the arid 1st Interglacial peroid we find P. trogonthertt distributed from southern England over middle Germany and eastward to southern Russia, as shown in the Forest Bed of Cromer, the sands of Mosbach, the gravels of Siissenborn, the sands of Petersdorf in Schlesien, and the clays of Jockgrim in Pfalz. Already in this /st Interglacial period migration towards the east may have begun. See p. 1056 below, where Parelephas trogontherii type is placed by Professor Osborn in the 2d Interglacial.—Editor. | THE MAMMONTINA: PARELEPHAS 1045 In the 2nd Interglacial period we observe practically the same geographic distribution of Parelephas tro- gontherii; its southernmost appearance in Germany at this time is at Steinheim on the river Murr; its northern- most appearance is at Rixdorf near Berlin. In Thuringia it is found in the Ilm gravels below the older travertines of Taubach, also in the stream gravels at Vieselbach near Erfurt, ete. For the better understanding of the geologic as well as the geographic distribution of the Parelephas trogon- therii phylum in Germany may be cited the following geologic correlation by Soergel (op. cit., p. 106): Dem I. ‘Interglazial’ gehoren an: Dem II. ‘Interglazial’ gehoren an: Dem III. Interglazial gehoren an: die Sande von Mosbach [n. die Bachkiese bei Vieselbach, der Travertin von Taubach- Baden], die Saalekiese von Uichteritz Ehringsdorf-Weimar, die Kiese von Mauer|b. Heidel- b. Weissenfels, der Travertin von Burg-Grae- berg], die IImkiese unter dem iilteren fentonna, die Sande von Petersdorf b. Travertin von Taubach- der Travertin von Bilzingsleben und verschiedene andere Trav- ertinvorkommen Thiringens. Gleiwitz in Schlesien, Ehringsdorf [b. Weimar], die Tone und grauen Rhein- die Schotter von Steinheim sande b. Jockgrim i. d. (II. Glazial-Interglazial). Pfalz, die Kiese von Siissenborn [bei Weimar], die allerdings bis in die II. Eiszeit hin- eingehen. [Plains (‘Steppe’) and forest die Sande von Rixdorf b. Berlin fiir eine Ablagerung des II. ‘Interglazial.’ [Plains and forest faunas not clearly [Plains and forest faunas again appear (‘“‘Wald’’) faunas, as listed above.] The Siissenborn deposits extend into II Gracia time (fide Soergel, 1912.) separated; Parelephas trogontherii and Hesperolorodon antiquus oecur in the same horizon.| partly separated, as in Mauer and Mosbach. Hesperoloxodonantiquus survives Par- elephas trogontherii, which disappears. In the tundra fauna appears Mam- monteus primigenius.] COMPLETE SEPARATION BY OSBORN OF THE GENERIC PHYLUM PARELEPHAS Osborn finally (1924.633, p. 2) concluded to cut the Gordian knot and terminate this confusion by distinguish- ing the generic phylum Parelephas throughout from both Mammonteus and Archidiskodon, as shown in the follow- ing citation: {p. 2} Much more difficult has been the separation of the third generic series of the Mammontine, which hitherto has been referred to the genus Elephas but which we now remove to the new generic phylum Parelephas. [p. 4] The eight or ten species included within this genus constitute a very ancient and wholly distinct generic phylum, for which the name Parelephas seems appropriate, because in certain characters these animals parallel the true Hlephas, although in profound cranial and dental structure they are closely related to and convergent with the mammoths Archidiskodon and Mammonteus. GENERIC CHARACTERS.—A phylum of the Mammontine. Llephas jeffersonii, genotypic species, LH. armeniacus, L. intermedius, E. trogontherii, FE. trogontherioides. Cranium intermediate in form between that of Archidiskodon and that, of Mammonteus, namely, brachycephalic, acrocephalic. Frontals concave, occipital crest elevated; occiput more or less convex. Molars in the Upper Pliocene and Lower Pleistocene stages with relatively few ridge-plates, i.e., M 3 +31; progressive Upper Pleistocene stages with multiple ridge-plates, i.e., M338. Ridge-plates compressed to 7-8-9 in 100 mm. Molar crowns broad, M? short, with enamel of intermediate thickness, more or less crimped or sinuous. The progressive ridge formule in Parelephas are distinct throughout from those of Mammonteus, and the final ridge formule in the two generic phyla are different, namely: Mammonteus primigenius compressus, M 3 $+= final stage. Parelephas jeffersonii progressus, M 3 $¢=final stage. The crania of Parelephas throughout are readily distinguishable both in frontal and lateral aspects, and especially in vertical section, from those of Mammonteus, as can be seen in all of Falconer’s beautiful plates of 2. primigenius and in Pohlig’s excellent figures of P. trogontheriz. The jaws of Parelephas and of Mammonteus are less readily distinguishable, but by more profound study they can also be separated from those of Mammonteus. The contrasts in the crania of the two genera may be summed up as follows: Mammonteus.—Cranium and jaws extremely compressed fore-and-aft (cyrtocephalic); extremely elevated and pointed above (hypsicephalic); extremely depressed and foreshortened below (bathycephalic). Parelephas.—Cranium moderately compressed fore-and-aft (eyrtocephalic); moderately elevated occipitofrontal borders (acrocephalic); moderately depressed molar-grinding area (bathycephalic). Thus, while the intermediate forms of crania and teeth of Parelephas and of Mammonteus may prove difficult to separate, the two finally progressive forms are readily separable, namely, Parelephas jeffersonii progressus and Mammonteus primigenius compressus. 1046 OSBORN: THE PROBOSCIDEA To this phylum certainly belongs the Elephas armeniacus of northern Asia Minor, as first observed by Fal- coner, as well as many specimens to the eastward described under other names. The connection of this phylum with the Parelephas jeffersonii of late Glacial times in the United States seems highly probable; the resemblance both of the teeth and cranium has been repeatedly observed between LI. trogontherii and the northerly type form- erly known as “Elephas columbi,”’ now known as Parelephas jeffersona. The southerly type (Georgia, South Carolina, Florida) of Hlephas [=Parelephas| columbi, with rather primitive ridge formula (M 3 }2,), has recently been reinforced by the large and more progressive Parelephas floridanus (M 3 22+), apparently a late Pleistocene stage distinguished by its larger size, more robust tusks, and coarser ridge-plates from the northerly (Indiana) Parelephas jeffersona type. A branch of the Parelephas columbi stage appears to have migrated to South America, as mentioned by Lull 1908, p. 204) and Freudenberg (1922, p. 159) on the authority of Lartet (1859, pp. 500, 505). Larrer, 1859, pe. 500! anp 505.—L’existence des Eléphants n’est encore indiquée dans Il’ Amérique méridionale que par un fragment de molaire A lames épaisses, rapporté de Cayenne par le capitaine Perret et donné par lui au Musée de Marseille. . . . Dans l’Amérique du Sud, deux formes spécifiques du genre Mastodonte se montrent dans des dépdts post-pliocénes; mais peut- étre se sont-elles aussi retrouvées dans des formations plus anciennes et rapportables a l’Age précédent ou pliocéne. Quant au type Eléphant, iln’y est encore indiqué que par le seul fragment déja cité d’une molaire 4 lames épaisses, rapporté de Cayenne par le capitaine Perret. FREUDENBERG, 1922, pp. 159, 160.—Wir haben Lartets Autoritit fiir den Fund eines Zahnes von Hlephas im unteren Pleistociin von Cayenne (Franzésisch-Guayana). Nach der Beschreibung der dickriickigen Schmelzplatten ist das Exemplar offenbar von El. imperator. Wahrscheinlich vollzog sich eine Wanderung nach dem Siiden, bevor die Bedingungen geschaffen waren, welche spiiter die Wanderung der Proboscidier nach Siidamerika erméglichten. . . . Es ist das einzige mir bekannt gewordene Beispiel eines wahren Elefanten im Siiden des Hochlands von Mexiko. Osborn, 1929: Through the extreme courtesy of Mm. Laurent and Repelin of the Faculté des Sciences de Marseille, this priceless molar fragment from French Guiana has been located in the Muséum d’ Histoire Naturelle de Marseille, photographs taken, and casts made, especially for the purpose of specific and generic identification for the present Memoir. As shown in figure 957, on a one-half scale, this specimen consists of three ridge-plates of coarse enamel, as described by Lartet, remotely resembling in crown and side views the posterior ridge-plates of a superior grinding tooth of Parelephas columbi (Fig. 952). In 1929 the present writer (Osborn, 1929.797, p. 20) made this the type of a new subspecies, namely, Parelephas columbi cayennensis. The recently discovered Parelephas floridanus includes parts of three full skeletons collected by the Holmes Florida Expedition of the year 1929. In this stage, intermediate in ridge-plate formula (M 3 33+) between Hlephas [Parelephas| columbi (M 3 *2,) and P. jeffersonii (M 3 3%), the ridge-plates are broad at the base and compressed at the summit; the incisive tusks extremely massive and relatively short. The males attain very large size. The femora measure, according to age, 1230 mm. to 1393 mm., as compared with 1250 mm. in the adult Indiana type of P. jeffersonii. SUMMARY OF MIGRATION The restudy of long-known specimens like Hlephas jacksoni (1838), Elephas columbi (1857), Hlephas armen- iacus (1857), and Elephas intermedius (1861), together with the recent recognition of Parelephas columbi cayen- nensis in French Guiana, builds up a Parelephas phylogeny and migration second only to that of Archidiskodon (Chapter XVI). We await the discovery (probably in northern Africa) of an ancestral stage more primitive than Parelephas trogontherioides of the Upper Pliocene? of Italy. 'Dr. Richard 8. Lull (letter, Sept. 24, 1928) writes: “From this description, mentioning the fragment of a molar with ‘thick plates,’ I drew my own conclusion as to the genus and species of that elephant. It seemed to me that that it would be Hlephas [= Archidiskodon| imperator.” *{See footnote 1 on page 1049 below.—Editor.} THE MAMMONTINA: PARELEPHAS Tig. 933. Geographic distribution (according to the numbers in the accompanying list) of species of Parelephas. 1047 Ly fro 40 - PARELEPHAS The white dots within the black areas represent the approximate localities where the types of these species were discovered. The white crosses represent referred specimens. It will be noted that Professor Osborn regarded Elephas texrianus Owen, Blake (1859-1861) as a synonym of Parelephas columbi, and Elephas roosevelti Hay (1922) as a synonym of Parelephas jeffersonii. 1838 1857-1868 1857 1859-1861 1861 1872 1872 1885 1891 1910 1913 1922 1922 1922 1923 1924 [1924 [1926 1926 1929 1929 16. Wife 18. ORDER OF DISCOVERY AND DESCRIPTION OF SPECIES OF PARELEPHAS . Jackson County, Ohio . Brunswick Canal, Georgia . Armenia, Asia Minor . San Felipe de Austen on Brazos River, Texas . France, 3d Interglacial loess . Indiana Indiana . Sussenborn near Weimar, Germany, 2d Interglacial sands Cromer Forest Bed, East Anglia, England Tiraspol, southern Russia . Piedmont (Villafranchian), Italy . Jonesboro, Grant Co., Indiana . Mexico, Tecamachalco, Puebla . Ashland, Cass Co., Illinois . Pine Creek, Whitman Co., Washington . Zanesville, Ohio Province of Kazusa, Japan Province of Kazusa, Japan Port Williams, Clallam Co., Washington Florida, near Bradenton, Manatee County Cayenne, French Guiana, South America See Figure 933 ORIGINAL NAME Elephas jacksoni Mather E. (Euelephas) Columbi Falconer Elephas armeniacus Falconer Elephas texianus Owen, 1859, Blake, 1861 Elephas intermedius Jourdan Elephas Indianapolis Foster. Later in same year changed to Elephas missis- sippiensis—see below Elephas mississippiensis Foster Elephas trogontherii Pohlig Elephas (antiquus) Nestii Pohlig Elephas Wiisti Pavlow Elephas antiquus var. trogontherioides Zuffardi Elephas jeffersonii Osborn El. Columbi var. Felicis Freudenberg Elephas roosevelti Hay Elephas washingtonii Osborn Parelephas jeffersonii progressus Osborn Euelephas (Parelephas) protomammon- teus Matsumoto Parelephas protomammonteus proximus Matsumoto Elephas eellsi Hay Parelephas floridanus Osborn Parelephas columbi cayennensis Osborn Speciric REFERENCE IN PRESENT Memoir = Parelephas jacksoni = Parelephas colwmbi = Parelephas armeniacus = Parelephas columbi = Parelephas intermedius = Nomen nudum = Indeterminate, possibly Parele- phas(?) mississippiensis(?) = Parelephas trogontherit =Parelephas(?) trogontherti nestii = Parelephas wiisti = Parelephas trogontherioides = Parelephas jeffersonii =Parelephas columbi felicis = Parelephas jeffersonii = Parelephas washingtonir = Parelephas progressus =Palzxolorodon protomammonteus—see Chap. XIX] = Palzxolorodon protomammonteus prox- imus—see Chap. XIX] = Parelephas(?) eellsr = Parelephas floridanus = Parelephas columbi cayennensis 1048 OSBORN: THE PROBOSCIDEA SuPERFAMILY: ELEPHANTOIDEA Osborn, 1921 FAMILY: ELEPHANTIDZ Gray, 1821 SuBFAMILY: MAmMontiIN«a Osborn, 1921 Genus: PARELEPHAS Osborn, 1924 Original reference: Osborn, Amer. Mus. Novitates, No. 152, pp. 4 and 5 (Osborn, 1924.633). GENERIC CHARACTERS (CF.OSBORN, 1924.633, p. 4).—Genotypic species, Hlephas jeffersonit Osborn, of Indiana; species referred to this genus, Elephas armeniacus, E. intermedius, E. trogontherti, and E. trogontherioides. Cranium intermediate in form between that of Archidiskodon and that of Mammonteus, namely, bathycephalic, acrocephalic. convex. en Md cece 14+) Miss of intermediate thickness, more or less crimped or sinuous. environment, grazing and browsing. APPROXIMATE PHYLOGENETIC ORDER OF SUCCESSION OF PARELEPHAS RipGe ForMULA Frontals concave, occipital crest elevated; occiput more or less Molars in the Upper Pliocene to Middle Pleistocene stages with relatively few ridge-plates, progressive Upper Pleistocene stages (P. progressus) with multiple ridge-plates, i.e., Ridge-plates compressed to 6-6.5-7-8-9 in 100 mm. Molar crowns broad, M® short, with enamel Adapted to continental plains or steppe LenetH BreaptH HricHtT Upper Pleistocene Ohio Parelephas progressus Osborn Type: M3 ae a = a Pleistocene Ohio Parelephas jacksoni Mather Type: M3 (Imperfectly known) M 2 ave Upper Pleistocene Indiana Parelephas jeffersonit Osborn Type: M3 aa ae 2 1 68 Pleistocene Illinois Elephas roosevelti Hay = syn. a 289 98 202 Parelephas jeffersoni Type: M3 aa 3040 92 154 Upper(?) Pleistocene French Guiana Parelephas columbi cayennensis So. America Osborn Type: M 3 superior—fragmentary Upper(?) Pleistocene Vlorida Parelephas floridanus Osborn Type: M3 ane ee 238 ; ie cl : 2 290 79 180 Upper(?) Pleistocene Georgia Parelephas columbi Faleoner ee i TESREEE one ee ee Pleistocene Washington Parelephas washingtonii Osborn Hoe wile , : : ; Type: M3 e348 88 THEORETIC PERIOD OF MIGRATION TO NORTH AMERICA Pleistocene France Parelephas intermedius Jourdan Ref.?: M3 0-32 §=6270 92 150 3d Interglacial Upper Pleistocene S. Russia Parelephas wiisti Pavlow Cotypes: M3 30 340e 100e 198 Pleistocene Asia Minor Parelephas armeniacus Falconer Type: M3 se) Ah9) 105 181 Mi sk iihg en =A ; ee - Peete P 15+ 220 101 138 Middle Pleistocene Germany Parelephas trogontherii Pohlig Type: M3 GEE 270+ 75 100 2d Interglacial (Siissenborn) : |Abundant, attaining greatest size (Mosbach) | Lower Pleistocene Germany Parelephas trogontherti Ref; M3 Not available 1st Interglacial (Basal Mosbach) (“kleinere Var.’’) 'reudenberg oweE isis : ae ere Sy: a 5 ‘ ; 18 302 80 167 ower Pleistocene Mngland Parelephas(?) trogontherii: nestit Cotypes: M3 AF 9434 83 103 1st Interglacial Forest Bed s : (Cromer) Upper Pliocene Italy Parelephas trogonthertoides Lectotype: M3 M18 210+ #110 150 Villafranchian! (type) Zuffardi Cotype: Sie eB 148 188 English Crag (ref.) '[See footnote 1 on opposite page.—Editor.] THE MAMMONTINA: PARELEPHAS 1049 Only an approximate phylogenetic order of succession may be given at the present time. In this phylogenetic list we include with some doubt the imperfectly known Elephas jacksoni Mather, also the recently described Elephas washingtonti Osborn. In phylogenetic order it would seem that this Parelephas phylum first appears in Elephas trogontherioides of the Upper Pliocene (Villafranchian)! of Italy. Doubtless there were many intermediate forms or ascending mutations between this Pliocene stage and the Mid-Pleistocene Elephas trogonthervi of Pohlig at the close of the 2d Interglacial period when this animal was quite abundant. The type of E. trogontherii of Germany is in fact in about the same stage of evolution of the third upper molar as the type molar of the EL. armeniacus of Asia Minor. The phylum is next represented in the 3rd Interglacial in the plateau loess of southern France in the form of Elephas intermedius of Jourdan. Depéret and Mayet consider the type of Elephas intermedius as more progressive than the type of H. trogontheriz. Pohlig also states that a form related to E. trogontherii survived into 3rd Inter- glacial times in Germany and was then replaced by Elephas primigenius; whereas Depéret and Mayet consider that the Hlephas trogontherti-intermedius phylum of Europe became extinct before 3d Interglacial time. This 3d Interglacial period may possibly mark (Fig. 795) its time of migration across Asia into North America.” Priority OF DEPERET IN SEPARATING THE E, TROGONTHERII PHyLUM.—To Charles Depéret and his colleague Lucien Mayet belongs the credit of clearly distinguishing the origin and geographic distribution of the “Rameau de l’Hlephas trogontherwt,” to which phylum Osborn has assigned the generic name Parelephas. Depéret and Mayet (1923, pp. 176-183) first distinguished the ‘A. Rameau de |’Elephas trogontherii Pohlig’”’ from “B. Rameau de VE. primigenius Blumenbach,” both of which they included within the “Groupe des Elephas trogontherii et E. primigenius (Mammouths).” They credit Jourdan with the recognition of E. intermedius as distinguishable from E. primigenius, followed by Gaudry (1876) and by Leith Adams (1877) who traced this form back to the Forest Bed of England, then by Pohlig (1889 [1888]) who described EF. trogontherii of Siissenborn near Weimar, but which Depéret and Mayet remark is absolutely identical with the #. intermedius of Jourdan. Finally they trace the “Rameau de I’ Elephas trogontheriz”’ down into the Pliocéne supérieur (Villafranchian of Italy), confirming a note by Pohlig (1889 [1888], p. 208) as to the tooth subsequently made by Zuffardi (1913, p. 174, Tav. x11 [v1], figs. 3a, 3b) the type of Hlephas primigenius var. trogontherii. Depéret and Mayet observe (p. 177) that the molars of this E. trogontherti var. are readily distinguished from those of the contemporary EF. [Archidiskodon] meridionalis Nesti and FE. [Hesperoloxodon] ausonius Major-Depéret. Zuffardi also described and figured another molar under the name of Hlephas antiquus var. trogontherioides (Zuffardi, Tav. rx [mu], fig. 6), and two other molars of the E. trogonthervi form (PI. rx [1], figs. 4 and 5a, 5b). They state (p. 179) that these four molars of E. trogontherii described from the “Villafranchien de l’Astésan” are the only ones thus far observed in Italian Museums, although there are probably others, and that they are clearly distinguishable from ‘H. meridionalis’ and ‘EF. antiquus,’ concluding (p. 180) as follows: On peut done dire seulement que la forme pliocéne de I’Z. trogontherii posséde des molaires de dimensions plutot un peu réduites, sans qu’il puisse étre question d’une véritable mutation de petite taille de V’espéce. On doit s’attendre 4 découvrir un jour dans le Pliocéne ancien de quelque région éloignée les types ancestraux nains du rameau de IZ. trogontherii. [There is a tendency among many geologists (see Chap. XXII of this Memoir by Dr. Edwin H. Colbert, where the views of Haug, Hopwood, Pinkley, et al are cited) to regard the Villafranchian as of Lowest Pleistocene (or Quaternary) rather than of Upper Pliocene age. Apparently one of the most reliable factors in the determination of the boundaries between the Pliocene and the Pleistocene, aside from the geological evidence, is a change in fauna, for example, the presence of Bos (including Bison), Elephas, or Equus, would indicate the Pleistocene age of the deposit in which they were found. Compare also Haug, 1911.1, pp. 1761 and 1767, Hopwood, 1935.2, pp. 46 and 47.—Kditor.] *ISee page 1071 below where Professor Osborn expresses the opinion that migration to America may have taken place during 2d Interglacial, and possibly as early as 1st interglacial time.—Hditor.} 1050 OSBORN: THE PROBOSCIDEA They also recognize (p. 180) the ““Rameau de |’Hlephas trogontheriv”’ in the Lower Pleistocene Forest Bed of Cromer, but find no trace of it in the Pliocéne supérieur Crags of England. From the Forest Bed they trace (p. 181) the phylum HL. trogontherii, with gradual increase in size, through the terraces of 30 meters (Chellean or Tyrrhenian stage) in the valley of the Thames, also at Gray’s and at Ilford where Lydekker (1886.2, pp. 127, 134) mistook F. trogontherii for FE. antiquus. In Germany observations fail to indicate the Phocene age of E. trogontherii, but this species appears in a Ist Interglacial stage corresponding to the Forest Bed (Sicilien, Lower Pleistocene), also according to Wiist (1901, p. 61) in the ancient gravels of the Ilm at Siissenborn near Weimar, where it was very abundant. This level they regard as contemporaneous with the 1st Interglacial sands of Mosbach near Wiesbaden and of Mauer near Heidel- berg (Chellean). They also state (p. 182) that Pohlig records F. trogontherii from Siissenborn, Heinthurm and Denstedt, near Weimar; Brucksdorf near Halle; Sulza, Angelhausen, near Armstadt; Apolda, Mosbach, near Wiesbaden; the lower gravels of Taubach, and of Rixdorf near Berlin; the gravels of the terrace of 30 meters at Mauer near Heidelberg, and the travertines of Taubach and of Weimar. It became extinet, without extending higher, in the Quaternary. Osborn, 1929: Observe the constantly ascending ridge formula in M 3 +4+ (in the Upper Pliocene) to M 3 # (in the Upper Pleistocene), a period of perhaps 1,250,000 years. Observe also that during 3d Interglacial times Parelephas disappears in western Europe and in early Pleistocene times it appears in North America. SKULLS OF MAMMOTHS All fiqures one-twentieth natural size } } 7 fi fase kes)? / \ ‘ 1 y / y | Powe UCU ‘ / | E. PRIMIGENIUS Ref. er 2 = Cy) Falc., 1847, Pl. XLV, Fig. XXIV (rev.? E. PRIMIGENI ar NTHERTI f rr ree ee ee Ae a ‘ay E. PRIMIGENIUS Ret. Pohlig, 1891, p, 384, Fig. 120 (rev) __ Ponlig, 191. r) "386, Fig. 121 £. PA!IMIGENIUS Ref —S Pontig, 1891. p 384, Fig. 120 E. PRIMIGENIUS Ret Falc,, 1847, Pi. XLII, Fig, XXIV ve PRIMIGENIUS TROGONTAE AI vena Pohlig, 1891, p. 386, Fig t2t Fig. 984. Cranrau Prorites or MAMMONTEUS PRIMIGENTIUS (1,2,5) AND PARELEPHAS TROGONTHERI (3,4,6) OF EuROPE One-twentieth natural size. The plate lettering is incorrect (compare Fig. 865, upper row, and key, p. 977) 3. Parelephas trogontherii (female); front view of cranium. After Pohlig, 1891. Parelephas trogontherti (female); side view of same cranium. After Pohlig, 1891. 6. Parelephas trogontherii; adult male cranium in side view. After Falconer, 1847. 5. Mammonteus primigenius; adult cranium. After Pohlig, 1891. 1. Mammonteus primigenius; front view of cranium (after Pohlig, 1891), which resembles Parelephas in certain characters. 2. Mammonteus primigenius, front view of cranium (after Falconer, 1847), which appears to be typical of ‘H. primigenius.’ ao > Observe that the Parelephas trogontherii cranium, both in the male (6) and in the female (3, 4), is relatively lower, broader, and less compressed fore and aft than the Mammonteus primigenius male (5); in other words, it is less hypsicephalic and less bathycephalic, with broader space between condyle and orbit. Compare especially (5) M. primigenius and (6) P. trogontherii, both males. THE MAMMONTINA: PARELEPHAS 1051 DIsTINCTIVE CRANIAL CHARACTERS OF PARELEPHAS Up to the time that Pohlig separated the grinding teeth of Parelephas trogontherwi from the grinding teeth of Mammonteus primigenius the crania in these two distinct lines of generic descent were confused, as well as the grinding teeth, yet, as shown in figures 865, 934, 935, and 937, the crania of Parelephas are readily distinguishable from those of Mammonteus. The principal characters by which we distinguish the Parelephas crania are the following: (1) In frontal aspect the crania of Parelephas are relatively broader, more spreading, and more brachycephalic than those of Mammonteus, which are deeper and more bathycephalic; (2) in lateral aspect (a) the orbit is more widely separated from the occipital condyle, (b) the occiput is much more convex, thus throwing the occipitoparietal apex farther forward, (c) the height from the occipital apex to the superior molar crowns is less deep, i.e., less bathycephalic, (d) the apex formed at the summit of the cranium is less acute, (e) the facial front is shorter and more deeply concave, (f) the maxillo-premaxillary sockets are less vertical and the tusks emerge in a less vertical plane; (3) in frontal aspect (a) the premaxillary sockets are more expanded at the extremities, whereas in Mammonteus they are more elongate and more closely compressed, (b) the transverse diameter of the frontals is relatively broader than in Mammonteus, (c) the anterior nares are proportionately broader transversely and less deepened vertically ; (4) in brief, the proportions of the cranium of Parelephas throughout are harmonious with those of the grinding teeth, i.e., less compressed anteroposteriorly, less bathycephalie and less hypsicephalic than those of Mammonteus , Profiles (Fig. 935) of juvenile, adult, and middle-aged crania of Parelephas reveal a contour which is readily distinguishable from that of the larger Archidiskodon cranium, on the one hand, and that of the smaller and much more compressed Mammonteus cranium, on the other; but it is not at all surprising that Falconer confused the Parelephas and Mammonteus crania, because they present so many points of subfamily (1.e., Mammontine) ort, 15 16 M3 .} 5 16, M3 nou you rh . now M3 y tl R ai ~anentll u NN 8 A) u ly INS) A) au8 7 Sk : 3 e x 28 Ewe z kK a Sg 3 RR = Sk =o oSaiss w Ow 298 a SS SES QBo®& Zz / ZX SEEN ry ace es 5 = iS ZN S BS 8 B98 v S) oe a 9 Q NS Be ZR 2 eS ES rhs gx roy EXL urgs cay wo oY 3 as ra mS Seow 28 Ee aS a8 3% we us fare ats az a 3X aon ane ant ane = Ww SOS ES — —S SS Ort PS) = Lerr LaTerau PRoriLEs OF SEVEN SPECIES OF PARELEPHAS WITH PROGRESSIVE Ripar l'orMULa One-thirtieth natural size Fig. 935. This series is remarkably uniform in its progressive characters: 1) M34¢ to M 3 34; superior ridge-plates exceeding the inferior ridge-plates in number. 2) The orbits lie just below the level of the occipital condyles, with relatively wide separation. 7 3) The occipitofrontal profile is restored in P. intermedius (C), P. columbi (E), and P. floridanus (I), but is perfectly preserved in P. trogontherti (A, B), female and male of Mid-Pleistocene time, also in P. washingtonii (D), P. jeffersonii (G), aged type, and P. jeffersonii (H), giant male in the Nebraska Museum. 4) All the profiles present a concave forehead, a moderately elevated summit, and a rather angulate occipital contour. 5) As shown by comparison with figure 934, the hypsicephaly, acrocephaly, and bathycephaly of the cranium are much less acute than in Mammonteus. 1052 OSBORN: THE PROBOSCIDEA resemblance. The relation (Figs. 806, 816) of the basis cranii to the fronto-occipital contours reveals a very profound difference between the fully adult Parelephas jeffersonii (Nat. Mus. 10261) and the adult Hlephas indicus profile, on the one hand, and the Loxedonta africana profile, on the other. In all stages Parelephas agrees both with Archidiskodon and Mammonteus in the concave frontal plane or forehead. PARELEPHAS TROGONTHERII! 4500MM..14’9v¥e” e PARELEPHAS FLORIDANUS 3721MM.,12’2%" e GERMANY MOSBACH FLORIDA a ud $ ° PARELEPHAS TROGONTHERI|! 4054mm..13/3%" e PARELEPHAS JEFFERSONI! 3402mm., 11’ 146” GERMANY TAUBACH INDIANA ELEPHAS INDICUS BENGALENSIS 3200MmM.,10’6” PARELEPHAS COLUMBI 3646 mMmM.,11/1 ve” INDIA GEORGIA Fig. 936. PARELEPHAS OF EvROPE AND AMERICA IN COMPARISON WITH ELEPHAS INDICUS BENGALENSIS Restorations to a one-fiftieth scale by Margret Flinsch Buba, under the direction of Henry Fairfield Osborn. The crania conform in size with the three skeletons more or less fully known, namely, of Parelephas inter- medius of France (Fig. 944), said to attain a skeletal height of 3750 mm., or 12 ft. 34 in., of P. jeffersonii in the American Museum (Fig. 966), with an estimated height of 3200 mm., or 10 ft. 6 in., of the giant P. jeffersonii of the Nebraska Museum (Neb. Mus. 1-4-15), the skeleton of which is unknown, and of P. columbi of the Amherst Museum (Fig. 955), with an estimated height of 3430 mm., or 11 ft. 3 in., finally of the giant P. floridanus, known THE MAMMONTINA: PARELEPHAS 1053 from an associated humerus and femur in the American Museum to attain an estimated height of 3584 mm., or 11 ft. 9 in., probably influenced by the favorable climatic conditions of Florida in contrast to the more severe northern conditions which produced P. jeffersoni?. For estimated height in the flesh, compare figure 936 on opposite page. A. IMPERATOR Ref. M. PRIMIGENIUS Ref P. JEFFERSONII 7yoe ~ WASHINGTONII Los Angeles Mus. Var. \7us 8580 Amer. 75.9950 Amer. “us. 863/ A Yoo natural size Fig. 937. CranraAL PROFILES OF THE MAMMONTIN® OF AMERICA One-twentieth natural size (A) Archidiskodon imperator, young male in the Los Angeles Museum. (B) Mammonteus primigenius, adult male, dwarfed, in the U. S. National Museum. (C) Parelephas jeffersonii, type aged male in the American Museum of Natural History. (D) Parelephas washingtonii, referred young adult male in the American Museum of Natural History, the least progressive of the series. Observe extreme progressive hypsicephaly, cyrtocephaly, and bathycephaly as we pass from right (D) to left (B, A); thereby the occipital crest is elevated, the grinders are depressed, the orbits are approximated to the condyles (B), the face is flattened or concave. LecroryPE AND Coryprs oF PARELEPHAS TROGONTHERIOIDES About two-fifths natural size Vig. 938. Lectotype and cotypes of Blephas antiquus Vale. var. trogontherioides Zuffardi, 1913, Tay. rx (11), figs. 3a-6b, namely: Lectotype from Nizza della Paglia (Astésan), third left superior molar, 1.M* (6a, 6b), with 14+ ridge-plates; cotypes from near San Paolo de Villafranca, third right inferior molar, r.Mg (4), with +14-+ ridge-plates, third left inferior molar, 1.Mg (5a, 5b), with 4514 \ ridge-plates and talon, and second left inferior molar, ].Me (3a, 3b), with at least 10 ridge-plates. Compare Zuffardi, op. cit., pp. 1380, 155-158, also Depéret and Mayet, 1923, pp. 177-179. These figures are designated by the author as follows: “{ig.] 83a.—Molare probabilmente quinto inferiore sinistro (@),—pag. 155 [35]. 3b.—Lo stesso visto dal fianco esterno. “4 .—Molare ultimo inferiore destro (‘8’),—pag. 156 [86]. [Mus. Palais Carignan, Turin.] “ 5a.—Molare ultimo inferiore sinistro (‘9’),—pag. 156 [36]. 5b.—Lo stesso visto dal fianco interno. 6a.—Molare ultimo superiore sinistro (‘5’),—pag. 158 [38]. [Geol. Mus. Turin.] 6b.—Lo stesso visto dal fianco esterno. [Geol. Mus. Turin.]’’ 1054 THE MAMMONTINA: PARELEPHAS 1055 2. SYSTEMATIC DESCRIPTION OF EUROPEAN AND ASIATIC SPECIES IN ASCENDING PROGRESSIVE ORDER Parelephas trogontherioides Zuffardi, 1913 Figure 938 Type: Upper Pliocene (Villafranchian stage), Piedmont, northern Italy, Nizza della Paglia and San Paolo de Villafranca (Astésan). Referred: Red and Norwich Crags of East Anglia. Third superior and inferior molars, M*, M:, described as amutation of small size of Hlephas trogontherii; grinding teeth of reduced dimensions; ridge-plates of similar character to 2. trogontherii; laminar frequence 6 to 6.5 in 10 em.; total esti- mated ridge-plates in M 3 “*** as preserved in the lectotype (Fig. 938, 6a, 6b), in the cotype M 3 s-ya-y (Fig. 938, 4), in the cotype M 3 y-44-y (Fig. 938, 5a, 5b); estimated ridge-plate formula M 3 7-44 somewhat inferior to that of EH. trogontherii, as we should anticipate. We may depend upon the observations of Falconer, Pohlig, Zuffardi, and Depéret as to the resemblance of these cotypes to EH. [=Parelephas| trogontherti, which, if cor- roborated by future discovery, make this the most primitive species. As to size, Osborn observes that whereas the typical molars of Parelephas trogontherioides exhibit fewer ridge-plates (M 3 ++4*), they are about equal in size (length, M*, 210-+-mm., breadth 110 mm., height 150 mm.) to Pohlig’s type (Fig. 939) of P. trogon- thervt (M°, length 225 mm., breadth 101 mm., height 138 mm.), also superior in size to the smallest M* of P. trogontherii from Stissenborn near Weimar (length 213 mm., breadth 74 mm., height 118 mm.); consequently P. trogontherioides cannot be described as a “‘mutation de petite taille”; it is, however, inferior in size to one of the largest superior molars of Siissenborn, namely, length 317 mm., breadth 76 mm., height 183 mm. (see Soergel, 1912, Pl. vir). EH. |Elephas] antiquus var. trogontherioides Zuftardi, 1913. “Klefanti Fossili del Piemonte.’ Palaeont.-Italica, XIX, pp. 130, 155-158. LEcTOTYPE AND Coryprs.—Lectotype, third left superior molar, 1.M* (Fig. 938, 6a, 6b); cotypes, third right inferior molar, r.M; (4), third left inferior molar, 1.M; (5a, 5b), and second left inferior molar, 1.M> (8a, 3b). Horizon AND Locauity.—Upper Pliocene (Villafranchian stage),' Nizza della Paglia and San Paolo de Villafranea (Astésan), Piedmont, northern Italy. LecroryPr AND CoryPr Ficurrs.—Zuffardi, op. cit., 1913, Tay. rx (11), figs. 3a-6b. Descrietion.—(Zuffardi, 1913, p. 130): ‘‘Perd pensando che il Jourdan considerava la sua specie EH. intermedius come specie di passaggio dall’ EZ. antiquus all’ E. primigenius, per evitare dannose confusioni, proporrei ora il nome di H. antiquus var. trogonthervoides alla variet&é in questione, in ricordo del nome specifico dapprima dato al materiale attualmente ad essa attribuito. ...[p. 155] Ricordo prima un dente [cotype 1.M.] (Tav. rx [11], fig. 3a, 3b) che si trovava in raccolta senza indicazione alcuna e che distinguo colla lettera G. La corona larga con fianchi non molto convessi, termina anteriormente con una faccia ristretta e arroton- data, e posteriormente é tronca. La superficie di abrasione é concava e di forma ovale. . . . Lunghezza della corona m. 0,156. Larghezza massima della corona, m. 0,105. Altezza massima della corona m. 0,094. Indice dentale 0,015. Frequenza laminare 6%. Simbolo [ridge-plate formula] « 10—.” Zuffardi states on page 156 that the two third inferior molars [cotypes] (Tav. Ix, figs. 4, 5a, 5b) were [erroneously] referred to Elephas antiquus Falconer and Cautley, from the Pliocene lacustrine of Astigiana S. Paolo, according to the label attached to the specimens in the Turin Museum, numbered 8 and 9 respectively. No. 5 [lectotype] of the collection (Tav. rx, figs. 6a, 6b), also [erroneously] referred to Hlephas antiquus, is from Nizza della Paglia (cf. p. 158); length of crown 210 mm., maximum breadth of crown 110 mm., maximum height of crown 150 mm., laminar frequence 6! to 7, ridge-plate formula—1l15 x. Depéret and Mayet (1923, p. 178) estimate the total ridge-plates of this type specimen at about 20. Zutiardi’s full type description is as follows: (Zuffardi, 1913, p. 158) “La stessa discussione pud farsi in riguardo di un altro grosso molare superiore (Tav. rx [1], fig. 6a, 6b). Esso porta Vindicazione ‘5’ Huelephas antiquus Cautl. et Fale. Nizza della Paglia—Dono del sig. cav. Pio Corso Bosen- asco’ ed é accompagnato da un biglietto cosi concepito ‘n.° 20— Rinvenuto nel 1819 in un podere della famiglia Corso Bosenasco dal sig. Pio Corso Bosenasco Sindaco di Nizza Lapaglia, a Nizza Lapaglia.’ E un dente alto e massiccio, troncato davanti, arro- tondato e assottigiiato in dietro. I fianchi della corona, ancora rivestiti dal cemento, sono verticalmente quasi diritti, e longitu- dinalmente convesso interno, concavo l’esterno. La superficie di abrasione, quasi rettangolare-arrotondata, 6 composta di dodici lamine, tortuose, larghe come gli interspazi di cemento e tendono ad allargarsi bruscamente nel mezzo in modo da originare una espansione ovolare. Le corna piegano pili o meno sentitamente all’ indietro, le lamelle sono leggermente pachiganali e fittamente increspate. Della 1.* lamina é presente solo la meta interna della lamella di smalto posteriore, e della 2.* manea la meta esterna della lamella anteriore. Sino all’8.4 compresa, le lamine hanno figura completa; la 9%, 10%, 11.8 hanno tre elementi laminaroidi di cui il mediano pit. sviluppato, e assai meno l’estremo interno ridotto quasi a un semplice anulo. La 12.* é composta di quattro cercini appena usati. Segue l’altra parte della superficie di abrasione non usata che forma con la precedente un angolo di poco superiore al retto. Essa quantunque ricoperta ancora da cemento, dalle costo- lature laterali pud ritenersi constituita da cinque lamine delle quali Vultima, strettissima e arrotondata, pud considerarsi come tallone prossimale.”’ “Le radici, per la massima parte perdute, si iniziano in un brusco restringimento laterale della corona e vi si vedono i tronconi basali di un paio di robusti rami anteriori probabilmente liberi, e i resti dell’ammasso delle altre radici. Dall’andamento di queste e dalla forma della corona sembra manchino al massimo altre due amine.” 'See footnote 1 on p. 1049 above regarding the possible Lowest Pleistocene age of the Villafranchian.—Hditor.] 1056 “Valori metrici: lunghezza della\corona.?: . 2.2.2... 1s. eee m. 0,210 Larghezza massima della corona............. SOO Altezza massima della corona............... 0150 Indicerdentalere s.r ct ee eee «0,015 Hrequenza laminare: op. cee eae eee 65-7 LEN DOLOME ee GER ere acer cs hcg ee a Aaa ers —1l5x” “F d’uopo notare pero che il Falconer ((40], vol. II, pag. 249-250) si riferiva forse anche a questo dente quando parlando del suo #. armeniacus, che credeva ravvisare in un grosso molare segnato col n.° 7 nella nostra raccolta, e che ci occupera pitt avanti (pag. 174 [54]), affermava che altri ve n’erano tra i molari di S. Paolo.” PLIOCENE oF IraLy (DEephRET AND Mayet, 1923, pp. 176- 183).—According to Falconer (1868), Pohlig (1889 [1888]), and Zuffardi (1913), this Parelephas trogontherioides branch |i.e., phylum] although rare first appears in the Upper Pliocene (Villa- franchian) of Italy. As discovered near San Paolo de Villafranea this [lectotype] M® exhibits so closely all the characters of the cor- responding tooth of Hlephas trogontherti that it can only be separat- edasa variety. Falconer (1868) mentioned San Paolo as one of the localities in Italy containing teeth resembling the type of E. armeniacus of Asia Minor, which he accordingly referred to the Armenian species (see p. 1060 below). For this and other Upper Pliocene teeth found at or near San Paolo and described and figured both by Zuffardi and by Depéret and Mayet, the subspecific name Elephas antiquus var. trogontheri- oides was given, the characters being summed up as follows (Depéret and Mayet, 1928, p. 179): “Ces molaires présentent tous les caractéres essentiels de lH. trogontherii quaternaire, savoir: une couronne trés large et trés haute, avee des lames épaisses et assez écartées (fréquence laminaire de 6 4 6, 5); un émail tortueux et fortement pliss¢, orné sur sa longueur de quelques sinus aigus irréguliers, mais ne ressemblant en rien au sinus loxodonte médian des EB. meridionalis et antiquus. . . . [p. 180] On peut done dire seulement que la forme pliocéne de |’F. trogontherii posséde des molaires de dimensions plutét un peu réduites, sans qu’il puisse étre question d’une véritable mutation de petite taille de V’espéce. On doit s’attendre 4 découvrir un jour dans le Pliocéne ancien de quelque région éloignée les types ancestraux nains du rameau de VE. trogontherii.” First INTERGLACIAL OF GERMANY (F'REUDENBERG, 1926).— In the basal layers of the Mosbach sands, near Wiesbaden (Fig. 932, 12), Freudenberg records small mutations (“kleinere Vari- ante”) which he refers to Hlephas trogontherii, associated with artificially broken off (‘‘kiinstlich abgetrennte’’) portions of the horns of Cervus verticornis, charred wood (‘‘Holzkohlen’’), and OSBORN: THE PROBOSCIDEA earth colored by fire (‘rotgebrannte Erde’’), also bones belonging to an ancient camping place of primitive man which might well have belonged to Homo heidelbergensis; of the same geologic age as the Cromer Forest Bed which is quite as old as the deposits of Jockgrim (Freudenberg, 1909). The very ancient characteristics of this small F. trogontherzi mutation at once confirm the great antiquity of this basal Mosbach sands horizon, which also contains the Etruscan rhinoceros (Dicerorhinus etruscus), Trogontherium cuvieri, and Hippopotamus. In this lower Mosbach stage of this small #. trogontherti mutation, the third inferior molar, Mg, ex- hibits 8 ridge-plates in 10 em., exactly as in the primitive trogon- thervi molars of Jockgrim, whereas the true Hlephas primigenius Mg exhibits 7 ridge-plates in 10 cm. These basal sands of Mosbach cannot be more recent than the /st Interglacial (Giinz-Mindel) stage of the Alps, or the Elephant Bed of Cromer (Forest Bed series). Still more ancient are the underlying layers at Mosbach containing Hlephas [=Archidiskodon] meridionalis and an Upper Pliocene fauna with Mastodon [= Anancus] arvernensis and the Red Crag species. Osborn, 1928: The above note by Dr. Wilhelm Freudenberg is deeply interesting as tending to confirm the great antiquity of the Parelephas phylum in western Europe and the possible co- existence of a primitive species of Parelephas (?sp. trogontherioides) with Heidelberg man (Homo heidelbergensis). Parelephas trogontherii Pohlig, 1885, 1888-1891 Figures 794, 865, 871, 929, 934-936, 939, 940, Pl. xxi1 Type: 2d Interglacial deposits at Siissenborn near Weimar (Fig. 932, 11), northern Germany. Referred: 1st-2d Interglacial. Pohlig (1887.2, p. 274). “vom mir zuerst in den alten Flusskiesen Thiiringens zu Siissenborn etc. bei Weimar in grésserer Menge aufgefundene altdiluviale Elephanten- zahnform beschrieben.” This very important species, widely distributed in 2d Inter- glacial times in western Europe, is, with the more progressive Elephas intermedius of Jourdan, a member of the distinct phyletic series Parelephas. Parelephas trogontherii belongs to a phylum by itself which appears sparsely in Europe at the close of Pliocene time and reappears in the /st and 2d Interglacial periods. We do not agree with Pohlig’s opinion that the species H. trogontherii is a con- necting ancestral link between Hlephas meridionalis and E. primi- genius. With Depéret we regard it rather as a distinct phylum. The estimated ridge-plate count of the type, M*® (Fig. 939, upper) is +2*, somewhat exceeding the ridge-plate count of H. [P.] trogon- therioides Zuffardi. Soergel (1912.2, Tab. vir, vir) records three inferior and five superior molars from the type locality of Siissenborn (/s¢ to 2d Interglacial deposits), in the Rebling and Weimar Museum collections, which vary widely in length, breadth, and height, as follows: SUssENBORN: ReBLInG AND WemmaR Mus. Ridge formula Length Breadth Height R.Mz, No. 46 (Weimar) x18x (x17x) 270 62 160 17 (Weimar) 16x 255 83 105 10 (Weimar) x21x 316 73 137 R.M® No. 80 (Rebling) x18x 302 91 192 69 (Rebling) 20x 291 88 160 32 (Weimar) x 20x 296 66 137 89 (Rebling) x 18x 264 71 162 87 (Rebling) x 20x 274 85 171 THE MAMMONTINA: PARELEPHAS Elephas trogontherit Pohlig, 1885. ‘Ueber eine Hipparionen- Fauna von Maragha in Nordpersien, iiber fossile Klephantenreste Kaukasiens und Persiens und iiber die Resultate einer Mono- graphie der fossilen Elephanten Deutschlands und _Italiens.’’ Zeitschr. deutsch. geol. Ges., 1885, XXXVII, Heft IV, p. 1027. Typrn.—Apparently a last superior molar, M*, of the right side, also a last inferior molar, Ms, of the same side. Hort- ZON AND Locauiry.—2d Interglacial deposits, Siissenborn near Weimar, northern Germany. Typr Ficure.—Pohlig, 1888, p. 193, fig. 79, and p. 195, fig. 82 (Fig. 939 of the present Memoir). Schwabe Coll., Weimar. REFERRED SPECIMENS.—The Siis- senborn stage of the upper and lower grinding teeth of Elephas trogonthervi, illustrated by Wiist, 1901, Taf. n, figs. 4-12 (Fig. 940), appears in comparison with the stage erroneously referred to ‘Elephas’ [Hesperoloxodon] antiquus in figures 3 and 18. RELATIONSHIPS OF ELEPHAS TROGONTHERII POHLIG (1885, 1887, 1888-1891, 1907, 1912) AND SOERGEL (1921) RELATIONSHIPS.—(Pohlig, 1885, p. 1027): “6. Unter der Bezeichnung ‘Elephas trogontherii Pohl.’ fihre ich in meiner Monographie eine europadische Molarenform auf, welche zwischen denjenigen des FE. primigenius und E. meridionalis zoologisch, wie ihrer geologischen Lagerstitte nach, in der Mitte steht, dem E. antiquus in der Lamellenformel am naichsten kommt, aber durch die Kronenform und geologische Lagerstitte von dieser Art weit schirfer gesondert ist als von den anderen beiden Species. Das Verhaltniss von FE. trogontherti zu EH. armeniacus Fale. und auch E. namadicus Fale.-Cautl. bleibt noch genauer zu untersuchen.— E. meridionalis und E. primigenius stehen nach Kranologie und Dentition iiber H. trogontherii hin in directer Verwandtschaft.” Description (PoHLIG, 1887.2, p. 274): ‘Unter diesem Namen Elephas trogontherii habe ich in meiner Monographie eine, wie es scheint, in Europa allgemein verbreitete, von mir zuerst in den alten Flusskiesen Thiiringens zu Siissenborn ete. bei Weimar in grosserer Menge aufgefundene altdiluviale Hlephantenzahnform beschrieben, welche geologisch sowohl als zoologisch die beiden Arten EH. meridionalis Nesti aus dem Pliocaen und EL. primigenius Blum. aus dem Plistocaen véllig mit einander verkettet: in der Form der Krone und Kaufliche und in dem Charakter der ein- zelnen Schmelzfiguren auf letzterer stimmt jene altdiluviale Rasse mit den zuletzt genannten beiden Arten tiberein, steht jedoch, was die Anzahl der Schmelzlamellen und die Dicke der Schmelzwande anbetrifft, zwischen diesen beiden Species in der Mitte,—in den unterplistocaenen Schichten mehr dem E. meri- dionalis, in den mittelplistocaenen Lagerstiitten mehr dem Mam- muth in dieser Beziehung angenihert.”’ Pohlig (1885-1887) from the first expressed the opinion that Elephas trogontherii forms a connecting ancestral link between Elephas meridionalis and Elephas primigenius, its earlier represen- tatives resembling the former, its more recent representatives re- sembling the latter, accordingly he wrote the specific name at the time EF. primigenius trogontherii. He held that this species entered Europe during the second glaciation (Saxonian-Mindel) and sur- vived during the 2d Interglacial period (Helvetian, Mindel-Riss) ; thus #. trogontherii was contemporary with the appearance of the Chellean industry in France (Penck). Pohlig (1888-1891, p. 458) pointed out that while Elephas trogontherii resembles E. antiquus 1057 in ridge formula, in the structure of the crowns of the molar teeth it is sharply separated, because the grinders of EH. trogontherii are different in proportion from those of EF. antiquus; the enamel Fig. 79. Letzter Maxillenmolar von Elephas trogontherit. Original zu Weimar (Dr. Schwabe), */ rh ies Fig. 82. Letzter Mandibelmolar von Elephas trogontherii. Original zu Weimar (Dr. Schwabe), 1/,. Types OF PARELEPHAS TROGONTHERI. COMPARE SOERGEL, 1912.2, Tas. vir One-third natural size Fig. 939. Type third superior and inferior molars of Elephas trogontherti Pohlig, 1885. After Pohlig, 1888-1891, p. 198, fig. 79, and p. 195, fig. 82. These are the specimens first mentioned by Pohlig (1885, p. 1027) and figured by him in his Memoir of 1888-1891: “Fig. 79. Letzter Maxillenmolar von Elephas trogontherit. Original zu Weimar (Dr. Schwabe), }5.”” This is apparent- ly a last superior molar of the right side, r.M°; it is complete and displays 15 ridges. “Fig. 82. Letzter Mandibelmolar von Elephas trogontherii. Original zu Weimar (Dr. Schwabe), }4.’’ This is a last inferior molar of the right side, t.M3; it is broken in front and displays 16+ ridges. The incomplete ridge formula of these types of Hlephas trogontherii is: M 3+¢1. See text below. 1058 OSBORN: bands rise above the cement not vertically but obliquely, after the manner of the Asiatic elephant. 2. trogontherii bespeaks a relative- ly moist climate but is also occasionally mingled with remains of the reindeer (Rangifer) where the forest bordered on the open country, as attested by the appearance of both animals in Siissen- born and Steinheim. Pohlig (1888-1891, p. 386, fig. 121) figures the female skull as Hlephas (primigenius) trogontherti (reproduced in Fig. 865, Nos, 3, 4 above), in which the superoccipital region is imperfect and restored. Pohlig’s opinion as to the transitional character of this animal between LP. genius is indicated in the names he has successively assigned to it, meridionalis and EB. primi- namely: Hlephas trogontherti Pohlig, 1885, p. 1027. EKlephas (primigenius bezw. meridionalis) trogonthervi, 1887, p. 274. Elephas (primigenius) trogontherii Pohlig, 1887 [1888"], p. 799. Soergel (1921.2, p. 68) in his comprehensive review entitled ‘Blephas columbi Faleoner’ likewise adopts Pohlig’s opinion and places Hlephas meridionalis-trogonthervi-primigenius as successive phyletie stages in the same west European phylum. He assigns (op. cit., p. 55) the following collective ridge formula to the Elephas trogonther?i specimens of Siissenborn and Mosbach: M 1323 M2 Dp 2% Dp 3 =f Dp 1 i 11-13 aaaae Soergel is of the opinion that Llephas trogontheria (western Iurope) and the typical Hlephas columbi (southern U.S.A.) are in similar stages of evolution. He points out that the typical Llephas THE PROBOSCIDIEA cit., pp. 69-97) concurs with Pohlig in the opinion that Hlephas trogontherii is ancestral to Hlephas primigenius. Hapirar.—tIn a letter of June 30, 1922, Soergel states: According to my interpretation 1’. trogontherii lived during a period of limited rainfall, in a half-arid climate, not migrating during glacial times into the colder ice-clad regions of middle Hurope (compare also Soergel: ‘Die Ursachen der diluvialen Aufschot- terung und Erosion,’ Verlag Borntriiger, 1921). 2. trogontherii therefore occupied the region later traversed by the mammoth. We still regard /. trogontherii as the direct ancestor of 2. primi- genius. We possess in the museums of Europe all the transition forms in large numbers and are unable to make any sharp dis- tinction between the two species. If one seeks a complete under- standing of the chief literature of value on this subject, it may be found in the following works: M. Pavlow, ‘Les éléphants fossiles de Ja Russe,’ Nouv. Mém. d. |. Soc. Imp. des Nat. de Moscou, T. XVII, 1910; W. Soergel, ‘Hlephas trogontherii und Hlephas antiquus ete.’ Paleontographieca, 1912; KH. Wiist, ‘Das Pliozéin und das iilteste Pleistoziin Thuringens. Abhand. d. naturf. Gesellsch. zu Halle, Bd. XXIII, 1900. I rejoice that in the princi- pal points we agree as to the diluvial elephants and their phylogeny. PHYLOGENETIC CONCLUSION (OsBORN, 1924).—Osborn does not agree with either Pohlig or Soergel that the typical Hlephas trogontherii is a descendant of . [Archidiskodon] meridionalis; nor does he agree that it should be regarded as transitional or ancestral to the typical #. [Mammonteus] primigenius. The pro- found difference between the broad-plated molars of the Archi- Vig. 940. After Wiist, 1901, Taf. 1. type locality (Siissenborn near Weimar). “Nr. 3. EHlephas antiquus Falc. “6. Desgl. M.IIT max. dextr. “18. Hlephas antiquus Fale. meridionalis may be regarded as the direct ancestor of the Mlephas columbi-imperator phylum: (Op. cit., 1921, p. 68) “Diese Tatsache berechtigt zu der Annahme, ‘dass die nach Amerika iiberwandern- den Formen der Meridionalis-trogontherti-Reihe schon beim Ueberwandern resp. kurz vorher besondere Charaktere gegentiber dem europiischen Fl. trogontherii meridionalis ausgebildet hatten, Charaktere, die in der weiteren Entwicklung sich zum ‘Jmperator- Typus’ steigerten. Is war also im iiltesten Diluvium im Kreis der kontinentalen Elefantenformen Polen, El. trogontherii meridionalis im Westen und dem direkten eine Variationsbreite mit zwei Vorfahren des El. imperator im Osten vorhanden.” Soergel (op. M.III max. dextr. M.II max. dextr. Referred molars (r.Mg with 18+ ridge-plates) of ‘Blephas trogontherit’? Pohlig from the 2d Interglacial of Siissenborn and Weimar, Germany. Wiist erroneously refers r.M® (Nr. 3) and r.M? (Nr. 18) to Elephas antiquus; they are Parelephas trogontherii molars from the Siissenborn. .. . Siissenborn. . . . Weimar.” diskodon planifrons-meridionalis-imperator phylum widely distin- guishes this phylum from Elephas trogontherii; Osborn accepts rather the view of Depéret and Mayet that /. trogontherii repre- sents a distinct branch of the mammoths. Moreover, the extreme hypsicephalic, acrocephalic, bathycephalic cranium of Elephas primigenius, with corresponding hypsodont, finely compressed, and laminated molars, cannot be derived from the cranium and molars of the Hlephas trogontherti type. Osborn agrees entirely with Depéret and Mayet that we have to do with three distinet phyla, certainly separate from the close of Pliocene time. Again Osborn does not accept the opinion of Soergel that 2. trogontherti 'In the Bibliography of the present Memoir (p. 795), it is possible that 1887.1 should read 1888. THE MAMMONTINA: PARELEPHAS is related to the true 2. [=Parelephas| columbi of the southern United States, although it may well prove to be related to the animal described by Cope, Hay, Osborn, and others as ‘‘Hlephas columbv”’ but which now proves to represent a distinct species, namely, the Parelephas jeffersonii of Osborn. Consequently, Osborn, as cited above, has placed EF. trogontherii in the distinet phylum Parelephas, the generic characters of which are defined by cranial and dental characters in the chapter on the ‘Classification of the Elephantoidea”’ (Chap. XY). Parelephas (?) trogontherii nestii Pohlig, 1891 Figure 941 Type locality: ‘Jungpliocaenen,” Forest Bed (Norfolk), Walton (Essex), and Southwold (Suffolk), England. Regarded by Depéret and Mayet (1923, p. 163) as a synonym of Hlephas [Hesperoloxodon| ausonius F. Major. Cotypes regarded by C. W. Andrews as Lower Pleistocene. belonging to 2. {= Parelephas| trogontherii. Pohlig (op. cit., p. 303) in reviewing Leith Adams’ Monograph on Elephas antiquus observed that certain teeth of the Thames Valley (Adams, 1877-1881, p. 33), of Norfolk (p. 40) and of South- wold (p. 38), do not agree with the continental molars [of Germany| jung- . Forestbed stammenden” molars do belong to the referred to 2. antiquus. Pohlig continues, that part of the pliocaenen . . typical 2. antiquus, but that the majority belong to an inferior V3 Nalural S7ZE - Brit. Mus. 27915 ; NE wd, A 1059 race; he then separates this race as 1. Nestii. Pohlig’s types or colypes of EB. Nestii are therefore molars of Forest Bed age. Coryprs.—As shown above, the subspecies Llephas antiquus Nestii was erroneously based on cotype specimens selected from Leith Adams’ Monograph of 1877-1881, pp. 37 and 38, namely, srit. Mus. 33327, a dredged lower molar from Happisborough with +16+ridge-plates in 11.5 inches [catalogued by Lydekker, 1886.2, p. 128, as a “right third upper true molar’’|; Brit. Mus. 27915 [given as No. 27515 by Leith Adams], a 14+ ridge-plated third left upper true molar from Walton (Essex), with 13+ ridge-plates in 8.5 inches, and Brit. Mus. 39463, a half-worn third left lower true molar from Southwold (Suffolk), compare Lydekker’s “Catalogue of the Fossil Mammalia in the British Museum,” 1886.2, pp. 128, 129, and 135. All these cotypes are of Lower Pleistocene Cromeri- an or Forest Bed age; their generic and specifie reference to ‘B. antiquus’ is doubtful; C. W. Andrews refers Brit. Mus. 27915 and 39463 to ‘Hl. trogontherii; Hopwood (letter, August 6, 1929) kindly furnished photographs and detailed measurements which tend to confirm Andrews’ reference to ‘#. trogontherii.’ Happisborough (Forest Bed), England. Brit. Mus. 33327, r.M*, length 268 mm., breadth 92 mm., height 196 mm.; ridge- plates +16+, 6.3 in 10 em. Walton England. Brit Mus. 27915, 1.M?’, length 302 mm., breadth 80 mm., height 167 mm.; ridge-plates 18, 61 in 10 em. (referred to ‘F. trogontheriv’ by C. W. Andrews). Southwold (Suffolk), Brit. Mus. 39463, 1.Ms, length 243+- mm., breadth 83 mm., height 103 mm., ridge-plates (Essex), Iengland. # 0098 7 6 5 4.327 | pie 4 a ie Ge ey Brit Mus. 39463 Cotyrres or ELEPHAS ANTIQUUS NESTIT POHLIG NOW TRANSFERRED TO PARELEPHAS(?) TROGONTHERI NESTII Fig. 941. Cotypes or syntypes of Elephas antiquus Nestii Pohlig, 1891, after photographs kindly furnished by Dr. A. Tindell Hopwood of the British Museum, August 6, 1929. From the Lower Pleistocene of eastern England. One-third natural size. (Left) Brit. Mus. 27915. Left third superior molar, |.M’*, from Walton (Essex), England, exhibiting 1-18 ridge-plates; also crown view of same. (Right) Brit. Mus. 39463. Incomplete left third inferior molar, ].Mz, from Southwold (Suffolk), England, exhibiting 1-14 posterior ridge-plates; anterior ridge-plates broken off. Lateral and crown views of same. 1060 OSBORN: 14+, 7% to 8 in 10 em. (referred to ‘EH. trogontheriv’ by C. W. Andrews). Unfortunately these specimens were not figured by Leith Adams or Pohlig, but they were described by Leith Adams as belonging to his ‘Hlephas antiquus.’ It is to be regretted that the name Elephas antiquus Nestii has been adopted and widely quoted in the literature, for we cannot agree with Depéret and Mayet (1923) that ‘Hlephas antiquus Nestii’ Pohlig, erroneously based on Lower Pleistocene cotypes, is synonymous with ‘Elephas ausonius’ Major-Depéret-Mayet, correctly based on an Upper Pliocene type of the Villafranchian stage.! Hisrory.—In 1891 Pohlig imperfectly established this sub- species (named after Filippo Nesti) as of Upper Pliocene age, whereas the cotypes selected by him, as mentioned by Leith Adams in his Monograph of 1877-1881, are said to be from Walton, South- wold, and Norfolk, the latter from the Cromer Forest Bed level which is of Lower Pleistocene age. This subspecies, founded on a misconception as to its Pliocene age, is invalid; it is certainly not in the same geologic stage as the Upper Pliocene ‘Hlephas’ ausonius Major-Depéret-Mayet, from the Villafranchian of Italy.! Although not figured, the ridge formula was said by Pohlig (1891, p. 305) to agree with that of #. antiquus: ‘In der Lamellenformel wiirde £. Nestii sowohl mit E. antiquus typus, als auch mit EL. trogontherii iibereinstimmen.” E. Nestii Pohlig, 1891. ‘“Dentition und Kranologie des Elephas antiquus Fale. mit Beitragen tiber Elephas primigenius Blum. und Elephas meridionalis Nesti. . . . Nachtrige,”’ pp. 303, 304, 465. On page 465 the name appears as EHlephas antiquus Nestii nf. Typr.—(Letter, Pohlig, Sept. 10, 1924): ‘As type specimens of H. Nestii from the forest bed are named in my mono- graph (p. 304) those of L. Adams’ [monograph], p. 38, from Walton and Southwold, and the Nr. 33327 of the British Museum, from Norfolk.” Locauity AND Horizon.—Forest Bed (Norfolk), Walton, and Southwold, England; Lower Pleistocene. Typxr Vicure.—Not figured. See figure 941 of the present Memoir after photographs. Referred skull of northern Italy figured by Pohlig, op. cit., 1891, p. 350=Cranium B of Nesti (Hlephas meridionalis) ; referred to Hlephas antiquus by Weithofer 1890, and erroneously referred by Pohlig to his subspecies L. antiquus Nestit. Tyer Descriprion.—(Pohlig, 1891, pp. 303-305): “b. Kin Theil der letzteren—die aus dem jungpliocaenen [Footnote: ‘Vel. u. (sub HE. meridionalis).’| Forestbed stammenden—lassen die Zugehérigkeit wenigstens zu der Species H. antiquus zu, manche mogen selbst als typische Molaren derselben gelten; die meisten haben jedoch Abweichungen von der typischen Gleich- missigkeit der Kronenbreite nicht nur, sondern auch von dem charakteristischen Lamellenbau der diluvialen Ausgangsform, be- sonders von dem ausgepriigten Loxodontismus, der complicirteren Festonirung und der grésseren Dicke des Schmelzes bei jener. Da nun diese Kigenthiimlichkeiten zum Theil mit den im ersten Ab- schnitt erérterten, ebenfalls jungpliocaenen Erfunden von der Species aus dem Arnothal iibereinstimmen, so scheint es mir nun- mehr doch geboten, diesen pliocaenen /. antiquus von dem dilu- vialen durch eine Rassenbezeichnung, etwa #. Nestii, vorlaufig zu trennen. ... Es miisste danach die in dem ersten Abschnitt aus- THE PROBOSCIDEA gesprochene Vermuthung, dass Loxodon von unbekannten Masto- dontenformen sich ableite, fallen gelassen werden; denn ganz wie E. Nestii zum typischen E. antiquus verhialt sich nach Mitgetheil- tem das Wenige, was man vom £. priscus weiss... , zum modernen E. africanus,—es zeigt den Charakter des letzteren minder markirt, mit Eigenthiimlichkeiten archidiskodonten Gepriges, wenn auch in geringem Grade, vermengt. . . . In der lamellenformel wiirde E. Nestii sowohl mit EL. antiquus typus, als auch mit E. trogontherii iibereinstimmen; und da an mehreren Punkten der englischen Ostkiiste, wie bei Mundesley-Cromer, Clacton etc. interglaciale Schichten mit #. antiquus und EF. primigenius—an ersterem sogar mit ganz ihnlichem Erhaltungszustand—die jungpliocaenen mit E. meridionalis, E. Nestii und E. trogontherii iiberlagern, also Molaren aller dieser Formen in die Brandung und von da in die Museen gelangen: so war die Schwierigkeit, alles das nunmehr Vermengte in der rechten Art wieder voneinander zu sondern, anfangs begreiflicher Weise selbst fiir den Geiibten zu gross. Und doch sind eben grade diese Verhaltnisse von grésstem Interesse.”’ Osborn, 1930: We are therefore obliged to regard Elephas antiquus Nestii as comparable to the species ‘Hlephas’ [= Par- elephas(?)| trogontherii Pohlig (ef. Parelephas trogontherioides Zuffardi). Parelephas armeniacus [alconer, 1857 Vigure 942 Pleistocene, near Khanoos, Province of Erzerum, Armenia. We note that Falconer in 1868 (Vol. II, pp. 187, 192, 193, 248, 249, 250) observed a strong resemblance between his type of ZL. (Buelephas) armeniacus and Upper Pliocene molars from the locality of San Paolo, or near it, Nizza della Paglia, Italy, recently chosen by Zuffardi as the types of Elephas antiquus var. trogon- therioides. It is probable, therefore, that the species Hlephas armeniacus of Faleoner belongs in the generic phylum Parelephas, but we cannot be certain of this reference until the cranial charac- ters of the species become known. The specific distinctions are: Ridge-plates of M 3 *17 (Fig. 942), worn anteriorly. This appears to be a more progressive stage than Parelephas trogontherii and much more progressive than P. trogontherioides. f). (Hueleph.) Armeniacus Falconer, 1857. ‘On the Species of Mastodon and Elephant occurring in the fossil state in Great Britain. Part I. Mastodon.” Quart. Journ. Geol. Soc. London, Vol. XIII, Synoptical Table opp. p. 319. Typr.—(Faleoner, 1863, p. 74): “. .. two last upper molars in fine preservation, and a portion of a lower, all apparently of the same individual.” Brit. Mus. 32250, 32251, and 32252; also fragment of tusk, head of femur, and portion of tibia (Brit. Mus. 32256, 32254, 32253), which Lydekker (1886.2, p. 174) regards as belonging to the same individual as the teeth. Horizon AND Loca.iry.—Pleisto- cene. Near Khanoos, Province of Erzerum, Armenia. TYPE Figure.—Falconer, 1863, Pl. 11, fig. 2. Hisrory.—Falconer first named this species in his “Synoptical Table” of 1857, facing page 319: “Spec. 14. EH. (Eueleph.) Armeniacus ... Armenia: Erzeroom ... In the Brit. Mus. Coll. Discovered between Erzeroom and Moosh in 1856. The molar '[See footnote 1 on page 1049 above regarding the possible Lowest Pleistocene age of the Villafrancbian.—Editor.] THE MAMMONTINA: PARELEPHAS plates closely approximated, and the enamel-edges very undulat- ed.” In 1863, pp. 74-76, he renewed his description as follows: “These Khanoos molars are intermediate in character, between the Mammoth and the Indian Elephant, but more nearly allied to the latter... . In the synoptical table appended to my Memoir on the ‘Species of Mastodon and Elephant, &c.,’ the Khanoos fossil form is ranged between E. indicus and EF. primigenius, under the provisional name of H#. Armeniacus. [Footnote: ‘Quart. Journ. Geol. Society, 1857, vol. xiii, p. 319.’] . . . [p. 76] while 2. Armeniacus, as stated above, approaches nearer to the existing Indian species.” Falconer also speaks of the remains of fossil elephants on the banks of the Bosphorus and the northern shores of the Black Sea and of the Sea of Azof (op. cit., p. 75). On page 74 he gives what may be called the type description. Typr Description (FALCONER, 1863, Pp. 74).—‘“The speci- mens presented by Colonel Giels to the national collection, consist of two last upper molars in fine preservation, and a portion of a lower, all apparently of the same individual. These molars strike a practised eye, at the first glance, as presenting something intermediate between the Mammoth and the existing Indian ONE oF THE TypE Motars OF PARELEPHAS ARMENIACUS One-half natural size Vig. 942. Third superior molar, 1.M*, of Elephas Armeniacus Falconer, 1857. Falconer, 1863, Pl. u, fig. 2, p. 114: “Represents the erown-aspect of the last true molar, (m.3) upper jaw, left side, of #. Armeniacus, from a specimen in the British Museum, No. 32,250, procured by Col. Giels, in the province of Erzeroum, in Armenia.” Elephant. The case is of so much interest, that I shall venture on some of the details. The left upper molar (m.3, being No. 32,250 Museum Regist. Paleont. Gallery) is entire from behind the large front fang, the portion borne upon which had been ground down by protracted wear. [Footnote: ‘Pl. 1, fig. 2, shows the crown-view of the tooth.’] The anterior part of the crown to the extent of 2.7 inches is also worn out, presenting merely a smooth surface of ivory, behind which there are seventeen ridges and a posterior talon. Of these, fifteen are more or less worn. The anterior nine form transverse narrow discs; the next six are divided nearly equally by two rather wide longitudinal channels into three divisions, consisting each of a flattened elliptical disc. The trans- verse dises, in their general character, bear a close resemblance to those of the Indian Elephant, the enamel-plates being rather thick, with very pronounced close-set crimping in the middle, but dimin- ishing towards the cornua. These discs are narrower than is com- See footnote 1 on page 1049 above.—Hditor.] 1061 monly seen in the existing species, less open and less parallel The crown is broad, and the enamel plates are high. To render these descriptive details more appreciable and available for comparison, I append the principal dimensions [1.M*]:— across. {In.] [Mm. Extreme length of crown............. 11.75 299 Length of crown surface in use (partly WOLDSOUG) Re rriemcece aie tae rere 9.5 242 Space occupied by the anterior ten discs measured at top of crown......... 59.7 146 Ditto ditto, at base of crown...... 6.1 156 Width of crown at 3rd ridge (greatest). . 4.1 105 Ditto at i thyGittormeereeeer Ball 95 Height of crown at 12th ridge......... Gf gull 181)” (?) E. ARMENIACUS IN ITALY (FaLconeR, 1868).—The same description is found in the “Paleontological Memoirs,” 1868, Vol. II, p. 247, in which Falconer again treats Elephas armeniacus as intermediate between EHlephas primigenius and LE. indicus but more nearly allied to the latter. He adds (p. 249): “I now entertain a strong suspicion that remains of #. Armeniacus, or of a form closely allied to it, occur in Italy. This impression is founded upon specimens which I observed in the Natural His- tory Museum of Turin, in the University Museum of Pisa, in the private collection of the Marchese Carlo Strozzi at Leghorn, and in those of Professor Ponzi and Signor Ceselli at Rome, the satisfactory specific identification of which puzzled me greatly. . . [p. 250] The space occupied by the twelve dises of wear, meas- ured along the summit of the crown, is 7 inches, yield- ing an average of about .6 inch to each, which comes very near that indicated above in the #. Armeniacus of Khanoos, 7.e. .57. The principal dimensions are:— Length of crown (not quite entire), 13.75 in. Extreme width of crown, 4.5 in. Height of crown at 12th ridge, 8. in. Space occupied by the 12 dises of wear, 7. in. I have detailed notes of numerous other molars, ex- humed on the same occasion, from the same locality, St. Paolo, or near it, ‘Nizza della Paglia,’ which yield similar characters. ... I have been led to identify the ‘Khanoos’ and St. Paolo molars as being of the same species, #. Armeniacus, and to consider that they are not referable either to L. primigenius or EL. antiquus|Foot- note: ‘See antea, pages 187 and 192, note-—[Ep.]’].” Osborn, 1928: (1) It is now known that these Upper Pliocene! Italian specimens thus referred by Falconer to H. armeniacus actual- ly belong to Elephas antiquus var. trogontherioides Zuffardi, as confirmed by Depéret and Mayet (see above). (2) Leith Adams (1877-1881, p. 241) mentions EZ. armeniacus but does not add to our knowledge. (3) Lydekker (1880.1, p. 285) erroneously gives the ridge formula of H. armeniacus as M 3 **, whereas Falconer’s type figure shows: M 3 +*7%. (4) Lydekker (1886.2, p. 174) remarks: “‘Hlephas armeniacus, Falconer [lFootnote: ‘Quart. Journ. Geol. Soc. vol. xiii, table facing p. 319 (1857).’|. Syn. After 1062 OSBORN: THE PROBOSCIDEA Buelephas armeniacus, auct. This species is definitely known only by the following [six] specimens, which belong to an animal of very large size. The structure of the molars indicates a species intermediate between EF. primigenius and EF. indicus, which is probably an ancestral form closely connected with both [ootnote: ‘Ibid. vol. xlii, p. 174 (1886).’]. The crowns of the molars are extremely wide, the ridges approximated, with their extremities curving backwards, the enamel is moderately thick and much plicated, and the wear of the crown irregular. In their great width these teeth agree with those of E. primigenius, but in the other characters with those of #. indicus [Footnote: ‘See Leith-Adams, British Fossil Elephants (Mon. Pal. Soe.), p. 241.’].” (5) Lydekker (1886.3, p. 174) remarks: ‘From strata of un- known age at Erzerum, in Armenia, Dr. Falconer many years ago described some elephant molars under the name of Elephas armeniacus; and as Erzerum is comparatively near to Tabriz, it may be suggested that some of the Maragha elephants’ teeth may not improbably belong to this species; but be this as it may, the Erzerum and Maragha faunas may be geographically grouped together.” PHYLOGENETIC CONCLUSION (OsBoRN): The extremely able and acute observations of Falconer (1857-1868) on this species from Armenia and on allied forms from Italy, confirmed by Zuffardi’s observations on the resemblance of this animal to Elephas trogontherii of Pohlig, as signified by the specific name trogontherioides, also the recent observations by Depéret and Mayet (1923), all emphasize the intermediate character of this species and its phyletice relationship to the EHlephas intermedius of Jourdan and to the H#. trogontheriz of Pohlig. The ridge formula, M 3 **7", indicates that EL. armeniacus Faleoner is in about the same stage of evolution as the type of L. trogontherii Pohlig, in which the ridge formula is said to be M 3 42%. While Falconer repeatedly described the type of 2. armeniacus as intermediate in character between FH. primigenius and EF. indicus, but approaching the latter species more closely, Osborn regards it rather as belonging to the same phylum as F. intermedius and EF. trogontherii and refers it to the independent genus Parelephas. PARELEPHAS ERRONEOUSLY RECORDED IN CHINA AND JAPAN Irroneous are the records of Parelephas in China and Japan. The supposed Hlephas armeniacus (Brit. Mus. 29007) is related (Lydekker, 1886.2, p. 169, Hopwood, letter, 1928) to Hlephas [Palxoloxodon| namadicus. Matsumoto’s types of Parelephas protomammonteus and P. protomammonteus proximus of Japan also belong to Palzolorodon. The following literature may be quoted: LypreKxer (1886.3, p. 174).—‘‘There is in the British Mu- seum an elephant’s molar from China (No. 29007), which has been suggested to belong to this species [Hlephas armeniacus]; and if this were correct it would seem that the range of #. armeniacus was somewhat the same as that of Rhinoceros Blanfordt, i.e. that it extended from western Asia through the regions lying to the north of India and China; I am, however, disposed to refer the specimen to #. namadicus.” PARELEPHAS ARMENIACUS REF.—(Leith Adams and G. Busk, 1868, pp. 496-499): “In the British Museum there is a large part ‘MUSee page 1334 below.—LEditor.] of a fossil molar tooth from China, which, on comparison with the teeth presented by Colonel Gills [Giels] from Armenia, so exactly resembles them in every respect, that no doubt can be entertained as to its belonging to the same species, namely, H. Armeniacus of Dr. Falconer; and as such, I believe, it was regarded by that eminent paleontologist. The occurrence, therefore, of the same form in Japan would not have been very surprising; but, so far as I am able to judge, it is impossible to identify Dr. Duggan’s specimen with #. Armeniacus.”’ Japan.—The Japanese tooth found by Dr. Duggan, in com- pany with Mr. Hodgson, in 1859, is referred to Palxoloxodon (see figure by Leith Adams, 1868.1, p. 497).! Parelephas intermedius Jourdan, 1861 Figures 935, 943, 944 Pleistocene plateau loess, near Lyons, Rhone Valley, France. Type locality believed to be more recent (i.e., ?3d Interglacial) than the 2d Inter- glacial deposits of Siissenborn, containing the type of Hlephas [= Parelephas| trogontherit. Jourdan applied this name to the species of mammoth which he regarded as intermediate or transitional between Elephas primi- genus and EH. indicus. The name is appropriate today for the specific stage more progressive and geologically younger than Parelephas trogontherii. The history of this stage is very interest- ing, as detailed below. In referred grinding teeth from the same locality (Fig. 943), the ridge-plate formula is: M 1 js;3 M 322. Depéret (letter, Sept. 10, 1924) assigns the following ridge formula: M3 aacan: In his original description of this species, Jourdan remarked that of all fossil elephants this approached most nearly to Hlephas indicus. He applied the name intermedius to certain broad molars resembling in breadth the molars of #. primigenius but with thicker and more widely separated lamell, in distinction from the narrow molars of 2. antiquus. Unfortunately the type molars of /. cnter- medius have been misplaced and no type figure was published; consequently the specific name Hlephas intermedius Jourdan, 1861, although repeatedly cited, has not gained acceptance; none the less it is certainly valid. The species is related to but not identical with the widely known Elephas [=Parelephas| trogontherti of Pohlig, which in turn resembles the Parelephas jeffersonii of Osborn. Depéret and Mayet (1923) describe this animal throughout as Elephas trogontherit. Elephas intermedius Jourdan, 1861. ‘Des terrains sidérolit- iques,’’ Compt. Rend. Acad. Sci., Tome LITT, 1861, p. 1013. Tyrer Derscrrprion.—(Op. cit., p. 1013): “La faune du sidérolitique du néocéne ou ¢tages les plus supérieures des terrains tertiaires se caractérise done dans ses couches inférieures par |’ Hlephas meri- dionalis, dans les couches moyennes par |’Hlephas antiquus, et dans les supérieures par |’Hlephas intermedius, qui, de tous les fléphants fossiles, est celui qui présente le plus de rapport avee lV Eléphant actuel des Indes.” Hisrory.—Lortet and Chantre (1872, p. 79) observe: “Blephas intermedius (Jourdan, mss.).—Cette espéce, eréée par M. Jourdan sur une forme de l’Hlephas antiquus de Falconer, est celle qui est la plus commune dans le bassin du Rhéne lyonnais; elle parait méme y avoir été cantonnée. Les caractéres transitoires THE MAMMONTINA: entrel’Hlephas primigenius et Elephas antiquus, faciles 4 constater sur de nombreuses piéces, ont engagé M. Jourdan 4 élever au rang d’espéce cette forme, et de lui assigner le nom spécifique d’inter- medius. Ces ecaractéres sont trés-apparents sur certaines dents; cependant, parmi celles qui ont été considérées comme appartenant a lV Elephas intermedius, un grand nombre de piéces peuvent étre rangées dans la catégorie de |’Elephas antiquus.” VALIDITY OF THE SPECIES ELEPHAS INTERMEDIUS (OSBORN, 1924).—Inasmuch as Jourdan’s type description (1861, p. 1013) characterizes the species as being closely related to Hlephas indicus, and inasmuch as Lortet and Chantre (1872, p. 79) cited the name, amplified the specific characters, and referred to this species the important skeleton of Lyons (Fig. 944), it appears best to retain the specific name Hlephas intermedius, atleast for the more progressive speci- mens which occur in this region of France. PARELEPHAS 1063 It appears best also to conserve the name Llephas trogontherti Pohlig for less progressive specimens from Pohlig’s type locality, especially since according to Depéret, Mayet, and Pohlig this phylum (i.e., ‘“‘rameau de 1’ Hlephas trogontherii Pohlig’’) is of long duration. Typr Locauity or E. Lyons remains of this species are found abundantly in the loess covering the plateaus (Depéret and Mayet, 1923, p. 182): “Aux environs de Lyon, il abonde dans le loess des plateaux (Saint-Cyr, Saint-Romain-au-Mont-d’Or, la Duchére, Fourviére, Caluire, Margniole, Croix-Rousse, Saint-Clair, le Vernay, ete.), ot Jourdan l’a décrit sous le nom d’E. intermedius. C'est A cette espéce qu’ap- partient le beau squelette entier reconstitué au Muséum de Lyon, et provenant du lass de la montée de Choulans, 4 Lyon. Les molaires du loess de la région lyonnaise représentent une variété ou peut-étre une mutation plus évoluée, 4 lames un peu plus serrées (fréquence laminaire entre 6, 5 et 7) que dans les molaires d’Alle- magne et d’Angleterre (fréquence laminaire 6 4 6, 5).” This is the level of the type locality of Elephas intermedius Jourdan, which Depéret states may represent a variety, or perhaps a more advanced mutation with laminge somewhat more con- tracted, namely, between 6, 5, and 7, than the German and English Consequently Depéret regards INTERMEDIUS.—In the environs of varieties with laminz 6 to 6, 5. Elephas intermedius as more progressive than the typical Hlephas trogontherw of Pohlig. Rererrep Mouars oF PARELEPHAS INTERMEDIUS, Lyons MusgeuM One-half natural size Vig. 943. Superior molar teeth referred to Elephas intermedius Jourdan from the collection of the Muséum de la Ville 4 Lyon. After photographs, prepared under the direction of Doctors Depéret and Mayet (see letter of September 10, 1924). (Left) This beautifully preserved r.M® exhibits 22 ridge-plates, of which the 12 anterior are worn. (Right) This first, inferior molar, |.My, exhibits 12 worn ridge-plates, but the formula appears to be (with an additional unworn ridge-plate): 1M 1q9. 1064 OSBORN: THE PROBOSCIDEA DrpPERET (TRANSLATION OF A LETTER DATED Lyons, AUGUST 26, 1921).—The Elephas intermedius was created by Jourdan, a former paleontologist of Lyons, who applied the name, unfor- tunately without a figure, to certain broad molars, which he ob- RESTORED SKELETON OF PARELEPHAS INTERMEDIUS, Lyons MusEUM One-sixtieth natural size Fig. 944. Restored skeleton referred by Jourdan to his species Elephas intermedius. The plate was prepared under Jourdan’s direction but was first printed by Lortet and Chantre in the frontispiece of their Memoir of 1872[1876], from which it is here reproduced one-sixtieth natural size. The height is said to be 3 m. 75 em. =12 ft. 34 in. (Lortet and Chantre, op. cit., p. 79): ‘“Hlephas intermedius (Jourdan, mss.). — Cette espéce, eréée par M. Jourdan sur une forme de |’Elephas antiquus de Falconer, est celle qui est la plus commune dans le bassin du Rhone lyonnais; elle parait méme y avoir été cantonnée. Les caractéres transitoires entre l’ Elephas primigenius et |’ Elephas antiquus, faciles 4 constater sur de nombreuses piéces, ont engagé M. Jourdan A élever au rang d’espéce cette forme, et de lui assigner le nom spécifique d’intermedius. Ces caractéres sont trés-apparents sur certaines dents; cependant, parmi celles qui ont été considérées comme appartenant a l’Elephas intermedius, un grand nombre de piéces peuvent étre rangées dans la catégorie de l’Elephas antiquus. Dans la vallée de la Sadne on a trouvé des ossements d’Hlephas intermedius, dans le lehm ou A la limite des alluvions, 4 Lyon et aux environs. (Saint-Rambert- ’Ile-Barbe, le Vernay, Caluire, la Duchére, Vaise, la Demi-Lune, Saint-Just, Fourviére et la Quarantaine.) C’est Ala Quarantaine que M. Jourdan a trouvé, en 1862, le squelette de Il’individu qui a pu étre remonté dans les galeries du Muséum de Lyon par les soins de M. Charles Révil, l’un de nos préparateurs. Ce squelette, un des plus grands et des plus complets que l’on puisse voir en Europe, mesure 3 métres 75 centimétres de hauteur au garrot. I] est surtout remarquable par la courbure trés-prononcée des ses énormes défenses et par les formes trapues et massives de ses membres.” This animal, discovered in the Pleistocene loess of the hill of St. Foy, Lyons, France, is regarded by Depéret and Mayet as similar to Hlephas trogontherit Poblig. Plateau loess (“provenant du loess de Ja montée de Choulans, 4 Lyon.’’ Depéret et Mayet). Depéret and Mayet remark (1923, p. 176): “Lortet et Chantre (1876) n’ont figuré de cette espéce que le squelette reconstitué du loess de Choulans qui se trouve au Muséum de Lyon et forme le frontispice de leur ouvrage; mais ils n’ont donné malheureusement aucune figure des molaires de |’2. intermedius, devenu ainsi une espéce purement nominale, qui ne saurait étre retenue.”’ The skull is said to be erroneously restored at the summit of the occiput. The rounded summit of the cranium in the above figure agrees fairly well with the rounded summit characteristic of Parelephas trogonthervi; the tusks appear to be recurved rather than in- curved as in Parelephas jeffersonit. served to be very distinct from the narrow molars of the Hlephas antiquus type, molars resembling in breadth those of Elephas primigenius, but with thicker and more widely separated lamelle. Lortet and Chantre in their memoir on the Mastodons and the Elephants (Archiv. Mus. Lyon, T. I., 1872, et T. II, 1879) cited this species and figured other pieces which could be related to it. I believe that HL. intermedius is probably the same animal as that named by M. Gaudry Elephas primigenius var. i lames écartées and by Pohlig Elephas trogontherii. We have at Lyons in the University Museum also in the Municipal Museum numerous molars of this species and even a complete skeleton discovered in the loess of the hill of St. Foy at Lyons. It is this complete skeleton [Fig. 944] which has been figured by Lortet and Chantre, but in the absence of figures of the type itself I fear that it may be necessary for us to adopt the name of trogontherii which appears to have priority. Dergrer AND Mayet, 1923, p. 176.—Depéret and Mayet, while regarding FH. intermedius as conspecific with E. trogontherii, treat this animal as follows: “III Groupr prs ELEPHAS TRO- GONTHERI ET EK. PRIMIGENIUS (MAmmoutTHs). Le groupe des Mammouths comprend des Eléphants de taille moyenne, au crane allongé, au vertex trés élevé et étroit en haut (cranes en déme), aux défenses fortement spiralées, aux molaires généralement larges et hautes (type hypsélodonte), avec lames nombreuses plus ou moins serrées dont les bandes d’émail sont paralléles et ne présentent pas de sinus loxodontes médians. Nous y distinguerons deux rameaux ayant vécu parallélement depuis le Pliocéne supérieur jusque dans le Quaternaire: A. Le rameau de l’F. trogontherii Pohlig. B. Le rameau del’ 2. primigenius Blumenbach.”’ Depéret and Mayet continue: “A. RamMuau DE L’ELEPHAS TROGONTHERII.— Jourdan avait reconnu le premier, voici plus d’un demi-siécle, parmi les molaires de Mammouth de la région lyon- naise, une forme aux lames plus épaisses, plus écartées et moins nombreuses que dans le Mammouth normal et lui avait donné le nom d’F. intermedius. Lortet et Chantre (1876) n’ont figuré de cette espéce que le squelette reconstitué du loess de Choulans qui se trouve au Muséum de Lyon et forme le frontispice de leur ouvrage; mais ils n’ont donné malheureusement aucune figure des molaires de l’ £. intermedius, devenu ainsi une espéce purement nominale, qui ne saurait étre retenue.”’ Gaudry (1876, p. 40 also Pl. rx) and Leith Adams (1877, p. 31) observe similar intermediate forms of teeth in Louverné (Mayenne), France, and in England in the Forest-bed, to the middle of the Pleistocene. Finally (Depéret and Mayet, 1923, p. 177): ‘“C’est 4 Pohlig que nous devons d’avoir distingué, parmi les Eléphants quater- naires d’Allemagne, sous le nom d’F. trogontherdi (1889, p. 189), une forme particuliére du groupe des Mammouths, dont les molaires ont une couronne large, comme chez les 2. meridionalis et primi- genius, mais avec un nombre total de lames (14 4 22 aux Mui) supérieur & la moyenne de l|’H. meridionalis (12 4 14), mais in- férieur 4 celle de l’E. primigenius (18 a 27). Cet EH. trogontherit est d’ailleurs absolument identique 4 1’. intermedius de Jourdan, comme nous avons pu nous en assurer sur les nombreux sujets du loess de la région lyonnaise.”’ Thus Depéret and Mayet first concluded that the Hlephas trogonthervi of Pohlig is absolutely identical with the Hlephas inter- medius of Jourdan, but subsequently stated (p. 183) that it belongs THE MAMMONTINA: PARELEPHAS to a more progressive stage, thus agreeing with Osborn’s opinion in the present Memoir. GEOGRAPHIC DISTRIBUTION OF I). INTERMEDIUS-TROGONTHERII Puytum.—Depéret and Mayet (op. cit., p. 180) declare that with the exception of the rare molars of the Upper Pliocene of Astésan, Elephas trongontherii represents a very distinctive Quaternary branch or phylum: (1) “La distribution géographique s’étend de l’Est 4 Ouest depuis la Russie jusqu’a |’Europe occidentale et du Sud au Nord depuis l’Italie jusqu’aux Iles Britanniques. Laissant de cdété les détails de cette dispersion géographique, nous essayerons seule- ment de préciser l’étendue de sa répartition stratigraphique dans le terrain quaternaire.”’ (2) In England the species [i.e., phylum] appears very abun- dantly at the beginning of the Quaternary period in the Forest-bed of Cromer, or Sicilian stage, where it has been described and figured under different names by Leith Adams (1877); it persisted in England to higher Quaternary levels, namely, to the gravels of the ‘terrace of 30 meters (Tyrrhenian stage of Depéret, Chellean industry). (3) In Germany the species FE. trogontherti was first! observed in the ancient gravels at Siissenborn near Weimar, contemporane- ous with the deposits of Mosbach, and of Mauer near Heidelberg; this is 2d Interglacial time of the Osborn-Reeds Table (Pl. xxtv) of the present Memoir; at this time the species appears to be very abundant in Europe. (4) In Europe the species [i.e., phylum] #. trogontherii be- comes extinet in 3d Interglacial time and is replaced by the true mammoth Hlephas primigenius during the period of the Fourth Glaciation. (5) In southern France EZ. trogontherii does not occur in the Sicilian stage (Durfort, Solilhac, Malbattu, Roziéres) in which Elephas meridionalis is abundant. RinGe FormMuta (LETTER, DEPHRET, SEPTEMBER 10, 1924).— “Ces jours derniers, nous avons pu enfin faire avec M. Mayet quelques recherches au Muséum de la ville of se trouvent environ une trentaine de molaires de cette espéce. Malheureusement il y en a fort peu qui soient des M®* intactes et susceptibles de fournir des mensurations. De l’ensemble de mes mesures vous pouvez en toute sécurité inserire le chiffre 4°"=*" pour la formule des M* de Il’ Elephas inter- medius: c’est done une mutation de passage entre le trogontherii et votre H. roosevelti [P. jeffersonii|. J’ai déja eu cette impression dans mon Mémoire sur les Eléphants pliocénes et j’ai été trés sensible a |’éloge que vous voulez bien faire de ce travail.” Parelephas wusti Pavlow, 1909 Figure 945 Upper Pleistocene gravels of Tiraspol (gouv. Kherson), southern Russia. This most welcome discovery by Pavlow (1909.1? — supple- mented by letters of May 31 and August 22, 1924) probably repre- sents an Upper Pleistocene stage about as progressive (M3 5;) as 1065 the Elephas intermedius of Jourdan, namely, M3 3%. It is the most easterly record of this phylum, with the possible exception of the tooth erroneously recorded from China, as noted above, page 1062. This specific stage of Parelephas is certainly more progressive than the P. trogontherii of the 1st Interglacial gravels of Siissenborn, a level containing Dicerorhinus etruscus and Equus stissenbornensis (see Table, p. 1044 above) and now regarded as of Lower Pleisto- cene age, and the ridge-plate formula agrees closely with that of P. intermedius of the plateau loess near Lyons. The upper level of Siissenborn, containing Hlephas trogonthervi type, is attributed to the 2d Interglacial stage, that is, more recent than II Guactat. Pavlow (1910, pp. 6-18), in describing the dentition of Elephas wiisti of Tiraspol, bases her description on a series of superior and inferior molars (Dp 3-M 3) from the Tiraspol collection of Moscow, excellently figured (as reproduced in our Fig. 945) from photographs in PI. 1, figs. 1-12 inclusive. In the same plate (fig. 23) appears a third inferior molar from Tiraspol in the Geological Collection of the University of Moscow which she refers to Hlephas hysudricus. Pavlow at first states (p. 7) that the second molars [= Dp 3] approach those of Elephas meridionalis of the Val d’Arno (Weit- hofer, 1890, Pl. x11) and of Hlephas antiquus of Weimar (Pohlig, 1891, Taf. 2 bis, fig. 8), but on the final page (p. 18) she con- cludes: ‘“‘Prenant en considération que les dents de Siissenborn, décrites par Wiist sous le nom de l’El. trogontheriz Pohl., se dis- tinguent des formes pour lesquelles Pohlig a créé ce nom, nous pro- posons un nouveau nom spécifique de lH. Wiisti pour nos formes, celles de Siissenborn, décrites par Wiist, et les formes de Tiraspol, décrites par Mr. Sinzow.” It is therein suggested that Hlephas wiisti, a species dedicated to Doctor Wiist, is not remote from the Elephas trogontherii Pohl. of Siissenborn. So far as we can judge from Pavlow’s figures (Pl. 1) and description (pp. 6-18) the cotype molars of either side have the following ridge formula: Parelephas wiisti: Dp 3 5% Dp 4 +8 M 1 7333 M 2 7248 a ae 13-14 15-16 Miaieeaat 70 78 210 35 M, 57 67 167 183 31 40 111 Dps 49 133 37 94 A/113707 Neotype _. = All % Natural size R. M9 (Rev) =< he A Florida Phosphate Beds of Charleston 5 C 4 ay L}- v After Leidy A.M 13708 A./4 /3708C R. Dp? /nner view R. M! (Rev) Holmes Coll. L. Dp? R. Op2 (Rev) Holmes Coll. R. Dp, (Rev) A. 41. /3708N R. My (Rev) A. /4./3708Y A./4. 137081 Key To PARELEPHAS COLUMBI Uprrr AND LOWER GRINDING TEETH Fig. 952. Characteristic superior and inferior molars selected from the Cohen Collection, phosphate beds, Charleston, S. C. (Amer. Mus. 13707, 13708 a-z, 13708 aa-kk, 13709), in all thirty-eight specimens, also milk teeth in part, r.Dp*, after Leidy, 1.Dp*, 1.Dpe, from the Holmes Collection, Florida. The key to the grinders in this figure is as follows: Neotype (Amer. Mus. 13707) r.M® (rev.), length 249 mm., breadth 96 mm., 19\be ridge-plates. Neotype (Amer. Mus. 13707) 1.Mz3, length 298 mm., breadth 91 mm., 16e ridge-plates. Referred (Amer. Referred (Amer. Referred (Amer. Referred (Amer. Referred (Amer. Referred 1.Dp*, Holmes Collection, Plorida, 3 ridge-plates. Referred r.Dp2, Holmes Collection, Florida, 3 ridge-plates. Mus. Mus. Mus. Mus. Mus. 13708-c) 1.M’, length 229 mm., breadth 90 mm., ridge-plates 13+ worn, 3 unworn, total 16e. 13708-i) 1.Me, length 210 mm., breadth 74 mm., very aged, preserving 12) ridge-plates. 13708-m) r.M!, length 169 mm., breadth 77 mm., half worn, preserving 1144 ridge-plates. 13708-y) r.My, length 183 mm., breadth 57 mm., one-fourth worn, preserving 13 ridge-plates, 9 worn, 4 unworn. 13708-n), r.Dp,, length 133 mm., breadth 49 mm., preserving 13 ridge-plates, 9 worn, 4 unworn. Referred r.Dp*, after Leidy, in the Wagner Free Inst. Sci., Philadelphia, 10 ridge-plates. The ridge-plate formula of Parelephas columbi of the phosphate beds and Florida, as illustrated in the above key figure, is as follows: Dp 2 3 3 Dp 32° Dp4yzs M171? M2 thts Mg t8hoM 1238 16+ We observe that the three intermediate molars, Dp 4, M 1, M 2, while increasing rapidly in size, exhibit a substantially similar number of ridge-plates below, namely, 13. THE MAMMONTINA: PARELEPHAS LAMINAR FReQquENCY.—The ridge-plates are typically diver- gent at the unworn summit of the crown (5-64 lamine in 10 em.), more contiguous as we descend toward the base of the crown (6-7}5 laminz in 10 em.); in mid-crown the ridge-plates are set widely apart as in Archidiskodon imperator, more widely apart than in Parelephas jeffersonii. In all degrees of age and wear the minimum laminar frequency is 5% in 10 em.; the typical laminar frequency is 6}4 in 10 cm., the maximum laminar frequency is 84% in 10 em. The above variations accord with the point at which the measurement is taken. This explains why certain of these fractured, broad-plated grinders have been mistakenly re- ferred to Archidiskodon imperator by Hay and others, while certain of the narrow-plated molars have been referred to Parelephas jefferson. The grinders (Fig. 954) of the Amherst skeleton belong to the narrow-plated variety. See also Florida specimens of P. floridanus, (Nat. Mus. 11808, 11806, 11810), page 1079 below. PARELEPHAS COLUMBI ABUNDANT IN THE SOUTHEASTERN UNITED STATES This is Hay’s interesting account of the geology and fauna of the important and distinet type of Hlephas columbi Falconer. (Hay, 1923.1, p. 157): “1. Brunswick, Glynn County [Georgia].— This is the type locality of Elephas columbi. This species was based by Faleoner (Quart. Jour. Geol. Soc. Lond., XIII, 1857, table opposite p. 219 [319]) on a part of a tooth received from the geologist Charles Lyell and which had been found in the Brunswick Canal. The specimen consisted of 10 median plates of a lower second or third molar. Falconer figured it in 1868 (Paleont. Mem., vol. 11, pp. 214, 221, plate x). Lyell (Second Visit, ete. vol. 1, p. 348) noted that an elephant had been found in excavating the canal. Richard Harlan, in 1842 (Proce. Acad. Nat. Sci. Phila., vol. 1, p. 189), stated that a large collection of bones of various animals had been presented to the Academy by J. Hamilton Couper, of Darien, Georgia. Among these were teeth of E. primi- genius. Couper, in 1848 ([1846.1] Hodgson’s Memoir, ete., p. 45), stated that two lower jawbones with teeth, several loose teeth, two tusks, and several vertebre of Hlephas primigenius had been collected in the canal during 1838 and 1839. These remains quite certainly belonged to Elephas columbi unless possibly some belong- ed to EH. imperator. Leidy (Jour. Acad. Nat. Sci. Phila., vol. vu, 1869, p. 254) records the presence in the collection of the Academy of a lower molar of #. columbi. The present writer has seen in this collection parts of four teeth of this species which had been sent from the Brunswick Canal, doubtless parts of the Couper collection. The species are listed on page 369. . . [p. 369] The most striking was the great ground-sloth, of the genus Megatheriwm, and which Leidy afterwards called Megatheriuwm mirabile . . . .[p. 370] Elephas columbi (p. 157). Mammut americanum (p. 120). Bison [latz- frons?] sp. indet. (p. 261)... . E. [Hquus fraternus] leidyi (p. 193)... . Megatherium mirabile (p. 36).”’ (Op. cit., p. 370): “J. Hamilton Couper (Hodgson’s Memoir, pp. 37-40) has given an account of the topography and geology of the region through which the Brunswick Canal was being con- structed (map 40). On one of the plates of the work is a section from the ocean westward 21 miles. About 10 miles west of St. 1077 Simon’s Island the canal passed through Six-mile Swamp. This is connected at its northern end with Altamaha River, at the southern with Turtle River. The swamp has thus the appearance of a lake which has become filled with alluvial deposits. These consist of a compact clay, usually yellow and impregnated with iron. There are thin strata of soft, chalky marl and many fragments of petrified wood. At the bottom of this deposit were found the bones of Megatherium, Elephas, Mammut, Equus, and Bison. Beneath the clay stratum was sand with marine shells.”’ From Hay’s faunistic tables of 1914.1, 1923.1, and 1924.1, the following species may be selected as characteristic of the Upper Pleistocene life zone, to which Parelephas columbi probably belongs: FaAuNA OFTHE Typrn LocaLiry FAUNA OF THE PHOSPHATE or EvepHas [= PARELEPHAS| COLUMBI NEAR DARIEN, BRUNSWICK CANAL, GEORGIA Parelephas columbi type Megatheriwm mirabile type Mastodon americanus ref. Equus [fraternus] leidyi type Bison latifrons (2) ref. BEDS NEAR CHARLESTON, 8S. C., WHERE ELepHas [= Par- ELEPHAS] COLUMBI OCCURS IN ABUNDANCE Parelephas columbi ref. Ocalientinus (Serridentinus) obliquidens type Mastodon americanus ref. Bison latifrons ref. Tapirus haysii ref. Uprrr Piersrocenr AGr.—Besides the above and numerous other species, Hay (1923, p. 363) includes many specimens referred to several other species found in South Carolina but not definitely recorded from the ‘Charleston phosphate beds,’’ for example, Archidiskodon imperator ref., which may have been washed in or dredged from an older horizon. Hay, 1914, 1923.—In his Memoir of 1914 and again in 1923 Hay grouped together as one species, namely, ‘“Hlephas columbi”’ (Hay, 1914, pp. 410-421, also 1923, pp. 430, 431, map 12), the mammoths now separated by Osborn into Parelephas columbi [southeasterly range in Fig. 953] and Parelephas jeffersonii [norther- ly range in same figure]. Those in the northerly range probably include some specimens referable to Mammonteus. Osborn, 1928: Osborn determines as Parelephas columbi two of the grinding teeth from the Charleston phosphate beds of South Carolina and from Ocala, Florida, described and figured by Hay, 1914, p. 413, Pl. x1, as follows: Dp? (Ocala, Florida), see Leidy, Trans. Wag- ner Inst. Sci., Vol. II, p. 17, Pl. m1, figs. 6 and 7, also Hay, 1914, p. 413, Pl. Lxt, figs. 2 and 3. Dp* (Ocala, Florida, fide Leidy), length 110 mm.; ridge-plates 8-9 (see Hay, op. cit., Pl. LXI, figs. 5, 6). Nat. Mus. 1614 Dp* (phosphate beds, South Carolina), length 101 mm., height fourth ridge-plate 94 mm. ; ridge-plates 8+ (see Hay, op. cit., Pl. uxt, fig. 4). 1078 OSBORN: THE PROBOSCIDEA PARELEPHAS COLUMBI' OF FLORIDA Mus. 11620), an aged individual recently described by Gidley (see FLoripa.—Parelephas columbi is fairly abundant in Florida; page 1005 above). The true P. columbi, relatively abundant, agrees relatively few specimens of Archidiskodon imperator are recorded very closely in size and ridge-plate formula with Faleconer’s type from Florida, the most perfect being the Venice mammoth (Nat. from Georgia (Fig. 948) and with Osborn’s neotype from the South MNEOTYPE Wisconsin Driftless NORTHERLY CIRCLES = DISTRIBUTION OF PARELEPHAS JEFFERSONIL AND MAMMONTEUS PRIMIGENIUS, SOUTHEASTERLY CIRCLES = DIsTRIBUTION OF PARELEPHAS COLUMBI AND ARCHIDISKODON IMPERATOR Fig. 953. Range of Parelephas jeffersonii and P. columbi, including the type locality (1) of P. columbi in Georgia, also the neotype locality (3) from the phosphate beds of South Carolina. Modified from Hay (1923, map 12, p. 431), who erroneously treated Parelephas jeffersonii and P. columbi as belonging to the same species. ‘Since this Florida section was written (1928) the more progressive Parelephas floridanus stage, M3 Pz , has been described (Osborn, 1930,837). The grinding teeth (Nat. Mus. 11806, 11808, 11810) belong to P. floridanus (op. cit., p. 17). THE MAMMONTINA: PARELEPHAS Carolina phosphates (Fig. 951). Recently described by Gidley, from Melbourne and Vero, Florida, presumably from the ‘No. 2” bed of Sellards, are three P. columbi'! molars: Nat. Mus. 11808. Third superior molar, r.M*, adult; laminar frequency 6-616 ridge-plates in 10 em. Total ridge-plates 19-20, 13 preserved (tooth not complete). Nat. Mus. 11806. A second superior molar, 1.M?; laminar frequency 8 ridge-plates in 10 em., +12+ ridge-plates preserved. Nat. Mus. 11810. A third superior molar, r.M*, aged individ- ual; laminar frequency 7-8 ridge-plates in 10 em. Total ridge-plates 18 to 20, 15 preserved, 3 to 5 missing. Gidley (June 14, 1928) observes that while in Parelephas columbi ref. (Nat. Mus. 11808) the laminar frequency is only 6 measured on the outer side and 6s measured on the inner side of the crown, in 10 em., at the same level, Nat. Mus. 11806 (P. colwmbi ref.) exhibits a laminar frequency at the base of nearly 8 ridge- plates in 10 cm., the compression being much greater. Can such a difference be accounted for on the ground of individual variation? Gidley (letter, December 6, 1928) observes in Nat. Mus. 11806 (P. columbz) 12 ridge-plates plus 7 to 10 missing, a total of 19 to 22, hence an M?*; in Nat. Mus. 11810 (P. colwmbz) he ob- serves 15 worn ridge-plates plus 3 to 5 missing, a total of 18 to 20. Loomis (1923-1928) collected in the No. 2 horizon, latest Pleistocene, near Melbourne, Brevard County, Florida, for the Amherst Museum, the invaluable specimens now assembled in the mounted skeleton (Amherst Mus. 25-1) of the typical Parelephas columbi represented in our figures 954 and 955 and fully described below. AMHERST SKELETON OF PARELEPHAS COLUMBI One of the most fortunate discoveries in the recent history of mammalian paleontology is that of the Amherst skeleton referred to Parelephas columbi, found in 1923 and exhumed in December of that year. It is recorded from the Sellards ‘‘No. 2” horizon, latest Pleistocene, Brevard County, Florida, near Melbourne. Through Prof. Frederic B. Loomis of Amherst College it came into the possession of the Amherst Museum; it bears the number Amherst Mus. 25-1. direction of Professor Loomis, and mounted, as shown in figure 955, about one fifty-sixth natural size. The materials were assembled under the TypicaL oF P. cotumsBi.—The beautifully preserved third superior and inferior grinding teeth in the Amherst specimen (Fig. 954) agree very closely in their ridge-plate formula (M 3 +33) with the typical Parelephas columbi of Georgia and the phosphate beds of South Carolina, in which the prevailing ridge-plate formula isM 3 Test ae SKELETAL CHARACTERS.—Professor Loomis writes (cf. letter, July 28, 1928): In regard to our skeleton of Archidiskodon |Parelephas] columbi the four posterior grinding teeth [Fig. 954] and about 4 feet of each tusk are original, but most of the skull is [See footnote on opposite page.—Editor.] 1079 Mo.aks OF THE AMHERST SKELETON (cr. Fia. 955) Fig. 954. Parelephas columbi (Amherst Mus. 25-1). (Upper figures) Third left superior molar, 1.M*, crown and internal views, Superior and inferior molars found in incomplete skull of About one-third natural size. displaying 18+ ridge-plates, of which 15+ are worn. Height 206 mm. (Lower figures) Third left inferior molar, 1.M3, crown and external views, displaying 16+ ridge-plates, of which 12+ are worn. Length 318 mm.= 12% in. 1080 restored. All the vertebral column, excepting four dorsals, and all of the ribs, excepting a few gaps, were found together. Of the forelimb, the right scapula is complete and half of the left scapula; both humeri were present, the right humerus lacking the head. The left radius and ulna were complete and fortunately the carpals, metacarpals, and phalanges of the forefeet, excepting three digits. Of the pelvic region, the ischium and pubes, and the lower portion of the ilium, including some 12 inches above the acetabulum, are present, but the upper part of the ilium is restored. The hindlimbs, including the femora, tibiz, and left fibula, are complete, also the lower end of the right fibula. It is very interesting to give the principal measurements in comparison with those of Archidiskodon imperator (Amer. Mus. 10598) and of A. imperator maibeni (Neb. Mus. 5—-9-22): OSBORN: THE PROBOSCIDEA var. silvestris, Hl. Columbi var. Falconeri, and El. Columbi var. imperator, based on type grinding teeth. Osborn consequently describes these four subspecies under Parelephas columbi (HL. Columbi var. Felicis) and Archidiskodon imperator, giving facsim- ile reproductions of the broad-plated type figures directly after Freudenberg. We also reproduce herewith (Fig. 901) the figure of a 13-plated grinding tooth after a photograph loaned by Sefiorita Reyes, referable to A. imperator. Osborn, 1928: Probably members of this phylum migrated into Mexico in Upper Pleistocene time, long after A. cmperator had become extinct. Reyes, 1923.—The Proboscidea collections from the Valley of Mexico were described by Senorita Alicia EX. Reyes in her paper entitled ‘Los Elefantes de la Cuenca de Mexico,” 1923, p. 227: NEW FORELIMB ESTIMATES OF PARELEPHAS AND ARCHIDISKODON P. columbi Amherst Mus. 25-1 Ft. Complete height at shoulder from summit of scapula to base of manus, as mounted 11 With fully extended manus l’orrELIMB— Vertical diameter Right scapula 3 Right humerus 3 Left radius 3 Left ulna 3 Manus and carpus fully extended 1 Manus and carpus, as mounted 1 Hinpuiue— Vertical diameter Right femur 4 Right tibia , Fibula 2 Complete length of hindlimb, acetabulum to base of pes, as mounted 8 The above comparative measurements (see also Vig. 912) indicate the following shoulder heights, that i A. imperator Amer. Mus. 10598 A. imperator maibeni Neb. Mus. 5-9-22 In. Mm. Ft. In. Mm. Ft. In. Mm. 3 3430 il 5 3482 12 1% 3699 12 656 3826 4% = 1037 3 4% 1017 3 6 1066+ 4% 10380 3 7% 1095 4 1%, 1251 1% 952 2 11 890 3 5% ~=1060 3 6%, 1085 559 yy, 412 1 645 480 1 454 424e 4%, 1340 [4 8 1422*] 8% 825 8% 825 1% 92477 is, summit of scapula and neural spines of anterior dorsals to the tip of the fully extended digits of the manus. Vt. Skeletal shoulder height hil Height in the flesh 11 PARELEPHAS COLUMBI RELATIVELY RARE IN MEXICO The true Archidiskodon imperator (see p. 1013 above) is evi- dently very abundant in Mexico, whereas Parelephas columbi appears to be relatively rare. Recent descriptions are those of l’reudenberg (1922) and of Senorita Reyes (1923). IREUDENBERG, 1922—We owe to this author a full history of this much misrepresented species (see his Memoir, p. 138. “T. Elephas Columbi Falconer und seine Rassen. Geschicht- liches.””). On pages 138-140 he clearly distinguishes this species but wrongly, as we believe, compares (Fig. 18, p. 140) Falconer’s types of Blephas columbi and FE. armeniacus with a second inferior molar from Mexico. After thus comparing the broad-plated grinding tooth of #. |= Parelephas| columbi with the narrow-plated grinder of #. [= Parelephas| armeniacus, he proceeds to describe a number of new subspecies, namely, Ll. Columbi var. Felicis, El. Columbi *lemur, Neb. Mus. 13-24-10-14 and 4-12-13. In. Mm. Ft. In. Mm. Tt. In Mm. 3 3430 11 5 3482 12 654 3826 114% 3644 12 1% 3702 13 AY, 4068e “Tntre los numerosos mamiferos fésiles de la Cuenca de México, quizA ninguno tan ampliamente repartido como el ele- fante. Sus restos se encuentran en mayoria en nuestros museos, y son incontables las instituciones extranjeras que conservan en sus colecciones molares de Hlephas procedentes de la cuenca. Estos restos, s6lo por excepcion han sido debidamente clasifi- cados, entre otras causas, porque las dos especies a que pertenecen, I). imperator y E.Columbi, presentan muchas formas de transici6n. Este hecho me ha inducido a revisar los ejemplares existentes en los museos de la Ciudad, con el objeto de intentar una clasificaci6én, basAndome en los datos asi obtenidos.”’ In her valuable paper she fully describes and figures five speci- mens of Hlephas imperator and five specimens of 2. columbi chiefly from the localities of Tequixquiac, from the region of Zumpango, at Tepexpan, in the deposits above the basalt emitted by the volcano THE MAMMONTINA: PARELEPHAS AMHERST MUS. 1081 AMHERST SKELETON OF PARELEPHAS COLUMBI FROM NEAR Mevpourne, Fiorma About one fifty-sixth natural size Vig. 955. Loomis of Amherst College in the No. 2 horizon, latest Pleistocene, Brevard County, Florida, near Melbourne. While the specimen was found in 1923 and taken up that December, it did not come into the possession of the Amherst Museum until 1925. Materials for the mounted skeleton of Parelephas columbi, bearing the number Amherst Mus. 25 -1, were collected by Prof. Frederic B. Skull mostly restored, also portions of both tusks, four dorsals, parts of the limb bones, ilium, manus, and pes; otherwise largely original (see below). Superior and inferior molars shown in figure 954. of Chiconautla, and in Villa de Guadalupe. Her observations relat- ing to localities and to #. columbi are as follows, those relating to FE. imperator are presented in abstract above. Elephas |= Parelephas| columbi. Escuela de Ingenieros No. 2. R.M?* with 18 ridge-plates, 14 in 25em.; length 280 mm., breadth 110 mm., height 200 mm. From Tequixquiac. Fig. 10, p. 236. Instituto Geologico (without number). R.Ms, with 1516 ridge- plates in 25 cm.; length 290 mm., breadth 102 mm., height 180 mm. From Zumpango. Instituto Geologico 213. Fragment of skull and erroneously restored left tusk; also both third molars, length of r.M* 350 mm., breadth 115 mm., height 190 mm. Total ridge-plates 1915; 16 in 25em. From Tequixquiac. Fig. 11, p. 237. Instituto Geologico 214. Inferior mandible with 1.M;; 16 to 17 ridge-plates in 25 cm.; length 268 mm., breadth 90 mm., height 190mm. From Villa de Guadalupe. Fig. 12, p. 238. Escuela Ingenieros No. 3. L.M®, with 21 ridge-plates, 20 in 25 cm.; length 275 mm., breadth 88 mm., height 190 mm. From Tequixquiac. Fig. 13, p. 239. GEOLOGIC AGE OF THE PROBOSCIDEA FAUNA OF MEXICO See Cope, 1884.2, “The Extinct Mammalia of the Valley of Mexico,” also Furlong, 1925, “Notes on the Occurrence of Mammalian Remains in the Pleistocene of Mexico,” ete., and Villada, 1903, “Apuntes Acerca de la Fauna Tésil del Valle de México.” Reyes, 1923.—This author concludes (1923, p. 239) with a comparison of Parelephas columbi and Archidiskodon imperator and a discussion of the observations of Leidy, Osborn, and Falconer. She dismisses the observations of Barcena and Castillo (1882-1884) as to the human remains being contemporaneous with the mam- moth at Tequixquiac, from which locality both P. colwmbi and A. imperator are recorded. It is not stated whether P. columbi is geologically more recent than A. imperator, as is probably the case. Reyes writes (June 12, 1924): “The little data which we have concerning the stratigraphy of the Valley [of Mexico] seem to confirm your hypothesis that H. columbi is more recent than ZL. imperator; but they are insufficient. It is not possible to correlate Quaternary strata of the Valley with “las Nordicas,’’ because glacial periods or extremely cold periods do not exist. Diaz Lozano informs me that the flora is constantly tropical. All the examples which I have examined seem to me to belong to the two species columbi and imperator. More intensive study would perhaps permit a subdivision into varieties.”’ FAUNAL Succression.—A brief summary of our present knowl- edge of the Proboscidea and other mammals of this region in Upper Pliocene and in Pleistocene times is as follows: PLEISTOCENE Parelephas columbi: Tequixquiac, Zumpango, Villa de Guadalupe. Archidiskodon imperator: Tepexpan, Tequix- quiac, Zumpango, Villa de Guadalupe. PLiIoceNE oR Cordillerion tropicus (cf. Mastodon humboldti’). PLEISTOCENE Cordillerion oligobunis Cope, Tequixquiac. Cordillerion oligobunis progressus Freudenberg, Canyon of Aculeingo. 1082 PLIOCENE Cordillerion oligobunis antiquissimus Freuden- berg, Hidalgo, Valley of Amajaque. Cordillerion(?) oligobunis felicis Freudenberg, Puebla. Cordillerion(?) oligobunis intermedius Freuden- berg, Mexico. Hipparion castilli [Protohippus| Cope. Rhynchotherium tlascale Osborn, type from Valley of Mexico, neotype from Sonora. PLIOCENE PLIOCENE OR PLEISTOCENE PLIOCENE Other Pleistocene and Upper Pliocene faunas are the following (Cope, 1884, Villada, 1903, Furlong, 1925): Equus {curvidens Owen], E. tau Owen, E. crenidens Cope, E. barcenei Cope, E. occi- dentalis Leidy, ref. Bison latifrons Harlan, ref. Camelops hesternus Leidy, ref. Platygonus (?)compressus Le Conte, ref. Tequixquiac: Capromeryx mexicana Furlong. Ainocyon dirus. Teleoceras (?)fossiger Cope, ref. [Pliocene]. Glyptodon |mexicanus]. Nothrotherium. Neotoma, near occidentalis, ref. Near Monte- rey: Furlong concludes (1925.1, p. 152): ‘“{1] The mammalian remains obtained at the four localities, Tequixquiac, Zumpango, Saltillo, and Monterey, in central and north-central Mexico, are indicative of the large and varied mammalian faunas that will be found in the Pleistocene deposits of Mexico when intensive work is carried on. [2] The two types of deposits recognized in the quarry at E] Tajo, near Tequixquiac, and the mammalian forms found in each of these accumulations, indicate apparently two distinct stages in the Pleistocene history of this region. [3] The presence of Anocyon dirus, a coyote-like form, and of Capromeryx mexicana in the brecciated fissure deposits suggests a faunal horizon compar- able perhaps to that found at Rancho La Brea, California. The assemblage of mammals thus far found in the beds overlying the fissure deposits indicates a somewhat later horizon in the Pleisto- cene. [4] The paucity of mammalian remains found in the cave- fissure deposits in Cerro de la Silla offers little opportunity to add to our knowledge of the upland mammalian fauna of the Pleis- tocene or to make comparisons with the faunas from the Cali- fornia caves. The presence of Nothrotheriwm sp., Equus sp., and Neotoma, near occidentalis, a form related to the recent wood rat, are close to the types found in Samwel Cave, Shasta County, California.” Parelephas columbi felicis Freudenberg, 1922 Figure 956 Pleistocene, Tecamachalco, Puebla, Mexico Freudenberg’s excellent figure and description of the type, an r.M*, which he named 4. Columbi var. Felicis, enables us to relate this type to the genus Parelephas, a species probably more progressive than the typical P. colwmbz. OSBORN: THE PROBOSCIDEA El. Columbi var. Felicis Yreudenberg, 1922. “Die Siiugetier- fauna des Pliociins und Postpliociins von Mexiko,’’ Geol. und Palaeont. Abhand., N. F., 1922, Band XIV, Heft 3, pp. 147- 152. Typrn.—Third right superior molar, r.M*. Original in Leipzig (Pal. Coll. Univ. Leipzig 4403), Coll. Felix. Steppenform. Horizon anp Locauiry.—Tecamachalco, Puebla, Mexico. Pleis- tocene. Typr Fiacurre.—Op. cit., Taf. vu (xvi), fig. 4: “Elephas Columbi var. Felicis nov. forma.” Type Derscription.—(1925.1, p. 148): “Ein von Tecama- chaleo . . . im Staate Puebla stammender Molar, No. 4403 der paliontologischen Sammlung der Universitat Leipzig, gleicht eben- falls sehr einem solchen von El. primigenius, unterscheidet sich jedoch durch eine etwas bedeutendere Dicke der Schmelzlagen und ist vielleicht schon der folgenden Art zuzurechnen; doch stehen die Schmelzbiichsen ausserordentlich gedringt. Das gleiche gilt fiir einen ebenfalls in unserer Sammlung befindlichen Zahn von Ejutla im Staat Oaxaca [Footnote: ‘Unsere Textfigur 19a und 19b. El. Columbi var. silvestris.’|. Herr Prof. Dr. Felix hatte die Liebenswiirdigkeit, mir aus dem paliontologischen Museum der Universitit Leipzig diesen Zahn neben einigen anderen aus Mexiko a CN 3 hseaa = Typr OF PARELEPHAS COLUMBI FELICIS Fig. 956. Type, r.M®, of Elephas Columbi var. Felicis Freudenberg, 1922, Taf. vin (xvi), fig. 4, one-half natural size. From Tecamachalco, Puebla, Mexico. Original in Leipzig (Pal. Coll. Univ. Leipzig 4403), Coll. Felix. Observe that of the 14+ ridge-plates exposed by wear there are 94 in 10 em., slightly exceeding the maximum number attributable to this species as noted in the specimens of the Cohen Collection from the Charleston phosphate beds. zur Untersuchung zu senden. Es handelt sich bei dem Elefanten- zahn von Tecamachaleo um einen M* des rechten Oberkiefers. Er trigt die Katalognummer 4403 des dortigen Museums. Der Zahn hat aussen weisse Farbe, welche auch das Innere aufweist, und oberfliichlich einzelne Himatitflecken. Diese Art der Erhaltung begegnete mir nur bei einigen mitteldiluvialen Fossilien, z. B. von San Luis.” CuHaraActERS.—Freudenberg (op. cit., pp. 147-152) points out the confusion by Cope and Felix of the races of Elephas columbi Fale. with those of 2. primigenius; he also points out the charac- ters of Falconer’s type of 2. columbi; following Falconer’s estimate of 1863, p. 56, he gives (p. 141) the ridge formula of H. columbi as: M 145, M 234, M 32°55. He concludes (p. 145) that H. columbi Fale. is related to 2. imperator Leidy; he then proceeds to describe a number of races or varieties of these species of “Hlephas columbi,” of which E. columbi var. Felicis is the first mentioned; he distin- guishes this subspecies by its closely compressed lamellae as most THE MAMMONTINA:: PARELEPHAS nearly resembling #. primigenius, in contrast to a second subspecies El. Clowmbi var. Falconeri (ef. A. imperator) in which the lamellze are less compressed. Osborn, 1930: The relatively compressed ridge-plates (see legend of Fig. 956) together with the estimated ridge formula of M 3 2° may range this subspecies Parelephas columbi felicis be- yond the limit of the typical P. columbi (M 3 ;*3;) into the ridge- plate estimates of the more progressive species P. floridanus (M 3 224) or P. jeffersonii (M 3 3%). Parelephas columbi cayennensis Osborn, 1929 Figures 957, 979 Cayenne, French Guiana, South America. Probably Upper Pleistocene. As set forth on page 1046 of the present chapter, the sole elephant so far known to have reached South America is a sub- species of Parelephas columbi reported by Captain Perret and successively mentioned by Lartet (1859), Lull (1908), and Freuden- berg (1922), located in the Marseille Museum by Doctors Laurent and Repelin in 1929, and finally described by Osborn (1929.797, p. 20) as Parelephas colwmbi cayennensis. Lartet (1859, p. 500) characterized this specimen as ‘‘un fragment de molaire 4 lames épaisses,’”’ which led Freudenberg (1922, pp. 159, 160) to refer it to El. [Archidiskodon] imperator, from which it proves to be positively distinguished by the very narrow molar crown as described below. Parelephas columbi cayennensis Osborn, 1929. ‘New Eurasi- atic and American Proboscideans.’”’ Amer. Mus. Novitates No. 393, December 24, 1929, pp. 20, 21. Typr.—‘Three and a half ridge-plates of a third right superior molar, r.M®, collected by Captain Perret in Cayenne (French Guiana), South America, and now preserved in the Muséum d’Histoire Naturelle, Marseille, France, as No. 8449 (cast Amer. Mus. 21933).”’ HorRIzON AND Locauiry.—Probably Upper Pleistocene. Cayenne, French Guiana, South America. Tyre Figure.—Op. cit., p. 21, fig. 20. SpeciFic CHARACTERS.—The superior type fragments, photo- graphs and casts of which have been kindly furnished the present writer through the courtesy of Director W. Laurent of the Muséum d’Histoire Naturelle of Marseille, and Prof. W. Repelin, Conserva- teur, is characterized by Professor Repelin (translation of letter of March 30, 1929) as follows: ‘‘They were in rather bad condition and so badly cemented together that I had to take away the larger part of the cement. One of these fragments, A—1, is represented in side view on one of these photographs. The lamellze are very worn and they show the plate of blackish or brownish enamel rather in relief and the ivory forms a small median depression. The cement which separates the lamelle is not very thick in this specimen. In the other photograph, A—2, the specimen is shown as seen from above. Finally another fragment, B, is also represented as seen from above. It has been worn in the direction of the lamellz, but this worn part shows only a very irregular section of ivory and enamel.” The fragment consisting of three and a half lamelle or ridge- plates is made the type of the new subspecies Parelephas columbi cayennensis. These ridge-plates appear to belong to the posterior portion of the crown of a third superior molar of the right side, r.M*; they are strongly concave posteriorly and are composed of 1083 coarse enamel, deeply grooved or crenulated on the sides; the external cement, formerly present, has been dissolved or worn away; the apices of the three ridge-plates present a convex profile, hence supporting the superior molar reference indicated by the posterior concavity. They correspond broadly with ridge- plates 16, 17, and 18 of an r.M$ of Parelephas columbi; in size they correspond with ridge-plates 19, 20, and 21 of floridanus; they seem relatively narrow, thus agreeing with the narrow ridge- plated P. columbz rather than with the broad-plated Archidiskodon imperator. Geographically the nearest representative of Parelephas columbi cayennensis to the north is the Parelephas columbi felicis Freudenberg, 1922, from Puebla, described on the opposite page 1082 and illustrated in figure 956 of the present Memoir, in which the posterior ridge-plates are broader, with less coarse enamel, than in the present type specimen. Type or PARELEPHAS COLUMBI CAYENNENSIS OSBORN Az POST. Al Fig. 957. Three aspects (Al, A2, B) of the fragmentary type, r.M%, after cast (Amer. Mus. 21933) taken from type speci- men in the Marseille Museum No. 8449. One-half natural size. Compare Osborn, 1929.797, fig. 20 (Fig. 979 of the present Memoir). Al, A2, Lateral and inferior or root aspect of supposed ridge-plates 16, 17, and 18-%. B, Crown view of ridge-plate, partly worn, exhibiting thick enamel. Ye Nat. size Parelephas jeffersonii Osborn, 1922 Figures 806, 810-814, 818, 892, 930, 931, 935-987, 958-968, 975, 988, 1009, 1029, 1084, Pl. xxm Upper Pleistocene, post(?)-Wisconsin, IV Guacrat till (fide Leverett and Hay, 1923); Jonesboro, Indiana. Syn.: Hlephas roosevelti Hay, 1922. Compare also Elephas [Parelephas]| jacksoni Mather, 1838, which may prove to antedate Parelephas jeffersonii. This well-known and very important specific stage, genotypic of Parelephas, is widespread throughout the Middle United States and gives us a complete knowledge of the skeletal and dental characters. The ridge-plate formula is: Dp 3 * Dp 413 M12 M 2222 M 3 33 9e 1084 Hisrory oF THE NAME.—This species is very abundant in the latter part of the Pleistocene period in the northern United States and southward into Mexico; the type may even be of Postglacial age; it is the animal erroneously described by Cope and Osborn as “Elephas columbi,” and by Hay as “FE. primigenius,” and finally separated by Osborn as Elephas jeffersonii. It now becomes the most fully known species of American mammoth. ‘This species was described by Osborn July 8, 1922. Hay subsequently (Sept. 30, 1922) proposed the name Hlephas roosevelti. Mather in 1838 gave the name Elephas jacksoni to a similar animal, the type of which has been lost rendering the species indeterminate at present. In 1924 Osborn made Elephas jeffersonii the genotypic species of the new genus Parelephas. Elephas jeffersonii Osborn, 1922. ‘Species of American Pleis- tocene Mammoths. Elephas jeffersonii, New Species.’’ Amer. Mus. Novitates No. 41, July 8, 1922, pp. 11-16. TyPE.— Skull, jaws, and greater part of skeleton (Amer. Mus. 9950). Horizon AND Locauiry.—Upper Pleistocene, post(?)-Wisconsin, IV Guactat till; Jonesboro, Indiana, on the farm of Dora 8. [E.] Gift. Tyre Ficgurr.—Osborn, op. cit., 1922.555, p. 11, fig. 10. ConFusion as TO ParatyprEs.—(1) A pair of upper and lower grinding teeth of both sides (Amer. Mus. Warren Coll. 10457) from Zanesville, Ohio, erroneously selected by Osborn (1922.555, pp. 11, 12, also figs. 11 and 12) as paratypes; (2) subsequently (1923.601, p. 4) referred by Osborn to Elephas washingtonii; (3) finally (1924.633, pp. 4 and 7) made by Osborn the type of a new subspecies, Parelephas jeffersonii progressus (=Parelephas progressus); (4) new paratype [ideotype, see foot- note on p. 1068] jaws (Amer. Mus. 13225, 21892) selected by Osborn in the present Memoir. History OF SPECIFIC AND GENERIC SEPARATION BY OSBORN.— The following type description is reprinted in part from Osborn’s paper cited above (Osborn, 1922.555). Osborn introduced the subject as follows (p. 1): “The present article relates explicitly to the type characters of Hlephas columbi, of EL. imperator, and of the American specimens referred to EL. primigenius, three species which have become more or less confused in all the previous literature because the characters of the type specimens have not been precise- ly determined and compared. The object of the present com- munication is to clear up this confusion and to propose Elephas jeffersoni as a new species of American Pleistocene mammoth. .. . [Op. cit., p. 15] The American elephant heretofore widely known as ‘Blephas columbi, the Columbian Mammoth, will hereafter be known as Elephas jeffersonii, the Jeffersonian Mammoth.” OSBORN’S SPECIFIC (1922) TYPE DESCRIPTION AND TYPE FIGURES OF PARELEPHAS JEFFERSONII (Osborn, 1922.555, p. 11): “The above diagnoses of the real specific characters of the grinding teeth of [the true] Hlephas columbi. .. . and of the true 2. primigenius leaves without a name the animal which previously has been described in all the literature (excepting Soergel’s recent Memoir) as Elephas ‘columbi.’ This animal is better known than either of the others; it is represented in all the collections of the principal museums of the United States, as described by Hay (1914), and particularly in the American Museum by four especially fine specimens. Of these we select as the type Amer. Mus. 9950, including the skull, jaws, and greater OSBORN: THE PROBOSCIDEA part of the skeleton (Fig. 10), found near Jonesboro, Indiana, on the farm of Dora 8. [E.] Gift; purchased for the Museum with the Jesup Fund in 1904, restored and mounted in 1906; described by Osborn in 1907 as Hlephas colwmbi, determined [and described] by Hay (1914) as Elephas primigenius.” (Op. cit., p. 15): “CRANIAL CHARACTERS.—Still more obvious are the differences between the relatively long, broad, and shallow crania of H. jefferson and the relatively short, narrow, and deep crania of #. primigenius, proportions which are correlated respec- tively with the corresponding proportions just described and figur- ed in the teeth.” “The four complete skulls of this species known to the writer are those of (1) in the type mounted skeleton (Amer. Mus. 9950); (2) the fine male skull associated with the jaws and a large part of the skeleton (Amer. Mus. Cope Coll. 8681) from Whitman County, Washington, and labeled ‘Hlephas colwmbi’ [since referred to Par- elephas washingtonii|; (3) the young male skull (Amer. Mus. Cope Coll. 14475) from Dallas, Texas, also labeled ‘Hlephas columbi’ [now referred to Archidiskodon imperator]. (4) To these should be added the very large male skull (Nat. Mus. 10261) collected near Cincinnati, Ohio; in this specimen the ridge formula is M 3 *4; seventeen plates were in use; the compression of the grinding teeth is greater, namely: 9 ridge-plates in 100 mm. on the outer [convex] side, at the worn edge, 9 ridge-plates in 100 mm. on the worn mid-coronal surface. The cranial characters of this specimen are entirely similar to those of the three skulls in the American Museum collections, except that it is larger and more robust.” (Op. cit., p. 12): “This typical American species is named in honor of the third president of the United States, Thomas Jefferson, in commemoration of his long-continued devotion to mammalian paleontology.” Typr Locauity AND post(?)-Wisconsin Aap (IV GuaciAL) OF PARELEPHAS JEFFERSONI Typr.—The geologic age of the type of Parelephas jeffersonii is a very important matter; according to Leverett and Hay, the Glacial or Postglacial deposit where it was found is of post(?)-Wisconsin age, i.e., [V GLActAL. (Hay, 1923, p. 139): “6. Fairmount, Grant County [Indiana]. Here was found, in 1904, the nearly complete skeleton of the mammoth mounted in the American Museum of Natural History in New York City. It has been described and figured [as Hlephas primigenius] by the writer (36th Ann. Rep. Geol. Surv. Indiana, p. 718, figs. 63, 64 [1912.2]; Iowa Geol. Surv., vol. XXIII, p. 396, fig. 133) [1914.1]. It was found on the farm of Mrs. Dora C. [E.] Gift, about 4 miles east of Fairmount. The location is in the southeast quarter of section 23, township 23 north, range 8 east. This information has been furnished by Mr. George Swisher, surveyor of Grant County.” “This whole region is mapped by Leverett as being occupied by ground moraine of till plains, and the animal must have lived after the Wisconsin ice cleared away. A tract more or less morainic, an extension of the Union City moraine, is indicated by Leverett on his latest map as passing further south than Fairmount. At the earliest it must have been after the withdrawal of the ice from the Union City moraine that the animal lived. Considering the character of the surrounding country, the nature of the deposit THE MAMMONTIN A: inclosing the skeleton, and the depth at which it was buried, it might be supposed that it was not long after the formation of the Union City moraine that this elephant existed.” GENOTYPIC SPECIES OF PARELEPHAS (OSBORN, 1924.633).— After the establishment (Osborn, 1922.555) of EHlephas jeffersonii as a species distinct from Hlephas columbi and from Elephas primigenius, Osborn finally (1924.633) reached the conclusion that Elephas jeffersonii could be placed neither in the phylum of Archidiskodon nor in the phylum of Mammonteus (e.g., Mam- monteus primigenius) and for the reasons above recited he selected it as the genotypic species of the new genus Parelephas. SPECIFIC CHARACTERS OF PARELEPHAS JEFFERSONITL Ossporn’s Error (1922.555) as tro RipGe FORMULA BASED ON ZANESVILLE PARATYPE (SINCE MADE THE TYPE OF PARELEPHAS JEFFERSONIL PROGRESSUS OsBoRN, 1924).—In the above original description of Hlephas (Parelephas) jeffersonii, Osborn erroneously PARELEPHAS 1085 and described by Warren in 1855 (p. 168, Pl. xxvin, fig. C) as Elephas ‘Primigenius.’”’ It appears (Osborn, 1924.633, pp. 4, 7) that these Zanesville ‘paratype’ molars are typical of a progressive mutation of Parele- phas with a higher ridge formula, namely, M 332. To this sub- species the name Parelephas jeffersonii progressus was assigned. It becomes Parelephas progressus in the present Memoir. DENTAL CHARACTERS OF PARELEPHAS JEFFERSONII Ripce FormuLts.—It has been a very difficult matter to determine the ridge formula of the species P. jeffersonii; as noted above, the high ridge formula (M 332) originally given by Osborn, as based on the original paratypes from Zanesville, Ohio, was erroneous; the finally corrected ridge formula of the type, new paratypes, and referred specimens of Parelephas jeffersonii is: M 332. PARELEPHAS JEFFERSONIL RIDGE-PLATE COMPRESSION AT THREE LEVELS OF R.M3 One-third natural size Fig. 958. Right third lower molar, r.M3, of Parelephas jeffersonit new paratype [ideotype] (Amer. Mus. 13225). Observe: (1) The arcuate arrangement of the typical twenty-three ridge-plates of this partly worn molar. By counting the ridge- plates at different levels, as indicated by the horizontal lines A, B, C, D, we observe that the upper plates are much more closely compressed than the lower plates, as follows: A, Summit of crown, 111% plates in 10 cm. B, One-sixth below summit, 10 plates in 10 cm. C, Four-sixths below summit, 7) plates in 10 cm. D, Near base of crown, 6% plates in 10 em. selected as a paratype two upper and lower grinders from Zanesville, Ohio, contained in the Warren Collection (Osborn, 1922.555, p. 11): “As the paratype of this species we select a pair of upper and lower grinding teeth of both sides (Amer. Mus. 10457) acquired with the Warren Collection in 1906 and described by Warren in 1855 (p. 163, Pl. xxviny, fig. C) as Elephas ‘Primigenius.” In 1923, Osborn corrected this error (1923.601, p. 4), but remained in doubt as to the true relationship of the Zanesville teeth: ‘‘Related to this species [Hlephas washingtonii]| may be a pair of upper and lower grinding teeth of both sides from Zanesville, Ohio (Amer. Mus. Warren Coll. 10457) acquired with the Warren Collection in 1906 The type dentition (Amer. Mus. 9950) is that of a very aged male (Fig. 959), as indicated first by the remarkable length (= 11 ft. 4\¢in. or 3.47 m.), incurvature, and crossing of the tusks (Fig. 966) ; second by the fact that in the third superior and inferior molars the anterior ridge-plates are worn off. Consequently Osborn was unable to give the type ridge formula correctly in his type descrip- tion (1922.555) and made the error of depending upon the errone- ously associated Zanesville paratype; in his second description, Osborn (1923.601, p. 4) wrote: ‘Since the original description of Elephas jeffersonii was written the molars [Fig. 959] in the aged type specimen (Amer. Mus. 9950) have been cut out of the jaw of the 1086 OSBORN: THE PROBOSCIDEA skull and carefully sectioned; it has thus been found that they are namely: M 335; [;34]. There can be no question that the type relatively short and deep and present a different ridge formula, of EF. jeffersonii has a shorter jaw and shorter and deeper molar teeth than those of the erroneously associated paratype which may now be referred to #. washingtonii.”’ The result of sectioning the molar teeth in the aged type speci- men proved that the anterior ridge-plates were entirely worn off; the aged type ridge formula may therefore be written: M 3 ait The adult ridge formula, however, is M 3 25. Al E. jeffersonii Osb. Amer. Mus. 9950 Type (outer view) L. M®* Lie Bere sre 34 a rats AN QW) E. jeffersonii Osb. Amer. Mus. 9950 Type ops SA © 7G = E. roosevelti Nat. Mus. 2195 Type E. jeffersonii Osb. SS Amer. Mus. 9950 Type \— . S SS fouter view) ~ NAN R. Ms (rev.) SSSR E. roosevelti Nat. Mus. 2195 Type ria 16 ) 7 We a us Ss we " > f E. jeffersonii Osb. Amer. Mus 9950 Type (inner) view R. M 3 All }4 nat. size E. jeffersonii Osb. Amer. Mus. 9950 Type Tig. 959. (Lerr) Acep Tyre Turrp Superior AND INFERIOR Mowars oF PARELEPHAS JEFFERSONIL (Riaur) THE SAME SUPERPOSED ON ELEPHAS ROOSEVELT! [=SyN, PARELEPHAS JEFFERSONII] (Left) Aged type superior and inferior molars of Parelephas jeffersonii (Amer. Mus. 9950), viewed upon the outer surface with the anterior portion of the crown to the left. Superior molar, M®, partly encased in the maxilla, consequently the fangs are not displayed. Inferior molar, Mg, completely removed from the jaw and sectioned to show the incomplete posterior ridge-plates. Al, Outer view of left superior molar, 1.M*, 1-4 anterior ridge-plates worn off; remaining ridge plates (5-17) exposed; 17 ridge-plates in use. Total estimated ridge-plates 25. Bl, Right inferior molar, r.Mg; (rev.), 4 anterior ridge-plates worn off; 18 ridge-plates in use (1-18); incomplete ridge-plates 19-20 demonstrated by vertical and horizontal sections of this tooth. Total estimated ridge-plates 24. B2, Inner aspect of right M3, showing 18 ridge-plates in use (1-18); incomplete ridge-plates 19 and 20, at posterior border. Total estimated ridge- plates 24. (Right) Diagram showing in heavy lines type superior and inferior molars of Parelephas jeffersonii projected upon the type superior and inferior molars of Elephas roosevelti (light lines). This diagram was constructed by superposing type figure 959, left, upon type figure 968. This diagram shows: (1) That the young adult molar teeth of Z. roosevelti (light lines) exhibit a ridge formula of M 3 $3, whereas the aged molar teeth of P. jeffersonii (heavy lines) exhibit a ridge formula of M 3 {4%; (2) the anterior ridge-plates in this species are worn off before the posterior ridge-plates attain their full length; (8) in the type of F. roosevelti 10-13 ridge-plates are in use, in the type of P. jeffersonii 17-18 ridge-plates are in use; (4) the long diameter of M*is oblique, while the long diameter of My is horizontal; (5) following the rule in the Elephantide, approximately the same number of ridge-plates is in use at the same time in the superior and inferior molars, namely, in the young ZF. roosevelti +3; in the aged P. jeffersonii ~7'y. (6) The worn ridge-plate surface of M* is convex; that of Mg is concave. THE MAMMONTINA: PARELEPHAS It is proven by comparison of the aged type teeth (Fig. 959) of Parelephas jeffersonii with the adult type teeth (Fig. 968) of its synonym Elephas roosevelti, that in the aged type of P. jeffersonii five to six ridge-plates have been worn off in front of the upper molars and four ridge-plates in front of the lower molars, by ex- treme use. This is shown diagrammatically in figure 959, in which the aged teeth of the type of P. jeffersonii are superposed upon the adult teeth of the type of EH. roosevelti. This diagram also illustrates that the anterior ridge-plates begin to be worn off while the posterior ridge-plates have not attained their full depth and are still growing. Correctep RipGr FormuL#.—Fortunately it is now possible to verify this interpretation of the ridge formule of these two types and positively to establish the adult ridge formula of Par- elephas jeffersonii (EF. roosevelti syn.) by comparison of five different specimens as follows: 1087 RipG@e-PLaTE Count at DIFFERENT Crown Levets (Fic. 958.—(1) As observed in the diagram and figure (Figs. 958, 959) the ridge-plates are arcuate and the ridge-plate arches converge toward the summit, the greatest number of ridge-plates may be counted at the swmmit of the unworn crown, e.g., 11}5 ridge-plates in 100 mm., a ridge-plate compression approaching that of Mam- (2) As we descend down the side of the crown (B), we may find 10 ridge-plates in 100 mm. (3) As we reach the point of maximum expansion of the ridge-plate arches (C), we may count 7}5 ridge-plates in 100 mm. In general the compression is greater in the short superior molars than it is in the long inferior molars; consequently we get the highest count in the superior teeth. (4) This difference in the compression of the arcuate ridges explains the discrepancy in the records of the “laminaire fréquence” of Depéret and the ridge-plate counts of Hay. Similar discrepancies are observed in the ridge-plate com- monteus primigenius. In Ms, 17 ridge-plates In Msg, 12 ridge- Ridge formula: M 3 sq;. In M3, Ridge formula: M 328. In M?, 17 ridge-plates Indiana Type skull and jaws of P. jeffersonii (Amer. Mus. 9950), aged ridge formula: M 3 {33. in use, 3 rudimentary. Kansas New paratype! jaw of P. jeffersonii (Amer. Mus. 21892), a young jaw. Ridge formula: M 3s. plates in use. Kentucky New paratype! jaw of P. jeffersonii (Amer. Mus. 13225), a middle-aged jaw. 14-15 ridge-plates in use. Ohio Referred skull of P. jeffersonii (Nat. Mus. 10261), adult male. in use. Illinois Type grinding teeth of Hlephas roosevelti Hay (Nat. Mus. 2195). Ridge formula: M 333. Thus from the recurrent evidence of five different specimens, young, middle-aged, adult, and extremely aged (i.e., type), we conclude as follows: Typical adult ridge formula of Parelephas jeffersonii type and four referred specimens: M 3 #2. TypicaL RipGE-PLATE Compression.—Osborn (1922.555, p. 12) carefully described the ridge-plate compression in the type inferior molars of Parelephas jeffersonii (Amer. Mus. 9950) as follows: 7 ridge-plates in 100 mm. on the convex internal surface. 7-714 ridge-plates in 100 mm., obliquely worn on mid-coronal surface; 17 ridge-plates in use. 8 ridge-plates in 100 mm. on the concave external surface. Thus the number of the ridge-plates in the type of Parelephas Jeffersonii is: 7, or 8, or 9 in 100 mm., depending upon where the count is taken. [See footnote on page 1068 above.—Editor.] pression of Archidiskodon and of Mammonteus. According to the observations of Hay (1914), as accepted or modified by Osborn, the ridge-plate compression characteristic of different specimens of Parelephas jeffersonvi may be summarized as follows: lod In the inferior molars: 7 ridge-plates (at base) to 11% (at summit of crown) in 100 mm. In the superior molars: 7 ridge-plates (at base) to 10 (at sum- mit of crown) in 100 mm. REFERRED RIDGE FORMULA OBSERVED BY HAY AND OSBORN The majority of the teeth from the middle and northern United States described as “Hlephas columbi”’ by Hay (1914, pp. 410-421) actually belong to Parelephas jeffersonii and are readily distinguishable from the teeth of the true type of Hlephas columbi Fale. of the Southern States as well as from the true teeth of Mammonteus primigenius and of Archidiskodon imperator. Flimi- nating six of Hay’s specimens from Afton, Oklahoma, which may belong to A. imperator Leidy, also three specimens from the phos- 1088 OSBORN: THE PROBOSCIDEA OTHER SPECIFIC CHARACTERS OF THE TrrntaH (Hay, 1914, p. 395, Ossporn, 1922-1924).—(1) Ridge-plates more widespread than those of #. primigenius. (2) Enamel plates thicker and more channeled and crimped. (3) Superior ridge-plates transverse or concave posteriorly; inferior ridge-plates transverse or concave phate beds of South Carolina, which may belong to the true Elephas columbi Fale., Hay’s detailed observations on teeth prob- ably or possibly (Alaska?) referable to Parelephas jeffersonii or to P. washingtonii (Washington, Oregon, Alaska) may be summarized as follows: Kentucky Phil. Acad. Dp‘ (Big Bone Lick, Ky.), length 145 mm., width 75 mm., height of first plate 143 mm.; ridge- plates 12, front and rear talons. ‘A line 100 mm. long passes across eight of these plates, a number greater than one might expect in this species;” Alaska Nat. Mus. 6669 Dp, (Alaska, Yukon), length 180 mm., width 67 mm., height 185 mm.; 13 ridge-plates. ‘“There are seven ridge-plates in a 100 mm. line.” Montana Nat. Mus. 6052 Dp, (Glendive, Mont.), length 155 mm., width 75 mm.; ridge-plates 8+. ? Nat. Mus. 287 M2 (Loe. unree.), length 300 mm., width 78 mm., height of thirteenth plate 195 mm.; ridge- plates 18 or 19; eight plates ina 100 mm. line; amount of cement between the plates small. Iowa Univ. lowa 167 M2? (Logan, Iowa), length 192 mm., width 70 mm., height of eighth plate 160 mm.; eight plates in a 100 mm. line. Alaska Nat. Mus. 6668 Mo (Alaska), length 277 mm., width 90 mm., height of sixth plate 140 mm.; ridge-plates 19; seven ridge-plates in a 100 mm. line. Oregon M$ (Mt. Angel, Clack. Co., Ore.), length 355 mm., height of thirteenth plate 193 mm.; ridge- plates 21+; 5} ridge-plates in a line 100 mm. long. Indiana Neb. Mus. M; (Tipton Co., Ind.), length 280 mm., height of fourth plate 152 mm.; ridge-plates 22+-. anteriorly, i.e., dise shaped or bent. (4) Rear or last superior molars, M°’, usually strongly arched. (5) Superior ridge-plates, M$, not known to exceed 25 to 26; inferior ridge-plates of M; not known to exceed 23 to 24. (6) Bony sheaths of superior tusks To the above eight specimens observed by Hay from Kentucky, Iowa, Indiana, Montana, Oregon, and Alaska, we may add the following six specimens observed and included by Osborn in the species Parelephas jeffersonii or P. washingtoni: Mus. 9950 (type skull) M*, M; (Jonesboro, Ind.), type aged male, anterior ridge-plates worn off. M 3{37. Mus. Indiana Amer. 8681 (referred Parelephas washingtonii skull) M?* (Whitman Co., southwest Washington), male: M?, 10+ ridge-plates; M’, 23 ridge-plates. (See Fig. 976.) Washington Amer. Kentucky Amer. Mus. 13225 (paratype jaw) M; (Twin Cr. near Sanders, Ky., Big Bone Lick region), middle aged, paratype of P. jeffersonii: Ms, 24 ridge-plates. Kansas Amer. Mus. 21892 (paratype jaw) Mz; (near Pendennis, Lane Co., Kan.), young adult, paratype of P. jeffersonii: Ms, 24 ridge-plates. Ohio Nat. Mus. 10261 (referred skull) M* (Cincinnati, O.): M/°, 26 ridge-plates. Illinois Nat. Mus. 2195 (type teeth) M*, M; (Ashland, Cass Co., Ill.). Type of Elephas roosevelti: M°, 25 ridge-plates; Mz, 24 ridge-plates. (See Fig. 968.) The type and referred ridge formula of Parelephas jeffersonii is: Dp 2? Dp 35* Dp 445 M 1277 M 2182" M 3 34. According to these fourteen specimens, as described by Hay and Osborn, the average specific ridge-plate frequency of Parelephas jeffersonii in 10 em., as compared with that in Mammonteus primigenius, Archidiskodon imperator, and Parelephas columbi, is as follows: Mammonteus primigenius and M. primigenius compressus: 8-10-11-12-13 Parelephas jeffersonii: 7—8- Archidiskodon imperator: 5- Parelephas columbi: 5-6 9 of shorter and more divergent than those of Hlephas primigenius. (7) Superior tusks in young males diverging rapidly, in middle age beginning to rotate, in adult males converging, so that in aged males the tips cross. DrTAILED STRUCTURE OF THE THETH.—The detailed structure of the superior and inferior teeth of Parelephas jeffersonii is beauti- fully shown in the following figures of this Memoir which have been prepared with the utmost care and precision: Fig. 959. Vertical view of the aged superior and inferior molars of the type of Parelephas jeffersonii (Amer. Mus. 9950). Fig. 960. Superior view of the type and paratype [ideotype] inferior grinding teeth and jaws of Parelephas jeffersoni. THE MAMMONTIN#: PARELEPHAS Tig. 968. Exterior and crown views of superior and inferior grinding teeth of the type of Hlephas roosevelti (Nat. Mus. 2195). Fig. 959. Diagram showing superior and inferior grinding teeth of type of Parelephas jeffersonii and of Elephas roosevelti. Fig. 967. Middle-aged paratype [ideotype]| jaw of Parelephas jeffersonvi (Amer. Mus. 13225). Elephas jeffersonii Osb Amer. Mus, 9950 Type All 1/8 nat. size Elephas jeffersonii \ Amer. Mus. 13225 Paratype \ 1089 the type jaw (Amer. Mus. 9950) also the internal aspect of the middle-aged paratype [ideotype] jaw (Amer. Mus. 13225). Comparisons.—These comparative figures, which have been prepared with the greatest care to a uniform one-eighth and one- fourth scale, will enable the student to distinguish these jaws of Parelephas jeffersonit very readily from those of Archidiskodon imperator and somewhat less readily from those of Mammonteus Elephas jeffersonii Amer. Mus. 21892 Paratype JAWws OF PARELEPHAS JEFFERSONII FROM INDIANA, KENTUCKY, AND KANSAS One-eighth natural size Fig. 960. Superior view of type and paratype [ideotype] grinding teeth and jaws of Hlephas jeffersonii. [=Parelephas] jacksoni (Fig. 946). Compare (B) with Mather’s type of Hlephas A, Ideotype young adult jaw (Amer. Mus. 21892) with Mg in situ, eight plates in use; M3 with twelve plates in use; total number of ridge-plates in M3 twenty-four. From near Pendennis, Lane Co., Kansas. Compare Amer. Mus. 14558 (Archidiskodon imperator ref.—Fig. 889). B, Ideotype middle-aged jaw (Amer. Mus. 13225), Me shed, M3 with fourteen to fifteen plates in use; total number of plates in Mg; twenty-four; jaws somewhat more massive. From Twin Creek, near Sanders, Kentucky, Big Bone Lick region. C, Type of aged Parelephas jeffersonii Osborn (Amer. Mus. 9950); jaw with M3; worn to the edge of the anterior fang; one or possibly two plates have been worn off, seventeen plates in use, three posterior plates shown in section (Fig. 959, right). Jaws still more massive. Vig. 892. Comparative figure of jaws of Parclephas jeffer- soniz and other species. Tig. 958. Right third lower molar paratype [ideotype] of Parelephas jeffersonii (Amer. Mus. 13225). JAWS OF PARELEPHAS JEFFERSONIT The first to distinguish the jaw of Hlephas [= Parelephas| jacksoni from the jaw of Elephas [= Mammonteus| primigenius was Mather in his description and figure (Fig. 946) of 1838. When all the jaws figured in the present chapter, both in superior and lateral view, are compared with the Jaws figured in the Mammonteus chapter (Chap. XVIII), the latter, it will be observed, are shorter, deeper, and more compressed anteroposteriorly, while relatively broader, with a broader groove above the rostrum. SPECIFIC CHARACTERS OF THE JAWs.—Certainly belonging to this species are the three specimens shown from above in figure 960, namely, the aged type (Amer. Mus. 9950), the paratype [ideotype] middle-aged jaw (Amer. Mus. 13225) from the Big Bone Lick region, Kentucky, and the paratype [ideotype] young adult jaw (Amer. Mus. 21892) from Lane County, Kansas. The lateral and superior views of the middle-aged paratype [ideotype] are shown in detail in figure 967. In figure 892 is shown the internal aspect of From Jonesboro, Indiana. primigenius. (1) The jaws in all these species of Mammontine are correlated with the brachycephaly and acrocephaly of the cranium, that is, the jaws are excessively short and deep. (2) The ramus in P. jeffersonii (see Fig. 892 D, D1, C, C1) is far less robust than the ramus in A. imperator (Fig. 892 B, B1, A, Al). (3) The ramus of P. jeffersonii with rounded inferior border differs from that of Elephas indicus (Fig. 893 D, D1) in which the inferior border is more pointed. (4) The deeply depressed ramus of P. jeffersonai is readily distinguished from the horizontal ramus of Loxvdonta africana (Fig. 893 C, C1) or from the relatively shallow ramus of the Elephas [= Parelephas] washingtonii type (Fig. 893 B, B1). Rostrum.—The most characteristic feature is the prominent rostrum perfectly preserved in the paratype [ideotype] of P. Jjeffersonii (Fig. 960 B) which projects conspicuously beyond the line of the chin. This prominent rostrum apparently distinguishes the Parelephas jeffersonti jaw from the Mammonteus primigenius jaw. This rostrum is the feature especially shown in Mather’s type figure (Fig. 946) of Hlephas jackson7; it is also pointed out in Mather’s type description which leads us to believe that #. jackson is more closely related to Parelephas jeffersonii than to the true Mam- monteus primigenius, although in certain specimens referred to E. primagenius the rostrum is quite prominent, so that we cannot place too great reliance on this character. 1090 TYPE AND REFERRED SKULLS OF PARELEPHAS JEFFERSONII We may fortunately compare five finely preserved crania of different ages referable to this species in American collections with a number of European crania referred to Hlephas intermedius (Fig. 935) and to E. trogontherii (Fig. 934). Thus we may establish the fundamental similarity of cranial structure which distinguishes all the species referable to the genus Parelephas. CRANIAL CHaraActTeRs (Fias. 934, 935, 961-963).—As point- ed out in the introduction to the Mammontine, superficially the profile aspect of the cranium of Parelephas jeffersonii (Fig. 962A) resembles that of Elephas indicus, but the midcranial section shows that it is fundamentally distinct. The cranial form of P. jeffersonii is actually intermediate between the cranial form of Archidiskodon imperator and that of Mammonteus primigenius. Close comparison with M. primigenius is afforded in figure 962, in which two crania, one of P. jeffersonii (A) and the other of P. washingtoni (B) are OSBORN: THE PROBOSCIDEA shown on the right side of the profile in comparison with a single cranium of M. primigenius (C,C!). In this comparison it is ap- parent that M. primigenius is even more brachycephalic and acrocephalic than P. jeffersonit or P. washingtonit. COMPARISON OF PARELEPHAS AND MAMMONTEUS CRANIA We observe (Fig. 961) that in the frontal aspect these three crania of Parelephas are remarkably similar and readily distinguish- able from crania of Mammonteus (Fig. 962 C). In frontal aspect: (1) The occipital crest, which is perfectly preserved in the aged type skull from Indiana (A) is broad and swelling at the summit; it is also well preserved on the right side in the adult skull of the specimen from the state of Washington (B); it is erroneously restored in the specimen from Ohio (C). (2) The anterior narial openings are very broad and widely open, whereas in Mammonteus they are smaller and more contracted. (3) The orbital sockets are FRONTAL Views or Ture Parevernas Crantra: (A, C) P. serrersonu, (B) P. wWASHINGTONII All figures one-twelfth natural size Tig. 961. Front view of the (A) type skull of Parelephas jeffersonii Osborn (Amer. Mus. 9950), from Jonesboro, Indiana, compared with one skull (B) in the American Museum and one (C) in the U. S. National Museum, as referred to in the original type description. A, Front view of the aged type skull of Parelephas jeffersonii (Amer. Mus. 9950); actual measurement 28.8 inches across the outside of the orbits. B, Referred skull of Parelephas washingtonii (Amer. Mus. Cope Coll. 8681) from Whitman County, Washington, heretofore designated as “‘Hlephas columbi,” and subsequently referred by Osborn (1922.555, p. 15) to Parelephas jeffersonii. This skull was never figured or described by Cope, it was found in an unopened box in his collection. For full description of this specimen, see pages 1101, 1103, and figures 937, 973, 974, 971, and 976 of the present Memoir. C, A very large male skull in the National Museum (Nat. Mus. 10261), collected near Cincinnati, Ohio, referred to P. jeffersonit in Osborn’s type descrip- tion (1922.555, p. 15): “The cranial characters of this specimen are entirely similar to those of the three skulls in the American Museum collections, except that it is larger and more robust.’”’ Occiput incorrectly restored. THE MAMMONTIN#: PARELEPHAS much more prominent than in M. primigenius, as perfectly shown in all three skulls (A,B,C). (4) The maxillo-premaxillary sockets of the superior tusks are relatively less elongate and less parallel than in Mammonteus; they diverge widely where the tusks issue from the skull. (5) In profile view (Fig. 962 B) the orbits are more widely separated from the occipital condyles than in M. primigenius (Fig. 962 C). (6) In profile view the occiput is more prominent and bulging (Fig. 962 B) than the relatively vertical occiput of M. primigenius (Fig. 962 C). (7) In its proportions the M. primigenius cranium is broader, shorter, higher, deeper, i.e., more brachycephalic and hypsicephalic, than the P. jeffersonii cranium. (8) In all these characteristics the cranium of P. jeffersonii appears to agree more closely with the crania of the European species Parelephas intermedius (Vig. 944), and P. trogontherii as shown in profile in figure 934, than it does with either profile or front views of the cranium of Mammonteus primigenius. SKULL OF PARELEPHAS JEFFERSONII (KNOWN AS THE FRANKLIN COUNTY MAMMOTH) IN THE NEBRASKA STATE MUSEUM Discovered in the loess of Wisconsin or late Pleistocene time. NEBRASKA SKULL.—Since the crania above described come from the states of Indiana, Ohio, and Washington, it is interesting C1 1091 to add the description and figures of the superb skull and tusks from Nebraska preserved in the Museum of the University of Nebraska (Neb. Mus. 14-15), photographs of which (Figs. 963 and 964) we are able to reproduce through the courtesy of Prof. E. H. Barbour. Like the two crania from Indiana and Ohio previously described, this Nebraska skull was originally referred by Professor Barbour to ‘‘Elephas columbi,” but it agrees in all particulars with the two crania above described as Parelephas Jeffersonvi and is the largest and in many respects one of the finest representatives of this type of cranium, the tusks exceeding in length and in circumference those of the type skull of P. jeffersonii from Indiana. The left profile view of this Nebraska cranium (Neb. Mus. 14-15) is shown in figure 963; the profile is closely similar to that of the type of Parelephas jeffersonii, except that the concave plane of the forehead is relatively more elongate; we observe that the orbital region is very prominent, the summit of the occiput is rounded, the postoccipital profile is very convex, the occipital condyle is elevated; the entire profile is similar to that of the male skull of Parelephas figured by Falconer (lig. 9384). The palatal view (Fig. 963) displays the third superior grinding teeth in situ and exhibits (lig. 965) nineteen ridge-plates, of which the anterior MAMMONTEUS PRIMIGENIUS (C, C!), PARELEPHAS JEFFERSONII (A), P. wAsHINGToNt (B) All figures one-twelfth natural size Pig. 962. Profile views of type and referred skulls of Parelephas jeffersonit (A), P. washingtonit (B), and Mammonteus primigenius (C) in the American Museum and United States National Museum, also front view of M. primigenius (C'). A, Type of Parelephas jeffersonii (Amer. Mus. 9950), reversed. An aged individual. Main portion of the tusks not included. Irom Jonesboro, Indiana. B, Referred skull of Parelephas washingtonii (Amer. Mus. Cope Coll. 8681). From Whitman County, Washington. C, Referred skull of Mammonteus primigenius (Nat. Mus. 8580) from Siberia, with jaws belonging to another individual (Nat. Mus. 8579), from Alaska. C!, Front view of same skull and jaws. c60T ‘yuossaffal svydajaivd 04 84881}109 PUB soouR[quiaser ay} aAresqQ, *(“uI"B'e) snyBoul AIOPPNB [BUII}X9 oY} PUB ‘AZIS [BINYBU YIYSIE-OUO Te Burztqryxe ‘(6TSE “BIL ‘qdaq ‘snyv, “loury) snoypur spydapgy JO WNIWBIO JO MOTA [BTEC “(44 31y]) “BUTUIEIOJ [BIUBIO 9} PUB SPITE]? qgueuodur0d 94} 008 Fe 8 Dit CRO EGIY OO Oy OMG ROMY OSS (OsCHONs CoC ETT CHAOS IS SLIT SSL soyoie aryeuosAZ sso10B {UTIAIMSBITA “UIUL CCST punoj oq [ITM 4390} ey} PUB UINTUBIO sty} jo wor ‘jorg Aq ydvrdojoyd Iezyy “ST6T -F-T SDI GN) syedse (eTPPIa d) V¥SVUEAN FO HLONWVINT XINDOXD soreu 1olseysod ay} SuIpnpout BO FOS CS IIS ACO ea Byixeurord Jo aseq 07 yndt900 jo ySato WOTF qUdUIdINSBIT] ydiosep Y ‘azis [eIMyeU YzUO}-9UQ “SHSNF rotedns ay} JO S}ax908 ssao] 9} UT pateAoosiq, JO G96 PUB F96 samay eredut0g “E601 PU® T6OL sesed uo ‘Tt Judy ‘eysiqen ‘AzuNOD aTyTeLT qeqduep eeu ) pereyey 4JeT pur (4301) yereyed ‘2uosuaffal soydajaivg “96 “BL NITHNVUY AHL AO VINVU) ‘uoumtoads ures otf} Arey] xeul pus Areyrxeureid pepuedxa Ayyeor3 ayy eAlasqQ §=“Moqieg “H “A ‘ajejduioo sysnq pue ‘e[qIpueul SISNWIVONGH SMOIGNI SVHAATH qmyg “qouureur Ayuno0H upyUBAy ayy se uMouy ‘(CT 40 GNV (IINOSUMMIaL! SVHIATAUY, 49H 6IBE SOW VEY (! | snoipur "3 THE MAMMONTINA: PARELEPHAS thirteen are in use. Behind these thirteen more or less worn ridge- plates (Fig. 963) are observed seven to eight unworn ridge-plates making an estimated total of twenty to twenty-three ridge-plates. The widely divergent maxillo-premaxillary region presents the usual contrast with the parallel and convergent bony sockets of the Mammonteus primigenius type of cranium. Comparison is afforded in figures 899 and 900 with the palatal view of two crushed skulls of Archidiskodon imperator from Hay Springs, Sheridan County, Nebraska (see Chap. XVI). Locauity AND GroLoaic Lrevet.—This skull was discovered from 11 to 12 feet underground while digging for the foundation of the new high school of Campbell, Franklin County, Nebraska; the date of discovery was April 1, 1915. The geologic deposit was a bed of loess which we may infer belongs to late Pleistocene times. As restored under Professor Barbour’s direction and mounted in Tue FRANKLIN County Mammotu oF NEBRASKA Fig. 964. Nepraska SkuLL.—Complete skull, jaws, and tusks of Parelephas jeffersonit (Neb. Mus. 1—4—15) discovered in the loess near Camp- bell, Franklin County, Nebraska. After photograph by E. H. Barbour. Skull essentially without restoration. Part of the coronoid region of the jaw restored in plaster; symphysis of the jaw perfect; both mandibular rami broken. Tips of the tusks weathered off and restored with characteristic curvature. One-tenth natural size. the University of Nebraska Museum, the skull and tusks are shown in side view in figure 964. The tusks are 12's feet in length and 29 inches in circumference at the thickest point; the cranium and jaws are essentially without restoration; the enormous depth of the skull from the occipital condyle to the inferior border of the jaw is shown by the seated figure of a student whose forehead rests just opposite the occipital condyle. TYPE SKELETON OF PARELEPHAS JEFFERSONILI See figures 931 and 966 of mounted skeleton The type skeleton of Hlephas jeffersonzi was found near Jones- boro, Indiana, in 1903. It was purchased for the Museum with the Jesup Fund in August, 1904, from Dora E. Gift and others. As found the skeleton was imbedded in a muck deposit of late Pleisto- cene age, fifteen feet below the surface; this deposit is considered of IV GuactaL, post(?)-Wisconsin age, by Leverett and Hay. The whole upper portion of the skeleton was complete and intact, including the remarkably incurved tusks, which were preserved for their entire length although requiring surface restoration and 1093 thorough soaking, and reinforcing with an internal steel rod in plaster to prevent them from crumbling away. Vigorous efforts were made to find the missing lower bones of the limb and the feet but without success. MounTING OF PARELEPHAS JEFFERSONII TypE.—The missing parts are the radius and ulna of both sides, the right tibia and fibula, the fore- and hindfeet. These parts are restored from the more massive limbs of two specimens of Archidiskodon and it is doubtful whether they are of the proper proportions for Parelephas. Thanks to the kind coéperation of Dr. Marcellin Boule of the Museum of Paleontology, Paris, casts were secured of the lower portions of the limbs and of the feet of the great skeleton of Elephas meridionalis (Fig. 866) in the Paris Museum; on arrival in the American Museum these casts were remodeled, recourse being had for comparison to remains of the feet and limbs of Archidiskodon Moar RIDGE-PLATES OF FRANKLIN County MAMMOTH OF NEBRASKA. AGED PARELEPHAS JEFFERSONIL Fig. 965. Franklin County Mammoth. Diagram drawn from casts showing the (13) superior and (16) inferior worn ridge-plates of Parelephas Jjeffersoni ref. (Morrill Coll., Neb. Mus. 1-4-15). Outlines furnished by Professor E. H. Barbour, August 29, 1924. (Upper) Crown view of M?, M%. (Lower) Portion of the crown of M3 of the same specimen. See figure 963 for cranium of this specimen before restoration, figure 964 after restoration. imperator; all this work of restoring and remodeling was done by Otto Falkenbach under the direction of head preparator Adam Hermann. Mr. Hermann himself restored the missing surfaces of the upper parts of the skeleton which were, however, in exception- ally complete condition. Posr.—The animal is represented with uplifted head and as if just starting to walk. For the pose of the skeleton the codpera- tion of Director Hornaday of the Zoological Park was secured; the tracks of the Indian elephant ‘“‘Gunda” slowly walking over a sandy surface fifty feet long were carefully plotted by park engineer Beerbower (Fig. 1243 of this Memoir); through this means it is believed that the fore- and hindfeet are correctly placed. The author also made a special study of the position of the bones of all parts in the living elephant. The result of all this work is a pose and gait which are believed to be true to life. The most striking features of this skeleton of an aged in- Type SKELETON OF PARELEPHAS JEFFERSONII IN THE AMERICAN MusEuM One twenty-fourth natural size Fig. 966. First published type figure of the aged skeleton of Elephas jeffersonii Osborn, 1922 (Amer. Mus. 9950), as mounted in the American Museum. This specimen, including the skull, jaws, and greater part of the skeleton, was found near Jonesboro, Indiana, on the farm of Dora E. Gift; purchased for the American Museum with the Jesup Fund in 1904; parts of limbs restored and mounted in 1906; described and figured by Osborn in 1907 (1907.295) as Elephas columbi; redescribed by Hay (1914) as “‘Elephas primigenius,” and again in February, 1923, p. 139, as E. primigenius of post(?)-Wisconsin age. This skeleton was finally selected and figured by Osborn in 1922 (1922.555, p. 11, fig. 10) as the type of Hlephas jeffersonii new species. (Cf. Osborn, 1907.295, p. 256): As found the skeleton was imbedded in a muck deposit of late Pleistocene age, fifteen feet below the surface. The whole upper portion of the skeleton was complete and intact, including the remarkably incurved tusks, which were preserved for their entire length although requiring surface restoration and thorough soaking, and reinforcing with an internal steel rod in plaster to prevent them from crumbling away. Vigorous efforts were made to find the missing [restored] lower bones of the limb and the feet but without success. The missing parts [restored] are the radius and ulna of both sides, the right tibia and fibula, the [both] fore-and hindfeet. Thanks to the kind cooperation of Dr. Marcellin Boule of the Museum of Paleontology, Paris, casts were secured of the lower portions of the limbs and of the feet of the Zlephas meridionalis (lig. 866) in the Paris Museum; on arrival in the American Muscum these were remodeled, recourse being had for comparison to remains of the feet and limbs of Elephas imperator; all this work of restoring and remodeling was done by Otto Falkenbach under the direction of head preparator Adam Hermann. The animal is represented with uplifted head and as if just starting to walk. For the pose of the skeleton the cooperation of Director Hornaday of the Zodlogical Park was secured; the tracks of the Indian elephant ‘(Gunda” (Chap. XXIII, p.1598) slowly walking over a sandy surface fifty feet long (Fig. 1243 of the present Memoir) were carefully plotted by park engineer Beerbower; through this means it is believed that the fore- and hindfect are correctly placed. The author also made a special study of the position of the bones of all parts in the living elephant. 1094 THE MAMMONTINA: dividual of Parelephas jeffersonii are the following: (1) The com- plete incurvature and crossing of the tusks, indicating that this animal is an old bull which had retired from the herd. (2) The relatively small size of the skull. (3) The abbreviation of the back and body in contrast to the vertical elongation of the limbs. Dietrich (1916, p. 76, 1924, p. 13 — p. 1276 below), in describ- ing Hlephas antiquus recki, discusses the size of the humerus of recent and fossil proboscideans and gives a number of measure- ments which we may compare with those of the type of Parelephas jeffersonii as follows: Estimated Shoulder Height Length of Humerus Parelephas trogonthervi of Mosbach 1480 mm.! 4500 mm. [Schmidtgen, 1926, p. 64] [Parelephas jeffersoniz type 1085+- 3200] Hesperoloxodon antiquus of Taubach 1300 Mammonteus primigenius 1270 Palxoloxodon recki 1235 4000 ca.? PARELEPHAS OF INTERMEDIATE SIZE VERTEBRAL CoLuMN.—The vertebral formula of this generic and specific type is here shown in comparison with that of speci- mens of Loxodonta, of Elephas, and of Mammonteus: Loxodonta africana Flower, 1885 “Jumbo” Amer. Mus. Dept. Mam. 3283 @envicalste ker fcc Sat oe octet 7 a DOT SAS eee rine cttco eee 19 20 RairsiofenibS sameeren tee a: um bars’ rc essac sete een er 4 3 Sacral smerarcense vcewta ties: 5 4 Caudal stays eres. ats se eesseio es 24+ 21 The size of the animal is indicated by the following compari- sons with the great Archidiskodon meridionalis of the Paris Mu- seum: Length, incurved tip of tusks to vertical line of tail......... Length, base of tusks to vertical line of tail................ Height at withersi(toitopofiscapula))).s...-....-5..-25 554: [cone throlmemuUlamryere tates ronnie ses saya seem Ane eer 11 ft. 4% in. (8470 mm.) Length of right tusk (outside curve)...... 440 mm., fide Schmidtgen, 1926, p. 63. PARELEPHAS 1095 Compare also the fully adult skeleton of Parelephas jeffersonti with the recently described titanic skeleton of Archidiskodon maibeni (Figs. 910, 912). We reach the interesting conclusion that the adult Parelephas of the north temperate regions of Eurasia and North America is midway in size between the giant Archidisko- don imperator of the more southerly latitudes and the relatively dwarfed Mammonteus primigenius of the northerly latitudes. Broadly speaking the proportions of these members of the sub- family Mammontine may be estimated from the extreme height at the withers as follows: 12 ft. 6% in. =3826 mm. 10 ft. 6 in. =3200 mm. 9 ft. 3 in. =2820 mm. Archidiskodon imperator maibeni Parelephas jeffersonaz Mammonteus primigenius ELEPHAS ROOSEVELTI HAY, 1922 This Pleistocene species from Ashland, Illinois, based on su- perior and inferior molar teeth, r.M,’ r.M; (Nat. Mus. 2195), was described by Hay September 30, 1922. Hay observes (op. cit., p. 100): ‘The peculiarity of all these teeth is the low elongated form and the approximate parallelism of the upper and the lower bor- ders.” ‘This species appears to represent a stage of evolution very close to that shown in the type and paratype teeth of Hlephas Jeffersonii described by Osborn about two months previous, namely, July 8, 1922. Consequently we may regard Elephas roosevelti as a synonym of Parelephas jeffersonii. Elephas roosevelti Hay, 1922. “Further Observations on Some Extinct Elephants,” Proc. Biol. Soc. Washington, Vol. XXXV, Sept. 30, 1922, p. 101. Elephas indicus —Mammonteus primigenius Parelephas Flower, 1885 Falconer and Felix, 1912 Jeffersonii Amer. Mus. Type 14559 (M. p. Amer. Mus. compressus) 9950 7 7 7 7 19-20 18-19 19 19 of 18-19 19 19 5-3 4-3 5 4 4 4-3 4 5 24-30+ 21 21(Zalensky) 12+ The author’s designation of the type specimen, locality, formation, and characters of Hlephas roosevelti, new species, is as follows (Hay, op. cit., p. 101): A. meridionalis Paris Mus. (Measurements given by Gaudry) 22 ft. 3%, in. (6800 mm.) 17 ft. 10% in. (5450 mm.) 12 ft. 6% in. (8830 mm.) 4 ft. 5% in. (1360 mm.) P. jefferson Amer. Mus. 9950 ee Eee 17 it. 9% in. (5430 mm.) Rear ofa BY, 13 ft. 35 in. (4050 mm.) screen eae 10 ft. 6 in. (8200 mm.) 4 ft. 1 in. (1250 mm.) 74030 mm., fide Schmidtgen, 1926, p. 64; see Osborn’s estimate of 3600 mm. (p. 1277 below). 1096 Elephas jeffersonit Amer. Mus. 13225 Ref. Elephas jeffersonit Amer. Mus. 13225 Ret. OSBORN: THE PROBOSCIDEA A 1/4 nat. size PARATYPE OF PARELEPHAS JEFFERSONII Fig. 967. Jaw of middle-aged paratype [ideotype] of Parelephas jefferson (Amer. Mus. 13225) from Twin Creck, near Sanders, Kentucky, Big Bone Lick region. ridge-plates (see Fig. 960B, showing 24 ridge-plates) of which the anterior The rudiment of Mg has been recently shed; M3 exhibits twenty-three fourteen are in use. “Type specimen. Upper and lower hindermost molars, No. 2195, U. S. National Museum. Type locality. Ashland, Cass County, Illinois. Type formation. Pleistocene. Diagnosis. Hindermost molars long and low, the base and the summit approxi- mately parallel, consisting apparently of 25 plates; of these 8 in A E. roosevelti Nat. Mus. 2195 Type Al E. roosevelti Nat. Mus. 2195 Type (outer view) / 15 dM ff it I if ha iy | Ld ipyt rh i Li f IV pe iy f if \ Lady BI E. roosevelti Nat. Mus. 2195 Type (outer view) E. roosevelt! Nat. Mus. 2195 Type 10 PARPLEPHAS JEFFERSONIL (SYNONYM ELEPHAS ROOSEVELT!) Turrp Supprior AND INFERIOR MOLARS Fig. 968. Figured for the present Memoir. Type superior and inferior molars of Elephas roosevelti Hay (Nat. Mus. 2195), from Ashland, Cass County, Illinois. One-fourth natural size. A, Al, Crown and external views of the right superior molar, r.M#, showing twenty-five ridge-plates (1-25 in the figure). B, B1, Crown and side views of the right inferior molar, r.Mg, showing ridge-plates 3-24, the anterior ridge-plates being broken away. THE MAMMONTIN/A: PARELEPHAS a 100 mm. line; enamel thin, delicate, and little folded. figure. None. See figure 968 of the present Memoir].”’ Type Description OF ELEPHAS ROOSEVELTI.—The type description was introduced by the following notes: (Hay, Sept. 30, 1922, p. 100): ‘Professor Osborn in his paper of July 8 [Osborn 1922.555] has published an interesting figure of upper teeth of an elephant (his fig. 8) [‘Fig. 8. Fourth and third superior molars of . . . Elephas primigenius’’| found in Indiana. On plate LIx of the twenty-third volume of the lowa Geological Survey, I published a figure of very similar teeth found at Milwaukee, Wis- consin, and preserved in the Public Museum of that city. The hindermost molar had just begun to suffer wear. In the U. 8. National Museum are right and left hindermost molars (No. 2195) of similar form, found at Ashland, Cass County, Illinois; also an upper left hindermost molar (No. 4761) hardly different, discovered in Wayne Township, Darke County, Ohio. The peculiarity of all these teeth is the low elongated form and the approximate parallel- ism of the upper and the lower borders. Inasmuch as the molar descends at a nearly right angle with the grinding face of the tooth in front it seems probable that the skull was short. Professor Osborn has referred his specimen to Hlephas primigenius; but I find no teeth from Alaska or the Old World which present similar characters. I believe that a hitherto unrecognized species is in- dicated. This I propose to call Hlephas roosevelt? in honor of another statesman and naturalist, one whose multifarious interest led him to pursue living elephants in their African wilds.”’ “The Ashland teeth are chosen because with them came the nearly complete lower right hindermost molar. The length of the molars is close to 300 mm., the height 170 mm., the width of the upper teeth 90 mm., of the lower 85 mm.” Osborn, 1924: Hay’s type description, in which he compares this animal with #. primigenius, was not accompanied by a figure and for the purposes of this Memoir the type teeth of Hlephas roosevelti (Nat. Mus. 2195) were kindly loaned to the American Museum by Curator Merrill for examination and execution of the type figure which is reproduced herewith (Fig. 968). This type figure, including its very careful enumeration of the ridge-plates as preserved, namely, 24-25 ridge-plates preserved above, 21 ridge-plates preserved below, agrees with the type de- scription of Hay. The mid-coronal ridge-plate count is 8 in a 100 mm. line. The diameters of the molars are as follows: [Type Left Right Third superior molars, length across QOMIALEe=plateShy +. eke eee ete 227 mm. 282 mm. Third superior molars, height of tallest ridge-plate............ 202 192 Third superior molars, max. width 95 98 Third superior molars, breadth- lengthyindexse seen eee ane Third inferior molars, length, esti- tooth incomplete mated, of 22 ridge-plates...... 304e Third inferior molars, height of tall- estimidce=platereener errr rare 154 Third inferior molars, max. width 92 Third inferior molars, breadth-length INGEX RRS eet he ea eee tooth incomplete 1097 The diameters, ridge-plate measurements, and indices of these young molars closely resemble those of the aged molars of the type of Elephas jeffersonii (Fig. 959, left) as clearly shown in the diagram (Fig. 959, right), in which the superior and inferior molars of the ag- ed E. jeffersonii type are projected to the same scale upon the type superior and inferior molars of the young adult LZ. roosevelti. Con- sequently we have been able to amplify the specific characters of E. jeffersonii by a study of these grinding teeth of EF. roosevelt, which compare so closely in all respects except in age and wear. The ridge-plate compression is as follows: Left Right M* ridge-plates in 100 mm. on convex externalisurfaces cere nhac ae 915 9 ridge-plates in 100 mm. on mid- coronallsuriacesseeee ae ace ee 10 gis ridge-plates in 100 mm. on mid- coronal concave internal surface. 8 9 M; ridge-plates in 100 mm. on mid- coronal concave external surface. 7 7-8% ridge-plates on worn mid-coronal surface, thatis,in100mm..... incomplete ridge-plates in 100 mm. on convex internalisuriaceme ssa ae cee 6-735 Parelephas progressus Osborn, 1924 Figures 969, 970 Upper Pleistocene, post(?)-Wisconsin (IV GuactaL) age, Zanesville, Muskingum County, Ohio. This very progressive species is based on a pair of superior and inferior grinding teeth described by Warren in 1855 as Elephas ‘Primigenius’; erroneously selected by Osborn (1922.555, pp. 11, 12) as a paratype of Elephas jeffersonii. The type teeth belong to a much larger animal than the type and referred specimens of Elephas jeffersonii and present a more progressive ridge-plate formula, namely: M328. Its descent from the P. jeffersonii stage is indicated by the similarity in the ridge-plate compression, indi- cating that the cranium of this species will prove to have the same distinctive proportions as the cranium of P. jeffersoniz. Parelephas jeffersonii progressus Osborn, 1924. ‘“‘Parelephas in Relation to Phyla and Genera of the Family Elephantide.” Amer. Mus. Novitates, No. 152, December 20, 1924, pp. 1, 4, and 7 (Osborn, 1924.633). Typr.—Amer. Mus. Warren Coll. 10457. A pair of superior and inferior grinding teeth of both sides. Horizon AND Locauiry.—Zanesville, Muskingum County, Ohio; Upper Pleistocene, Wisconsin or post(?)-Wisconsin (IV GuactAL) age. Typr Figurr.—Osborn, 1922.555, p. 13, fig. 11, and p. 14, fig. 12. Tyrer Descriprion.—(Osborn, 1924.633, pp. 4, 7): ‘These progressive grinders, described by Warren in 1855 as Hlephas primigenius, by Osborn in 1922 as a paratype of Hlephas jeffersonii, belong to a much more progressive stage than the type of Hlephas jeffersonii, and referred specimens, presenting a progressive ridge- plate formula of M 3 2%, as compared with the typical ridge-plate formula of Parelephas jeffersonii, namely M 3%%. It is interest- ing to observe that these type molars (Osborn, 1922.555, p. 14, 1098 fig. 12) show eighteen ridge-plates in use in both M* and M3; the superior molars show from four to six ridge-plates in excess of the inferior molars; thus they are readily distinguishable from those of Mammonteus primigenius compressus, as described above. In all other characters these type grinding teeth (Osborn, 1922.555, fig. 12) are related to those of Parelephas jeffersoniz. Consequently, we assign the subspecific name progressus to denote this extreme stage in the long Parelephas series.” Typn, ZANESVILLE, OnI0.—A pair of upper and lower grind- Al 1/4 nat. size Amer. Mus. 10457 (outer view) Amer. Mus. 10457 ‘outer view) Amer. Mus. 10457 (inner view) OSBORN: THE PROBOSCIDEA ing teeth of both sides (Amer. Mus. 10457) acquired by the American Museum with the Warren collection in 1906. It was described by Warren in 1855 (p. 163, Pl. xxvu, fig. C) as Hlephas ‘Primigenius’ and described and figured by Osborn in 1922 (1922.555, p. 13, fig. 11) as Elephas jeffersonii. This type affords a complete comparison with the species Parelephas jeffersonii (as defined above with a ridge-plate formula of M 3 $2), the ridge-plate formula in P. progressus being M 334;25,) the right M; presenting the last lower ridge-plates more fully developed than in the left lower M3. CHARACTERS OF Typn.—It is interesting to observe that these Amer. Mus. 10457 V4 nat. size (outer view) Type Movars oF PARELEPHAS PROGRESSUS (SIDE AND Crown Views) ORIGINALLY FIGURED BY OSBORN AS PARATYPES OF ELEPHAS JEFFERSONIL One-fourth natural size Fig. 969. Side views of type molars of Parelephas jeffersonii progressus (Amer. Mus. Warren Coll. 10457), Zanesville, Ohio, one- fourth natural size. After Osborn, 1922.555, p. 18, fig. 11. Al, External aspect of left superior molar, 1.M*, showing 30 ridge-plates, of which 18 are in use. B1, External aspect of left inferior molar, ].M3, showing 24 ridge- plates, of which 17 are in use. B2, Internal aspect of right inferior molar, r.M3, showing 26 ridge-plates, of which 18 are in use. Fig. 970. Crown views of type molars of Parelephas jeffersonit progressus (Amer. Mus. Warren Coll. 10457), the same individual as in figure 969, one-fourth natural size. After Osborn, 1922.555, p. 14, fig. 12. A, Crown view of left superior molar, ].M*, showing 30 ridge- plates, of which 18 are in use. B, Crown view of right and left third inferior molars containing respectively 26 and 24 ridge-plates, ef which 17 are in use. THE MAMMONTINA:: PARELEPHAS type molars show 18 ridge-plates in use in both M* and Ms, but that the superior molars show from four to six ridge-plates in excess of the inferior molars, the ridge formula being M 35,3255. The ridge-plate compression, however, corresponds with that of Par- elephas jeffersonzi and is directly intermediate between that of Parelephas columbi-Archidiskodon imperator and that of Mam- monteus primigenius, the internal, external, and coronal ridge- plate measurements being as follows: M?* 7-8 ridge-plates in 100 mm. on the external convex surface 8-8! ridge-plates in 100 mm. on the oblique mid-coronal erinding surface 9-10 ridge-plates in 100 mm. on the internal concave surface Ms, 64-7 ridge-plates in 100 mm. on the external concave surface 7-714 ridge-plates in 100 mm. on the oblique mid-coronal grinding surface 6-65 ridge-plates in 100 mm. on the internal convex surface This compression with averages of 6—-7—8-8}s-9-10 ridge-plates in 100 mm., according to the point of measurement of the ridge- plates, concurs with the measurements given by Osborn in the six type and referred specimens of Parelephas jeffersoni. Comparing the ridge-plate formula of Parelephas progressus (M 3 sqicog) With that of the most progressive form of Mam- monteus primigenius from Alaska and Indiana, namely, M. primi- genius compressus (M 3 s\23), We observe that P. progressus has a higher ridge-plate formula, but M. primigencus compressus (Al- aska) has a much closer ridge compression. As shown in the type figures of Parelephas progressus, here- with reproduced (Fig. 969), the ridge-plates in external and internal aspects have the sinuous and arcuate forms, such as are observed in Parelephas jeffersonii, more pronounced than the simple, arcuate ridge-plates of Mammonteus primigenius of Alaska (Fig. 1008). GroLocic Location AFTER Hay.—(Hay, 1923, p. 134): “2. Zanesville, Muskingum County, [Ohio].— . . . Zanesville is situated in the unglaciated part of the State [of Ohio]; but outwash from both the Illinoian and the Wisconsin glaciers has been de- posited along the river. For a knowledge of the Pleistocene epoch in that region, Leverett’s work may be consulted (Monogr. U. 3. Geol. Surv., vol. XLI, p. 158, plate nm)... . In 1853 (Amer. Jour. M® Total mid-length of crown including all the ridge-plates...... Maximum height of crown, height of tallest ridge-plate...... Maximum width of crown at the broadest portion.......... Breadth-lencthrindexs= ammeter nn neces a one ceteris: Meignt-len gti dexeprware peer tree eee ioe > 3 | 2& S8 S& 88 SR 88 oe, Sey Bey Cee Ben ae Seg ews, leases tee eS ee nes S202 42 44 72 75 x a aX a & AQ Cervical 1 220) 225 2 152 3 155 4 153 5 147 171 199 6 145 163 197 7 148 Dorsal 1 2 3 121 4 132 5 135 130 6 133 150 7 124 8 123 9 118 10 Naas alate ll 114 117 12 122, iil4 127 13 125 129 14 122) 127 15 123 135 16 126 120 17 126 127 =—s- 186 18 25 LO 26 137 19 127 121 136 Lumbar 1 142 154 127 144 2 150 143 3) 153 175 4 161 | Sacrals 1—4, length 377 ~=—- 320e 320e OSBORN: THE PROBOSCIDEA FIVE OF THE CRANIA OR JAWS. COMPARE TABLE XIV. series present the following vertical and transverse measurements of the centra: TYPE? Amer. Mus. 26833a, probably associated with type (Amer. Mus. 26820). Cervicals, including Cl, C5 (tr. 171 Xver. 154), C6 (tr. 163 X ver. 156). Dorsals, including D1, D11 (tr. 117 Xver. 126), D13 (tr. 129Xver. 116), D15 (tr. 185Xver. 118), D18 (tr. 119 Xver. 122), D19 (tr. 121 Xver. 122). Lumbars, including L1 (tr. 127 Xver. 118), L2 (tr. 148 « ver. 116). PARATYPE? Amer. Mus. 268336, probably associated with large aged paratype (Amer. Mus. 26821). Cervicals, including Cl, C2, C3, C4, C5 (tr. 199 X ver. 181), C6 (tr. 197 Xver. 182). Dorsals, D4, D5 (tr. 130 X ver. 156), D6, D7, D8, D9, D11, D12 (tr. 127Xver. 119), D14, D15, D16, D17 (Er. 136 X ver. 131). Lumbars, L2, L3 (tr. 175 Xver. 128). Sacrals 1-4, length 320+-mm. REFERRED? Amer. Mus. 26838e, including a few dorsals of larger measure- ment, namely, D17 (tr. 127 Xver. 128), D18 (tr. 126 X ver. 127), D19. Amer. Mus. 26833d, including four vertebrae of the largest measurement, namely Dorsals, D6 (tr. 150 ver. 165), D18 (tr. 137 X ver. 135), D19 (tr. 186 X ver. 139). Lumbars, L1 (tr. 144. ver. 136). Amer. Mus. 26833c, of the smallest measurement, Cervicals, including C1 (tr. 225 Xver. 122), C2, C3, C7 (tr. 148 X ver. 150). Dorsals, ineluding D6, D10 (tr. 117 Xver. 123), D12 (tr. 114Xver. 117), Dl4 (tr. 127Xver. 113), D16 (ér. 120 Xver. 110), D18 (tr. 115 X ver. 114). Lumbars, including L1 (tr. 154 X ver. 115), L38, L4. Sacrals, including sacrals 1-4, length 320e mm. COMPARISON WITH PARELEPHAS JEFFERSONII (SEE TABLE XIV).—The transverse measurements of the type (Amer. Mus. 9950) of Parelephas jeffersonii in general agree most closely with the smallest P. floridanus (Amer. Mus. 26833c), omitting C7, D1 and D2 which could not be obtained. In the linear measurement of the sacrals 1-4, P. jeffersonii (377 mm.) exceeds the smaller P. floridanus (820e mm.). Very significant are the steadily increas- ing transverse diameters in the posterior dorso-lumbar vertebre of P. jeffersonti D3 (tr. 121) Lumbar 4 (tr. 161). The anterior dorsals are wide transversely; in the middle region of the back the vertebre become quite narrow, and near the lumbar series they become wider; this rule was also found to hold with Hlephas indicus. THE MAMMONTINA: PARELEPHAS 1115 Whereas, as appears in Table XTV,the transverse measurements measurements of P. floridanus display great irregularity, owing to of Parelephas jeffersonii are fairly constant, similar transverse irregular disposition or distortion and to possible errors in our assemblage. ye ves FLa.Surv.V.4529 : a REFERRED Ricut TuorrpD INFERIOR MOLAR OF PARELEPHAS FLORIDANUS Mertuop or RipGkE-PLATE MEASUREMENT Fig. 989. Total length 274 mm., maximum height of fourteenth ridge-plate, 185 mm. A finely preserved right third inferior molar, r.Mg (Fla. Surv. V-4529) from Fig Island, Itchatucknee River, Florida, referred to Parelephas floridanus. Courtesy of the Florida Geological Survey. 9IIT ‘NVJ( dO GDY FHL dO TIVH ‘ANOLSI ‘(SQINADINIYd SAGLNOWWYJY) HLOWWYJY NUGHLYON YO ATIOONM AHL, “066 “O14 TWuoOLVN dO wodsojy NYOINGWY AHL NI LHOINY ‘Y SaTUVHD Ad ONILNIVd HRLAY CHAPTER X VIII THE GENUS MAMMONTEUS! (SUPERFAMILY ELEPHANTOIDEA), OF THE SUBFAMILY MAMMONTINAE, THE TRUE NORTHERN WOOLLY MAMMOTH EARLY DESCRIPTIONS OF THE MAMMOTH DURING THE SEVENTEENTH AND EIGHTEENTH CENTURIES (1696-1788) IN EURASIA. HISTORY OF EURASIATIC AND NORTH AMERICAN DISCOVERY. CHARACTERS OF MAMMONTEUS PRIMI- GENIUS AND OF SUPPOSED ANCESTRAL STAGES LEADING BACK TO UPPER PLIOCENE TIME. EXTREME STAGES OF EVOLUTION IN AMERICA. FEEDING HABITS AND GEOGRAPHIC DISTRIBUTION. I. Hisrorican INrRopucTION (1696-1788). Mammonteus primigenius. Names successively applied to the mammoth. Typical ridge formula. ee Native Siberian origin of the word Mammut Skulls and jaws of the true Mammonteus primigenius. “Adams skeleton.’ II. CHARACTERS OF THE SUBFAMILY MAMMONTIN®, INCLUDING 2 Z Primitive European stages of Mammonteus primigenius. THE GENUS MAMMONTEUS. Mammonteus primigenius leith-adamsi. 1 External characters and feeding habits. Mammonteus primigenius hydruntinus. 2. Skeletal characters of Mammonteus primigenius. Mammonteus primigenius fraast. 3. Historical order of naming of species of Mammonteus. Mammonteus primigenius astensis. 4. Geologie and provisionally ascending phyletic order of 3. American stages of Mammonteus. species and subspecies of Mammonteus. TWurasia and : Mammonteus primigenius americanus. North America. P J Mammonteus primigenius compressus. III. Systematic Description or Species OF MAMMONTEUS. Mammonteus primigenius alaskensis. 1. Typical progressive Eurasiatie stages of Mammonteus. 4. The frozen Mammoth of Siberia. I. HISTORICAL INTRODUCTION (1696-1788) The subfamily name Mammontinz Osborn, 1921, replaces the name Euelephantinz Osborn, 1918, because the subgenus Huelephas Falconer is invalid (see Chapter XIX, p. 1177); the term Mammontine, signifying ‘les mammonts,’ the ‘mammoths,’ was substituted in 1921. As set forth in the two preceding chapters, the Mam- montinee embrace three genera, namely, Archidiskodon of the southern and south temperate zones, Parelephas of the intermediate and north temperate zones, and Mammonteus' of the northerly and cireumpolar zones. It is probable that these northerly or woolly mammoths were the first mammalian fossils of northern Eurasia to be discovered and recognized as extinct. The earliest descriptions are naturally lost in obscurity. Of this early period Falconer (1846, pp. 11 and 12) writes: Next, in regard to the establishment of the species.—The fossil remains of the Mammoth had, during ages, attracted more or less attention in every country in Europe, having been found in England and in all parts of the Continent, from Italy to Siberia. But it was only towards the close of the last century that definite notions as to the species, were arrived at. Pallas, who had better opportunities for determining the point than any of his contemporaries, upon the perfect remains so commonly met with in Russia, erroneously considered the fossil teeth to be identical with those of the Indian species. A great advance was made in the inquiry through the discovery, by Peter Camper, of the specific difference between the teeth of the Asiatic and African Ele- phants [Footnote: ‘P. [A]. Camper, Déscript. Anatom. d’un Eléphant male, p. 16.’], when Blumenbach and Cuvier almost [The generic name Mammonteus (Mammonteum Camper, 1788, pp. 251, 259; Osborn, 1924.633, p. 2) for the northern Mammoth is of doubtful validity as Camper’s description has reference to an animal in America and not to the Mammoth, moreover he used the word “mammonteum” in an adjectival sense. Dr. A. Tindell Hopwood in his recent (1935) memoir on the ‘Fossil Proboscidea from China,” p. 97, adopted the generic term Mammuthus Burnett, 1830, genotype Mammuthus borealis. Yor a historical account of the names applied to the Mammoth and the Mastodon, see Chapter XXI below on Nomencla- ture.—Kditor.] 1117 1118 OSBORN: THE PROBOSCIDEA simultaneously entered upon the investigation, and arrived at the same result, viz., that the Mammoth was an extinet form, dif- fering from both of the existing species. Struck with the length of the cranium, and of the incisive sheaths in the Mammoth, as represented in the figures of Messerschmidt’s specimen attached to Breyne’s excellent remarks in the Philosophical Transactions [Footnote: ‘Phil. Trans. vol. xl. 1738 [1741], p. 124.’], and connecting these peculiarities with the great width of the crown, and the narrowness and number of the plates in the fossil grinders, Cuvier was conducted to his first happy conclusion. The prob- ability of a similar difference characterizing the species in other fossil genera, flashed across his mind, and opened to him new views respecting the theory of the earth. Great and important were the results; and after they had been achieved, the illustri- ous Anatomist reverted, in terms of the liveliest acknowledgment, to the long neglected figures of Messerschmidt, which had helped him to the first idea [Footnote: ‘Cuvier, Oss. Fossil. tom. i. p. 178.’]. ScELETO ELEPHANTINO TONN&, 1695, CONFUSED WITH THE MAMMOTH, BLUMENBACH, 1799 In 1695 Ludolf described the mammoth of Siberia. The earliest account of the fossil elephant in Germany is that of Wilhelmus Ernestus Tentzelius! (Phil. Trans. Roy. Soe., London, 1698, Vol. XIX, pp. 757-776): TeNTZELIUS (TENTZEL), 1698, pe. 757, 758.—II1. Wilhelmi Ernesti Tentzelia Historiographi Ducalis Saxonici Epistola de Sceleto Elephantino Tonne. ... Tonna inter Thuringiz Dinastias haud postrema, . .. quorum alter appellatur Burg-Tonna, . quam effodientes mense Decembri superioris anni ossa quedam maxima reperiunt, . . . Prior sententia mihi cumprimis placet, quam ita defendere aggredior, ut primo ostendam, sceleto nostro omnia convenire, que ad vera elephanti ossa requirun- tur; deinde evincam, non esse minerale fossile, sed animale petrefactum; denique inquiram, quomodo in has terras ipsumque locum pervenerit Elephantus. This classic or original deseription of the ‘elephant’ discovered in a sand-pit at Burgtonna near Gotha, Thuringia, is of the skeleton referred to by Blumenbach in his type description of Elephas primigenius cited below, but this is not to be regarded as the type specimen.’ It is addressed ‘“‘ad Virum toto orbe celeberrimum Antonium Magliabechium, Serenissimi Magni Hetrurte Ducis Bibliothecaritum & Consiliarium.” Following the dedication Tentzelius gives a description of the discovery in the month of December, 1695; he fully describes parts of the teeth, skull, and skeleton found, and discusses local opinion as to whether they were the remains of a petrified elephant or of a fossil unicorn (p. 758); he cites the descriptions of John Ray, of Pliny, and of Aristotle to prove that the specimen from Tonna was an elephant; he assures the most learned and celebrated Magliabechi that the bones and teeth correspond point by point with those of the elephant; he proceeds with a comparison with the Ceylon elephant (p. 771) and with Pigafetta’s descriptions of the African elephant; he cites Steno as to the habits and distribution of the Indian and African elephants, and concludes that the Burgtonna remains (p. 774) constitute proofs of the Flood, of the rise of the waters even to the height of the Thuringian mountains. The Editor of this classic contribution to the Philosophical Transactions, (Gothae, 1696, p. 776), concludes: The Author of this Letter has favour’d the Royal Society with some pieces of the Bones of the Sceleton of this Elephant, viz. part of the Skull, wherein appear its Cells, some of the Teeth both of those that grind, and such as are called Elephants Teeth or Ivory, with some other pieces of Bones, all which they found agreable to his Description, and ordered they should be carefully preserv’d in their Repository. Thirty-one years later Sir Hans Sloane (Phil. Trans., 1729, pp. 457-494, 497-514) refers to this discovery as follows (p. 508): The Skeleton of an Elephant which was dug up in a Sand-pit near J’onna in Thuringen, in 1695, is one of the most curious, and also the most compleat in its Kind, forasmuch as they found the whole Head, with four Grinders, and the two dentes exerti, or Tusks, the Bones of the fore and Hind-legs, one of the Shoulder-bones, the Back-bones, with the Ribs, and several of the 1Karl von Zittel (“History of Geology and Paleontology,” 1901, p. 184) refers to this discovery as follows: ‘The skeleton found at Burgtonna in 1696 was one of the most famous discoveries, as it gave rise to a dispute between Ernst Tentzel and the medical faculty in Gotha. The other professors saw in the large bones only sports of nature, but Tentzel proved to their discomfiture that the bones were real, and had belonged to elephants.” *The Burgtonna skeleton belongs (Dietrich, 1930) to Hlephas antiquus; while first mentioned, it is not the type. In defining and describing Hlephas primigenius (1799), Blumenbach had in mind the true mammoth of Siberia and Germany. THE MAMMONTINA: MAMMONTEUS 1119 Vertebre of the Neck. But the whole hath been so accurately described by Wilhelmus Ernestus Tentzelius, Historiographer to the Dukes of Saxony, in a Letter to the learned Magliabechi, printed in the Philosophical Transactions [Footnote: ‘No. 234, pag. 737.’], that it is needless to add any thing, the rather, as that Gentleman was pleased to oblige the Royal Society with some Pieces of the Bones of this Elephant, with Part of the Skull, wherein appeared its Cells, some of the Grinders, and Part of the dentes exerti; all which being produced at a Meeting of the Royal Society, were found exactly agreeable to his Description, and ordered to be carefully preserved in their Repository. [See footnote 1 below.] BREYNE’S DESCRIPTION (1735) OF THE ELEPHAS PRIMIGENIUS OF SIBERIA A second scientific description of the mammoth, from the river Indigirka, Siberia, with figure of the skull (Fig. 991), was a communication to Sir Hans Sloane, President of the Royal Society, by John Phil. Breyne, in a letter dated “Dantzick, Sept. 28, 1735,” from which the following excerpts are made (Phil. Trans. Roy. Soe., London, 1741, Vol. XL, pp. 124-138). A Letter from John Phil. Breyne, M.D.F.R.S. to Sir Hans Sloane, Bart. Pres. R.S. with Observations, and a Description of some Mammoth’s Bones dug up in Siberia, proving them to have belonged to Elephants. Sir, Your very learned and instructive Accounts of Elephants Teeth and Bones found under Ground, I saw with great Pleasure in the Philosophical Transactions, N° 403. and 404. In the same Year, to wit 1728, I was busied about the very same Matter, especially to prove, that the extraordinary large Teeth and Bones found under Ground, and digged up in several Places of Szberia, by the Name of Mammoth’s, or Mammut’s, Teeth and Bones, were, I. True Bones and Teeth of some large Animals once living; and, II. That those Animals were Ele- phants, by the Analogy of the Teeth and Bones, with the known ones of Elephants. III. That they were brought and left there by the universal Deluge. I made like- wise several useful Inferences about This Matter. At the same time there flourished in our City a Society of some learned and ingen- ious Gentlemen, who met once a Week ina certain Place: In one of those Meetings in the Month of March, I had the Honour to read and communicate my Thoughts and Observations about this Subject; which, as I believe, they will not be disagreeable to you, I have translated into the English Tongue, and joined to this present Letter. After that, viz. in the Year 1730, Dr. Messerschmidt returned to Dantzick, from his Travels thro’ Siberia, and was pleased to communicate to me some _ curious Draughts of a Part of a Skeleton, to wit, of a very large Skull, Dens exsertus & mo- laris, with the Os femoris, belonging to the Animal commonly called Mammoth, found in Szberia; by which our Assertion, that | hdofeph. Lianfact. WN? 446 nt oman Feetlo i 2 Visa EG. it). 00. ms \ 11 Yewiches a ZA . _AScale of2 anc rd Coriep BY CUVIER THe MESSERSCHMIDT CRANIUM OF SIBERIA. Fig. 991. Cranium of the mammoth, as described and figured by Dr. John Phil. Breyne in the Philosophical Transactions of the Royal Society of London, 1737 (printed in 1741, Vol. XL, No. 446, pp. 124-188, Pl. 1, figs. 1, 1). One-sixteenth natural size. This classical Breyne figure of the mam- moth cranium was reproduced by Cuvier in his ‘(Ossemens Fossiles’”’ of 1806 and used as the basis of comparison of the H. primigenius cranium with the crania of Z. africanus and E. indicus (Cuvier, 1806.1, Pl. xxxrx, fig. 1, and PI. xu, figs. 9.1, 10.A, 11.F.—our Fig. 992). Made by Brandt (1833.1,? p. XII) the type of Elephas giganteus. the Teeth and Bones, called in Rusland Mammoths Bones, are the true Teeth and Bones of Hlephants, is not only, as you wished in your first Account, put in a greater Light, but, if I am not mistaken, demonstrated beyond all Doubt. 1Hopwood (letter, May 20, 1930) finds that the Royal Society records (November 3, 1697) reveal that the Tentzel specimens were assigned the number 120B in various lists up to the year 1712: “It would seem, then, as though the specimens were lost between 1733 and 1763.” Fragments of the Burgtonna skeleton of Elephas antiquus, incisive tusks and other remains, are still preserved in the Gotha Museum (fide Dietrich, letter, April 14, 1930—see Chap. XIX, the Loxodontinz, p. 1181). *[See footnote on page 1136 regarding the date of description as 1832.—Kditor.] 1120 OSBORN: THE PROBOSCIDEA Observations on the Mammoth’s Bones and Teeth found in Siberia: Read in a Meeting of some learned Gentlemen at Dantzick in the Year 1728. by J. P. B. That learned and curious Gentleman Dr. Daniel Gottlieb Messerschmidt, who was sent some Years ago, by his late Czarish Majesty, Peter the Great, into Siberia, to search after the Products of Nature in this uninhabited and cold Country, was pleased to send mein the Year 1722, amongst some other Samples of Natural Things out of Siberia, two very large Teeth, called there, Mammoth or Mammut’s Teeth, with the following Inscription: Dens molaris, ut videtur, dilunianus, Bellue cujusdam hactenus incognite, nisi pro Elephantino habendus sit, cujus jam penes Te esto arbitrium, Russis Mammoth, repertus in Montium altissimis jugis ad Thomam fluwium. Alterwm est frustum aliud Eboris Denti exerto Elephantis non absimile, ab aliis repertum in Thome Montibus. [p. 137] The bones of this Skeleton, with the Ribs, Vertebrx, and others thereto belonging, were found in the sandy Side of a steep Hill, on the Eastern Bank of the River Indigirska [Indigirka], which falls into the Northern Ocean, not far from the Mouth of the Rivulet Wolockowoi ruczei. NAMES SUCCESSIVELY APPLIED TO THE MAMMOTH Among the first descriptions of species, to which Latin names were appended, are the following: 1696 Mammontova Kost [Mammotovoi kost in Ludolf, 1696.1,p. 92] (probably derived from the Tatar word mama signifying earth and kost signifying ivory =tusks) Ludloff [Ludolf]. Cited by Fischer de Waldheim, ‘“Oryectographie,”’ ITI, Fossiles du Gouvern. de Moscou, 1830-1837, p. 111. “T. Elephant. 1. Le Mammont... Hlephas mammonteus. m. [Fischer, 1830-1837]... Je conserve la dénomi- nation la plus ancienne, et je n’écris point Mammouth parceque ce nom n’a pris origine que par corruption ou une fausse lecture du mot Mammont [Footnote: Mammonimt en russe; le Yer (s) terminal a été changé par les anglais en h et le n a été pris pour un u. D ailleurs Ludloff est le premier qui en parle (1696) et appelle ces ossemens Mani- montova Kost et justifie ainsi ma dénomination systématique.’].” Cited by Cuvier (Ann. Mus. VIII, 1806, p. 45): “C’est sous le nom de cornes de mammont, mammontova-kost, quils désignent les défenses.”’ 1788 Mammontreum Camper (Nova Acta Acad. Sci. Imp. Petropol. Communicanda, II, 1787, p. 251). “". . os humanum petrifactum, aut fossile, etiamsi Mammonteorum, Elephantorum, . . . Adserere ex eodem principio audeo Mammonteum animal extinctum non modo esse, sed nullam omnino habuisse cum Elephanto similitudinem!”’ 1796 Mammoth, Cuvier [MS.] “Mémoire sur les espéces d’Eléphans tant vivantes que fossiles,”’ ete. (See Falconer, 1868, Vol. II, pp. 158, 159): ‘““‘Whatever may have been the approximation previously made by Merk or Blumenbach towards a distinction of the Mammoth from the two living species, Cuvier was undoubtedly the first to characterize the extinet species with exactness, in his joint memoir with Geoffroy, under the name of Elephas Mammoth, in the year 1796 [Footnote: ‘Mém. de l’Institut, 1" Classe, tom. ii.’]. Im the same year, he read a memoir at the first public meeting of the ‘Institute,’ but which was not published until 1806 [1799], in which the diagnostic marks are very pointedly expressed under the designation of Hlephas Mammonteus: ‘Maxilla obtusiore, lamellis molarium tenuibus, rectis,’ as distinguished from Hlephas Indicus: Fronte plano-concava, lamellis molarium arcuatis, undatis.’ Cuvier connected these dental and mandibular distinctions with others yielded by Messer Schmidt’s [Daniel Gottlieb Messerschmidt—see Breyne, 1737 (1741), Pls. 1, 1, 111] figure of the skull of the Mammoth, and combined the whole in the extended specific definition of the extinct form, which appeared in his memoir of 1806— ‘T/Fléphant a erdne allongé, A front concave, A trés longues alvéoles des défenses, & machoire inférieure obtuse, 4 ma- cheliéres plus larges, paralléles, marquées des rubans plus serrés.’"!! He abandoned the name EZ. Mammonteus of his memoir of 1796 [1799], and adopted the designation of Elephas primigenius, proposed by Blumenbach [Footnote: 'Cuvier’s definition (1806.1, p. 264) ‘“L’éléphant a crane allongé,... que nous nommons éléphant fossile (elephas primigenius, Blumenb.), est le mammont des Russes,” in contrast to (op. citl., p. 262) “L’éléphant a crane arrondi, . . . (elephas africanus),” also to (op. cit., p. 263) “L’éléphant a crane allongé. . . (elephas indicus),” is clearly illustrated in his Plates xxx1x and x1, in which he places Breyne’s figure of Elephas primigenius with his own figures of Elephas indicus and Elephas africanus. This definition was literally repeated in the three successive editions of the ““Ossemens Fossiles”’ (1812, 1825, 1834) and exert- ed a profound influence on other paleontologists, for it was not until 1825 that the mammoth of the south (Elephas meridionalis) was named by Nesti, in contrast to this elephant of the north (Zlephas primigenius) with curved tusks, also to Elephas antiquus Falconer (1847), with straight tusks. THE MAMMONTINA: MAMMONTEUS 1121 ‘Voigt’s Mag. 1803, Band v. p. 16.’], in 1803, which is that now generally accepted among paleontologists [see Blumenbach, 1799, p. 697, for first use of name Hlephas primigenius|. To this normal form, as already stated, Cuvier referred all the fossil remains of Elephants found over the whole of Europe, in Northern Asia, and in North America, however much at variance with the terms of his definition; and to the last he clung to the specific unity of the ‘Eléphant fossile’ with the jealous partiality of a discoverer for the earliest results of his most cherished labours. The distinctive characters in the molars of the Mammoth, as compared with those of the existing Indian Ele- phant, upon which Cuiver relied, may be expressed in the following terms:—1. Great narrowness or compression and approximation of the crown-ridges, involving both a larger number in the same length of crown and in triturating use at the same time. 2. Tenuity of, and absence of crimping in, the enamel-plates. 3. Greater width of the molar- crowns, both absolutely and relatively to their length.” 1796 Elephas mammonteus Cuvier (1796 [MS.], published August-September, 1799). (Cuvier, 1799, p. 21): ‘“‘les caractéres distinetifs des quatre espéces que ja’i décrites dans le cours de ce mémoire. Les voici: Especes qu'on sait exister. Elephas capensis [Cuv. = Loxodonta africana Blum.], fronte convexd, lamellis molarium rhomboidalibus. Elephas indicus [Linn. = Elephas indicus Linn.], fronte plano-concava, lamellis molarium arcuatis undatis. Especes qu’on ne connoit que fossiles. Elephas mammonteus [Cuv.=Elephas primigenius Blum.], maxilla obtusiore, lamellis molarium tenuibus rectis. Elephas americanus [Kerr= Mastodon americanus of the present Memoir], molaribus multi-cuspidibus, lamellis post detritionem quadri-lobatis.”’ A Fig.1. Fig.nF Léléphant & crane allongé, a front concave, & trés- longues alvéoles des défenses, & mdchoire inférieure ob= tuse, & mdchelieres plus larges, paralleles, marquées de rubans plus serrés , que nous nommons elephant f ssile ( ele- phas primigenius , Blumenb. ) , est le mammont des Russes, B Fig. 2. Fig. 9.1. L’éléphant @ crane allongé, a front concave , a petites oreilles , « mdchelieres marquées de rubans ondoyans que nous appelons eléphant des Indes (elephas indicus), estun quadrupéde qu'on n’a obseryé dune maniere certaine qu’au- dela de l' Indus. C Fig. 5. Fig. 10.4. Léléphant & créne arrondi, a larges oreilles , a mache- lieres marquées de losanges sur leur couronne, que nous appelons élephant d Afrique ( elephas africanus ), est un quadrupéde dont la seule patrie connue est jusqu’a présent T Afrique. Fig. 992. Comparison of crania of Elephas [Mammonteus] primigenius, EL. indicus, and E. [Loxodonta] africanus, after Cuvier, 1806.1, Pl. 39 (11), figs. 1, 2, 3, and Pl. 41 (rv), figs. 9.I, 10.A, and 11.F. One thirty-seventh to one-twentieth natural size. Figures 1 above and 11.F below are two views of the same skull (Messerschmidt, see Fig. 991), first figured by John Philip Breyne (1741.1, Pl. 1), subse- quently figured by Cuvier (1806.1, Pl. 39, fig. 1, and PI. 41, fig. 11.F), and finally made the type of Hlephas giganteus by Brandt (1833.1,! p. XII). The descriptive captions at the right are taken from Cuvier (op. cit., 1806.1, p. 262 (H. africanus), p. 263 (E. indicus), and p. 264 (E. primigenius). Cuvier’s designations were: “le Mammouth’” (Review, Cuvier, an 3 [1795], p. 90’); EHlephas mammonteus Cuvier, 1796, MS., published 1799, Fructidor, an VII [August-September, 1799], p. 21; not until 1806 did he adopt the specific name of Hlephas primigenius Blumenbach. [See footnote on page 1136 regarding the date of description as 1832.—Hditor.] 1122 OSBORN: THE PROBOSCIDEA 1799 Hlephas primigenius Blumenbach (‘Handbuch der Naturgeschichte,” Sechste auflage, p. 697)."! 3) Von einem ungeheuer grossen Elephanten (Hlephas primigenius?) [die vermeinten Riesenknochen** unsrer ehrlichen Alten]; unter andern auch in Menge in Deutschland***). So z. B. das beriichtigte Elephantengerippe das 1695 bey Burg-Tonna im Gothaischen ausgegraben worden ete. **s, Voigts Magazin. V. B.I. StS. 16 u.f. ***(Kriegsr. Merk) lettres sur les os fossiles d’elephans et de rhinoceros qui se trouvent en Allemagne &c. I-III. St. Darmst. 1788 u. f. 4. 43 Natural size Drawn from cast AM. 26981 22 23! 1 ' u ' ‘ U 1 V8 \17 \18 19 20°21 1 L.Mz f | ay Drawn from cast ] inca A. /1 26980 fe From Siberia From Osterode Lectotypes of ELEPHAS [= MAMMONTEUS] PRIMIGENIUS Azz. Originals in Blumentach Cll. Gottingen Mus. Fig. 993. Lecrotyprs or ELepHAs [MAMMONTEUS] PRIMIGENIUS BLUMENBACH After casts furnished by Director R. W. Hoffmann, Zoologische Institut der Universitat Gottingen (Left) Siberian lectotype (1.M3) from the Blumenbach Collection of the Zoological Institute of the University, Gottingen (cast Amer. Mus. 26980). Described by Dietrich (letter, July 4, 1930) as follows: ‘Es handelt sich bei dem Zahn aus Sibirien Ms sin., dist. niihere Provenienz und Acquisition unbekannt.” Interpreted by Osborn as a portion of a third inferior molar of the left side, namely, 1.M3, with 16 of the anterior ridge-plates preserved, of which at least 15 are more or less fully worn; 8 posterior ridge-plates are missing; 7) ridge-plates in 100 mm. on the convex internal side. Height of 14th ridge-plate (tallest) 112 mm. Drawn in crown and internal aspects. (Right) Osterode lectotype (1.Dp*) from the Blumenbach Collection of the Zoological Institute of the University, Gottingen (cast Amer. Mus. 26981). Determined by Dietrich (letter, July 4, 1980) as follows: ‘Der Zahn aus Osterode ist ein dritter oberer Milehmolar (m? [{sin.]); er ist Kurz beschrieben in Annalen der Physik, hersg. v. Gilbert 15, [45], 1813, p. 427/28. Er ist typisch fiir unsere spiitglacialen Mammute. Blumenbach diirfte ihn vor dem sibirischen Zahn erworben haben, so dass dieser m* aus Osterode als Type specimen des Hlephas primigenius Blumenbach, 1799, zu betrachten ist, wenn man nicht vorzieht, das Adams-Tilesius’sche Skelet zum Typus der Spezies zu machen.” Interpreted by Osborn as belonging on the left side, namely, as an 1.Dp‘, with 12} ridge-plates; 10-11 ridge-plates on the convex external side in 100 mm. The above passage in which Blumenbach, the pioneer of mammalian paleontology in Europe, refers to the discovery near ““Burg-Tonna”’ in the year 1695, is regarded as the type description of Elephas primigenius Blum. ; unfortunately the ““Burg-Tonna” skeleton, now in the Gotha Museum, belongs to ‘Elephas antiquus.’ On the advice of Dietrich (1930), we may select as lectotypes of FE. primigenius: (1) A grinder from Siberia in the Blumen- “The priority of publication of Blumenbach’s species Elephas primigenius (spring of 1799) is established over Cuvier’s Elephas mammonteus (August- September, 1799). Sherborn writes (letter, May 16, 1930): “It is rare that one finds so perfectly clear a statement in these old reviews. This comes from Gottingische Anzeiger von gelehrten Sachen, 18 Dec. 1799, p. 2057... . ‘Von dem Blumenbachischen Handbuch der Naturgeschichte ist schon vorige Ostern die 6te Auflage auf 708 S. herausgekommen.’” *See letter of February 8, 1930, from W. O. Dietrich, Geologisch-Paliontologisches Institut und Museum der Universitit Berlin: “Das zoologische Institut der Universitit G6ttingen besitzt nach Auskunft durch den Direktor Prof. Dr. R. W. Hoffmann nur zwei Stiicke fossiler Elefanten aus der Zeit Blumen- bachs 1. Einen Molar mit der Bezeichnung Hlephas primigenius. Diluvium, Sibirien. Coll. Blum. 2. Einen Molar mit der Bezeichnung Osterode (Harz) 1808.” THE MAMMONTINA: MAMMONTEUS 1123 bach Collection of the G6ttingen Museum, cast Amer. Mus. 26980; also (2) a molar from Osterode (Harz), Germany, original in same Museum; cast Amer. Mus. 26981 (see Fig. 993 on opposite page.) Blumenbach’s type description is in a rare document; no figure accompanies it; the sixth edition (1799) of his ‘Handbuch der Naturgeschichte,” in which the above type description appears, attracted the attention of the French naturalist Soulange Artaud, who visited Blumenbach and translated it under the title “‘Manuel d’ Histoire Naturelle,” 1803. In the manuscript of the French volume, Blumenbach inserted the name and definition of Elephas primigenius, consequently the species Elephas primigenius is of date 1799, 1803. 1799 Elephas mammonteus Cuvier (MS., 1796, published August-September, 1799). (Letter, Messieurs Emile Picard and A. Lacroix, April 15, 1930): ‘Nous n’avons pas trouvé trace d’unc telle publication, mais il est évident que le mémoire était écrit avant le 21 janvier 1796. ... Ce mémoire a été lu A la classe des sciences de I’Institut national, le pluvidse, an IV (21 janvier 1796). Cette lecture est mentionnée au procés- verbal de la séance (Procés-verbaux des séances de |’ Académie tenues depuis la fondation de |’ Institut jusqu’au mois d’aotit 1835, t.I page 6). Réservé pour étre imprimé, il a été publié dans le tome II des Mémoires de |’Institut national, en Fructidor, an VII (aott-septembre 1799).”’ Cited by Tilesius (1815, p. 470). 1806— Elephas primigenius Blum., Cuvier (“‘Oss. Fos.’’). 1834 1807 Elephas primevus Blumenbach (in M. Adams; translation (1808) from the French by Sir Joseph Banks, Phil Mag., XXIX, London, p. 152; cited by Tilesius, 1815, p. 452. 1807 EHlephas Mammouth Link, Beschr. Nat. Samml. Univ. Rostock (4) 1807,3 (fide Sherborn, 1928, Pt. XV, p. 3845). 1815 Elephas primordialis Blumenbach (in Tilesius, 1815, p. 470). 1820 Hlephas jubatus Schlotheim. 1829 Hlephas mammonteus Fischer de Waldheim. 1829 EHlephas paniscus Fischer de Waldheim. 1829 Hlephas periboletes Fischer de Waldheim. 1829 Elephas pygmaeus Fischer de Waldheim. 1829 Elephas campylotes Fischer de Waldheim. 1829 EHlephas Kamenskii Fischer de Waldheim. 18380 Mammuthus Borealis Burnett. 1832 Mammut Sibiricum (in von Meyer, Palaeologica, p. 64). 1832 EHlephas brachyramphus Brandt (‘Adams skeleton’). 1832 [Elephas| homotaphrus Brandt. 1832 Elephas giganteus Brandt. 1832 Elephas commutatus Brandt. 1832 EHlephas stenotoechus Brandt. 1832 Elephas platytaphrus Brandt. 1832 Hlephas affinis Brandt. 1834 Hlephas macrorynchus Morren. 1835 Elephas odontotyrannus Kichwald. 1837 DicycirorHertum E. Geoffroy St. Hilaire, Compte Rend. Acad. Sci., [V., No. 4, pp. 119, 120, fig. 1. 1839— E. |Elephas] primigenius sibiricus de Blainville (Atlas, “Ostéographie,” Pl. 11, 1845), also Depéret and Mayet, 1864 1923, pp. 183-201. sare E. [Elephas] primigenius germanicus de Blainville (Atlas, ““Ostéographie,” PI. 11, 1845). 1848 CuHeErRoLITEs von Meyer (in Bronn’s Handb. einer Gesch. d. Natur., III, Index Pal., p. 286). 1850 SyNoponTHERIUM Costa, ‘‘Paleont. del Regno di Napoli,’ Atti Acad. Pontaniana, V, Pt. I, p. 271, Tav. 11, figs. 1-4. 1885 PoLypIsKOoDONTEN Pohlig, p. 1027. 1888 PoLypiskopon Pohlig, Nova Acta K. Leop.-Carol. Deutsch. Akad., LIII, p. 252. 1913 Hlephas primigenius comune Issel, 1879 (in Zuffardi, 1913, p. 136)!. 'lZuffardi (1913, p. 136) attributes this subspecics to Issel (1879) who, in describing two molars from Camporosso near Ventimiglia, in the region of S. Andrea, Italy, states (p. 160) that these two teeth belong to the ‘comune Elephas primigenius,” using comune in the adjectival sense.—Editor. ] 1124 OSBORN: THE PROBOSCIDEA Out of this host of names, barbaric, semibarbaric, non-technically Latinized or technically Latinized, applied to the northern mammoth between the years 1696 and 1888, the species Hlephas primigenius of Blumenbach alone survives and is accepted in the scientific literature of the entire world. As to the genus, nearly a century and a half of research since 1799, when Blumenbach assigned the name Hlephas primigenius, demonstrates that the woolly mammoth belongs not to Elephas, but to a genus of its own, distinct by all the canons of nomenclature from the true modern Elephas. Technically the choice of a generic name rests between the inappropriate Dicyclotherium Geoffroy, 1837, Cheirolites von Meyer, 1848, Synodontherium Costa, 1850, and the first really appropriate generic name Polydiskodon Pohlig, 1885-1888. As Linnzeus chose many of his generic names, i.e., from those previously ill defined or not technically proposed, so in the present Memoir we may go back to the eminently appropriate and distinctive generic term Mammonteum, 1788' (changing the orthography to Mammonteus). NATIVE SIBERIAN ORIGIN OF THE WorD MAmMMutT Iprs, 1706.—The following is cited (pp. 25, 26) from the book of travels ‘Three Years Travels from Moscow over-land to China: thro’ Great Ustiga, Siriania, Permia, Sibiria, Daour, Great Tartary &ec. to Peking” of Eber- hard Ysbrant Ides, who was sent (“‘by the most Serene and most Potent Soveraign Czar, and Great Prince, Peter Alexewitz”’) on an embassy to the King of China in 1692, returning in the year 1695. Amongst the Hills which are situate North-East of, and not far from hence [village of Makofskoi near the river Keta], the Mammuts Tongues and Legs are found: as they are also particularly on the Shoars of the Rivers Jenize, Trugan, Mongamsea, Lena, and near Jakutsko7, to as far as the Frozen Sea. In the Spring when the Ice of this River breaks, it is driven in such vast quantities, and with such force by the high swollen Waters, that it frequently carries very high Banks before it, and breaks off the tops of Hills, which falling down, discover these Animals whole, or their Teeth only, almost frozen to the Earth, which thaw by degrees. I had a Person with me to China, who annually went out in search of these Bones; he told me as a certain truth, that he and his Companions found a Head of one of these Animals, which was discovered by the fall of such a frozen piece of Earth. As soon as he opened it he found the greatest part of the Flesh rotten, but it was not without difficulty that they broke out his Teeth, which were placed before his Mouth as those of the Elephants are; they also took some Bones out of his head, and afterwards came to his Fore-foot, which they cut off, and carried part of it to the City of Trugan, the Circumference of it be- ing as large as that of the wast of an ordinary Man. The Bones of the Head appeared somewhat red, as tho’ they were tinetured with Blood. Concerning this Animal there are very different reports. The Heathens of Jakuti, Tungust, and Ostiacki, say that they continually, or at least by reason of the very hard Frosts, mostly live under ground, where they go backwards and forwards; to confirm which they tell us, That they have often seen the Earth heaved up when one of these Beasts was on the March, and after he was past the place sink in, and thereby make a deep Pit. They further believe, that if this Animal comes so near to the surface of the frozen Earth as to smell or discern the Air, he immediately dies, which they say is the reason that several of them are found dead, on the high Banks of the River, where they unawares came out of the Ground. This is the Opinion of the Infidels concerning these Beasts, which are never seen. But the old Siberian Russians affirm that the Mammuth is very like the Elephant; with this only difference, that the Teeth of the former are firmer, and not so straight as those of the latter. They also are of Opinion, that there were Elephants in this Country before the Deluge, when this Climate was warmer, and that their drowned bodies floating on the surface of the Water of that Flood, were at last wash’d and forced into Subterranean Cavities: But that after this Noachian Deluge, the Air which was before warm was changed to cold, and that these bones have lain frozen in the Harth ever since, and so are preserved from putrefaction till they thaw, and come to light; which is no very unreasonable conjecture. Tho’ it is not absolutely necessary that this Climate should have been warmer before the Flood, since the Carkasses of the drowned Elephants were very likely to float from other places several hundred Miles distant, to this Country, in the great Deluge which covered the surface of the whole Earth. Some of these Teeth, which doubtless have lain the whole Summer on the Shoar, are intirely black and broken, and can never be restored to their former condition; but those which are found in good case, are as good as See footnote on page 1117 above regarding the doubtful validity of the genus Mammonteus.—Editor]. THE MAMMONTINA: MAMMONTEUS 1125 Ivory, and are accordingly transported to all parts of Muscovy, when they are used to make Combs, and all other such like things, instead of Ivory. The above mentioned Person also told me, that he once found two Teeth in one Head that weighed above 12 Russian Pounds, which amount to 400 German Pounds; So that these Animals must of necessity be very large, tho’ a great many lesser Teeth are found. By all that I could gather from the Heathens, there’s no Person ever saw one of these Beasts alive, or can give any account of it’s shape; so that the most that is said on this subject arises from bare conjecture only. Howort, 1882.1, pp. 30-32.—Howorth has given an excellent account of the name ‘‘Mammoth”’ in his article of 1882 in “The Field Naturalist,” page 30, from which we quote the following: In the year 1666, a learned Dutchman, named Cornelius Witzen, who became burgomaster of Amsterdam, visited Moscow. From the materials he there collected, and from various other sources, he compiled a famous work which he afterwards published at Amsterdam, entitled Noord en Oost Tatarye |Witsen, Nicolaes, 1692.1]... . It has never been translated into any other western language, and remains still buried to a large part of the scientific world in the original Dutch. In this work the name Mammoth appears for the first time, and is there written Mammout. Witzen describes how numbers of elephants’ teeth are found on the banks of the Siberian rivers, and says ‘by the Inlanders,’ 7. e. the Russian settlers in Siberia, ‘these teeth are called Mammouttekoos, while the animal itself is called Mammout.’—(Op. cit. ed. 1785, 742). In the Grammatica Russica of Ludolf, published in 1696, p. 96, these bones are called Mammotovoi Kost. Among the Russians the name is invariably written Mamont—Bull. St. Pet. Acad. x. 258. The insertion of the n in this form is no doubt a corruption, although an old corruption, since Captain J. B. Miller, in 1716, says ‘the inhabitants (7. e. the Russians) call the beast Mamant.’ Schiefier explains it as due to a confusion made by the peasants between the name Mammoth and that of Saint Mamas, called Mamant by the Russians, to whom the second of September is dedicated in the Russian calendar—(7d. note 2). Pallas derived Mammoth from the word Mama, which he says means earth in the Tartar tongue, but as Baer says, in the great Polyglot, edited by Pallas himself, in which the various dialects of Tartar are illustrated, no such name as Mama occurs as a synonym for earth. Baer himself suggests that the name is derived from ma, which means the earth in various Fin dialects, while mut or muit is the name by which the mole is still known to the Esthonians, whence he explains Mammoth as meaning the earth-mole—(Bull. St. Pet. Acad. x. 258). Not only is this an inconsequential etymology, since all moles live under ground, but the notion of finding an explanation of the name among a people so far removed from Siberia as the Esthonians is quite fanciful. MKlaproth was told by the Buriat and Mongol Lamas whom he consulted in 1806, that the name was of Tibetan origin. This again is a most unlikely quarter to go to for our etymology. A much more reasonable explanation of the name was given long ago by the Swedish exile to Siberia, Strahlenberg, who wrote his well-known description of the north and eastern parts of Europe at the beginning of the last century. In the 13th chapter of this work, under the heading Mamatova Kost, we read ‘As to the name it doubtless had its origin from the Hebrew and Arabick; this word denoting Behemot, of which Job speaks (in the xi. chapter), and which the Arabs pronounce Mehemot**** This is certain, that they (7. e. the Arabs) brought the word into Great Tartary; for the Ostiacks near the river Oby call the Mammuth Khosar, and the Tartars call it Khir; and although the Arabian name of an elephant is Fyhl, yet if very large they add the adjective Mehemodi to it; and these Arabs coming into Tartary, and finding there the relicks of some monstrous great beasts, not certain of what kind they might be, they called these teeth Me- hemot, which afterwards became a Proper Name among the Tartars, and by the Russians is corruptly pronounced Mammoth.*** The Russian Mammoth certainly came from the word Behemot; in which opinion I am confirmed by the testimony of an ancient Russian Priest, Gregory by name, Father-Confessor to Princess Sophia, who was many years an exile in Siberia, from whom I was told, that formerly the name for these bones in Siberia was not Mammoth, but Memoth, and that the Russian dialect had made that alteration.’—(Strahlenberg Eng. ed. 403). This view is curiously confirmed by the fact that Father Avril, a Jesuit, who went overland to China in 1685, and who is quoted by Witzen in the notices he gives of the Mammoth, never calls it Mammoth, but always Behemot.—(Aoril’s Travels, Eng. ed., 175, 177). Witzen himself also says that these teeth belong to the beast Behemoth, called Mammout, otherwise Mammona by the Russians.—(Op. cit., 742). The Turkish dialects habitually interchange the letters B and M, which are, in fact, used indifferently by them, so that the change from Behemoth to Mehemoth is perfectly regular and natural, and we can hardly doubt that Mammoth is in fact a mere form of Behemoth. At first it looks strange that the Arabs should have given a name to this beast which has become current all over the world; but this is easily explained when we remember the immense enterprise and energy of the Arabs in the ninth and two succeeding centuries, when, as we know, their traders and emissaries frequented the borderlands of Siberia, and probably first initiated the trade in fossil ivory in the west. ... Father Avril’s narrative shows that this trade still survived in his day, and he tells us the Persians and Turks put a great value on elephants’ teeth from Siberia, and preferred a scimitar or a dagger haft made of this precious ivory before a handle of massy gold or silver. There is, therefore, every reason to believe that the name Mammoth, as well as Mammoth ivory itself, were first introduced to the notice of the western world as an article of commerce by the Arabs, who were familiar with it probably as early as the ninth century. 1126 OSBORN: THE PROBOSCIDEA Pig. 994. Restoration of the migrating woolly mammoth (Hlephas [Mammonteus] primigenius Blum.) as it appeared on the river Somme, northern France. Details after the Upper Paleolithic etchings and paintings of Magdalenian time, especially in the caverns of Font-de-Gaume and Combarelles. Painted in 1919 by Charles R. Knight, under the direction of Henry Fairfield Osborn, for the Hall of the Age of Man, American Museum of Natural History. II. CHARACTERS OF THE SUBFAMILY MAMMONTINA, INCLUDING THE GENUS MAMMONTEUS SUPERFAMILY: ELEPHANTOIDEA Osborn, 1921 FamILy: ELEPHANTID& Gray, 1821 SUBFAMILY: MamMontTiINn& Osborn, 1921 (as defined above in Chapter XVI, Archidiskodon) Genus: MAMMONTEUS Camper, 1788! Original reference: Camper, 1788, pp. 251,259; Osborn, 1924.633, p. 2. Pleistocene of Europe, northern Asia, and North America. Syn. (partial list): Dicyclotherium Geoffroy, 1837; Chezrolites von Meyer, 1848; Synodontherium Costa, 1850; Polydisko- donten Pohlig, 1885, Polydiskodon Pohlig, 1888. MamMMONTEUM CampeER, 1788, pv. 251.—‘‘... os humanum petrifactum, aut fossile, etiamsi Mammonteorum, Klephantorum, .. . Adserere ex eodem principio audeo Mammonteum animal extinctum non modo esse, sed nullam omnino habuisse cum Elephanto similitudinem!”’ GENERIC CHARACTERS IN CRANIUM AND TEETH (OSBORN, 1928).—The Mammontewm of Camper, 1788 |Mammonteus of Osborn, 1924], includes Elephas primigenius Blum., also Hlephas americanus DeKay, Elephas primigenius astensis Depéret and Mayet, Hlephas primigenius fraasi Dietrich, and Mammonteus primigenius compressus Osborn.? (1) Cranium related to that of Archidiskodon and of Parelephas, but extremely acrocephalic, hypsicephalic, bathycephalic. Frontals concave, occipital crest greatly elevated, occiput slightly convex. (2) Molars, in Upper Pliocene to Upper Pleistocene stages, with relatively numerous ridge-plates; ae Pliocene stage, M. primigenius astensis, M 3 73303 typical Upper Pleistocene stage, Mammonteus primigenius, M 3 #4; final progressive stage, M. primi- genius compressus, M 3 ai. | Ridge-plates compressed in typical superior molars to 10-11-12 in 100 See footnote on page 1117 above for remarks on the doubtful validity of the genus Mammonteus.—Hditor.] *(To these should be added Mammonteus primigenius alaskensis described below. (pp. 1159-1161), this. chapter.—Kditor.] THE MAMMONTINA: MAMMONTEUS 1127 mm., in highly progressive superior molars 13 in 100 mm.; in progressive inferior molars 8-9-10 in 100 mm. Molar crowns broad, M* extremely short with enamel ridge-plates of minimum thickness, ridge-plates more or less crimped or sinuous. (3) Manus pentadactyl with five digital nails (fide Herz, 1902), manus and pes tetradactyl, not pentadactyl (fide Salensky, 1904, p. 86—see citation from Salensky! on page 1129 below); four digits (fide Pfizenmayer, 1926, p. 239), total phalanges in each foot reduced to nine in On2eE3 2 2 Phalanges, (OND NAO OER Ole ee (4) Dietrich (1912) records five digits in the manus and pes of the large and more primitive Upper(?) Pleistocene M. primigenius fraasi. (5) Tail abbreviate, caudals 21. (6) Habitat, northern tundras and steppes, grassy meadows; summer diet chiefly of grasses. All authors have included E. primagenius within the genus Hlephas, the genotypic species of which is Elephas indicus Linneeus, 1754 (= Hlephas maximus Linnzeus, 1758). It has now been demonstrated that the true northern mammoth (Hlephas primigenius) is one of the final members of a long series of species and of ascending mutations extending back through the entire period of Pleistocene time and first recognized in the Upper Pliocene of Italy as Hlephas primigenius mut. astensis, described by Depéret and Mayet in 1923. Throughout this long geologic period these ascending mutations and species exhibit fundamental dental or cranial characters and proportions which clearly distinguish them from Elephas indicus; consequently we have to do with a distinct generic phylum, comparable to the phyla of Archidiskodon and of Parelephas. For this ‘northern’ or ‘woolly’ mammoth the appropriate name Mammonteum Camper (= Mammonteus) was revived by the present author in 1924 (Osborn, 1924.633, p. 2). 1. EXTERNAL CHARACTERS AND FEEDING HABITS The true Mammoth is the only extinct proboscidean of which the characters of the soft parts and of the hairy and woolly covering, as well as the nature of the food both of the Mammoth and Mastodon, are fully known. The student is referred to the reports of Herz (1902) on the mammoth discovered on the Beresowka River; also to Salensky! (1903) who described in the greatest detail the external characters of the same individual. The most recent descriptions, however, are those of Quackenbush (1909) noting the discovery in 1907 at ‘Historic Bluff,” Eschscholtz Bay, Alaska, of part of a skeleton of the Mammoth, together with some skin and hair (Amer. Mus. 13749—paratype of M. primigenius compressus), and of Felix (1912) in his “Das Mammuth von Borna,”’ from which we may cite by translation: SuMMER Foop oF GRrAssES (FELIX, 1912, p. 11).—On uncovering the skull a portion of the animal’s food was found in the form of a wad lying between the upper and lower teeth. Its death, therefore, must have been so sudden that it did not have time to swallow this food. In addition, the entire stomach was discovered containing about 12 kilograms (24 lbs.) of undigested food. On examination this material proved to contain a flora of no great variety, but of exceeding interest, because it consisted of plants that are still native to the place, i.e., Bere- sowka River, northern Siberia. They are almost exclusively grasses. The needles of conifers occur very rarely. Here is the list: 1. Alopecurus alpinus Sm. [=alpine foxtail grass]. 6. Carex glareosa Wg. [=clustered sedge]. 2. Hordeum jubatum L. [=squirrel-tail grass]. 7. Carex ineurva Lightf. [=curved sedge]. 3. Agrostis borealis Hartm. [=red bent-grass]. 8. Thymus serpyllum L. [=wild thyme]. 4. Atropis distans Griseb. [=sweet-grass]. 9. Oxytropis campestris DC. [= field oxytrope]. 5. Beckmannia cruceformis Host. [=slough-grass]. 10. Papaver alpinum L. {=alpine poppy]. 11. Ranunculus acer L. var. borealis [=common buttercup]. 'See Zalensky, Vladimir Vladimirovich, in Bibliography of Volume I of the present Memoir.—Hditor.] 1128 OSBORN: THE PROBOSCIDEA (Op. cit., p. 12): All these kinds of plants are found in the same locality at the present day and form a char- acteristic meadow flora. With the exception of Alopecurus alpinus Sm. and Papaver alpinum L. which are also found in the tundra, there were no typical tundra plants. From other discoveries we know that the mammoth fed also on: (1) Betula nana L. [=dwarfed birch]; (2) Salix polaris Wahlnbrg. [=arctic willow]; (8) Cladonia rangiferina Hoffm. |=reindeer moss]. In addition to these the mammoth ate (probably in winter) the bark and twigs of conifers, chiefly larches. Proportions.—(Felix, 1913, pp. 18, 14, translation): (1) This discovery not only gave us complete knowledge of the skeleton of 2. primigenius but showed conclusively the proper position of the tusk in the jaw, and its inclina- COMPARISON OF THE Tip OF THE TRUNK OF THE MAMMOTH (MAMMONTEUS PRIMIGENIUS) WITH THAT OF THE INDIAN ELEPHANT (EHLEPHAS INDICUS) AND OF THE AFRICAN ELEPHANT (LOXODONTA AFRICANA) After Fleroy, 1931, Figs. 1, 2, and 3 Vig. 995. The following is cited from Doctor I'leroy’s interesting description (1931, pp. 863 and 869) of the MAMMONTEUS PRIMIGENIUS first discovery of the tip of a mammoth’s trunk: RECONSURGOITON In 1924 a well preserved mammoth trunk was found by an unknown Tungus in the everfrozen soil on the banks of am the Bolshaya Baranikha River in the Kolyma district. This region is very distant from any inhabited place, devoid of , regular means of communication and peopled with half-wild tribes. Five years elapsed before the news of a discovery, so exceptionally interesting for the Science, reached by a good chance some of our geologists working in the Kolyma district. The trunk in this time passed from person to person, its tip was cut off, then dried and the rest was thrown away and lost. Only in 1929 the dried tip of the trunk was handed over by Mrs. Kondratiev, a resident of Sredne- Kolymsk, a little town on the Kolyma River, to Mr. K. J. Pjatovsky, a geologist, who sent it to the Zoological Museum of the Academy of Sciences through Mr. M. J. Tkatchenko, an assistant of the Yakutsk Museum. TABLE OF MEASUREMENTS (CF. op. ciT., P. 869) mm mm Length of the tip of trunk on the external side 280 Length of the ‘lip’ on the exterior border 36 Maximal width 114 Width of the right nasal opening 23 Width above the base of the ‘lip’ 74 Width of the left nasal opening 17 Width of the ‘lip’ 71 ‘Thickness of the wall dividing both nasal openings 6 Length of the finger-like process 95 Thickness of the skin on the exterior side 2 Width of the finger-like process on the base 57 Thickness of the skin on the interior side 0.5 LOXODONTA AFRICANA tion. (2) As to the animal’s outward appearance, it involved a number of corrections in even the best reconstruc- tions that had been made. (3) The proportion of the length of skull and trunk in the mammoth is quite different from that of existing elephants. The length of the mammoth’s skull is more than half that of the trunk while the elephant’s skull [#. indicus] is always less than half the trunk length. The mammoth’s head, therefore, was larger in proportion to the body than that of the recent elephants, and in consequence the tusks could attain enormous proportions. The largest of the tusks in the St. Petersburg Museum measures no less than 4, 17 m. (13’8”) and in the Franzens-Museum of Briinn there is a tusk that actually exceeds 5 m. (16’5’’) in length! THE MAMMONTINA: MAMMONTEUS 1129 The trunk of the Beresowka mammoth was entirely missing, but probably differed little from that of recent elephants. The many representations of the mammoth found in the Paleolithic caves of France all show a strong- ly developed trunk. The ear was somewhat smaller than that of the Indian elephant. Its length was 38 em. (15’’), its breadth, 17 em. (6%’’). The ears, as well as the whole body, were covered with a thick coat consisting of short wool and longer hair. The massive body, rather short in proportion to its height, is joined to the mighty head by a short neck which seems still shorter on account of the great muscles. The taal—almost unknown until this diseovery—is (1) conical in form, sharply pointed at the end and about 36 em. (14’’) broad at the root. It ended in a bunch of bristles. (2) Salensky' gives the length—measured from the back—as 60 em. (23%’’). Pfizenmayer writes [1905, pp. 524, 525]: ‘The tail of this specimen—measured from the under side—is 35 em. long, decidedly shorter than that of living elephants. The number of [caudal] vertebree is only 21.’ (3) It would appear from this that the older reconstructions make it much too long. The difference in the measurement of Salensky and of Pfizenmayer can hardly be attributed to the mode of measurement. Proba- bly Salensky included the bunch of bristles at the end in his length, while Pfizenmayer did not. The skin was extraordinarily thick and underneath it was a layer of fat from 1 to 9 em. (3-34 in.) deep. The whole body was thickly covered with hair, even the legs down to the horny ends of the toes. The mammoth was thus particularly well fitted to withstand cold, while its outward appearance was quite different from that of its living relatives. The covering of the body consisted of three elements: (1) Fine, soft, woolly hair about an inch (20-25 mm.) long, in color varying from faded blond to yellow brown, and covering the entire body; (2) coarser and longer hair up to 20 inches (50 em.) in length, of a dark rust colored brown, covering the entire neck and trunk (of the body) perhaps forming a fringe of hair still heavier and thicker from the cheeks along the shoul- ders and sides to the rump, similar to that of the yak. (3) ‘“Bristles’’—so called on account of their stiffness—from 8 to 14 inches (20-35 em.) in length, much darker than the other kinds of hair, found only in the “brush” at the end of the tail. 2. SKELETAL CHARACTERS OF MAMMONTEUS PRIMIGENIUS These authors do not give a complete description of the osteological characters of the Beresowka mammoth. The most important note is that of Salensky' (1903, p. 86): “‘An osteological study of the Beresowka mammoth led to the unexpected conclusion that considerable differences exist between the Hlephas primigenius and the recent elephants [H. indicus], notably in the foot, which is tetradactyl and not pentadactyl. These differences absolutely exclude the possibility of the direct descent of the elephant from the mammoth.”’ One of the fullest comparative discussions of the mammoth skeleton is that of Dietrich (1912), in which the vertebral formula, covering our whole knowledge, is summarized below. ABBREVIATED VERTEBRAL CoLUMN.—Since the abbreviated backbone shares the extreme fore-and-aft com- pression of the cranium, EF. primigenius has relatively the shortest vertebral column of any of the Proboscidea. The reader is here referred to pages 930, 931, Chapter XV for a full treatment of vertebral distinctions in the genera of the Proboscidea. This abbreviation is indicated, however, in the shortening of the vertebral centra as well as in the number of dorso-lumbar vertebree (dorso-lumbars 24-23 in Elephas indicus, 23 in Loxodonta africana, 22-24 in 'See footnote on page 1127 above. SKELETONS OF MAMMONTEUS PRIMIGENIUS MounTED IN THE Museums oF Russia, GERMANY, AND BELGIUM Mounted skeletons of Kolyma-Beresowka, of Borna, of Steinheim, and of Lierre. Reduced to a uniform scale of about one-fiftieth natural size Fig. 996. Skeleton of Mammonteus primigenius from the Kolyma- Beresowka River, Siberia, as mounted in the Museum at Leningrad, Russia. Fig. 998. Skeleton of Mammonteus primigenius from Borna, Germany, as After Abel, 1925, p. 56, fig. 36, as reconstructed by W. Salensky [see footnote, mounted in the Museum fiir Vélkerkunde, Leipzig (Grassi-Museum). After Abel, 1925, p. 279, fig. 204 (see Felix, 1912, Taf. vim, for original). p. 1127 above]. )) Fig. 997. Skeleton of Mammonteus primigenius (=Elephas [Mammonteus| primigenius fraasi Dietrich, 1912) from Steinheim on the Murr, Wiirttemberg, Germany, as mounted in the Stuttgarter Ree ea T . . . = 5 AAEMIV@ETRS Doth T perro Naturalienkabinett. After Abel, 1925, p. 57, fig. 37, as reconstructed by E. Fraas and W. O. Dietrich. About one-fiftieth natural size. Fig. 999. Mammouth du Musée de Bruxelles découvert a Lierre (Province d’Anvers) en 1860. After Dupont ‘L’Homme pendant les Ages de La Pierre dans les Environs de Dinant-sur-Meuse,”’ Brussels, 1873, Pl. 1. One-fiftieth natural size. It will be observed that the acrocephalic, hypsicephalic, and bathycephalic proportions of the cranium and the harmonic abbreviation of the vertebral column are manifest in each of these reconstructions. The sharp notch behind the peaked skull in the Combarelles restoration (Fig. 1000) is a feature of every drawing of Paleolithic age. Compare figure 994. ‘ADAMS SKELETON,’ 1807 (sen Fia. 1014).—The first complete remains, known as the ‘Adams skeleton’ as mounted (Fig. 1014) in the Zoological Museum of the Academy of Sciences of the U. 8. S. R., Leningrad, were discovered in 1799 on the banks of the Lena River at the threshold of the Polar Sea (cf. Lang, 1925, p. 28): “Imbedded in ice, as it had been for thousands of years, its meat was still in such condition as to be eagerly devoured by polar bears, wolves, and other As time went on every warm season bared more of the body; only the natives contested the booty by securing some of the meat for their dogs through the following years of exposure. It was then that the intrepid explorer and botanist Adams happened to arrive in the neighborhood and, hearing of the famed monster, lost no time in reaching it. Most of the soft parts were gone, one limb had been carried away, and a native had sawed off both tusks and sold them for about fifty rubles. Through Adams’ energy and foresight practically all remaining bones were collected. He also took to Petrograd a piece of the hide with the hair in place. It was from the still frozen side upon which the mammoth lay, and so heavy as to tax the strength of ten men to drag it along the shore. A large amount of loose, coarse hair, evidently trampled into the snow by feasting polar bears, was long enough to be This ‘Adams skeleton’ as described and figured by Tilesius in 1815 was further described by Cuvier as cited below carnivores attracted from great distances. considered as having formed a mane.”’ (see Fig. 1014). [Made by Brandt (1832, pp. XI, XII) the type of Elephas brachyramphus.—Kditor.| 1130 THE MAMMONTINA: MAMMONTEUS 1131 Mammonteus primigenius, and 23 in Parelephas jeffersonii).: as shown in the following comparative table—see also Chapters XV and XIX: Loxodonta africana Elephas indicus . Parelephas Mammonteus primigenius a Jeffersonii Flower, “Jumbo” Flower; Hse LYPe 22% ' Faleoner and Felix, ‘Adams Beresowka 1885 Amer. Mus. 1885 Amer. Mus. Amer. Mus. 1912 Mammoth,’ Mammoth Dept. Mam. me §9504 ae. °F 14559 (M. p. Tilesius Pfizenmayer 3283 r! compressus),», Hf 1815 in Dietrich : ae 1912 @enviealsis--5 4. -- 7 iG 7 7 7 7 _ a 7 Worsals.......... 19 20 19-20 19 18-19 aye ota 19)7 18 Pairs of ribs...... Se 19 18-19 19, 17519? Tumbars........ 4 3 5-3 4 4-3 5 4? 5 SHON. co Go C0 Ebe 5 4 4 5 4-3 4 4 (@andalsi: 1.5.5: 24+ 21 24-30+ 12+ 21 21 (Sa- 8+ 21 lensky)! } 1001 Grotto of Combarelles, in which representations of the mammoth are next in Fig. The above outlines of the woolly mammoth are from the order of frequency to those of the horse. Compare Capitan, Breuil, and Peyrony, 1924, pp. 136, 137, figs. 120, 121. MacCurdy (1924.1, p. 275) records twenty or more caverns in France, Spain, and Germany in which drawings of Fig. 1000. Restoration of the woolly mammoth sketched on the wall of the cavern of Les Combarelles aux Eyzies (Dordogne), France. After Capitan, Breuil, and Peyrony, 1924, p. 137, fig. 121. Observe the very small feet, the short tail, the drooping hind quarters, the upcurved backbone, and the neck sharply incurved behind the summit of the occiput, characters which agree closely with those of the skeletons of Steinheim (Fig. 997) and of Borna (Fig. the mammoth occur, mostly in outline, while the famous painted murals of Font-de-Gaume are of a dark brownish tint. The mammoth was the only member of the elephant family in northern Europe during the period of the Fourth Glaciation or close of the Stone Age. In Spain, however, the straight- tusked Elephas [Hesperoloxodon| antiquus lingered to the very end of the Stone Age and its outline in the Cavern of Pindal, northern Spain, is shown in figure 998). 1047 of the present Memoir (Chap. XIX, the Loxodontinz). PrizENMAYER, 1926, p. 239.—While the skeleton of manus and pes in elephants has five digits, that of the mammoth has only four in both fore- and hindfeet [ef. Herz, 1920, five digits'in manus]. The first digit is com- pletely lacking, and the other digits even in older animals show three phalanges only in the third digit, while digits II, 1V and V have but two phalanges in fore- and hindfeet. All four extremities (feet) of the Beresowka mammoth were complete, and the left hindfoot from the knee joint down was preserved in alcohol with the skin and soft parts. This formula of four digits for the Siberian mam- moth was demonstrated when three extremities of the Beresowka mammoth were macerated by Professor Salen- sky,' director of the Museum, and myself as his assistant. There can be no doubt about this fact, as we also proved it to be the case in three older fore- and hindfeet which had been preserved with the soft parts intact. They belonged to the Maydell mammoth (1869) in the St. Petersburg Museum collection, and showed the same feature. Also the perfectly preserved fore- and hindfeet of the mammoth of Sanga-jurach agreed with the former See footnote on page 1127 above. 1132 OSBORN: THE PROBOSCIDEA discoveries in this respect. The phalangeal formula of the adult Siberian mammoth for both manus and pes is 2 2 Phalanges. 02 3 therefore = IV V Digits RESTORATION.—It will be observed in the Paleolithic restoration (Fig. 1002) that the apex of the cranium is extremely short and pointed (cyrtocephalic, acrocephalic), while the facial profile of the cranium (Fig. 1000) is rounded, a feature probably due to the storage of fat during the winter beneath the woolly dome. All these drawings indicate also a sharp depression between the head and the back, due to the extremely sharp elevation of the cranium; they also indicate the rapid downward slope (usually exaggerated) of the back toward the pelvis. Consequently the drawings fully confirm the evidence afforded by the fossilized skeletons, that the mammoth was wholly different in bodily contour from either of the existing elephants, Elephas indicus or Loxodonta africana. Another characteristic is the rapidly sloping hind quarters, as observed in all the numerous drawings and paintings of Mammonteus primigenius by the Cro-Magnon artists of Upper Palzolithic age, the most striking and Fig. 1002. Charging mammoth incised on a tusk of Elephas primigenius discovered in 1865 in the Magdalenian cultural level of the great rock shelter of La Madeleine (Dordogne), France, and first described by Lartet in the Comptes Rendus des Séances de |’Académie des Sciences, Vol. LXI, 1865, pp. 309-311, subsequently figured by Lartet and Christy in the Reliquie Aquitanice, 1875, B. Pl. xxvu, and amplified by them in the Descriptions of the Plates, p. 168, as follows: “It is a thin oblong piece, convex from side to side with the roundness of the tusk, and somewhat concave in the longitudinal direction, owing to its curvature. The outer surface presents what at first appears to be a medley of faintly scratched lines; but, on closer and more careful inspection, they re- solve themselves into a characteristic outline of a hairy Elephant, with some of the lines doubled and redoubled apparently by the old artist’s repeated attempts to sketch out the main features of his sub- ject. The lofty skull and hollow forehead are recog- nizable as striking features, characteristic of the Siberian Mammoth at St. Petersburg [Footnote: ‘See Le Hon’s “L’Homme fossile,”’ 1867, p. 70, woodcut.’], of the skull of the Mammoth from Ilford, Essex, preserved in the British Museum [Footnote: ‘Much modern information on the features, distri- bution, and general natural history of the Mam- moth, together with references to other authors, will be found in the Memoirs by Mr. Henry Woodward, F.R.S., of the British Museum, in the Geological Magazine, 1864, vol. i. p. 241, and woodcut; 1868, vol. v. p. 540, plates 22 and 23; 1869, vol. vi. p. 58; and 1871, vol. viii. p. 198, pl. 4.’], and of the Belgian Mammoth at Brussels [Footnote: ‘See Dr. E. Du- pont’s ‘“L’Homme pendant les Ages de la Pierre dans les Environs de Dinant-sur-Meuse”’ (pl. 2), 8vo, 2nd edit. 1872.’]. The small eye and long trunk of the Elephant, and the great curved tusks and shaggy hair peculiar to the Mammoth, are easily recognized. The upper and more convex sketch- lines of the back agree with the high withers of the Mammoth; and the lower and sloping dorsal lines probably had reference, in the draughtsman’s mind, to some special attitude of the animal, with which also the outstretched portion of the hind leg, and the elevated tail, may be associated. “Mr. H. Woodward, F.R.S., F.G.S., has kindly communicated the interesting suggestion that the attitude of the animal, together with the vertical position of the trunk, would well accord with the idea of one of a herd of Elephants coming down by moonlight to drink, and that the confusing double lines might then be explained as an attempt, on the part of the artist, to represent the rest of the herd. In running, or when alarmed, the trunk of the Elephant is always raised. And he adds that there can be little or no doubt that the sketch, rude as it is, was the result of a life-study of the animal, and is consequently of the highest importance as attesting the actual presence of the living Mammoth in France when the Caves of Périgord were occupied by Man.” In the present figure the other outlines, probably designed to indicate a charging herd of mammoths, are represented by dotted lines only (cf. Osborn’s “Men of the Old Stone Age,” p. 384, and Fig. 199): This classic engraving, now preserved in the Museum d’Histoire Naturelle, Paris, is one of the most realistic pieces of Palwolithic engraving which has ever been found; observe especially the outline of the ear, the elevation of the highly peaked (acrocepahlic) head, also the remarkably lifelike action of the limbs and body. There are indications that the artist used this relatively small piece of ivory for the representation of three mammoths; the tusks and trunks of two other elephants appear in the distance. ‘ i ‘ ‘ ' 1 ; i ' 4 ‘ ‘ THE MAMMONTINA: MAMMONTEUS 1133 probably the most accurate representation of the mammoth being the famous charging mammoth as engraved on an ivory tusk (Fig. 1002), the original of which is now in the Museum d’Histoire Naturelle, Paris. GEOGRAPHIC DistrRIBUTION.—(1) The geographic distribution of Mammonteus primigenius in the United States, in the recent valuable memoirs by Hay (1923, 1924), is unfortunately marred by the fact that many of the recorded specimens attributed by him to ‘‘Elephas primigenius”’ really belong to the animal described in the present Memoir as Parelephas jeffersonii. (2) The same confusion of the remains of Parelephas with those of Mammonteus doubtless applies to the plotted distribution’ by Osborn (Fig. 1005) in the Northern Hemisphere, designed to show the former distribution of the true mammoth; the European plotting (solid black) is largely after Boule and certainly includes both the ‘Hlephas primigenius’ and the ‘Elephas trogontherw’ range. (3) The Asiatic plotting (deep bars) is largely conjectural, and the same doubt exists as to the distinction between the ‘E. primigenius’ and the ‘E. trogontherii.’ The American plotting (solid black) is also after Hay and, as above noted, doubtless includes the range of species of both Mammonteus and Parelephas. (4) The distribution of MAPS AND THEIR EXPLANATIONS. MAPS AND THEIR EXPLANATIONS. Mar 12. Map 11. +++ sats eal Wisconsin lowan Mlinoian Kansan Prewisconsin Driftless Groarapuic DIsTRIBUTION OF MAMMONTEUS AND PARELEPHAS. AFTER HAy Fig. 1003. Geographic distribution of Mammonteus primigenius in the central (left) and eastern (right) regions of North America, after plotting by O. P. Hay: (Left) Hay, 1924, map 12, pp. 340, 341; (right) 1923, map 11, pp. 428, 429. A point of great significance is this indubitable occurrence of Mam- monteus primigenius in the Wisconsin and pre-Wisconsin ‘drift’ (IV Guactat), in the areas of the ‘middle drift’ (Illinoian, III Guacrau), and Iowan?, IV Guacrat), and in the ‘old drift’ (Kansan, II Guacrau). See Plate vi opposite page 348 of Volume I of the present Memoir. This plotting by Hay (1924) also includes finds of Parelephas jeffersonii and of P. progressus (listed under the names ‘Elephas boreus,’ 1924, pp. 47-56, and ‘Elephas columbi,’ 1924, pp. 57-84). As shown in the present Memoir, the crania and grinding teeth of the various species of Mammonteus and Parelephas, after years of confusion by all paleontologists including the present author, are now readily distinguishable. The plotted distribution (Fig. 1005) is entirely out of date (1938); it is now superseded by figure 795 in Chapter XV above, in which the range of the southern ‘trogontherians’ and northern ‘mammoths’ is theoretically plotted.—Editor.] 1134 OSBORN: THE PROBOSCIDEA Pleistocene life in the very important geographic area of Alaska, on the confines of North America and Asia, has been treated in recent years by Maddren (1905), by Gilmore (1908), by Quackenbush (1909), by Osborn (1910. 346), and by Frick (1930). A synthesis of their observations is recorded in the accompanying map of 1930 (Fig. 1004.) Alaska, broadly connected with the Asiatic mainland, was a peninsula for free migration between Eurasia and America; the fossil fauna thus far discovered, chiefly of Pleistocene times, is of boreal or cireumpolar char- acter but includes mammals of the north temperate latitude, such as the horse, the bison, a single foot bone of KEY @ MAMMONTEUS PRIMIGENIUS © M.COMPRESSUS CEQUUS ALASKAE, E.LAMBE! © CAMELOPS @RANGIFER SU SYMBOS TYRELLI | MARCTODUS YUKONENSIS @< GENERA NOT SPECIFIED | OMASTODON AMERICANUS XX BOOTHERIUM BOMBIFRONS A BISON CRASSICORNIS, BISON OCCIDENTALIS || @ OVIBOS MOSCHATUS, 0. YUKONENSIS, 0. MAXIMUS | BISON, MAMMONTEUS, EQUUS, FELIS ATROX ALASKENSIS J) | | Aicwccan DIRUS ALASKENSIS — (FRICK- KAISEN 1929) 130 a SSS 160 = —— SS = — Fig. 1004. Fossil mammalian fauna of Alaska-Yukon and location of principal discoveries to the year 1929. Modified after Quackenbush (1909, see full bibliography, pp. 128-130, and map), including discoveries and records of Maddren (1905) and identifications of Gerrit S. Miller, Gidley, Hay, Lambe, also of Frick (1930). As shown by the symbols, Mammonteus primigenius was extremely abundant; M. prim. compressus, represented by the single paratype specimen from Eschscholtz Bay; Hquus surprisingly abundant, there being recorded thirty-three localities, including (Wainwright) the north coast; Mastodon americanus very rare, being recorded only in the upper Canadian Yukon, on the Indian River, Long. 140, Lat. 64; and six records of Ovibos. In the very rich explorations, made through the cooperation of the Fairbanks Exploration Company, Frick, Bunnell, and Kaisen (1929) discovered the two new types Felis atrox alaskensis and 4nocyon dirus alaskensis, resembling like forms from the Rancho La Brea, southern California, intermingled with very numerous remains of Bison cras- sicornis, Equus, Mammonteus,” ete. Compare Mastodon americanus alaskensis Frick, 1933, on page 176 of Volume I of the present Memoir.—Hditor.] *[Mammonteus equals Mammonteus primigenius alaskensis sp. nov., from near Fairbanks (Long. 148.10 W., Lat. 64.59 N.) the description of which will be found on page 1159 below.—Editor.] THE MAMMONTINA: MAMMONTEUS 1135 a camel, and the recently discovered lion" or tiger (Felis atrox alaskensis Frick). Alaska was free from glaciers , except in its central mountainous belt. Scattered remains of mammals occur: (1) In the frozen tundras along the northern coast, (2) in the black muck accumulated in gulches and valleys of the smaller streams, (3) in the fine elevated clays of the ‘Yukon silts’ and the ‘Kowak clays,’ (4) in the more recent fluvial and alluvial deposits. In the northerly clays of Kotzebue Sound, Eschscholtz Bay (1907-1908), Quackenbush discovered a female mammoth skeleton (Amer. Mus. 13749) with portions of hair and wool perfectly preserved. Recent stripping operations of the U.S. Smelting and Mining Company, on Gold Stream and Clery Creek near Fairbanks (Fig. 1004), yielded the following (Frick, 1930, p. 73): “. . . during the four summer months some twenty-eight large cases of skulls, jaws, and bones—rare and important evidence on the prehistory of Alaska which otherwise would have been lost to science. The great percentage of this material, interestingly enough, came from three restricted areas, ‘bone pits,’ scattered between several widely separated operations of the Company, the re- mainder of the worked areas being, for the purposes of the bone hunter, nearly barren.” Pre-Tunpra or Fosst. Fauna or ALASKA-YUKON Mammonteus primigenius Blum. Symbos tyrelli Osgood Bison alleni Marsh Equus lambei Hay Mammonteus primigenius compressus Osb. Bodtheriwm sargenti Gidley Ovis sp. Camelops sp. Gidley Mastodon americanus Kerr Ovibos yukonensis Gidley Oreamnos sp. Arctodus yukonensis Lambe Rangifer sp. Ovibos moschatus Zimm. Equus alaske Hay Anocyon dirus alaskensis Frick Bison crassicornis Rich. Felis atrox alaskensis Frick S NS ZZ Amer: Geogr: Soc. NY) GEOGRAPHIC RANGE OF MAMMONTEUS AND OF PARELEPHAS Superseded by figure 795, Chapter XV, above Vig. 1005. This diagram shows that these two mammoths, confused in all previous descriptions, were north temperate and circumpolar in distribution. Closest to the Pole and to the northern ice-sheets during the Fourth Glaciation (Wisconsin-Wiirm) was doubtless the range of Mammonteus primigenius; closest to the 40th parallel was doubtless the chief range of Parelephas trogontherti, P. intermedius, and P. jeffersonii. It is possible that Parelephas trogon- therit and its successor P. intermedius migrated (diagonal) at the close of 1st or 2d Interglacial time across northern Asia, thence across Bering Strait into Alaska and southward to the 40th parallel of the United States. ‘According to Frick’s researches (letter, March, 1930), the cranium of Felis atroxr alaskensis resembles that of the lion (Felis leo) rather than that of the tiger (Felis tigris). OSBORN: THE PROBOSCIDEA TINAE: Mam “0 eee (1) BEE ~ EQUATOR Fo. ena Both | = 1) Fig. 1006. Geographic distribution of the principal species of Mammonteus. where the types of these twenty-nine species and subspecies were discovered. The white crosses represent referred specimens. 3. HISTORICAL ORDER OF NAMING OF SPECIES OF MAMMONTEUS EXCLUSIVE OF SPECIES WHICH ARE NOW KNOWN TO BELONG TO PARELEPHAS TROGONTHERIT], ETC. Late Pleistoce ne uf. 1799 1799 1807 1807 1815 1820 1829 1829 1829 1829 1829 1829 1830 1832 1832! 1832 1832 1832 1832 1832 1832 1834 See Figure 1006 SPECIES AS NAMED IN EUROPE AND ASIA Elephas primigenius Blumenbach, Osterode (Harz), Germany, and =Mammonteus primigenius Siberia Elephas mammonteus Cuvier (1796 MS.), Siberia, ete. = Mammonteus primigenius Elephas mammouth Link (fide Sherborn, 1928, Pt. XV, p. 3845) = Mammonteus primigenius Elephas primevus Blumenbach (in Adams, p. 152), near mouth of the = Mammonteus primigenius Lena River, Siberia Elephas primordialis Blumenbach (in Tilesius, p. 470) = Mammonteus primigenius Elephas jubatus Schlotheim, Diisseldorf, Germany = Mammonteus primigenius Elephas mammonteus Fischer de Waldheim, Russia = Mammonteus primigenius Elephas paniscus Fischer de Waldheim, Volga, Russia = Mammonteus primigenius Elephas periboletes Fischer de Waldheim, Podolia, Russia = Mammonteus primagenius Elephas pygmaeus Fischer de Waldheim, district of Calomna and of = Mammonteus primigenius Zwenigorod, Russia Elephas campylotes Fischer de Waldheim, borders of the Bug, Russia = Mammonteus primigenius Elephas Kamenskii Fischer de Waldheim, Siberia = Mammonteus primigenius Mammuthus Borealis Burnett (name only) [= Mammonteus primigenius] Mammut Sibiricum von Meyer (p. 64) = Mammonteus primigenius Elephas brachyramphus Brandt (pp. XI, XII), near mouth of the Lena River, Siberia. Type: ‘Adams skeleton’ = Mammonteus primigenius [Elephas| homotaphrus Brandt [Nomen nudum] Elephas giganteus Brandt, Indigirka River, Siberia. Type: Skull discovered by Messerschmidt (see Fig. 991 above) = Mammonteus primigenius Elephas commutatus Brandt, Volga River, Russia = Mammonteus primigenius Elephas stenotoechus Brandt, Russia = Mammonteus primigenius Elephas platytaphrus Brandt, Russia = Mammonteus primigenius Elephas affinis Brandt, St. Petersburg, Russia Elephas macrorynchus Morren, Tamise, Belgium = Mammonteus primigenius The white dots within the black areas represent the approximate localities {In the Bibliography (Vol. I, p. 766, of the present Memoir) the following species described by Brandt (nos. 4, 13-18 ine.) are given as of date 1833. It has been found since this Bibliography was compiled that a separate publication, containing these names and descriptions, appeared in 1832; hence the adop- tion of this date. Editor.] THE MAMMONTINA: MAMMONTEUS 1137 20. 1835 EHlephas odontotyrannus Kichwald, banks of the Nieman, dist. of Novogrodek, gouv. Vilna, Russia = Mammonteus primigenius 21. 1845 2H. [Elephas] primigenius sibiricus' de Blainville (Atlas, ‘“Ostéographie,” Pl. 1) = Mammonteus primigenius 22. 1879 Elephas primigenius comune Issel [in Zuffardi, 1913, p. 136], Campo- rosso, near Ventimiglia, Italy = Mammonteus primigenius (fide Issel, 1879, p. 160) Upper to Lower 23. 1888 Hlephas (primigenius) Leith-Adamsi Pohlig, Dornap (Fuhlrott),= Mammonteus(?) primi- Pleistocene Thuringia, Germany genvus leith-adamsi 24. 1891 LHlephas primigenius Blum. var. hydruntinus Botti, Otranto, Italy = Mammonteus primigenius hydruntinus 25. 1912 Hlephas primigenius Fraasi Dietrich, Steinheim, Germany = Mammonteus primigenius fraasi 1913 EHlephas primagenius Blum. var. trogontheriz Pohl. Zuffardi = Mammonteus primigenius mutation astensis (see No. 26 below) Upper Pliocene’ 26. 1923 EHlephas primigenius mutation astensis Depéret and Mayet, San Paolo == Mammonteus primigenius de Villafranea, Italy astensis 2. SPECIES AS NAMED IN NORTH AMERICA Upper Pleistocene 27. 1842 EHlephas americanus DeKay, Monroe County, near Rochester, = Mammonteus primigenius New York americanus 28. 1924 Mammonteus primigenius compressus Osborn, Rochester, Indiana = Mammonteus primigenius COMPTeSSUS 29. 1933 Elephas primigenius alaskensis Osborn (in Frick, nomen nudwm),= Mammonteus primigenius vicinity of Fairbanks, Alaska alaskensis sp. nov. Possibly twenty or more of the above species are synonyms of Mammonteus primigenius and probably belong to the Upper Pleistocene drift and Postglacial deposits of the Fourth Glaciation. Four of the above, more primi- tive, of greater geologic age, and with probably lower ridge formule, are as follows: 1888 Hlephas primigenius (?) Leith-Adamsi Pohlig, Nova Acta Leop. Carol., LILI, Nr. 1, pp. 229, 282. 1891 Elephas primigenius var. hydruntinus Botti, Bol. Soc. geol. Ital., IX, p. 709. 1912 Hlephas primigenius fraasi Dietrich, Jahresh. Ver. vaterl. Naturk. Wiirttemberg, LX VIII. This species is fully annotat- ed below. 1923 Elephas primigenius mut. astensis Depéret and Mayet, ‘Les Eléphants Pliocénes,” Deuxiéme Partie, pp. 183, 184. This species is fully annotated below. 4. GEOLOGIC AND PROVISIONALLY ASCENDING PHYLETIC ORDER OF SPECIES AND SUB- SPECIES OF MAMMONTEUS. EURASIA AND NORTH AMERICA Falconer and Hay have referred to Elephas primigenius the American specimens of the true woolly mammoth, the ridge formule of which remain to be more precisely determined. Falconer attributed to the American speci- mens the same ridge formula, namely, M 3 31, but remarked that the ridge-plates were more compressed or closer together; this compression reaches the highest possible stage in Mammonteus primigenius compressus, in which See footnote on p. 1391 below where it is stated that 2. primigenius sibiricus and other subspecies “perhaps may be regarded as geographic designations rather than as .subspecies.’’—Editor.] *{Possibly Lower Pleistocene (see footnote 1 on p. 1049 above).—Editor.] 1138 the ridge-plates increase to M 3 37. OSBORN: THE PROBOSCIDEA six! species and subspecies have been described as follows: Upper Pleistocene Alaska, Indiana IV Guactau and Postglacial Alaska and the Upper Pleistocene United States IV Guacrau and Postglacial Siberia and west- Upper Pleistocene ern Europe IV Guactrau and Postglacial 3d(?) Interglacial Western Europe Mammonteus primigenius compressus Osborn Mammonteus primigenius americanus DeKay Mammonteus primigenius primigenius Blumenbach Mammonteus primigenius fraasi Dietrich Altogether, including Upper to Lower Pleistocene and Upper Pliocene stages, = The most extreme or progressive subspecies known M 3 27 M27 =Slightly progressive American stages M 33¢: =Typical or true mammoth M 3 34 = Ancestral Upper(?) Pleistocene subspecies of Mammonteus primigenius M 3 a5 3d(?) Interglacial Thuringia, Mammonteus primigenius Leith-Adamsi =Small primitive branch of the Germany Pohlig main stem of Mammonteus primigenius =(Cf. Faleoner, 1868, II, p. 170): Small “slightly thick” ... pre-glacial variety of Elephas primigenius from the Norwich coast (ef. Fig. 1020). = Ridge formula: (ef. Fig. 1019) Forest Bed, East Anglia Mammonteus primigenius astensis (?) ref. Depéret and Mayet Lower Pleistocene Italy Mammonteus primigenius astensis type M 38 7+2=3¢ Upper Pliocene? Depéret and Mayet As shown above, the typical or true mammoth was found in Siberia long before it was recognized in western Europe. Blumenbach had in mind both Siberian and North German specimens in defining Elephas primigenius in 1799 (1799.1, p. 697), also Cuvier had the Siberian mammoth in mind in defining EHlephas mammonteus in 1796 (MS., published in 1799). Unfortunately Blumenbach in his type description mentioned as an example the Burgtonna skeleton, which we now know belongs to ‘Elephas antiquus’ and is not the type. The exact evolutionary stage of ‘Elephas primigenius’ was finally defined and determined by Falconer in 1863 (pp. 64,65), with a typical ridge formula of M 34. Attributed, or referred, to the typical species Elephas primigenius have been all the Pleistocene mammoths of western Europe and North America including, as we have seen, all the species of Parele- phas also all the primitive and geologically ancient species and subspecies of the true mammoth. The American subspecies Mammonteus primigenius americanus has the same ridge formula but the plates are more compressed (fide Falconer). Consequently the recognition of the geologically ancient forms with relatively low ridge formule, as listed above, is a matter of the utmost importance and interest. We regard it as somewhat problematic whether the Upper Pliocene Mammonteus primigenius astensis Depéret and Mayet, 1923, is ancestral to Mammonteus; certainly it shares characters of Mammonteus and of Parelephas; its ridge formula of M 3 +*:2° is higher than that of the contemporary Parelephas species P. trogon- therioides. The Forest Bed stage, previously determined by all authors as the true Mammonteus primigenius, more similar to M. primigenius 9-20 probably belongs to a primitive stage with fewer ridge-plates, perhaps M 3 722%, astensis. From among the considerable number of Forest Bed specimens probably a correct ridge-plate count can be made. To these should be added the new subspecies Mammonteus primigenius alaskensis (see pp. 1159 to 1161 of this chapter for description).—Editor.] *(See footnote 1 on page 1049.—Editor.] THE MAMMONTINA: MAMMONTEUS 1139 AURIGNACIAN Mammoru Hunters or Moravia.—The following passages may be freely cited from Doctor Absolon’s recent description in the Illustrated London News (Nov. 23, 1929) of his remarkable discoveries in the years 1924-1929 of mammoth-hunting stations of Moravia: Of these stations we now know about one hundred in Moravia, but only a small part of their area is explored as yet... . It is because Moravia was a kind of passage, by way of which the fossil mankind of the increasing Aurignacian tribe penetrated from Asia through Russia to the West of Europe. ... They are: Véstonice, Piedmosti, Petfkovice, Pekarna, and Ondratice. Pekarna isa cave (Fig. 1); the rest of them are in loess on the slopes of hills. PekArna is the most important, Véstonice the largest of them... . This was the first culture of the Old Aurignacian invasion of mammoth- hunters coming from Asia, the cradle of mankind. .. . One thing is certain—namely, that the mammoth-hunters killed these huge pachyderms in hundreds, and that in diluvial Moravia, a great tragedy, like the destruction of elephants in Africa, took place. ... In the refuse-heap discovered in 1925, no tusks were found in the whole area of 45 square metres; but in that encountered in 1926, on the other hand, three heaps of tusks were piled one over another, between which was left a narrow path (Figure 13). In the same year an imposing sight opened before our astonished eyes, when we discovered a field of huge pelvic bones (Fig. 14) of adult mammoths. The skulls are usually broken to pieces, because mam- moth-brain was appreciated by primitive hunters as a delicacy. Nevertheless, we found an intact skull of strikingly large propor- tions in 1928 (Figure 15). Long bones (femurs, tibiz) were found also in strange position forming a half-circle, so that their broken ends all pointed in one direction: evidently the fire was kept alight by the fat which flowed out of the ends of the burning bones into the flames... . Lower jaw-bones lie generally apart, and the teeth have often keen knocked out and piled up in heaps. In 1927 we found at Piedmosti a jaw-bone within which was a red-painted stone club (Fig. 9) which might have been used for striking out the teeth from the jaw-bones—a unique discovery. From 1924 to 1929 we count- ed as many as sixty mammoths, all of them caught and killed by man, on the area of 1600 square metres. There cannot be the least doubt that the hunters did not attack these powerful animals ‘face to face,’ but caught them by cunning, enticing or driving them into large pitfalls. The picture (Figure 16) shows a stratum dipping abruptly downwards. It must have been purposely made; dug in diluvial times. We intend to try to open this pitfall, for such I take it to be. Mam- moths trapped and caught were killed by large stones, trimmed to serve such a purpose. These stones might have been suspended in strong leather straps and thus let down on the animals by the united efforts of several men, in the same way that navvies drive piles into river-beds by means of rams. I have found one such stone, trimmed like a big pear, or bomb, 1 metre long, and weighing over 120 pounds (Fig. 11). [See Fig. 1037, this chapter.—Editor.] Fig. 1007. the diluvium of Moravia, now in the Moravian Government Museum of Brno (Brinn), Czechoslovakia. After photograph furnished by Prof. Karel Absolon Skeleton of the Mammoth (Mammonteus primigenius) from of the University of Prague and Curator of the Museum at Briinn. This skeleton was found nearly complete at Piedmont in Moravia, whereas all the skeletons in the kitchen-middens of the mammoth hunters of Moravia were scattered and partly destroyed for a food supply. It measures 3 m. or 9 ft. 10 in. in height and 4.42 m. or 14 ft. 6 in. in length. III. SYSTEMATIC DESCRIPTION OF SPECIES OF MAMMONTEUS Speciric Reviston.—(1) All the earlier writers even up to the time of Lydekker’s ‘‘Catalogue of the Fossil Mammalia in the British Museum” (1886.2) included within the typical species EKlephas [= Mammonteus] primi- genius all the progressive and primitive mammoths with fine-plated molars discovered in Pleistocene deposits, from the typical horizon of the Fourth Glaciation downwards to the Forest Bed stages of the Lower Pleistocene. (2) They also included within E. primiagenius, as we have seen above in the Parelephas chapter, many stages properly belonging to Parelephas trogontherii and its ascending and descending mutations. (3) Within the last half century, however, Pohlig (1888) and Dietrich (1912) have separated the stages of 2d and 3d Interglacial times in Germany as Elephas primigenius leith-adamsi Pohlig, 1888, and as Elephas primigenius fraasi Dietrich, 1912; (4) also Depéret and Mayet (1923) believe they have recognized an Upper Pliocene (Villafranchian) stage, to which they have given the name Elephas primigenius mut. astensis. Thus there have been four great steps in specific revision, concluding with the present revision by Osborn. As shown in the table above, the highly specialized Mammonteus conforms with the principle of ridge-plate addition as we pass from Upper Pliocene to the close of Pleistocene times, so beautifully manifested in the two more generalized mammontines Archidiskodon and Parelephas. Wemay select the third superior and inferior molars to show this parallel ridge-plate progression of M 3 in two of these three great lines of ascent. Mammonteus Parelephas Final Pleistocene M. primigenius compressus: ML Dies P. progressus: M 3 3% ?Postglacial M. primigenius americanus: M 3 $$ P. jeffersonii: M 3 3 IV GUuAcIAL or typical M. primigenius primigenius: M 3 P. columbi: M 3 1333. . ; M. primigenius fraasi: M3; P. intermedius: M 3 spor Upper Pleistocene f premigenaus sre : dee 2 ange _ i 4M. primigenius leith-adamsi: M 8 +324 P. trogontherii: M 3 it to Upper Pliocene |m . primigenius astensis: M 3 +8=3% P. trogontherioides: M 3 #24423 We observe that: (1) Mammonteus even in Lower Pleistocene time has a much higher ridge-plate formula than Parelephas; (2) but that Parelephas finally surpasses Mammonteus in the species P. progressus; (3) in Parelephas the ridge-plates are added more rapidly in the superior than in the inferior molars, a case of imperfect parallelism. Much further research is necessary to verify the ancestral relationships of these supposed early geologic stages to the typical Mammonteus primigenius primigenius. Also further research is essential to complete and verify the ridge formule of M. prim. fraasi and M. prim. leith-adamsi. In the meantime we may present the conclusions independently reached by Pohlig, Dietrich, and Depéret and Mayet that true ancestral stages of Elephas primigenius are to be found in the early Pleistocene and Upper Pliocene and that the ridge formule con- form with the general law of ridge-plate progression through the very long period of Pleistocene time, now esti- mated at not less than 1,000,000 years. 1. TYPICAL PROGRESSIVE EURASIATIC STAGES OF MAMMONTEUS There can be no doubt that during late Pleistocene (IV GuactAL and Postglacial) time there existed all over northern Eurasia a single species of mammoth to which the name Mammonteus primigenius has been assigned, since it furnished the type to which the earliest as well as the original and later descriptions were applicable. 1140 THE MAMMONTINA: MAMMONTEUS 1141 SYSTEMATIC DESCRIPTION OF SPECIES Mammonteus primigenius Blumenbach, 1799, 1803 Figures 794, 816, 818, 865, 934, 937, 962, 990-999, 1000-1002, 1007-1014, 1082-1034, 1042, 1062, 1084, 1226, Pl. xx1r Upper Pleistocene, [V Guactrat drift(?), Siberia and northern Germany. For synonyms, see page 1136. Tyre DESCRIPTION OF ELEPHAS PRIMIGENIUS, 1799.— (Blumen- bach, 1799, p. 697): ‘‘3) Von einem ungeheuer grossen Elephanten (Elephas primigenius?) [die vermeinten Riesenknochen** unsrer ehrlichen Alten]; unter andern auch in Menge in Deutschland***). So z. B. das beriichtigte Elephantengerippe das 1695 bey Burg- Tonna im Gothaischen ausgegraben worden etc.” Lecroryres.—Blumenbach’s type description (1799, p. 697) is in a rare document; no type figure accompanies it. Blumen- bach had the Siberian and German mammoth in mind. The ‘Ele- phantengerippe’ of Burgtonna is not the type. Dietrich and Osborn select (1930) as lectotypes: (1) A Siberian grinder in the Blumenbach Collection of the Zoological Institute of the Uni- versity of Géttingen, and (2) a molar from Osterode (Harz), Germany. The sixth edition of his ‘““Handbuch der Natur- geschichte,”’ in which the type description appears, attracted the attention of the French naturalist Soulange Artaud, who visited Blumenbach and translated his “Handbuch” into French under the title “Manuel d’Histoire Naturelle.’ In the manuscript of the French volume of 1803, Blumenbach inserted the name and defini- tion of Elephas primigenius, consequently the species Hlephas primigenius is of date 1799, 1803. Seconp Description, 1803 (BLUMENBACH-ARTAUD).—Many authorities, namely, Trouessart (1897, p. 711), Lydekker (1886.2, p. 175), and Hay (1902, p. 713), erroneously cite 1803 as the date of the type description of this species, ie., ‘“Manuel d’Histoire Naturelle, traduit de |’Allemand, de J. Fr. Blumenbach, . . . par Soulange Artaud,” 1803, Vol. II. The explanation of the definition of a new species in the translation is found in the ‘‘Préface du Tra- ducteur,” p. xvi: “C’est que j’ai travaillé sous les yeux de M. BLUMENBACH lui-méme; qwil a eu la complaisance de revoir mon manuscrit, et qu’ainsi je puis presque répondre de la fidélité de la traduction.”’ This shows that Blumenbach inserted the name and definition of EH. primigenius in Artaud’s MS. In Volume II, p. 407, of Artaud, we find the following definition: “3.° D’un éléphant énormément grand (elephas primigenius) (ce sont les prétendus os de géants de nos bons aieux); il se trouve des os fossiles de cet animal, en grande quantité, en Allemagne .. . (Le squelette d’éléphant, par exemple, qui a été trouvé, en 1695, prés de Burgtonna, dans le pays de Gotha).”’ In Artaud’s translation no figure of Hlephas primagenius oc- curs, although in the explanation of the figures appears the Ameri- can Mastodon designated as ‘28. Fossile Ohio incognitum”’ and on page 408 designated as “le mammouth de Ohio (mammout Ohioticum),” also the ‘“‘Table Alphabétique des Noms Latins, de Genres et d’Espéces” of Artaud’s translation, p. 443, gives the orthography as “Mammut Ohioticum.” Trovessart (1904-1905).—Trouessart (Quinquennale Sup- *f. Voigts Magazin. V. B. I. St S. 16 u. f. - plementum, 1904-1905, p. 600) remarks: ‘‘Le nom barbare de ‘Mammout,’ basé, par Blumenbach, sur une erreur grossiére (Videntité. du Mastodonte de V' Ohio avec l’Elephas primigenius), n’a aucun droit, malgré sa priorité, a étre substitué 4 celui de ‘Mas- todon,’ genre bien caractérisé par Cuvier.” Osborn, 1924: The above paragraphs from the quaint writings of Blumenbach and of Artaud firmly establish the specific name Elephas primigenius as of date 1799, 1803; they show that Blumenbach erroneously selected the Burgtonna skeleton! as an example of EL. primigenius; they show that there was no type figure. Finally, as pointed out by Trouessart, Blumenbach applied the barbaric name “Mammut” to the American Mastodon of Ohio under the erroneous idea that the latter was related to Elephas primigenius. TYPICAL UNIFORM RIDGE FORMULA OF MAMMONTEUS PRIMIGENIUS OF EUROPE (FALCONER, 1863).—As to the ridge formula of the mammoth, Falconer (1863, p. 64) remarks that in the European and American specimens the characters are constant, as defined by Cuvier and as witnessed in the British Museum collections from Siberia and in the American collections from Eschscholtz Bay. He concludes (op. cit., p. 65): “They all present, in the main, the same characters: a uniform ridge-formula; the same obtuse form of the lower jaw, and the same broad crowned molars, composed of closely com- pressed colliculi, with numerous digitations and attenuated un- crimped enamel-plates. .. . One of the most essential points, is to determine’ the constancy of the ridge-formula italics Osborn], which, after the examination ‘of a very large quantity of materials, I believe in the Mammoth to be thus: [Dp 2] 4, [Dp 3] 8, [Dp 4] #8, [M 1] 48, [M2] 44, [M3] #4.” Of this ridge formula he observes (op. cit., p. 65): ‘The plates advance by quaternary increments in each series, bearing in mind, that the first true molar, although of larger dimensions, commonly repeats the number of ridges presented by the last milk-molar, and that the last true molar in all the Elephants and Mastodons is more composite than the others. ... The formula in the North American Mammoth is identical with that of the Siberian and European forms. Exceptions are occasionally met, in which an unusual number of plates is presented.”’ COMPARISON WITH HELEPHAS (EUELEPHAS) INDICUS.—Fal- coner (1863) considers and repeatedly states that the ridge formula of E. primigenius is closely similar to that of the subgeneric group as typified by HE. (Huelephas) indicus in which he corrects his formula of 1857, p. 315, and substitutes (op. cit., p. 65) for E. indicus the following collective formula: Dp 24, Dp3$, Dp 413, M145, M246, M3 sea7- ***(Kriegsr. Merk) lettres sur les os fossiles d’elephans et de rhinoceros qui se trouvent en Allemagne &e. I-III. St. Darmst. 1783 u. f. 4. See footnote 2 on page 1118. 1142 OSBORN: THE Errors oF LeirH Apams, LypeKKrr, Hay.—Falconer, with unerring eye and logical mind, dwelt upon the constancy of the All 1/4 nat. size E. primigenius 22 Amer. Mus. 13752 Ref. A2 E. primigenius Amer. Mus. 13752 Ref. E. primigenius Amer. Mus. 14371 Ref. (inner view) B E. primigenius Amer. Mus. 14371 Ret REFERRED MAMMONTEUS PRIMIGENIUS MoLars OF ALASKA WITH TYPICAL M. pRIMIGENIUS RinGE FoRMULA Fig. 1008. Third superior and inferior molars (twenty-three to twenty- four ridge-plates) of Mammonteus primigenius, Alaska, of less compressed type (Amer. Mus. 13752, 14371). After Osborn, 1922.555, p. 10, fig. 9. The ridge formula of these typical specimens of M. primigenius corresponds very closely with that given by Falconer (1863) as characteristic of western Europe, namely, A, Crown view, left M*, twenty-three ridge-plates. A 2, Outer view of same tooth. It is apparent that one or possibly two of the anterior ridge-plates 23+ ig ou B 2, Inner view of right Mg of another individual, twenty-four ridge- B, Crown view of same. have been worn off, consequently the formula should be written: M plates. PROBOSCIDEA dental and cranial characters of Elephas primigenius. He did not confuse with this true species of mammoth (#. primigenius) the ridge formule of other species of elephants, as was done by sub- sequent writers, i.e., Leith Adams, Lydekker, lelix, Soergel, and Hay, who assigned to FE. primigenius the collective ridge formula derived from a considerable number of species belonging, as shown in this Memoir, to two distinct phyla, namely, to Mammonteus and to Parelephas. As to the ridge formula of Elephas primigenius, Felix (1912) erroneously follows Leith Adams in giving the ridge formula of EB. primigenius as M3 45757 aaa): This erroneous col- lective ridge formula first appears in Leith Adams (1879),! then in Lydekker (1886.2, p. 175), and finally reappears in Hay (1914, p. 395), as follows: Collective ridge formula: Dp 2 + Dp 3 $3 Dp 4 235 M124 M 224° M3 14-16 A 5 18-27° The above partly erroneous formula is undoubtedly based on the ridge-plate formule of species of Parelephas combined with the ridge-plate formule of species of Mammonteus. This confusion throughout Hay’s observations on the grinding teeth of Hlephas primigenius (1914, pp. 395-410) unfortunately renders his observa- tions of little systematic or phylogenetic value. Hay’s observations had been summarized for extensive citation in the present Memoir, but for the reasons given we are obliged to omit them. OsBorN’S OBSERVATIONS (1922-1924) on AMERICAN MusreuM SPECIMENS REFERRED TO MAMMONTEUS PRIMIGENIUS Osborn (1922) supports Faleoner’s observations of 1863. CoMPARE OSBORN, 1922.555, pp. 7-11.—Elephas primigenius is the name assigned to a series of species which ranged through the entire Pleistocene epoch, from the Lower Pleistocene Forest Bed? deposits of East Anglia to the southerly range of this animal in the middle United States. It is a collective species embracing an undoubted progressive evolution and intensification of specific characters extending over a very long period of time. It exhibits various extremes of fore-and-aft compression of the cranium with related fore-and-aft compression of the grinding teeth (hyp- sicephaly and bathycephaly). The cranium is high, pointed at the summit, relatively narrow, and relatively deep. The forehead from the peak of the cranium to the extremity of the nasals is rela- tively elongate and slightly concave. RipGe-PLATE CoMPRESSION.—As compared with F. colwmbi and F#. imperator, 10 ridge-plates of M. primigenius are compressed into a line 100 mm. in length in the typical M. primigenius of Germany; the most highly compressed tooth observed by Osborn is an M® from Alaska (Amer. Mus. 13749 [paratype of M. primi- genius compressus}) in which 18 ridge-plates are compressed into a line 100 mm. in length (Fig. 1024); a similar condition prevails in a female skull from Indiana (Amer. Mus. 14559 [since made the type of M. primigenius compressus]), in which 13 ridge-plates are compressed into a 100 mm. space, the total number of ridge-plates rising to 27 (Mig. 1022). 1See Bibliography of the present Memoir, Vol. I, p. 762, under Leith Adams, 1877-1881. °The Forest Bed Mammonteus primigenius astensis(?) is diagrammatically figured (Fig. 871, Archidiskodon, Chapter XVI, the present Chapter, Fig. 1020) in comparison with the contemporaneous molars of Parelephas (C, D) and with Archidiskodon (E-H). This figure by the author shows (A) the thin, fine-plated or “slightly thick” (fide Falconer, 1868, Vol. II, p. 170) enamel characteristic of Mammonteus at the beginning of Pleistocene time. THE MAMMONTINA: MAMMONTEUS The prevailing condition, however, is of the kind shown in figure 1008, namely, Amer. Mus. 13752, from Alaska, in which the ridge formula is M 3 *5*, here figured with a lower molar (Amer. Mus. 14371), in which the ridge formula is M 3 5;; in these speci- mens there are 10 plates in a 100 mm. line; these less compressed molars are arcuate, thus the count of the ridge-plates is greater on the concave side of the tooth and smaller on the convex side. For example, in a superior molar, M* (Amer. Mus. 10656), from Germany, the compression is as follows: M?® 10 ridge-plates in 100 mm. on the external convex surface, 11 ridge-plates on the horizontal mid-coronal surface, 12 ridge-plates on the internal concave surface. In the lower molars, these M* conditions are reversed in count- ing the ridge-plates in which the external surface is concave and the internal surface is convex. For example, in an M; from Alaska (Amer. Mus. 14343) the count is as follows: Ms; 10 ridge-plates in 100 mm. on the external concave surface, 10 ridge-plates in 100 mm. on the horizontal mid-coronal surface, 8-9 ridge-plates in 100 mm. on the internal convex surface. In the highly compressed superior tooth, M* (Amer. Mus. 13749 [paratype of M. primigenius compressus]), from Alaska (Fig. 1024), the count on all three measurements is the same, namely: M® 13 ridge-plates in 100 mm. on the external convex surface, 13 ridge-plates in 100 mm. on the horizontal mid-coronal surface, 13 ridge-plates in 100 mm. on the internal concave surface. It follows from the above observations that the horizontal mid-coronal section gives, as a rule, an average between the internal and external sides; the average in M. primigenius is 10 ridge-plates in 100 mm., the minimum observed is 8 in 100 mm., the maximum observed is 13 in 100 mm.! This variation in compression applies to fifteen specimens ranging through England, Germany, Siberia, Alaska, Ohio, and Indiana, and characterizes the typical M. primi- genius M? of Thuringia, Germany (cf. Soergel, 1912.2, Tab. vir, VIII). The worn mid-coronal surface is sometimes fully horizontal, ie., at right angles to the perpendicular ridge-plates; in such case it registers the exact distance between the plates. In other cases the wear is obliquely horizontal; in which case it increases the actual distance between the plates. It is important to note also that the ridge-plates are arcuate and more closely compressed towards their summits; thus more ridge-plates may be counted in 100 mm. at the summit of the crown than at the base of the crown, and as a rule the ridge-plate count should be taken midway between the summit and the base, both on the internal and external sides. In Mammonteus primigenius compressus Osborn.—Editor.] 1143 COMPARISON WITH PARELEPHAS AND MAMMONTEUS PRIMIGENIUS COMPRESSUS By the above observations of Osborn, the typical ridge formula of M 3 of Mammonteus primigenius agrees precisely with that es- tablished by Falconer in 1863, namely: M 3 34. This relatively low ridge formula agrees approximately with that of Parelephas Jeffersonti, namely, M 3 3%, but the M® of M. primigenius is relatively shorter and deeper than the M? of P. jeffersoniz; conse- quently the ridge-plate compression is much closer in M. primi- genius. The ridge-plate compression is still closer in Mammonteus primigenius compressus, in which M 3 is the broadest and deepest proboscidean molar known, the formula being M 3 ;55, the ridge- plate compression rising to 13 in 100 mm. Undoubtedly inter- mediate ascending mutations will be found between these two stages of ridge-plate evolution; meanwhile they may be distin- guished as follows: Typical Mammonteus primigenius primigenius of northern Europe, Alaska, and the United States: ridge formula M 3 43+; ridge-plate compression in the superior molars 10-11-12 in 100 mm. Mammonteus primigenius compressus of Alaska and Indiana: ridge formula M 3 g;2;; ridge-plate compression 13 in 100 mm. A fresh survey of the true Mammonteus primigenius grinding teeth, of Alaska and the northern United States, will probably reveal a series of intermediate ascending mutations between these two extremes. When the grinding teeth which have been errone- ously referred to Mammonteus primagenius are eliminated, namely, all of those molars actually belonging to Parelephas, we shall prob- ably discover these transitional mutations. SKULLS AND JAWS OF MAMMONTEUS PRIMIGENIUS Compare Figures 865, 962, 1009, 1010, 1023 The true cranium of Mammonteus primigenius (Figs. 865, 962 C, C) is readily distinguishable from that of Parelephas trogonthervi (Fig. 865) and of P. jeffersonii (Fig. 962 A) with which it has been confused except by Pohlig. The jaws of M. primigenius (Fig. 962 C, Cl, and Fig. 1011 C, B, A) are also clearly distinguishable. In this connection observe the characters pointed out by Falconer, Leith Adams, Pohlig, and others enumerated above. CraNIAL Marertats Figurep HrrewituH (Figure 962).— Crania of the true Mammonteus primigenius, as distinguished from crania of Parelephas jeffersonii, are very rare in the United States. Fortunately there is a finely preserved cranium from Siberia in the United States National Museum (Nat. Mus. 8580), photographs of which are reproduced in figure 962; the lateral aspect of this cranium (Fig. 962 C) is perfect, but the frontal aspect (Fig. 962 C) is foreshortened and thus gives an erroneous impression of the height of this cranium. CRANIAL CHARACTERS.—AIl the distinctive characters of the true Mammonteus primigenius skull arise from the maximum fore- and-aft compression, resulting in bathycephaly and hypsicephaly, which exceeds by far that of any other proboscidean or other mammalian skull thus far known. (1) This compression brings the anterior rim of the orbits much closer to the occipital condyles (Fig. 962 C) than in either P. jeffersonzi (Fig. 962 A), or P. wash- ingtonii (B); (2) it elevates the occipital crest, which is relatively higher, more elevated, and more pointed, i.e., acrocephalic, in M. 1144 OSBORN: THE PROBOSCIDEA Comparison OF MAMMONTEUS PRIMIGENIUS (C, C!) AND or PareLErHas (A, B) Crania See also comparative figure (Fig. 865) of the European Crania Fig. 1009. Profile views of type and referred skulls of Parelephas jeffersonii (A), P. washingtonii (B), and Mammonteus primigenius (C) in the American Museum and United States National Museum, also front view of M. primigenius (C!). All figures one-twelfth natural size. A, Type of Parelephas jeffersonii (Amer. Mus. 9950), reversed. B, Referred skull of Parelephas washingtonti (Amer. Mus. Cope Coll. 8681). An aged individual. Main portion of the tusks not included. From Jonesboro, Indiana. From Whitman County, Washington. C, Referred skull of Mammonteus primigenius (Nat. Mus. 8580), from Siberia, with Jaws belonging to another individual (Nat. Mus. 8579), from Alaska. C!, Front view of same skull and jaws. primigenius (Vig. 962 C) than it is in P. jeffersondi (Vig. 962 A) or P. washingtonii (Fig. 962 B). (3) This fore-and-aft compression brings the auditory foramen of M. primigendus close to the occipital condyles; it increases the hypsicephaly, whereby the apex of the occipital crest is raised relatively higher above the grinding surface of M*; and (4) it accounts for the extreme hypsodonty of M?°, which is the highest and the shortest of all proboscidean molar teeth. (5) This compression or bathycephaly extends to the lower jaw which has an excessively short, deeply depressed ramus, terminating in a deflected and extremely narrow rostrum (Tig. 962 C, C'). (6) This vertical compression or hypsicephaly extends to the vertically placed tubular maxillo-premaxillary insertions of the incisor tusks (Fig. 1023), which are relatively longer, narrower, and deeper than those of any species of Parelephas. (7) The frontal or facial aspect of Mammonteus primigenius compressus (Fig. 1023) is quite distinct from the frontal or facial aspect of Parelephas (Vig. 961); the arches supporting the orbits appear relatively broader, because the frontals above and the maxillaries below are laterally compressed. Figures 865, 934, 937.—Comparison with crania referable to the typieal M. primigenius of western Kurope is afforded in figure 865, especially the frontal view (No. 2) inseribed “ 2. primigenius Ref., Fale., 1847, Pl. xum, Fig. xxtv’; this west European cranium may be considered a “typical Mammonteus primigenius.”’ As far as we can judge from Ialeoner’s figure (1847, Pl. xin, fig. XXIV) this cranium shows exactly the same characters throughout as the crania above described from Alaska and from Indiana; we observe especially the small anterior nares, the tubular tusk insertions, the extremely elevated occiput. Another “typical Mammonteus primigenius” eranium (Tigs. 865, 934) is that figured after Pohlig (Pohlig, 1891, p. 384, fig. 120—reversed); this cranium in profile view may be compared with that of the female skull (Mammonteus primigenius compressus) from Indiana (lig. 1023) in its extreme acrocephaly, hypsicephaly, fore-and-aft compression, deep M® insertion, and relatively plano- frontal and occipital surfaces. In brief, the west EKuropean crania of the true M. primigenius exhibit precisely the same characters as the true M. primigenius crania of Alaska and of Indiana. - They THE MAMMONTINA: completely confirm the specific and generic separation of the cranium of Mammonteus from the cranium of Parelephas and still more from the cranium of Hlephas. Doubtless other true crania of M. primigenius will be discovered in the United States and in Alaska; it is of the utmost importance that these should be care- fully exhumed and that they should find their way permanently to some museum. Figure 1010.—It is through the courtesy of Charles W. Gilmore of the United States National Museum that we reproduce herewith (Fig. 1010) a superb cranium and tusks of M. primigenius discover- ed in the Yukon Territory, Canada, and described by Gilmore (1908, Pl. vit); it is difficult to determine from the photograph whether this cranium belongs to the true Mammonteus or to Parelephas; we are inclined to relate it to Mammonteus. A profile photograph of this fine skull would at once determine the question of its affinity. CRANIUM OF MALE MAMMONTEUS PRIMIGENIUS OF THE YUKON Fig. 1010. Male skull and tusks probably referable to Mammonteus primigenius, found in gravel 42 feet below the surface on Quartz Creek, near Dawson, Yukon Territory, Canada, March, 1904. After photograph by Gilmore appearing in his important paper entitled, “Smithsonian Exploration in Alaska in 1907 in Search of Pleistocene Fossil Vertebrates, Second Expedi- tion,” 1908, Pl. vir. As observed in the text, the generic and specific reference of this skull is somewhat uncertain; the very prominent antorbital tuberosities resemble those in the type female skull of Mammonteus primigenius compressus of Indiana. Figure 1023.—Fortunate also is the preservation of the skull of a female mammoth (M. primigenius compressus type) in the American Museum (Amer. Mus. 14559) from Rochester, Indiana, which more correctly shows the frontal plane of the true Mam- monteus primigenius cranial type. JAWS OF THE TRUE MAMMONTEUS PRIMIGENIUS We have enumerated above (pp. 1141-1144) the chief obser- vations of Falconer, Leith Adams, Pohlig, Soergel, and other writers on the jaws of the European specimens of the true Mam- monteus primigenius. The characters observed in the American specimens are as follows: The first jaw (Nat. Mus. 6666—shown in Fig. 1011 C) is from Alaska, twenty-five miles north of Anvik. The second jaw (Nat. MAMMONTEUS 1145 Mus. 8579—TFig. 1011 B) is also from Alaska, and indicates twenty- four ridge-plates; the third jaw (Nat. Mus. 4162—Fig. 1101 A) is from Elephant Point, Alaska. We owe to Gidley exact information regarding the American crania and jaws referred to Mammonteus primigenius in the present Memoir. E. primigenius Nat. Mus, 6666 Ref. Cl @® LEED E. primigenius Nat. Mus. 8579 Ref E. primigenius Nat. Mus, 4162 Ref. All 1/8 nat GrowTH STAGES IN THE JAWS AND TEETH OF MAMMONTEUS PRIMIGENIUS Fig. 1011. Internal aspect of three jaws of typical Mammonteus primi- genius from Alaska, in the United States National Museum. All one-eighth natural size. Sections Cl, Bl, Al. C, Juvenile jaw of Mammonteus primigenius (Nat. Mus. 6666) from Alaska, twenty-five miles above Anvik. Ridge formula: M 2 734, M 3 gaz. In My, at least one plate is missing in front of the alveolus of the anterior root, therefore there may be four or more plates missing; total preserved thirteen, maximum seventeen ridge-plates in My. In Mz no ridge-plates in use; 22+ ridge-plates developed out of the typical twenty-four ridge-plates characteristic of this species. B, Adult Mammonteus primigenius jaw (Nat. Mus. 8579) from Alaska, showing M3 in situ; seventeen plates in use, five additional plates, total twenty-two plates; apparently the two anterior ridge-plates of the typical number twenty-four have been worn off in this adult jaw. A, Aged Mammonteus primigenius (Nat. Mus. 4162), from Elephant Point, Alaska, M3 in situ; seventeen to eighteen plates in use, two additional plates unworn, nineteen to twenty plates present, apparently four anterior ridge-plates worn off out of the maximum twenty-four characteristic of this species. 1146 Jaw Cuaracrrers.—As in the ease of the skull, the jaws of the true Mammonteus primigenius have doubtless been partly confused by Faleoner and by all subsequent writers with those properly be- longing to Parelephas both in Eurasia and in North America. As shown in figures 962 and 1011 of this Memoir, the true Mammonteus jaws, with their true Mammonteus inferior grinding teeth, are harmonie with the extreme hypsicephaly and bathycephaly of the cranium. In proportion they are shorter anteroposteriorly, and deeper from the mandibular condyle to the symphysis of the mandibuli; the rostrum is deeper and more deflected; the section of the ramus is more rounded. These hypsicephalic proportions of the jaw are clearly dis- played in the referred adult jaw of Mammonteus primigenius (Nat. Mus. 8579) from Alaska, which is shown in side view in figure 962C and in mid-section in figure 1011 Bl; this adult jaw, both in ex- ternal aspect (Fig. 962) and in section (Fig. 1011 B, B1) displays the deeply depressed rostrum, also the relatively abbreviated third inferior molar, M3. In the young M. primigenius jaw (Nat. Mus. 6666) the rostrum is less depressed and as a whole is relatively longer and partakes more of the character of the young jaw of Hlephas indicus; in fact, all young jaws of Elephas, of Mammonteus, and of Loxo- donta are relatively elongate, and the deep, hypsicephalic pro- portion is acquired only in the adult. Fortunately three growth stages of the typical Mammonteus primigenius jaws are well displayed (Fig. 1011) in three specimens in the United States National Museum, as carefully drawn to a uniform one-eighth scale. COMPARISON OF MAMMONTEUS WITH PARELEPHAS AND ARCHI- DISKODON.— Exactly similar mid-sections of the jaws of Archidis- kodon and of Parelephas are shown in figures 892 and 893, by which it appears that the adult jaw of Mammonteus primigenius may readily be distinguished from adult jaws of Archidiskodon and of Parelephas in mid-section; also from the adult jaw of Elephas indicus (Fig. 893, D, D1) by the following bathycephalic characters: (1) Depression of the rostrum; (2) elevation of the coronoid and of the mandibular condyle; (3) narrowness of the space between the anterior border of the condyle and the posterior border of the angle; (4) relative abbreviation and depth of Ms, which is compressed into a much smaller, shorter space antero- posteriorly than that of Parelephas jeffersonii or of Archidiskodon imperator. This confirms the statement above, that the jaws and inferior grinders of the true Mammonteus are shorter, deeper, that is, more bathycephalic, in proportion, than the jaws of Hlephas, Parelephas, or Archidiskadon. FiscHER DE WALDHEIM, 1829.1, p. 285.—“C’est A Mr. le Baron Cuvier que nous devons le développement des caractéres anatom- iques qui font distinguer le Mammont ou l’espéce fossile d’Eléphant, de celles qui existent encore. Je me suis porté longtemps avec Vidée, que j’ai énoneée dans ma Zoognosie [Footnote: ‘Fischer, Weolh, INNE, joy 320.’|, que parmi les ossemens fossiles d’Eléphant pourraient étre Zoognosia tabulis synopticis illustrata. Mosquae. cachées plusieurs espéces, que nous confondons dans une seule, savoir celle du Mammont. La forme de la mAchoire inférieure, surtout celle des dents molaires et de leur lames nous conduisent OSBORN: THE PROBOSCIDEA i des différences, qu’on ne saurait admettre dans une et la méme espéce. Voici le résultat de mes recherches sur beaucoup de dents molaires que j’ai pu examiner et qui me fait distinguer les espéces suivantes: 1. EHlephas mammonteus, dentibus molaribus rectis, laminis numerosis angustis, parum elevatis, anguste fim- briatis. 2. Elephas paniscus, dentibus molaribus rectis, laminis latis elevatis, parum fimbriatis, latere longe distinctis. 3. Hlephas periboletes, dentibus molaribus rectis, laminis elevatis, profunde fimbriatis, oblique projectis. 4. Hlephas pygmaeus, dentibus molaribus similibus mam- monteo; sed magnitudine, plus quam dimidio minori- bus. 5. Elephas campylotes, dentibus molaribus subarcuatis, laminis angustis, numerosis, arcuatis, parum elevatis. Later in the same year in a Bulletin of the Société Impériale des Naturalistes de Moscou, Fischer (1829.2) reviews the above five species to which he adds a sixth (p. 276) under the designation of Elephas Kamenskii, as follows: ‘‘D’aprés cette maniére de voir, il faudrait nécessairement indiquer comme espéce distincte celle, Fig I Fig I A TypicaL TutrD SupER1ioOR MOLAR OF MAMMONTEUS PRIMIGENIUS Fig. 1012. Type of Zlephas odontotyrannus Kichwald, 1835, Pl. ux, figs. 1 and 2. One-third natural size. Last superior molar of the “right’’ [left] side, 1.M*. From Russia. THE MAMMONTINA: MAMMONTEUS i la quelle a appartenue la machoire inférieure de Siberie que nous devons a8. E. Mr. Bantich-Kamensky.”’ “6. EHlephas Kamenskii, molaribus subarcuatis utrinque attenuatis; laminis parum elevatis, numerosis, medio annulatis.”’ History AND SynonyMy.—Following the early descriptions of Ludolf (1696) and of Ides (1706), the mammoth waited nearly a century for the generic name Mammonteus Camper (1788)! and another decade (1799) for the specifie name primigenius Blumen- bach, after which it was fairly deluged with the host of generic and specific names listed above, which rank with those applied to the Mastodon among the curiosities of scientific literature. Cuvier (1806.1) traces the discovery of fossil elephants back to Theo- phrastus (born 372 B. C., died 287 B. C.), a pupil of Aristotle, and follows (op. cit., pp. 4, 5) with discoveries in Greece, Crete, and North Africa, continuing with a long account of the introduction of the domestication of the recent elephant in Europe and Africa and closing with a summing up of all the previous fossil discoveries of elephants of Upper Pliocene and Miocene times in all parts of Europe and northern Asia, as well as of the Mastodon in America. In the first and all subsequent editions of the ““Ossemens Fossiles”’ he recognizes only three species of elephants, living and fossil, namely, Hlephas africanus, Elephas indicus, and Elephas primi- genius, which he clearly separates by cranial and dental characters, as cited above (caption to Fig. 992). ApaMs (1807), TrLEestus (1815).—Adams described the animal as the mammoth; he states on page 152 that Blumenbach actually called the animal Elephas primxvus. Cuvier assigned the name Elephas mammonteus to the northern Mammoth (1799). Brandt named it Elephas brachyramphus (read 1831, published 1832). From the time of Camper (1788) and of Cuvier to the time of Tilesius (1815) a classical feature of the various descriptions is the constant use of ““mammonteum,”’ ““mammonteus,” and “mam- monteo”’ as the Latin designation of the mammoth, first as an adjective and then as a specific name, i.e., Hlephas mammonteus Cuvier, as compared with Elephas indicus Linneus. Among the outstanding points in Tilesius’ 100-page deserip- tion in Latin of the ‘Adams skeleton’ are the measurements given in “pedum anglicum”’ [=about 9 inches] and ‘‘pollicum anglicum”’ [=about 1 inch]. Tilesius gives the entire length of the skeleton, from the curvature of the tusks to the tip of the tail, as “‘viginti pedum” {[=about 15 feet in our measurement]. The vertebral column, according to the Adams-Tilesius descriptions of 1807, 1815 (written immediately on Adams’ return from the field), includes 28-30 presacrals, that is, from the first cervical to the last lumbar, as follows: Cervicals 7 Dorsals 17-19? Lumbars 4? Caudals 8+ Only the eight anterior caudals are preserved [see Tilesius’ figure reproduced herewith (Fig. 1014), which includes 7 cervicals, 1147 17-19? rib-bearing dorsals, and 4 lumbars, or 28-30 presacral vertebra]. Adams, in describing this specimen, used the quaint measure- ments and language of the time (Adams, 1807, ‘Relation d’un Voyage a la Mer Glaciale et Découverte des restes d’un Mam- mouth,” Journ. du Nord, St. Petersburg, translated by Sir Joseph Banks, Philosophical Magazine, 1808, Vol. XXIX, p. 148): “The parts least damaged are a fore foot and a hind one; they are covered with skin, and have still the sole attached. According to the assertion of the Toungouse chief, the animal had been so large and well fed, that its belly hung down below the knee joints. This mammoth is a male, with a long mane at his neck, but it has no tail and no trunk. The skin, three-fourths of which are in my possession, is of a deep gray, and covered with a reddish hair and black bristles. The humidity of the soil where the animal has lain so long, has made the bristles lose some part of their elasticity. The entire carcase, the bones of which I collected upon the spot, is 4 archines and a half high by 7 long, from the tip do é LOXODONTA AFRICANA OXYOTIS CENTRAL AFRICA 3568 MM..11/ BY2 ” INDIA BENGAL any NV AS Wl A , yah fl MAMMONTEUS PRIMIGENIUS N. EURASIA . 2999MmM.,9/10” Fig. 1013. Restorations by Margret Flinsch Buba, under the direction of Henry Fairfield Osborn, of Loxodonta africana oxyotis, Elephas indicus bengalensis, and Mammonteus primigenius. All figures to a one one-hundredth seale. See footnote on page 1117 above regarding the validity of the genus Mammonteus.—Editor.| 1148 OSBORN: THE PROBOSCIDEA WT ) y 7 4 Wipy ds Anacenwtu Slayns Cacrrs ( Flac’ Ut Clemocres de bcteadi mee Tome Sab X harmert abanterions parte Ou hameri a poiteriore fem Fig. 1014. ‘Adams skeleton’ of Elephas primigenius Blum. in the Zoological Museum of the Academy of Sciences of the U. 8. 8. R., Leningrad, Russia. Reproduced after Tilesius, 1815, Tab. x. About one twenty-sixth natural size. This was also reproduced by Cuvier in his “Recherches sur les Ossemens Fossiles,” Quatriéme Edition, Atlas, 1836, Pl. 17 (x1), one-fiftieth natural size. (Cf. Cuvier, op. cit., 1836, Explanation of Plates, p. 9): ‘Planche 17. Eléphans. Pl. x1. Fig. 1. Squelette entier d’éléphant fossile, rapporté par M. Adams de la mer Glaciale et copié d’aprés Tilésius. L’oreille a b a conservé quelques parties molles, et les pieds sont encore couverts de peau, et garnis de leurs semelles.”’ of the nose to the coceyx [Footnote: “An archine is a little more Cuvier (op. ciT., 1834, Vou. II, pp. 131, 1383, 204, 208, 231).— than two feet English measure.’|; without however comprehend- Cuvier describes this famous skeleton as follows: ‘‘La deuxiéme ing the two horns, each of which is a toise and a half long, and both est celle de |’éléphant rapporté 4 Pétersbourg par M. Adams, et together weigh 10 pouds [Footnote: ‘A poud is 40 pounds.’]. dont la conservation allait presque jusqu’au merveilleux. Le fait The head alone weighs eleven pouds and a half.”’ fut annoneé d’abord en octobre 1807, dans le Journal du Nord, Length of vertebral column 12-14 “pedum anglicum” {=9-10 feet] Neck short 17 “pollicum anglicum”’ [=17 inches| Entire length of skeleton, from forward arch of tusks to tuber- osities of ischium 20 “pedum anglicum’”’ {=15 feet] Height of skeleton 4 “archines” and a half |[=about 9 ft. 3 in| Length of tusks 1'4 “‘toise”’ {=about 9 feet] Length of humerus 40 “pollicum anglicum’” [=about 3 ft. 4 in.] Length of cubitus or ulna 35 “pollicum anglicum” [=about 2 ft. 11 in.] Length of femur 46% “pollicum anglicum” [=about 3 ft. 10% in.] Length of tibia 28 “pollicum anglicum” [=about 2 ft. 4 in.] Height of scapula 29 “pollicum anglicum’” [=about 2 ft. 5 in.] ‘{Lang in his article in Zoologica (1925.1, p. 28) gives a height of 9 ft. 11 in. at the shoulder, as mounted in the Leningrad Museum.—Editor.] THE MAMMONTINA:: recueil imprimé 4 Pétersbourg, n° xxx, et ce morceau, qui a reparu depuis en divers journaux allemands, a été réimprimé en 1815, dans le tome V° des Mémoires de |’Académie de Pétersbourg. MAMMONTEUS 1149 Measurements (Compare op. cit., p. 208) Crane de Mes- Crane du squel- Nous en tirons les détails qui suivent. En 1799, un pécheur serschmidt, ette de M. Tongouse remarqua sur les bords de la mer Glaciale, prés de Transact. philos., Adams, Pl. 17, Vembouchure de la Léna, au milieu des glagons, un bloc informe vol. xl, pl. I. [Fig. 1014] qu’il ne put reconnaitre. L’année d’aprés il s’apergut que cette Pl. 8, fig. I masses était un peu plus dégagée, mais ne devinait point encore ce [Fig. 991] que ce pouvait étre. Vers la fin de l’été suivant, le flanc tout entier Depuis le sommet jusqu’au bord de animal et une des défenses étaient distinctement sortis des des alvéoles 1,178 1,300 glacons. Ce ne fut que la cinquiéme année que les glaces ayant Depuis le sommet jusqu’aux fondu plus vite que de coutume, cette masse énorme vint échouer condyles occipitaux 0,663 0,770 Des condyles aux bords alvéolaires 0,946 4 la ete sur un bane de sable. Au mois de mars 1804, le pécheur enleva les défenses, dont il se défit pour une valeur de cinquante roubles. On exécuta, A cette occasion, un dessin grossier de l’animal, dont j’ai une copie que je dois 4 l’amité de M. Blumenbach. ... L’animal était male; ses défenses étaient longues de plus de neuf pieds en suivant les courbures, et sa téte, sans les défenses, pesait plus de quatre cents livres. . . . [p. 204] Encore plus récem- ment j’ai retrouvé ces longs alvéoles dans le crane des bords du Volga, gravé par M. Tilésius, et dont je donne une copie pl. 15, fig. 7. Il est vrai que ce caractére ne se montre pas sur le grand squelette de M. Adams, mais M. Tilésius nous dit expressément que les Tongouses avaient mutilé les bords des alvéoles lorsqu’ils en arrachérent les défenses, et qu’ensuite M. Adams, pour les y rajuster, fit encore rogner et égaliser ces bords [Footnote: Mém. de l’Acad. de Pétersb., t. v. (1815), p. 511.].”’ (Cuvier, op. cit., p. 231): “La peau est semblable 4 celle de Véléphant vivant, mais on n’y distingue pas les points bruns qu’on remarque dans l’espéce des Indes. M. Adams assure que la peau dont il avait conservé les trois quarts était d’un gris foneé. .. . [p. 232] M. Adams nous dit qu’une des oreilles de son individu était bien conservée et garnie d’une touffe de crins; mais dans son état actuel, comme on peut le voir, pl. 17 ab, fig.1, elle est fort altérée et n’a plus aucun poil. Les pieds du squelette de Péters- bourg sont encore couverts de peau et garnis de leurs semelles. M. Tilésius dit que ces semelles sont arrondies, et comme dilatées et foulées par le poids du corps; en sorte qu’elles remontent sur les bords du pied et les recouvrent. Il y avait quelque chose de semblable dans |’éléphant de la ménagerie de Versailles, décrit par Perrault. NiM. Adams, ni M. Tilésius ne nous parlent du nombre des ongles.”’ 2. PRIMITIVE EUROPEAN STAGES OF MAMMONTEUS PRIMIGENIUS As listed above (pp. 1136-1138), European stages of dwarfed dimensions, or supposedly more primitive than the typical Mammonteus primigenius of late Glacial and Postglacial times, were described in the following order: 1888 Mammonteus primigenius leith-adamsi Pohlig, a dwarfed 3d(?) Interglacial race of Thuringia, Germany. 1891 Mammonteus primigenius hydruntinus Botti, a dwarfed race of the cavern of Cardamone, Otranto, southeastern Italy. 1912 Mammonteus primigenius fraasi Dietrich, an Upper(?) Pleistocene stage of Steinheim on the Murr, central Germany. 1923 Mammonteus primigenius astensis Depéret and Mayet, an Upper Pliocene stage of Piedmont, northern Italy. Between the typical Mammonteus primigenius Blum. of northern Germany and the Upper Pliocene’ M. primigenius astensis of northern Italy there are doubtless numerous intermediate ascending mutations which only by very close monographic research can be clearly distinguished from each other. It must be remembered, how- ever, that Mammonteus was not a permanent resident like the members of the Hesperoloxodon antiquus or Parelephas trogontherii_ phyla, but a southerly and northerly migrant during the advances and retreats of the four great glaciations. Pohlig regards his subspecies leith-adamsi (Fig. 1015) as a diminutive variety of 3d(?) Interglacial time in the forests of Thuringia, while Botti describes also as a dwarfed subspecies his hydruntinus’ (Fig. 1016) from the caverns bordering the Otranto Straits, southeastern Italy. Dietrich clearly distinguishes his subspecies fraasz (Fig. 1017) as a full-sized race, with pentadactyl manus and pes, of Middle [Upper?] Pleistocene time, living on the Possibly Lower Pleistocene (see footnote 1 on p. 1049 above).—Editor. } Bearing on the presence of Mammonteus primigenius hydruntinus in southeastern Italy is the unique note of James Smith (Proc. Geol. Soe., London, 1847, p. 52): “The fragment which accompanied the specimen was given to Mr. Smith by Mr. St. John of Valetta, who stated that he found it, encrusted with stalagmite and adherent to the rock, in the island of Gozo. According to Dr. Falconer, it consists of two plates of a young grinder of the true Elephas primi- genius. The occurrence of so large an animal in a locality of such limited extent seems to point to a period when it was connected with a continent.” 1150 OSBORN: THE PROBOSCIDEA borderland between the forests and steppes of central Germany. Of great phyletic significance, if confirmed, is the very primitive stage MW. primigenius astensis (Fig. 1019) of the Upper Pliocene of northern Italy, described by Depéret and Mayet, and also observed in the Forest Bed, Lower Pleistocene, of Anglia (Fig. 1020); this primitive M. p. astensis is distinguishable by its thicker enamel and fewer ridge-plates (M 3 ,,4;) as compared with M 3 34 of the typical Upper Pleistocene M. primigenius. The supposed phyletic order of descent of these primitive species is given above on page 1138; the following are the type descriptions and figures in chronological order. Mammonteus primigenius leith-adamsi Pohlig, ingia, Dornap (Fuhlrott), Germany. Type Frgurn.—Op. 1888 cit., p. 229, fig. 101, c, d. Figure 1015 Type Derscription—Op. cit., p. 232: “. . . Noch fremd- artiger ist ein pachyganaler Mandibelzahn aus den Steinbriichen von Dornap (Fuhlrott), weleher—15 x in mindestens 0,22 ca. x 0,08 m aufweist; . . . Den vorstehend angefiihrten Belegen der typischen, der angusticoronaten und der pachyganalen rheinischen Mammuthform reihen sich in Folgendem einige héchst bemerkens- werthe Beispiele der diminutiven Primigeniusvarietat in den Bonner Sammlungen an; es mag geeignet sein, dieses Zwergmammuth durch eine besondere Bezeichnung, etwa E. (primigenius) Leith- Adamsi n.f. von der Stammform zu unterscheiden.” Trouessart (1897) cites Hlephas (primigenius) Leith-Adamsi as a subspecies of primigenius. His citation is as follows (op. cit., p. 711): ‘“d.—Leith-Adamsi, Pohlig (Var. minor), loc. cit., 1889.” Pohlig’s letter (Sept. 10, 1924) does not authenticate var. minor as a subspecifie term. Following the original description of the species in 1888, Pohlig in 1892 refers again to this species as perhaps one of the dwarf elephants which found its way to the mainland (op. cit., 1892.1, pp. 260, 261): “Es ergibt sich also der inter- essante Schluss, dass gleich dem Zwergmammuth, Hlephas Lezth- Adamsi, auch der mediterran insulare Zwerg-Urelephant Elephas Melite stellenweise wieder mit dem Festland communiciren konnte; —dieser jedoch offenbar, nach der grossen Seltenheit seiner Ueber- reste daselbst zu schliessen, nur auf ganz kurze Zeit.’”* 3d(?) Interglacial Travertines of Thuringia, Dornap (Fuhlrott). In 1888, Pohlig described this diminutive variety of Mammon- teus primigenius as a branch of the main stem of the true Mam- monteus primigenius. It was found in Thuringian deposits of 3d(?) Interglacial time and is consequently more ancient than the typical Mammonteus primigenius, which belongs to IV Guactau and Postglacial times. E. (Elephas| (primigenius) Leith-Adamsi Pohlig, 1888. “Dentition und Kranologie des Elephas antiquus Fale. mit Bei- triigen tiber Elephas primigenius Blum. und Elephas meridionalis Nesti,’”’? Nova Acta Leop. Carol., LIII, Nr. 1, pp. 229, 232. Typr.—Third left inferior molar, 1.M;. Original in Bonn. Horizon AnD Locauiry.—3d(?) Interglacial travertines of Thur- Mammonteus primigenius hydruntinus Botti, 1891 Figure 1016 La Grotta Ossifera di Cardamone in Terra d’Otranto, Italy. This subspecies is based upon a twelve ridge-plated first su- perior molar of the left side, 1.M! (Fig. 1016); the validity of this subspecies is dependent upon its geologic age. The title of Botti’s paper is ‘‘La Grotta Ossifera di Cardamone in Terra d’Otranto” and consists of descriptions of fossil bones found in this cave, including a molar and other portions of an elephant to which he assigned the name H. primigenius Blum. var. hydruntinus. The specific reference and type description are as follows: E. primigenius Blum. var. hydruntinus Botti, “La Grotta Fig. 1015. Type figure (c,d) of EB. (primigenius) Leith-Adamsi. After Qgsifera di Cardamone in Terra d’Otranto,” Boll. Soe. geol. Ital., ee aaaor BBO Vol. IX, 1891, p. 709, Tav. xxv1. Fig.101. Ein maxillarer und zwei mandibulare letzte Molaren von Elephas primigenius (in e—d von E. pr. Leith- Adamsi). Originale in Bonn (*/,). 1The type figure (Fig. 1015c, d) and description of the type third left inferior molar do not convince Osborn (1980) that this tooth belongs in the true Mam- monteus primigenius phylum. It appears too long and narrow, the ridge-plates are too widely separated, and the enamel is too thick. A reéxamination of the type would settle this question. °As noted above, the type third left inferior molar of ‘F. (primigenius) Leith-Adamsi’ in the Bonn Museum does not appear to clearly establish its re- lationship to the Mammonteus phylum. THE MAMMONTINA: Typr Description.—(Botti, op. crt., p. 705): “Uno di questi, un 1° molare vero, superiore, sinistro (tav. XxvI, fig. 1, la) presenta una corona alta 90™™ e la lunghezza di 122™™; la superficie triturante, ’estremita posteriore non essendo ancora scoperta, si riduce alla lunghezza di 105™™, la sua larghezza massima, misurata nel terzo anteriore 60™™; questa superficie inclina leggermente verso interno, ma non é convessa, bensi piana, particolarita che ricordo avere osservata in tutti gli altri molari superiori di Carda- mone, la quale, sebbene oppugnata da coloro che vogliono i molari superiori a superficie convessa, concorda coll’assioma dato da Fal- MAMMONTEUS 1151 lamine anteriori. Di tallone anteriore non scorgo traccia, sebbene una certa depressione, la quale potrebbe dipendere dal contatto del dente che lo precedeva nella serie.”’ “(P. 706] L’altro, molto consumato dall’uso, alto 55™™, (tav. XXVI, fig. 2, 2a), ha la superficie triturante lunga 97™™, la quale presenta la maggior larghezza di 46™™ nel terzo posteriore, concava dal davanti all’indietro, leggermente inclinata verso il lato interno. Ha parimenti dodici lame, manca affatto di radici e ritengo essere Vultimo molare di latte o premolare, inferiore, sinistro, della stessa sunnominata specie.” “T dischi resultanti dal consumo delle lamine per effetto della masticazione, sono, in ambedue i molari, quasi rettie stretti, niente allargati nel centro: le lamine di smalto, leggermente flessuose, corrono senza notevoli restringimenti, quasi parallele, ma note- volmente increspate.”’ “(Pp. 708, 709]. In conclusione, questi piccoli molari di Car- damone, che non trovano riscontro, che io sappia, se non in un molare (modello) del Museo di Firenze, asserto proveniente da Roma, ed in quello del Museo di Torino, proveniente dalle alluvioni del Po, in Italia, e dipoi in due individui del Museo nazionale di Budapest, hanno un facies affatto differente da tutti gli altri in TypE oF MAMMONTEUS HYDRUNTINUS Fig. 1016. Type 1.M! of Elephas primigenius Blum. var. hydruntinus Botti, 1891, Tay. xxvi, figs. 1, la, 2, 2a. From the Grotta Cardamone near Otranto, Italy. Scale (left figure) one-half natural size, height of crown 90 mm., length 122 mm.; (right figure) a milk tooth, height 55 mm., length 97 mm., breadth 46 mm. coner: ‘Constant character of mammoth’s molars of all ages and regions: worn surface nearly flat’ [Footnote: ‘Falconer H. Paleonto- logical Memoirs and Notes. Vol. I, p. 285.’}.”’ “Ed infatti undici lame in uso, le ultime quattro a piccoli dischi non ancora divenuti confluenti, ed una non ancora scoperta mi fanno ritenere per questo dente la formula di 12 lame, non com- preso il tallone, formula appartenente all’ultimo dente di latte (secondo Owen e Falconer) o premolare (secondo Blainville) ovvero ad un antepenultimo (ossia primo) molare vero di 2. primigenius Blum.” “Mi fanno inclinare a quest’ultima opinione la forma massiccia del dente e la presenza di robuste radici, sebbene troncate nel- Yesemplare in esame, in specie di una anteriore, isolata dalle suc- cessive, che sono fra loro saldate, la quale sostiene le due prime grandissimo numero esistenti nei Musei colla determinazione di L. primigenius Blum.; hanno lo smalto pit fino e pit finamente in- crespato, le lame pit serrate e pitt avvicinate, le dimensioni della meta pit: moderate.” “‘Nuove specie furono create con minor divario di forme e l’ele- fante di Cardamone differisce certamente dal primigenius tipico pit di quello che il primigenius differisea dall’indicus; ma con- siderando che il molare di Roma e quello delle alluvioni del Po, non che quelli di Budapest, furono raccolti prima di quelli di Carda- mone e nessuno penso a separarli dalla specie primigenia, e riflet- tendo che in sostanza e poi un solo e medesimo piano di struttura che si sviluppa, sebbene in grado diverso, nell’ HE. primigenius Blum. e negli elefanti di Cardamone, cosi stimo miglior partito di lasciar questi ultimi nella specie primigenia, tutt’al pit proponendo 1152 OSBORN: THE di distinguerli, poiché sono in fatto distinti, in ragione della dimen- sione di una meta minore e della pit sensibile ristrettezza dei loro elementi, come una varietai, che potrebbe chiamarsi, dalla regione di provenienza: E. primigenius Blum var. hydruntinus.” “Questa distinzione d’altronde non mi appartiene del tutto.” “Gia fino dal 1873, faceva notare Leith Adams alla Zoological Society considerevoli discrepanze nei molari a lame ordinariamente sottili del Mammuth, ed allegava una autorita altamente compe- tente, quella di M. Davies del British Museum, il quale, dall’esame di numerosi molari di Elephas primigenius, trovati in Inghilterra ed altrove, era da assai tempo venuto nella opinione che esistano due distinte varieta, facili a riconoscere, i molari dell’una essendo formati di lame sottili separate da ristretti strati di comento interposti, l’altra composta di lame pid dense con intervalli pit larghi, soggiungendo quest’ultima forma essere pil. frequente Meee dell’altra [Footnote: ‘Leith Adams A. op. cit. p. 6, 7. Mammonteus primigenius fraasi Dietrich, 1912 Figures 1017, 1018 hy wal “Mitteldiluvialen Schottern in Steinheim a.d. Murr,” Germany. This subspecies, named in honor of Eberhard Fraas by Profes- sor Dietrich in 1912, is based on a type skeleton mounted under the direction of Professor Fraas in 1911 in the Museum of Stuttgart. As displayed in the type figure (1912, Taf. 1, 11), it appears to represent a Middle [Upper?] Pleistocene race or subspecies of Elephas primigenius clearly characterized by its large size, fewer ridge-plates in lower molars, and five digits in the manus and pes, probably a typical phalangeal formula. Elephas primigenius Fraasi Dietrich, 1912. ‘Elephas primi- genius Fraasi, eine schwibische Mammutrasse.” Jahresh. Ver. f. vaterl. Naturk. in Wiirttemberg, Jahrg. LXVIII, pp. 42-106. Typr.—Skeleton, adult male. Mounted under the direction of Professor Fraas in the K. Naturalienkabinett, Stuttgart (No. 12837). Horizon AND Locatiry.—Found in 1910 in ‘‘Mittel- diluvialen Schottern in Steinheim a. d. Murr.” Type Fic- URE.—Op. cit., Taf. 1 and 01, also text figs. 2, 4, 11-14, 16-21, 24-26. Tyrer DescripTion.—(Op. cit., pp. 105, 106): “E. primigenius Fraasi, wie ich die neue Rasse Herrn Professor Fraas zu Ehren nenne, ist also keineswegs synonym mit der eben erwihnten Be- zeichnung, worunter lediglich eine Anzahl bestimmter Molaren- formen zu einer Mutationsreihe zusammengefasst werden. 2. vielmehr innerhalb Reihe eine distinkte, osteologisch vollstiindig bekannte Form, in deren Skelett primigenius Fraasi ist dieser eine Reihe von Merkmalen vereinigt sind wie es bisher an keinem anderen Mammut beobachtet ist. Ausserdem ist der 2. primi- genius Fraasi eine charakteristische Lokalform, denn die Stein- heimer Funde zeigen immer wieder das gleiche gewaltige Ausmass der Glieder.” Observe the relatively primitive structure of M, (ef. Dietrich, 1912, p. 76): M; max. sin. Fig. 12. Lamellenformel x 22 x [Footnote: “Die PROBOSCIDEA 13. Lamelle ist nur an der Labialseite entwickelt.’| in 27 em Linge, 22 em Hohe [Footnote: ‘An der 16. Lamelle.’], 10,4 em Breite, Abkauungsgrad x—17. Lamelle. ... Distanz der Lamellen 4—8 mm., Linge der Lamellen 7—8 mm (an der labialen Seite), Form des Lamellenumrisses, unten verschmilert, Dicke des Schmelzes 1, 4—1, 5 mm (starke Krauselung), Typus der Ver- schmelzungsfigur der Lamellen, median lamellar, lateral annuliir. (P. 106): “Diagnose: Grosse hoch- beinige, fiinfzehige Mammutrasse mit kurzem und schlankem Rumpf. Im Schiidel und den Molaren mit primiti- ven Merkmalen. Schiidel verhaltnis- miissig niedrig, Molaren weitlammellig- er als bei den jiingsten Mammuten. nach Fig. 1017. Type cranium of the subspecies Hlephas primigenius fraasi Dietrich, 1912, Taf. 11, figs. 1 and 2 [=Mammonteus primigenius fraasi of the present Memoir]. This cranium, associated with the skeleton now mounted in the K. Naturalienkabinett in Stuttgart (No. 12837) illustrates the extreme hypsi- cephaly and acrocephaly, fore-and-aft compression, of the profile of the fronto- occipital region and the marked concavity of the frontals, agreeing closely with the cranial characters of all species of Mammonteus and differing widely from the cranial profile of Parelephas. THE MAMMONTINA: Carpus und Tarsus hochentwickelt, der erste aussen rein serial, innen nahezu serial. Die Minnchen gehéren zu den Riesenformen; sie tragen starke, gebogene, aber nur wenig spiral gedrehte Stoss- zihne, deren Wachstum einer Hemmung unterliegt. Die Stoss- zihne werden gebraucht.” “Typus: Das Skelett No. 12837 (co? adult) im K. Naturalien- kabinett zu Stuttgart.” Hasits oF ExerHas [MAMMONTEUS] PRIMIGENIUS FRAASI (ef. Dierricu, 1912, p. 42).—In FL. [Mammonteus| primigenius the enamel folds barely rise above the cement, affording a relatively 5 Vig. 1018. present author by Dr. W. O. Dietrich. Compare Dietrich, 1912.1, Taf. 1. least 4 m. or 13 ft. 1}4 in. (see Dietrich, op. cit., p. 64). MAMMONTEUS 1153 even surface which is admirably designed for the comminution of The mammoth is thus chiefly a grass-eater, It also is an open country animal, but we cannot of course deduce the finer grasses. a fact which is also attested by the contents of the stomach. from the structure of its teeth whether this country was warm or cold, for while it is occasionally found mingled with an arctic fauna this is not invariably the case. The remains at Steinheim prove that it lived on the borderland between the forests and the steppes because it is found in association with H#. [Hesperoloxodon| antiquus. Mounted skeleton of Mammonteus primigenius fraast (Naturalienkabinett, Stuttgart, No. 12837), after photograph kindly forwarded to the The skeletal height, from the sole of the foot to the top of the cranium, is at 1154 OSBORN: THE Mammonteus primigenius astensis Depéret and Mayet, 1923 Figures 871, 1019, 1020 Upper Pliocene (Villafranchian stage),! San Paolo de Villafranca, northern Italy. If confirmed by more material, the recognition by Depéret and Mayet of an Upper Pliocene stage in the evolution of the genus Mammonteus is a step of the utmost importance in the phylogeny of the Proboscidea and especially in the phylogeny of the genus The type grinders (Fig. 1019) appear to Osborn to If, however, we Mammonteus. be relatively narrower than we should expect. may depend upon the highly trained and acute observations of Charles Depéret and Lucien Mayet then Elephas primagenius mut. astensis actually belongs in the Mammonteus phylum and is quite PROBOSCIDEA sont celles de M* d’un sujet de petite taille d’#. primigenius, M* dont la longueur moyenne est, d’aprés Pohlig, de 300 millimétres et atteint parfois 385 millimétres. La fréquence laminaire est de prés de 8 lames pour 10 centimétres de couronne; c’est le chiffre normal del’. primigenius. Les bandes d’émail sont trés minces, bien plus que chez l’E. trogontherii du méme gisement; elles sont tout aussi onduleuses, mais beaucoup moins plissées, Vémail étant presque uni d’un bout 4 l’autre des bandes, sauf quelques plis dans les lames antérieures trés unies. II n’existe aucune trace du sinus loxodonte.”’ Fatconer, 1868. Vou. II, Pp. 170.—Faleoner described as follows a perfect 1.M* “of the pre-glacial variety of Elephas primi- genius. . . . The matrix is indisputably of the forest-bed of the Norfolk Coast.” Length 11.5 in. (292 mm.), breadth 4.1 in. (104 Fie. 1019. (Left) Type of Elephas primigenius astensis from San Paolo de Villa- franca. After Depéret and Mayet, 1923, Pl. x1, fig. 5, p. 221: “Fig. 5.— Elephas primigenius, mut. astensis de San Paolo de Villafranca. M® droite. (Voir p. 184). Musée du Palais Carignan, 4 Turin, figurée par Zuffardi, 1913, pl. v1, fig. 2a [as Elephas primigenius Blum. var. trogontherti Pohl.].”” Less than one-half natural size; actual length 216 mm., breadth 94 mm., height 186 mm.; nineteen ridge-plates present. separate from what these authors have named ‘“‘Groupe des Elephas Trogontherit.” EB. |Elephas| primigenius mutation astensis Depéret and Mayet. ‘Les Eléphants Pliocénes,’”’ Deuxiéme Partie, 1923, pp. 183, 184. Typre.—Right third superior molar, r.M*. Musée du Palais Carignan, 4 Turin. Horizon AND Locanity.—San Paolo de Villafranea. Typr Ficure.—Op. cit., Pl. x1, fig. 5, p. 221; figured by Zuffardi, 1913, Tav. x11 [vi], fig. 2a, as Hlephas primigenius Blum. var. trogontherti Pohl. Description oF Typr.—(Op. cit., p. 184): “Molaire supéri- eure M*.—ll existe au Musée de Turin les deux M* d’un méme sujet. Nous figurons (pl. x1, fig. 5) la dent du cdté droit qui est la mieux conservée. L’animal était déja assez 4gé, car la couronne est trés arasée en avant par la mastication et ne compte plus que 19 lames présentes, il doit en manquer quelques-unes, 2 ou 3 peut-étre. La longueur conservée de la couronne est de 216 millimétres et on peut estimer la longueur totale probable 4 249 millimétres. La largeur de la couronne est de 94 millimétres et la hauteur au milieu atteint le chiffre considérable de 186 millimétres. Ces dimensions (Possibly Lower Pleistocene (see footnote 1 on p. 1049 above).—Editor.] TypE AND PARATYPE GRINDERS OF MAMMONTEUS PRIMIGENIUS ASTENSIS (Right) Paratype of Hlephas primigenius astensis. After Depéret and Mayet, 1923, Pl. x1, fig. 6, p. 221: “Fig. 6.—Elephas primigenius, mut. astensis. Pliocéne fluvio-lacustre de l’Astesan, 4 San Paolo. Mz droite. (Voir p. 185). Piéce du Musée géologique de Turin, n° 15, figurée par Zuffardi, 1913, pl. v, fig. 8a [as Elephas primigenius Blum. var. trogontherti Pohl.|.’’ Less than one- third natural size; length 280 mm., breadth 84mm., height 138 mm.; nineteen ridge-plates plus talon. mm.), height 7 in. (178 mm.); ridge-plates 18+, fourteen in a space of 7}; in. (190 mm.); enamel slightly thick. This 1.M? is either of Mammonteus primigenius or of Parelephas trogontherir (ef. Fig. 1020). ParatyPE.—(Op. cit., Depéret and Mayet, p. 185): ‘Les Ms figurées par Zufiardi (1913, pl. v, fig. 8-9) ont une couronne rela- tivement étroite et fortement arquée comme dans la derniére molaire inférieure des Eléphants. Nous figurons (pl. x1, fig. 6) la dent du cété droit qui est la mieux conservée. La longueur de la couronne est de 280 millimétres mesurée en ligne droite pour une largeur de 84 millimétres et une hauteur de 138 millimétres. On compte 19 lames plus le talon et des traces de quelques lames usées ila partie antérieure. La fréquence laminaire est de 7, 548 comme chez EB. primigenius. Les ecaractéres de |’émail sont identiques 4 ceux des molaires précédentes: émail mince, uni et non plissé, sans sinus loxodontes.” Osborn, 1924: The relatively high ridge formula, M 3 234, is what we should expect in an ancestral stage of the phylum Mam- monteus, which in M. primigenius, at the close of the Pleistocene THE MAMMONTINA: age, rises to M 3 34, and finally in M. primigenius compressus to M327. From theabove very accurate description and admirable figures may be deduced the ridge formula of Mammonteus primi- MAMMONTEUS PRIMIGENIUS7ASTENSIS MAMMONTEUS PRIMIGENIUS?7ASTENSIS est Bed of Cromer, Norfolk PARELEPHAS ? TROGONTHERI! SP.? aE Ipswich Mus Forest Bed PARELEPHUS 7? TROGONTHERI! SP? Sayin Mus No./240 Forest Bed Bones aed. A. MERIDIONALIS CROMERENSIS Ipswich Mus ARCHIDISKODON MERIDIONALIS CROMERENSIS v £ - ss) S v G 9 ra Be o ~ Q = 9 9 & = cy Q 9 | 1 NAT. SIZE Ipswich Mus Red Crag Close of Pliocene time Fig. 1020. Mammonteus primigenius (?)astensis (A, B). Crowns of referred superior (M*) and inferior (Mz) molars, from the Forest Bed of Cromer, Norfolk, England, compared with (?) Parelephas (C, D), Archidiskodon (E, F), and Archidiskodon or Hesperoloxodon (G, H) molars. All figures one- fourth natural size, reproduced from direct scale tracings by the author. For details see figure 871, Chapter XVI. MAMMONTEUS M 3 72:56. This is a much higher ridge formula than that of the typical Parelephas trogontherii of Pohlig, namely, M 3 iss, although the type of P. trogontherii belongs to a more recent geologic stage than the type of M. primigenius astensis. The ridge formula of M. primigenius astensis is thus higher than that of any contemporary species of mammoth or elephant, much exceeding the contemporary ridge formule of species of Archidisko- don and of Parelephas (see Fig. 1020). genius astensis: MAMMONTEUS PRIMIGENIUS (?)ASTENSIS, ForEST BED, ENG- LAND (OsBorN, 1922, 1924).—The Cromer Forest Bed of England includes a sub-arctic faunal phase of the northern latitude of East Anglia, Lat. 53, during the period of the first Scandinavian glacia- tion, which has no parallel in southern France or in northern Italy, Lat. 44-46. The Forest Bed contains two distinet faunas, namely: I. Survivals.—Upper Pliocene Red and Norwich Crags warm types which survived in the Forest Bed and belong to the /st Intergleaial stage. II. Arrivals—Pleistocene cold types which first appear in the Forest Bed and may belong to the I GLActIAL stage. From the studies of Geikie, Prestwich, Clement Reid, and from the collections of A. C. Savin, Osborn in 1922 (1922.563, p. 439) summarized these two faunas as given below. In brackets are indicated the possible references of the Forest Bed proboscideans to the north Italian species, but to determine the exact specific and subspecific references of the Forest Bed proboscideans, namely, ‘‘Hlephas meridionalis,” “‘Elephas anti- quus,” and “Elephas primigenius,”’ very close examination and comparison will be required: (1) The ‘‘Elephas primigenius’” listed below may prove to correspond in its ridge formula with that of the Hlephas primi- genius mut. astensis Depéret and Mayet (M 3 {3-55), of Asti, northern Italy. (2) It is also possible that the ““Hlephas antiquus Nestii type” of the Forest Bed may prove to correspond with members of the Parelephas trogontherii phylum.! FOREST BED, OR CROMERIAN, FAUNA OF EAST ANGLIA J. SuRVIVALS Warm Upper Pliocene Red Crag and Norwich Crag Types which Survive in the Forest Bed of Cromer Elephas [Archidiskodon| meridionalis |cromerensis?| type Elephas antiquus Nestii type [= Parelephas(?) trogontherii nestii| Elephas |Hesperoloxodon| antiquus (typical) Hyena striata Hyena antiqua Rhinoceros etruscus Equus stenonis Cervus carnutorum Macherodus sp.? Trogontherium Cuvieri Macacus sp.? Rhinoceros megarhinus Hippopotamus amphibius II. ARRIVALS Cold and northern Pleistocene Types which Fst A ppear in the Forest Bed of Cromer Elephas [Mammonteus| primigenius [= astensis? | Equus caballus fossilis Sus scrofa Bison bonasus Caprovis Savinii Ovibos moschatus Alces latifrons Capreolus capreolus Cervus elaphus Ursus speleus Felis spelxa Varied forest rodent fauna Northern forest small Carnivora After this text was written, Professor Osborn referred ‘Hlephas’ antiguus Nestit Pohlig, cotypes, to Parelephas(?) trogontherit nestit (see Chap. XVII, above, p. 1059.—Editor.] 1156 OSBORN: THE PROBOSCIDEA 3. AMERICAN STAGES OF MAMMONTEUS In the full description and discussion above of the typical species Mammonteus primigenius, with a ridge-plate formula of M 3 #, as determined by Falconer, this species appears to range completely across Eurasia into Alaska, with the same ridge-plate formula. In Alaska also occurs! the much more progressive species Mammonteus 2 primigenius compressus (M 3 ;5;), which ranges southward into Indiana, probably subsequent to the time of the appearance of the typical M. primigenius. Unfortunately the type of M. primigenius americanus DeKay, Upper Pleistocene of New York, has been lost and further research is necessary to determine precisely the ridge formula of this stage, which is estimated in the present Memoir as M 3 24. Owing to the confusion by Hay and others of species of the true Mammonteus with species of Parelephas 3.0 jeffersonii, in which the ridge-plate formula ranges from M 3 #4 to 28, much careful research remains to be done in connection with the revision and determination of the species and subspecies of Mammonteus characteristic of the northern regions of North America. The tentative arrangement is as follows: Mammonteus primigenius: M 3 34, typical of Alaska and ranging southward. Mammonteus primigenius americanus: M 3 $88, typical of New York. I 7 Mammonteus primigenius compressus: \ So far as our present evidence goes, Mammonteus primigenius reached its final and most progressive stages of evolution in North America, probably in Postglacial time. Mammonteus primigenius americanus DeKay, 1842 Figure 1021 Upper Pleistocene, IV Guacrau (Wisconsin diluvium), Irondiquoit River, Monroe County, near Rochester, New York. The type of this species was in the Cabinet of the Lyceum of Natural History, New York, which was destroyed by fire. The name was alluded to by Leidy (1858.2, p. 29), also by Adams, Cope, and Pohlig. As DeKay mentions thirteen plates in five inches (=126 mm.), and also gives an excellent figure, it seems desirable to retain this subspecific name as Mammonteus primi- genius americanus DeKay. E. [Elephas| americanus DeKay, 1842. “Natural History of New York. Part I. Zoology: Zoology of New York, or the New York Fauna,” 1842, p. 101. Typr.—Portion of an upper molar tooth with thirteen ridge-plates. HorizoN AND Locauiry.—Irondiquoit River, Monroe County, near Rochester, New York. Tyre Ficurrn.—Op. cit., Pl. xxxu, fig. 2. Type Descriprion.—“It is with some hesitation that I ven- ture to designate, under a new name, a species founded on speci- mens of teeth, which appear to differ widely from any hitherto met in this country. ... The specimens above alluded to were found in Tyre OF MAMMONTEUS PRIMIGENIUS AMERICANUS a diluvial formation near the Irondiquoit river in Monroe county, Fig. 1021. Type of Hlephas americanus De Kay, 1842, Pl. xxxu, fig. 2, three-fifths natural size. Portion of an upper molar tooth with thirteen ridge- plates. Irom near the Irondiquoit River, Monroe County, in the vicinity of ; sae 5 Rochester, New York. Type formerly in the Cabinet of the Lyceum of Natural a tusk and two molars, one of which is in the Cabinet of the History, New York, destroyed by fire. Sig Nal six ten miles east of the city of Rochester. According to a writer in the American Journal, Vol. 32, p. 377, these remains consisted of [See new subspecies Mammonteus primigenius alaskensis from near Fairbanks, Alaska (pp. 1159 to 1161 of the present chapter).—Editor.] THE MAMMONTINA: MAMMONTEUS 1157 Lyceum, and is that figured in the plate. This is six inches in its greatest depth; and, as nearly as can be conjectured from the part which remains, it must have been about eight inches long, and three in breadth on its grinding surface, which is, however, too much injured to exhibit the ends of theenamel. There are thirteen plates in a space of five inches, and they are more compressed than in any fossil species with which I am acquainted, being almost in contact, with very little interstitial substance. It is altogether different from any fossil elephant hitherto described, and merits the distinct appellation of Z. americanus.” Fatconer (1863).—Elephas primigenius [americanus]. Fal- Al Amer. Mus. 14559 (outer view) anterior MAMMONTEUS PRIMIGENIUS COMPRESSUS. TYPE. FROM INDIANA Fig. 1022. Type second and third superior molars of female Mammonteus primigenius compressus (Amer. Mus. 14559), from Indiana, one-fourth natural size. The teeth (A1) belong on the right side of the beautiful female skull also represented in figure 1023. After Osborn, 1924.633, p. 6, fig. 2 (originally figured as Elephas primigenius, see Osborn, 1922.555, p. 8, fig. 8). Ridge formula: M 3 27 (2) 27° A 1, External view showing twenty-seven ridge-plates of M’. A, Crown view showing M® with eight ridge-plates in use, closely com- pressed to M? with eight ridge-plates in use. coner observed (1863, p. 66) that a constant character of the North American mammoth is that the ridges and their constituent ele- ments are more attenuated and condensed. Thus, one tooth with 14 ridges shows an average of .86 in.; another presents 17 discs with an average of .46in. Taken singly, the difference between the higher ridge condensation in American as distinguished from European specimens seems inconsiderable, but when it extends over a length of crown comprising 16 or 24 ridges, it is perceptible ataglance. It gives a certain amount of distinctive physiognomy to the molars of the North American mammoth. Falconer, how- ever, does not regard this as indicating more than a slight geo- graphical variety, as the other characters remain constant to the true mammoth type. Osborn, 1924: Faleoner’s observation that a constant char- acter of the North American mammoth is that the ridges and their constituent elements are more attenuated and condensed conforms with De Kay’s type description that “there are thirteen plates in a space of five inches, and they are more compressed than in any fossil species with which I am acquainted, being almost in contact, with very little interstitial substance.” Assuming that De Kay’s measurement of thirteen plates in a space of five inches was ac- curately made, we find that the ridge-plate compression of the type of Mammonteus primigenius americanus is as follows: 10+ ridge- plates in 100 mm. This would relate this subspecies more closely to the typical Mammonteus primigenius than to M. primigenius compressus, but while De Kay’s specific name may be retained, we can hardly consider that this can be raised to a higher rank than a subspecies, namely, Mammonteus primigenius americanus. Mammonteus primigenius compressus Osborn, 1924 Figures 806, 819, 1022-1024 Type from Rochester, Indiana; paratype from Alaska. Postglacial or retreat period, [V (Wisconsin) GLAcrIAL. Probably of the This subspecies is typified by the female skull (Amer. Mus. 14559), from Rochester, Indiana, and the paratype from Alaska (Amer. Mus. 13749), fully figured and described in this Memoir. It is readily distinguished from the typical Mammonteus primi- genius of Eurasia (M 3 34) by the very high compression of its dental ridge-plates, namely, M 3 q;s;- The original description by Osborn (1924.633) is as follows: Mammonteus primigenius compressus Osborn, 1924. ‘“Par- elephas in Relation to Phyla and Genera of the Family Elephanti- dex.”’ Amer. Mus. Novitates No. 152, Dec. 20, 1924, pp. 5-7. Typr.—Skull of a female mammoth including both tusks, also superior grinding teeth. Horizon AnD Locatiry.—Roches- ter, Indiana; Upper Pleistocene of Alaska and of the Central United States. Tyre Ficgure.—Op. cit., 1924.633, fig. 2, p. 6; and figure 1023 of the present Memoir. PARATYPE.—A third right superior molar, r.M*, from Alaska (Amer. Mus. 13749). PARATYPE FiGurE.—See Osborn, 1922.555, p. 7, fig. 7 (Fig. 1024 of the present Memoir). Type Description (OsBoRN, 1924.633, pp. 5-7).—The origi- 1158 OSBORN: THE PROBOSCIDEA nal description of this subspecies is in part as follows: “(1) Ex- typical Mammonteus primigenius of Eurasia and of North America treme fore-and-aft compression and vertical elevation (hypsiceph- displays a very constant ridge formula, as especially observed by aly, bathyeephaly), correlated with extreme hypsodonty and Falconer, namely: fore-and-aft compression of the twenty-seven ridge-plates which compose M*. (2) Measurement across outside of orbits, 262 mm.; from top of skull to bottom of premaxillaries, 393 mm.” Falconer himself remarked that while American specimens of the “After careful and prolonged examination of the specimens true H. primigenius display a similar formula, namely, M 3 34, and the descriptions of Faleoner (1863), we conclude that the the grinding teeth in general show more closely compacted ridges.”’ Dp 2¢ Dp 3s Dp 443 M1743 M276 M 3 34. MAMMONTEUS PRIMIGENIUS COMPRESSUS. ‘TyrE I'EMALE SKULL FROM INDIANA Fig. 1023. Type skull of female mammoth (Mammonteus primigenius compressus), Amer. Mus. 14559, from Rochester, Indiana, acquired by purchase in 1921. See occipitofrontal section of same skull (Fig. 806); also teeth of same skull (Fig. 1022). This skull is the type of the species Mammonteus primigentus compressus Osborn. One-twelfth natural size. Observe in the three aspects of the skull, frontal, palatal, and lateral, the following important points: (1) The extreme fore-and-aft compression (cyrto- cephaly, hypsicephaly or acrocephaly) correlated with the extreme compression of the twenty-seven ridge-plates which compose M® and are displayed in detail in figure 1022; (2) the slender parallel female tusk insertions and deep extension of the incisive alveolar plates; (3) this is the most extreme fore-and-aft com- pression of the skull, grinding teeth, and alveolar processes thus far recorded; (4) the measurement across outside of orbits is 262 mm.; from top of skull to bottom of premaxillaries 393 mm. The cranium of this type, as beautifully displayed in the figure at the right, exhibits the utmost extreme of vertical elevation (hypsicephaly) of the occiput and of vertical depression (bathycephaly) of the grinding teeth. In harmony with the almost straight vertical line of the forehead and of the condylar pro- cesses are the maxille and the sharply downturned tusks. The apex of the skull, half complete, would terminate in a sharp peak (acrocephaly). This harmonious fore-and aft compression and deepening of the entire cranium, which brings the occipital condyles relatively close to the nares, apparently exhibits the absolute- ly finest stage in the evolution of the Mammonteus cranium and dentition. Certainly this remarkable foreshortening, doubtless initiated in the M. primi- genius astensis of the Upper Pliocene [Lower Pleistocene?], could go no farther and we may regard this species as the final stage in the evolution of the phylum Mammonteus. THE MAMMONTINA: MAMMONTEUS “Tt is owing to the excessively high compression (hypsodonty) and multiplication of the plates (polydiskodonty), amounting to M 3 @s7, that the new subspecific name Mammonteus primigenius compressus is now defined and illustrated by figure 2 [Fig. 1022 of the present Memoir.” The type and paratype superior grinding teeth are clearly illustrated in figures 1022 and 1024 of the present Memoir, and the cranial characters of the type are also shown in figure 1023, a female cranium in which the summit of the occiput is unfortunately broken away. We do not at present know of any jaw in which the ~ grinders display this extreme stage of compression. Consequently the ridge formula of the lower teeth is not certainly known. The ridge formula of M 3 is probably @,7. E. primigenius A2 Amer. Mus. 13749 Ref. (inner view) ; 27 36789 1418 E. primigenius A Amer. Mus. 13749 Ret yy) 2 3S 5-Ge7-6.0- 1/4 nat. size MAMMONTEUS PRIMIGENIUS COMPRESSUS. PARATYPE. FROM ALASKA Fig. 1024. Paratype figure of right third superior molar, r.M*, of Mammon- leus primigenius compressus (Amer. Mus. 13749), from ‘‘Historic Bluff,” Eschscholtz Bay, Alaska, showing the maximum compression of the ridge- plates; ridge-formula M 3 Gree One-fourth natural size. After Osborn, 1922.555, p. 7, fig. 7. Mammonteus primigenius alaskensis sp. nov.! Figures 1025, 1026 From vicinity of Fairbanks, Alaska. Pleistocene. {In anticipation of Professor Osborn’s description of a series of crania of the northern Mammoth, discovered in 1929 near Fairbanks, Alaska, Mr. Childs Frick, who generously accorded Professor Osborn the privilege of first description, used (1933.1, pp. 506, 632) the manuscript name Elephas primigenius alaskensis. Mr. Frick not only listed the probos- cidean material collected in the Fairbanks region by the joint Alaska College-Frick American Museum parties, but he figured (Pl. 12A) a palatal specimen (A. C.—F: A. M. 27010) which he referred to this subspecies. While it was Professor Osborn’s intention to describe in Novitates this new ascending mutational stage, prior to the article by Mr. I’rick in the Bulletin of the American Museum of Natural History, the text was never completed; in order, however, that such portion of it as he had prepared may be recorded, we are inserting it herewith, including certain essential additions.—Editor. | Elephas primigenius alaskensis Osborn (in Frick, 1933). ‘New 1159 Remains of Trilophodont-Tetrabelodont Mastodons,” Bull. Amer. Mus. Nat. Hist., LIX, pp. 506, 632. Coryrrs.—Four crania (infantile, with deciduous dentition, A. C.—F: A. M. 26991; ju- venile, with deciduous dentition, A. C.—F: A. M. 26990; adult female, with left maxillary tusk, A. C—F: A. M. 26989; and aged male, Alaska College). Horizon AND Locauiry.—‘‘Bone pits’ scattered between several widely separated stripping opera- tions of the U. 8. Smelting and Mining Company, vicinity of Fairbanks and the Tanana River, Alaska. Pleistocene. Corypr Ficures.—Figures 1025 and 1026 of the present Memoir. Referred figure, see Frick, 1933.1, Pl. 12A. Frick, 1933.1, p.632.—“ Elephas primigenius alaskensis Osborn. Among the twenty-six tusks secured of the Northern Mammoth there is one of unusual size, the same weighing in the neighborhood of 300 pounds. The average number of lamine in the normal m is twenty-five and the length of the crown in use varies between 157-207 mm. A mandible of an aged cow is remarkable in that the last molars have been extruded and the alveoli healed over. Another mandible exhibits a somewhat produced symphysis.” 4 Natural size Wiolar plone vas MAMMONTEUS PRIMIGENIUS ALASKENS!S ‘ Corype CraniA OF MAMMONTEUS PRIMIGENIUS ALASKENSIS. MincrANIAL SECTION All one-eighth natural size Fig. 1025. Superposed cranial outlines: (1) Infantile, (2) juvenile, (3) adult female, and (4) aged male showing relations to molar plane (attrition). This superposition reveals the growth phases in the four chief cranial planes, namely, molar plane, basi-occipital plane, occipital plane, frontal plane, which culminate in the lofty occiput and depressed palate (bathycephaly), in the foreshortened occiput to anterior nares (brachycephaly), and in the concave frontal plane characteristic of all the adult Mammontine. 'Imasmuch as Mr. Frick in 1933, pp. 506, 632, quoted the name Elephas primigenius alaskensis from Osborn’s manuscript, without diagnosis or desig- nation of type, this subspecific name in Frick is essentially a nomen nudum and its unequivocal establishment dates from the present Memoir. (Note by Dr. George Gaylord Simpson, 1939).—Editor. | 1160 OSBORN: THE PROBOSCIDEA “List oF MATERIAL FROM THE VICINITY OF I'AIRBANKS, ALASKA 2 complete and 12 partial radii (largest 61 em., smallest 26 tusks (largest, on curve 12 ft. 10 in., base circumference 56 cm.) Sle) 8 complete and 8 partial femurs (largest 113 em., smallest Large bull skull with m*s and tusk (9 ft. 8 in. x 18.5 in.), 96.5 em.) Alaska College Collection 12 complete and 14 partial tibiz (largest 68 em., smallest Large cow skull with m’s and tusk (6 ft. 5 in. x 13 in.) 44 cm.) { smaller skulls, including 2 of calves 3 complete and 2 partial fibule 6 partial palates 37 metapodials Some 24 mandibles or partial mandibles 17 astragali 204 detached molars or partial molars 16 caleanea 31 partial seapule 60 carpal or tarsal bones 7 complete and 5 partial humeri (largest 98 em., smallest 6 patelle 75 em.) Numerous vertebrae 16 complete and 5 partial ulne Several ribs, partial ilia”’ All 4 Nat size Aged male (left side reversed ) Alaska College Infantile MAMMONTEUS PRIMIGENIUS ALASKENSIS Oss. Amer. Mus. Frick Collection CoryrE CraNnia oF MAMMONTEUS PRIMIGENIUS ALASKENSIS All one-twentieth natural size Mig. 1026. Cotype crania of Elephas [Mammonteus) primigenius alaskensis: (1) Infantile (A. C.-F':A. M. 26991); juvenile (A.C.-F':A.M. 26990); adult female (A.C.-F’:A.M. 26989); aged male (Alaska College, through the courtesy of President Charles E. Bunnell). Compare figure 1023 above. THE MAMMONTINA: MAMMONTEUS ASSOCIATED FAUNA (ef. Frick, 1930.1, p. 79): PRE-TUNDRA FauNA OF ALASKA- YUKON Large herds of the super-bison, Bison crassicornis Rich, ref. A new Alaskan lion, Fels atrox alaskensis, Frick A new wolf, Aenocyon dirus alaskensis, Frick An arctothere, Arctodus yukonensis Lambe, ref. A camel, Camelops (?) Three species of musk-oxen, Ovibos, Symbos tyrelli Osgood, and Bodétherium sargenti Gidley A horse, Equus alaskxe Hay, ref. A mastodon, Mastodon americanus Kerr, ref. A Mammoth, Mammonteus primigenius alaskensis The association of these mammals points to a Pleistocene phase which Frick inelines to compare with the Sheridan fauna of the Aftonian or /st Interglacial stage of Nebraska. Ossporn’s Manuscrirt (1931).—The chief result of intensive researches on the genus Mammonteus is to establish in the evo- lution of the genus a number of successive ascending mutations ancestral to, or descending from, a typical ‘Elephas primigenius Blum.’ (1799) of Siberia and north Germany to which Falconer (1863.1, p. 65) rightly attributed a very constant ridge formula, namely: D524 Dp3$ Dp422 M112 M212 M324 12 12 1161 In 1924 Osborn described (1924.633, p. 5) the most progressive form Mammonteus primigenius compressus: M 3 @s;. Recent explorations in Alaska have revealed to President Charles FE. Bunnell of the Alaska Agricultural College and School of Mines and to Mr. Childs Frick a more primitive ascending mutation to which the name Mammonteus primigenius alaskensis may be assigned. The Alaska College and the American Museum are indebted to the United States Smelting and Mining Company for the privilege of making a very careful survey of the fossils revealed during the recent stripping operations of the company on Gold Stream and Clery Creek near Fairbanks; the survey was conduct- ed under the supervision of Mr. Peter Kaisen during the seasons of 1929 and 1930. On this material Frick (1930.1, p. 73) reports as “.. the joint Museum and College party sueceeded in harvesting during the four summer months some twenty-eight follows: large cases of skulls, Jaws, and bones—rare and important evidence on the prehistory of Alaska which otherwise would have been lost to science. The great percentage of this material, interestingly enough, came from three restricted areas, ‘bone pits,’ scattered between several widely separated operations of the Company, the remainder of the worked areas being, for the purposes of the bone hunter, nearly barren.” TABLE XV. hypsicephaly) Width across summit of parietals (brachycephaly) Incisive premaxillary alveoli, length (bathycephaly) s ce ss width (hypsicephaly) Glenoidal breadth (brachycephaly) Incisive tusk, length on outer curve “ee iv maximum diameter me “maximum circumference Third superior molar, M°, length, anteroposterior ‘“ breadth, transverse “ height, vertical (e.) CRANIAL AND DentTAL MEASUREMENTS Occipital condyle to front of orbit (ecyrtocephaly, brachycephaly) Occipital summit to attritional molar plane (bathycephaly, | ues) 2 ao | > ~~ 2S NS = == = & 5S) a oS Ss | weiss 2% 2s tha a ait eet ce ao 6) 2O aa a <4 ee > 340e 515 550 600 357 618 705 815 287 | 453e 548 650 x 320e 550 520e x 195 318 380e 280e 410e 465 540 x <», Mamimonteus =Northern Mam moths Farele phas =Trogonthenan Mammoths Patlaeoloxodon = Straight-Tusked Elephants OSBORN: THE PROBOSCIDEA discovered the ivory tusks by digging in the earth, the word ‘mammoth’ is certainly derived from the alleged Tatar word mamma, signifying ‘earth.’ As early as 1696 this was combined with another Tatar word kost, signifying ‘ivory,’ and the two words were Latinized by Ludolf into Mammotovoi Kost. The buried tusks were sometimes mistaken for horns; Cuvier alludes to these words of Tatar origin as follows: ‘C’est sous le nom de cornes de mammont, mammontova-kost, quwils désignent les défenses.’”” Thus in the Latinization of the Tatar word mamma into the Gallic mammont there was assigned by Camper in 1788 the name Mammonteus, which is now applied by Osborn to the woolly mam- moth as a genus distinguished in many external and internal characters from the genus EHlephas which is typified by the elephant of India. Thus, as cited in full above (p. 1124) from Ides, originated the word ‘mammoth,’ variously spelled in different languages mammot, mamant (Russian), mammouth (French), mammuth (German), from the Tatar mamma because the remains of these animals were found imbedded in the earth, the natives therefore believing that the animals burrowed like moles. Un- doubtedly the woolly mammoth was in its time the colossus of living mammals of northern or western Europe; in consequence the word Stegodonts GrEoGRAPHIC DisTRIBUTION OF MAMMOTHS IN UPPER PLIOCENE AND PLEISTOCENE TIMES Fig. 1028. Partly theoretic geographic distribution (1930) of the southern, intermediate, and northern mammoths. See figure 795 for distribution map of 1938. SouTHERN MAMMOTHS: INTERMEDIATE MAMMOTHS: NORTHERN MAMMOTHS: genius alaskensis. Parelephas trogontherii, P. jeffersonit. Relatives of the southern imperial mammoth, Archidiskodon imperator, are now known to have Archidiskodon planifrons, A. meridionalis, A. imperator. Mammonteus primigenius, M. primigenius compressus, M. primi- ‘mammoth’ has become a convenient adjective to signify gigantic, immense, of great com- parative size. As a matter of fact, the woolly mammoth is a dwarf in comparison with its relatives, the Jeffersonian and imperial ele- phants, as is shown in the illustrations by Knight which have been brought to a rela- tively uniform seale (Figs. 1029, 1030, 1034). ranged from South Africa to India and from Nebraska to Mexico. [Since this was written, a speci- men referable to Archidiskodon cf. planifrons has been described from Shansi, China, by Dr. Arthur Tindell Hopwood (1935.1, pp. 87-90), who states as follows: “It is clear that there is a wide gap in the known distribution of the genus, and that the forms from the Old and New Worlds are separated by most of Asia. Hitherto no species of Archidiskodon has been described from the Far East, and the specimen described below is the first evidence that this gap may be expected to close with the march of knowledge.’’—Editor.] The temperate Trogontherian and Jeffersonian mammoths of the genus Parelephas ranged along the 40th parallel from southern France to central United States. The northern mammoth, of the genus Mammonteus, ranged about the Arctic Circle from western Europe to eastern North America. word Elephas means ivory. Aristotle (384-322 B.C.) in his (1926), and Tolmachoff (1929). Long before the elephant was known in Europe, Phoenician traders brought ivory from the South to Greece; in the North ivory was also procured in the tusks, the average weight of which was 288 pounds, of the more or less fresh or partly frozen bodies of the woolly mam- moths. This northern source of the primitive ivory trade was summarized by Herbert Lang (1925); it has been thoroughly studied in re- cent years by Basset Digby (1926), Pfizenmayer In Tolmachoff’s recent and History of Animals! used €Xépas in its modern generic sense as applying to the elephant of India, in referring to its courage in combat: ~. .. Elephants also fight fiercely with each other, and strike with their tusks; the conquered submits entirely, and cannot endure the voice of the victor: and elephants differ much in the courage they exhibit.’’ The word ‘mammoth’ has a similar non-classie and indirect origin. In allusion to the fact that the Tatars of Siberia first ‘Translation by Richard Cresswell, London, 1887, p. 234. comprehensive memoir (1929) is found a map (Fig. 1027 above) and a list of not less than thirty-nine localities where frozen car- casses have been found, beginning with Ysbrant Ides (1692) and ending with Andrews’ discovery of 1923. The author gives due credit to the works of Basset Digby and of Pfizenmayer. He also gives a most detailed and interesting history of the progress of discovery of the woolly mammoth, Hlephas primigenius, and of its woolly companion, Rhinoceros antiquitatis, the ‘rhinoceros of antiquity.’ The export ivory industry of Siberia, dating back THE MAMMONTINA: MAMMONTEUS to very ancient times, furnishes a very good idea of the immense number of mammoths that have been discovered by exploring par- ties in the frozen ground of Siberia, estimated by Middendorf (1885) at 20,000 during the past two centuries and by Nordenskiéld (1882) at a very much higher figure. The highest estimate is 46,750 for the last two and a half centuries, although one estimate even puts the number at 250 specimens annually, a total of 62,500 for the two hundred fifty years. The northern ivory trade to China is now traced back as far as 500 B.C. Doubt- less the value of northern ivory rose as the Chinese gradually exterminated their own native or imported breeds of elephants from India. Herbert Lang (op. cit., 1925) notes that Ides, the famous Duteh traveler and am- bassador to China, seems to have been the earliest to gather first-hand information re- garding the frozen Siberian mammoths.' In traversing northern Siberia between the years 1692 and 1695, Ides learned that many of the Yakuts, Tunguses, and Ostyaks stead- fastly believed that these huge monsters spent their lives deep underground, moving about easily in spacious tunnels even though the earth was thoroughly frozen. Should they become particularly active, the whole ground might rise above them, caving in later as they passed on; but should they come to the surface and breathe the warm air, they instantly died. It seems as if the very gradual recog- nition of the woolly mammoth as actually quarters. oir). JEFFERSONIAN MAMMOTH OF INDIANA. ABOUT ONE FORTY-FIFTH NATURAL SIZE Fig. 1029. The Jeffersonian mammoth was probably hairy, with a fine undercoating of wool in the winter season. Its tusks are incurved exactly as in the remotely related woolly and imperial species. Its concave forehead is quite distinct from that of the modern elephant. As found in Indiana, it measured 104 feet at the shoulder. See also Witsen, Nicolaes, 1692.—Editor.] ImpertAL Mammotu or NEBRASKA. 1165 an extinct elephant extended well over a whole century; in 1696 the Russian explorer Ludolf described the mammoth of Siberia under the name Mammotovoi kost; two years later the German scholar Tentzelius defended against all sceptics the ABOUT ONE FIFTY-FIFTH NATURAL SIZE Fig. 1030. This superb animal, first named Hlephas imperator by Leidy, stands fully 13 feet at the shoulder—4 feet taller than the woolly mammoth but with similar curved tusks and sloping hind The best skeleton is in the Nebraska State Museum (Archidiskodon imperator maibeni) ; the finest cranium is in the Geological Institute of the City of Mexico (Fig. 902 of present Mem- The animal ranged from Nebraska to Mexico. little hair and no wool. Like the Indian elephant, it probably had discovery of a really fossil elephant at Burg-Tonna near Gotha, but this classic skeleton has recently proved to belong not to the mammoth family but to the straight-tusked elephant family known as the ‘elephant of antiquity’; in 1728 Sir Hans Sloane confirmed the finding of Tentzelius by reference to the discovery; seven years later (1735) John Phil. Breyne wrote to the Royal Society as follows: “... I was busied . . . to prove, that the extraordinary large Teeth and Bones found under Ground, and digged up in several Places of Szberia, by the name of Mammoth’s or Mammut’s, Teeth and Bones, were, I. True Bones and Teeth of some large Animals once living; and, II. That those Animals were Elephants, by the Analo- gy of the Teeth and Bones, with the known ones of Elephants. III. That they were brought and left there by the universal Deluge. I made likewise several useful Inferences about this matter.” The first finely etched figure of the cranium of the mam- moth brought by Messerschmidt from Siberia was published 1166 OSBORN: by Doctor Breyne in 1737 [1741?]. Thus step by step the way was prepared for Blumenbach to name the animal in 1799 and for the great Cuvier to thoroughly describe it in a series of memoirs THE PROBOSCIDEA County near Rochester, New York. Finally, in 1924, the present author described his Mammonteus primigenius compressus, the most highly specialized and perfected mammoth thus far found, as ARCHED TUSKS OF THE AFRICAN ELEPHANT Fig. 1031. Giant arched tusks of the African elephant, 11 feet in length, presented by Charles D. Barney to the Heads and Horns Collection of the New York Zoological Park. between 1796 and 1834. Since that time our knowledge has advanced by leaps and bounds. The first bones of the mastodon found on the banks of the Hudson River in 1705 and of the Ohio tiver in 1739 were naturally confused in Europe with those of the mammoth when Blumenbach first labeled them Ohio incognitum and then Mammut. A century later, in 1842, DeKay described his Elephas americanus, a fossil mammoth tooth found in Monroe SKULL AND TusKS OF THE WooLLy Mammortu Fig. 1032. Mammonteus primigenius found on the Yukon River, Alaska Skull and half-grown tusks of a middle-aged male specimen of With advancing age these tusks would curve inward and cross in the middle line CircuLaRr TUSKS OF THE WOOLLY MAMMOTH OF SIBERIA 1033. Contrast these tusks, 8 feet in length, of the male woolly mam- moth of Siberia with the arched tusks of the African elephant shown in opposite figure. Vig. circle-shaped known both near Rochester, Indiana, and in Alaska (see p. 1157 above). Beginning with the discovery of a complete frozen mammoth skeleton—known as the Adams’ mammoth—near the mouth of the rm E Wootty Mammornu, SomME River, FRANCE ONE-FIFTIETH NATURAL SIZE Fig. 1034. This mammoth is a little more than 9 feet at the shoulder, whereas the Jeffersonian mammoth is 1044 feet. and the imperial mammoth 13 feet. The painting closely follows the drawings in the cavern of Combarelles (p. 1169, tailpiece). The rapidly sloping hind quarters of the animal serve as a watershed for rain and sleet. THE MAMMONTINA: MAMMONTRUS Lena River in 1799, other complete skeletons were unearthed and mounted in the museums of St. Petersburg, Leipsic, Stuttgart, and Brussels, all closely similar in size, in the wheel-like curvature of the great tusks, in the extraordinary foreshortening of the skull (p. 1158 above), in the rapid falling away or sloping of the hind quarters, in the shortness of the backbone and of the tail, in the reduction or complete loss of one of the digits of the hand—briefly, in the truly marvelous adaptation, in every part of the skeleton as well as in the teeth, to the very severe conditions of boreal life. In 1912 Felix described a mammoth found near Borna, whose death must have been so sudden that the animal did not have time to swallow a mouthful of food which lay in the form of a wad between the upper and lower teeth; in the stomach of this animal were about twenty-four pounds of undigested plant food, exceed- ingly interesting because it consisted of plants that are still native to the place (Beresowka River, northern Siberia), the tundra flora which the mammoths stored up during the short summer season for the long winter. These plants are almost exclusively grasses and form the characteristic meadow flora; the needles of conifers occur very rarely. From other discoveries we know that the mammoth fed during the winter on the arctie willow, Salix polaris of Wahlenberg, and on other northerly dwarf plants. The description by Felix not only gives us complete knowledge of the skeleton of Hlephas primigenius but shows conclusively the proper position of the tusk in the jaw and also itsinelination. As to the animal’s outward appearance, it involved a number of cor- rections in even the best reconstructions that had been made; the proportion of the length of skull and trunk in the mammoth is quite different from that of existing elephants: in the mammoth the skull is more than half the trunk length, in the elephant (2. indicus) it is always less than half. The mammoth’s head, there- fore, was higher in proportion to the body than that of recent elephants and in consequence the tusks could attain enormous proportions. The largest of the tusks in the Leningrad Museum measures no less than 13 ft. 8 in. and in the Franzens-Museum of Briinn there is a tusk that actually exceeds 16 ft. 5 in. in length, including the long basal portion of the tusk which was ineased within the exceptionally long tube-like premaxillary bones shown in our figure (p. 1158) of the female mammoth skull of Indiana, the type of Osborn’s M. primigenius compressus. The trunk of the mammoth was extremely well developed. The ear was somewhat smaller than that of the Indian elephant, being about 15 in. in length and 6.7 in. in breadth, and, like the rest of the body, was covered with a thick coat consisting of short wool and longer hair. The tail is conical in form, about 14 in. broad at the root, sharply pointed at the end where it terminates in a bunch of bristles. The skin was extraordinarily thick and underneath it was a layer of fat up to 344in. deep. The whole body was thickly covered with fine soft hair about an inch long, varying in color from faded blond to yellow brown; coarser and longer hair, sometimes 20 in. in length, of a dark, rust-colored brown, covered the entire neck and trunk, perhaps forming a fringe of hair still heavier and thicker from the cheeks along the shoulders and sides to the rump. The above details regarding the external appearance of the 1“ George Grant MacCurdy: Human Origins. Vol. I, 1924, p. 265.” 1167 mammoth are given very fully because they enhance the reputation of the Old Stone Age artists not only as close observers but as portrayers with marvelous fidelity of the external appearance of the woolly mammoth. Among the very numerous etchings, drawings, and paintings there is one which possesses high artistic merit as truthfully depicting the characteristics of the charging mammoth (p. 1132 above). The majority of these etchings and sculptures, as fully enumerated! in MacCurdy’s encyclopedic volumes, are of the modern comic supplement order; they give the impression that our Stone Age ancestors were struck only with the humorous side of the mammoth as he appeared in the full panoply of his winter coat. But our increasing knowledge of these animals, derived from study of the frozen skeletons of Siberia and from the extremely close studies made by the present author in the pre- paration of this memoir on the Proboscidea, assures us that even the crude outlines on the walls of the cavern of Combarelles (p. 1169 tailpiece) are not of the comic supplement order but are very truth- ful portrayals of the woolly mammoth as he actually appeared dur- ing the winter season, rounded out with his huge thickness of fat (three and a half inches in places) over the entire body and with a woolly covering extending down to his very hoofs and masking the muscular outlines of the limbs beneath. His extremely elevated head was followed by a deep nick or indenture of the neck, then by the rising hump of the back which sloped rapidly downward into the depressed region of the pelvis and terminated suddenly in a short, blunt tail. Thus the apparent caricatures of the Stone Age artists are realities; the mammoth in his winter pelage was even more dis- guised than the yak of Tibet. The sloping hind quarters served admirably as a watershed for the torrents of sleet and rain and the whirlwinds of snow that raged during the northern blizzards. Not only this, but the apparently weak and sloping hind quarters were all that the mammoth needed for the forward propulsion of his body, since, unlike all modern elephants, he never used his tusks for digging or uprooting purposes; consequently the hind limbs were not propellers of the body as they are in the African and Indian elephants of today. Another important feature is the bulbous or well-rounded forehead, which rises like a sloping dome at the top of the otherwise pointed head; this swollen forehead, we are sure, was a food reservoir for the winter season which dis- appeared as winter advanced into spring, when the deep fatty covering all over the body was exhausted and the animal began to assume the normal outlines and proportions of other elephants. This interpretation is to our mind absolutely demonstrated by the contrast between the comic outlines of the Combarelles mam- moths of midwinter and the wonderfully spirited charging mam- moth of midsummer engraved on a section of mammoth ivory tusk discovered beneath the Magdalenian shelter of La Madeleine along the Vézére River (p. 1132 above). Here we see that the round fatty outlines of winter give place to those which conform very closely to the lean profile of the summer season; the high, peaked skull especially, with its characteristic concave forehead, affords us a view of the actual summer profile of the mammoth which corre- sponds with that of the Indiana mammoth shown on page 1158. The classic Magdalenian engraving is one of the most realistic 1168 pieces of Paleolithic art that has ever been found; there are indications that the artist used the relatively small piece of ivory for the representation of three mammoths, for the tusks and trunks of two other elephants appear in the distance. Observe especially the outline of the ear, the elevation of the highly peaked, acrocephalic head, and the remarkably lifelike action of the limbs and body. Toward the close of the Old Stone Age there began the wide-spread custom of the ceremonial burial of the dead to which we owe our really remarkable knowledge of the great hunting races that swarmed over cen- tral and western Europe during closing glacial time, succeeding the last of the Neanderthal : VESTONICE, MORAVIA race which seems to have had full possession ps 1036. View of the river shores and caverns at the height eS Mammoth skeletons are strewn over the floor. (Photo- graph by Doctor Ab- solon.) of the fourth or last descent of the Scandinavian Véstonice. glaciers. Largely known by their flint imple- ments found in the burials of hunters and warriors of Neanderthal time, we know little of their prowess; the Neanderthal flint imple- ments are relatively small and clumsy and it would appear that the Neanderthal hunters were not equipped to pursue the large and formidable woolly mammoths of their time. Succeeding the Neanderthals, however, was another race, probably from the Far East of central Asia, first found buried in STRATA OF THE PEKARNA Cave, Moravia Fig. 1035. New and Old Stone Age strata, from historic time downward to Lower Aurignacian altogether ten distinct layers, representing a period which Doctor Absolon believes extended over 100,000 years. The mammoth layer 1s contemporaneous with the Lower Aurignacian. by Doctor Absolon.) (After photograph THe Mammoru Pir or of the mammoth pit of OSBORN: THE PROBOSCIDEA the cavern of Aurignac and hence known as man of the Awugnacian stone culture. This race is known as the Briinn or Piedmost; they are long-headed, with a narrow, short face and rather promi- nent brow ridges, and with brain development inferior to the Giant KILLiIna STONE or THE MoORAVIAN HUNTERS Fig. 1037. Doctor Absolon interprets the large stones found in the cave as the weapons with which the mam- moths were killed after being trapped. Photo- graph by Doctor Ab- solon. See ee broad-faced, artistic Cro-Magnons who drew and painted the mammoth. The Briinn people were apparently more interested in eating the mammoth than in depicting it, but they have left some sculptures in bone and ivory of both animal and human figures. This was one phase of the Briinn Aurignacian culture, but their culture is more largely known by their stone implements adapted to the making of clothing and to the killing of smaller kinds of animals—none of the Aurignacian implements so far found in western Europe was adapted to the chase of larger Evidence of the killing and consumption for food of a great number of the wild horses of the period has recently been found near Solutré in southern France, but, so far as I know, without the accompaniment of very large flint implements of the chase. The discovery of giant killing flints by Dr. Karel Absolon of game. Ivory Fiaurtnr From BrassEMPOUY ig. 1038. Back and profile views of a woman’s head carved on ivory. (After J. Pilloy in Edouard Piette’s L’ Art pendant L’ Age du Renne, 1907.) THE MAMMONTINA: MAMMONTEUS the University of Prague and the Museum of Briinn, one of the most distinguished archeologists of central Europe, is therefore a revelation not only of the pursuit of the woolly mammoth for purposes of food, but of the killing methods employed, whereby the EQuiINnE Ivory STATUETTE FROM LOURDES Tig. 1039. This statuette, found in the Grotte des Espélugues at Lourdes, is carved from mammoth ivory. Twice natural size. du Renne, 1907). animals were driven into great pits and then felled by giant stones let down by their captors. These stones appear (Fig. 1037) like greatly magnified coups de poings of the long bygone Chellean and Acheulean age. Citations from the description of his remarkable discoveries in the years 1924-1929 of mammoth-hunting stations of Moravia are given on page 1139 above. The woolly mammoth appears to have ranged almost ex- clusively north of the 40th parallel; it loved the borders of the retreating glaciers of the close of the Glacial Age both in Europe and North America. In Alaska it was extremely abundant and was occasionally found in frozen form along the ancient shores of Eschscholtz Bay. Alaska, broadly connected with the Asiatic mainland by a great isthmus bounded on the south by the present mountain peaks of the Aleutian Islands, yields abundant remains not only of the true woolly mammoth closely similar to the ZL. primigenius of Blumenbach, but also an extremely rare and highly specialized grinder (Fig. 1022) to which the specific terminal com- pressus was recently applied by Osborn in descriptive relation to the exclusively compressed and fine-plated grinding teeth which attain the very high number of 27 compressed ridge-plates above and below, in comparison with the 24 ridge-plates above and below in FE. primigenius of Siberia and western Europe. This proves that the marvelous adaptability of the mammoth did not cease when he Kriighel Na 1U- (After J. Pilloy in Edouard Piette’s L’ Art pendant L’ Age 1169 entered the continent of America but reached even a higher point of specialization in his essential grinding tooth mechanism, just as we Americans pride ourselves today on our mechanical achieve- ments. This grinding tooth is a perfect marvel of adaptation and it is lodged below the very highly peaked cranium with a correspondingly deep depression and foreshortening of the jaw and approximation of the front and back planes of the skull which are so extreme as to appear almost artificial. It is this high, narrow, and deep skull to which the Greek ad- jectives hypsicephaly, acrocephaly and bathy- cephaly especially apply as absolutely unique in the animal kingdom [See Fig. 1023 above]; in adaptation to grassy diet it is the extreme antithesis to the long and relatively flatheaded cranium of the American mastodon which was the contemporaneous forest dweller of Alaska and as far south as Florida. The woolly mammoth, however, like the polar bear, aretic fox, ptarmigan, and arctic reindeer, was at its best when in the severe climate of the far North, defying with its woolly covering the coldest arctic blasts. During the summer season it lost not a moment’s time in laying in its grassy hoard for the coming winter, in comparison with its less hardy relative, the Jeffersonian mammoth of the mid-tem- perate region, and its subtropical and more remote relative, the imperial mammoth of the south temperate zone. Like many other fossil mammals the mammoths appear to have become rather suddenly extinct after the climax of the Old Stone Age, namely, during the slow northward recession of the final great glaciers of Scandinavia and North America. The cause of their disappearance at the very moment when they reached the highest degree of specialization and perfection of their grinding teeth (Fig. 1022) is a mystery; it is attributed by Tolmachoff to further specialization and by Howorth to the universal flooding which accompanied the sullen northward retreat of the great glaciers. A more likely explanation is that during this unfavorable period the herds may have become numerically reduced by under- feeding through lack of grassy food during the summer season. During these decades the underfed mothers were probably unable to protect their young from the attacks of wolves and other carnivorous mammals. Fortunately for us, during the height of their supremacy in western Europe they had been superbly drawn, modeled, and painted by the artists of the closing period of the Old Stone Age. In fact, it was not very long after the recognition of the woolly mammoth as a true fossil that man was also discover- ed in the fossil state. eT ea age “es oe. & « ath , s 4 ¥ * FT ty ae td Pig. 1040. A young adult bull elephant (Loxrodonta africana) in the bush of the Lake Paradise region, east Central Africa, as photographed by Mr. and Mrs. Martin Johnson in 1923-1924, and shown in the film “Simba.” Reproduced through the courtesy of Mr. Daniel E. Pomeroy. 1170 CHAPTER XIX THE GENERA LOXODONTA, PALAZSOLOXODON, AND HESPEROLOXODON OF THE SUPERFAMILY ELEPHANTOIDEA, SUBFAMILY LOXODONTINA LOXODONTA, PALHOLOXODON, AND HESPEROLOXODON, UNITED IN THE SUBFAMILY LOXODONTIN® BY SIMILAR CRANIAL AND DENTAL CHARACTERS. SIMILAR ESPECIALLY IN THE BROAD PREMAXILLARY ROSTRUM AND THE WIDELY SPREADING SUPERIOR INCISIVE TUSKS WITH UPROOTING FUNCTIONS TO THE END OF LIFE. RANEAN, AND EURASIATIC DISTRIBUTION. I. CLASSIFICATION AND History or Discovery OF THE LOXODONTIN®. 1. Difficulties of generic nomenclature. 2. History of discovery and separation of European, Indian, Mediterranean, African, Japanese, and Javanese species. 3. Order of discovery and description of the fifty-three type species of the extinct Loxodontine. II. Systematic REVISION OF THE LOXODONTIN®A. Subfamily characters. Loxodonta: Generic characters. 1. Order of description of eighteen living African species and subspecies. 2. Systematic description of species of Loxodonta: Loxodonta africana, L. cornaliae. Il]. Eurastatic Species OF PALAOLOXODON AND HESPEROLOXO- DON. Palxoloxodon: Generic characters. Palzxoloxodon namadicus. Hesperoloxodon: Generic characters. Hesperoloxodon antiquus. Upnor elephant (H. antiquus) Hesperoloxodon antiquus nanus. Hesperoloxodon antiquus platyrhynchus. Hesperoloxodon antiquus ausonius. Hesperoloxodon antiquus germanicus of Rumania. Hesperoloxodon antiquus italicus. Hesperoloxodon antiquus germanicus of Steinheim. IV. Exrincr Dwarrep SPECIES OF MerpDITERRANEAN ISLANDS. Palzxoloxodon melitensis. Palzxoloxodon falconeri. Palzxoloxodon mnaidriensis. Palzxoloxodon lamarmorae. Palzxoloxodon cypriotes. Palxoloxodon creticus. THE ALSO RELATIVELY SIMPLE GRINDING TEETH AND MORE OR LESS PRONOUNCED MESIAL ‘LOXODONT SINUS’ ADAPTED TO BROWSING RATHER THAN TO GRAZING. SURVIVING IN THE EXISTING LOXODONTA AFRICANA. OF EXCLUSIVE AFRICAN, MEDITER- V. Les EvéeHants Natns pus Ines MépITeERRANBENNES ET LA QUESTION DES ISTHMES PLEISTOCENES (VAU- FREY, 1929). VI. ANCESTRAL STAGES OF PALASOLOXODON IN AFRICA. Palxoloxodon atlanticus. Palxoloxodon jolensis. Palexoloxodon recki. VII. PALaoLoxopON AND LOXODONTA OF SouTH AFRICA. ?Palzoloxodon andrews. Palzxoloxodon hanekomi. Palxoloxodon yorki. Palexoloxodon wilmani. Palzxoloxodon kuhni. Palxoloxodon archidiskodontoides. Palzxoloxodon transvaalensis. Palxoloxodon sheppardi. Loxodonta zulu. Loxodonta prima. Loxodonta africana var. obliqua. Loxodonta subantiqua. VIII. LoxoponTINEs or JAPAN AND JAVA. Palxoloxodon namadicus naumanni. Palxoloxodon namadicus namadi. Palzoloxodon protomammonteus. Palzxoloxodon tokunagai. Palzxoloxodon protomammonteus proximus. Palxoloxodon namadicus yaber. Palxoloxodon tokunagat junior mut. Parelephas protomammonteus matsumotor. Palxoloxodon yokohamanus. Palxoloxodon hysudrindicus, Java. IX. ALONG SUMMARY OF (GEOGRAPHIC DISTRIBUTION THE EASTERN Coast oF AsIA. The cranial, dental, and incisive tusk resemblances of members of the Loxodontinz to each other and the profound distinctions in cranial profiles, sections, and proportions from the Mammontine (Archidiskodon, Parelephas, Mammonteus) types are fully set forth in Chapter XV and synoptically in figure 1041, which illustrates the profile and frontal aspects of the chief specimens known of Palzoloxodon namadicus and Hesperoloxodon antiquus in comparison with Loxodonta africana. The typical species of Palzoloxodon exhibits the broadly diver- 1171 SKULLS OF LOXODONTS FROM ASIA, EUROPE, AND AFRICA All figures one-twentieth natural size L. AFRICANUS Ref. L. AFRICANUS Ref. . AFRICANUS Ret. td Falc., 1847, Pl. XLIV, Fig. XVII (rew.) Faic., 1847, Pl. XIN A, Fig. 8 Faic., 1847, Pi. XLII, Fig. XVII ep \ is ih pS | p | Cina} \ ‘ — Oe L. NAMADICUS Ref. Faic., 1847, Pl. XXIV A, Fig. 4# L. NAMADICUS Type Falc., 1847, Pl. XII B, Fig. 1 (rev.) L. NAMADICUS Type Faic., 1847, Pl. XA L. NAMADICUS Ret. be 7, Pr Vv Fig. 4 = oe ae Sal L. NAMADICUS Ret. L. NAMADICUS Ref. Fajc., 1847, Pl. XLII, Fig. XXII Faic. 1847, Pl. XLIV, Fig. XXII (rev.? L. (ANTIOUUS) MELITAE Ret. Pohtig, 1893, Tef. |, Fig. 1 L. (ANTIQUUS) MELITAE Ref. + Pohlig, 1893, Taf. |, Fig. 1* he ANTIQUUS (NAMADICUS) Ref, Y L. ANTIQUUS (NAMADICUS) Ret Piigrim, 1905, Vol. XXXII, Pl 11 Pilgrim, 1905, Vol. XXXII, Pt. 12 L. ANTIOUUS Pontig, 1891, p. 350, Fig. 10° L. ANTIOUUS Ret. L. ANTIQUUS Weitnofer, 1890, Tat. tl, Fig. 2 Ponlig, 1891, 9. 350, Fig. 109 Fig. 1041. Cranta or LoxopONTA AFRICANA, PALHOLOXODON NAMADICUS, AND HESPEROLOXODON ANTIQUUS All to the same scale, one-twentieth natural size. Assembled in the year 1922. Compare figure 1069 of the year 1929 Upper Row. Lozodonta africana, after Falconer and Cautley, 1847, Pls. XLU, XLIV, XMI.A. Second Row. Palxolorodon namadicus Falconer and Cautley, type, 1847, Pls. x11.a, x18, front and side views. Two referred skulls of P. namadicus in front and side views, after Falconer and Cautley, Pls. xxIv.A, XLII, XLIV. Third Row. Referred skulls of Palxolorodon melitensis, after Pohlig, 1893, front and side views, island of Sicily, and of P. namadicus, after Pilgrim, 1905, front and side views. Fourth Row. (Left) Referred rostrum of Hesperoloxodon antiquus ausonius?, after Weithofer, 1890. (Middle and right) Rostrum of H. antiquus ausonius erroneously referred by Pohlig (1891, p. 350, fig. 109) to Elephas (antiquus) Nestit. Recorded as of Upper Pliocene age. 1172 THE LOXODONTIN#: CLASSIFICATION AND HISTORY OF DISCOVERY 1173 gent premaxillary rostrum similar to that of Loxodonta africana but differs very widely in the broadly rugose parietofrontal crest which extends like a Phrygian cap down over the frontals almost to the nasals; this is seen even in the dwarfed species L. (antiquus) melite! {= Palxoloxodon melitensis]. It is obvious that in Palzxoloxodon the cranium is somewhat more hypsicephalic and bathycephalic than in Loxodonta, but this cannot be shown in longitudinal section at present. The function of the gigantic rugose Phrygian cap is doubtless for the support and retraction of enormous divergent tusks and a greatly broadened and enlarged proboscis. A Fig. 1. Fig.uF Léléphant & crane allongé, @ front concave, & trés~ Tongues alvéoles des défenses, & mdchoire infericure ob- tuse, a mdchelieres plus larges, paralleles, marquées de rubans plus serrés , que nous nommons €léphant f ssile ( ele- phas primigenius , Blumenb. ) , est le mammont des Russes. B Fig. 2. Fig. 9.1. L’éléphant a crane allongé , a front concave , @ petites oreilles , & mdchelieres marquées de rubans ondoyans que nous appelons éléphant des Indes ( elephas indicus), est un quadrupéde qu’on n’a obseryé dune maniere certaine qu’au- dela de Indus. C Fig. 35. Fig. 10.4. L’éléphant & créne arrondi, a larges oreilles , @ mdche- licres marquées de losanges sur leur couronne, que nous appelons éléphant d Afrique ( elephas africanus), est un quadrupéde dont la seule patrie connue est jusqu’a présent T Afrique. Fig. 1042. Cuvier’s figures and definitions of Elephas primigenius (Messerschmidt’s cranium, ef. Fig. 991), EB. indicus, and E. africanus. After Cuvier 1806.1, Pl. 39 (1), figs. 1, 2, 3, and Pl. 41 (rv), figs. 9.I, 10.A, and 11.F. One thirty-seventh to one-twentieth natural size. See also figure 992 above. I. CLASSIFICATION AND HISTORY OF DISCOVERY OF THE LOXODONTIN/E (Continued from Chapter II, p. 32, of Vol. I, and Chapter XV of the present Volume) As repeatedly observed in the present Memoir (e.g., Chap. I, p. 5, Chap. II, pp. 17, 18), it is impractica- ble in palzontology to apply all the principles of nomenclature established in zoology and botany, because the classification of the imperfectly known fossil forms is ever changing with our increasing knowledge of origins, adaptive radiations, and phyletic successions. Such mutability does not disturb the nomenclature of living animals and plants, in which priority of adequate description, figure, and definition is the chief concern of systematists. Zoological principles of classification, as in the classic works of Cuvier and of Falconer, must therefore sur- render to phylogenetic principles of classification (cf. Chap. I, pp. 5-13) and names and classifications of the dawn period of paleontology must be reworded and rearranged on modern principles and knowledge (cf. Chap. XV, pp. 911, 912). Even certain technical rules of nomenclature, of orthography, and of priority, as revised by zoologists,” must give way to paleontologic standards; otherwise we find ourselves repeatedly compelled to The lettering in figure 1041 was executed at a time when all these species were united in the single genus Lozodonta, that is, before Professor Osborn had adopted Palzolorodon Matsumoto (1924) for members of the namadicus group and had created the new genus Hesperoloxodon for certain other species.—Editor. ] *The International Commission on Zoological Nomenclature, Congress held at Monaco in 1910. 1174 OSBORN: THE PROBOSCIDEA sacrifice the spirit to the letter and to bury as nomina nuda the classic terms of vertebrate paleontology proposed by such great founders as Blumenbach,' Cuvier, and Falconer. In no subfamily is this historic recognition and common-sense spirit more essential than in the Loxodontinz, as shown in the following revision. 1. DIFFICULTIES OF GENERIC NOMENCLATURE The generic nomenclature of the Loxodontine is full of confusion, which can only be cleared up by a common- sense interpretation and reéxamination of the original definitions and types. The history of nomenclature is as follows: LoxoponteE, F. Cuvier.—Frédérie Cuvier (1825) was the first to apply the appropriate term “Loxodonte” (signifying slanting toothed) to distinguish the African from the Indian elephants, but despite the profound differ- ences in dental and cranial characters and in geographic distribution, the generic term Elephas continued to be widely used as late as 1886, e.g., Lydekker, ‘Elephas africanus,’ and even in 1928, Andrews, Forster Cooper, and Bather adhered to the generic term Elephas in describing the Upnor elephant as ‘Elephas antiquus.’ In the case of the term ‘“‘Mastodonte”’ of G. Cuvier, 1806, it was not until 1817 that G. Cuvier adopted the Linnean classic system of generic orthography: (1) Accordingly F. Cuvier’s original term of 1825 was “Loxo- donte,’’ as G. Cuvier’s original term of 1806 was ‘‘Mastodonte.’” (2) In an unsigned review of F. Cuvier’s description in the Zoological Journal of London (1827, 1828, Vol. III, p. 140), the generic term Loxodonta was sub- stituted for his ““Loxodonte” of 1825; therefore the generic term Loxodonta should be credited to F. Cuvier as the generic term Mastodon should be credited to G. Cuvier. (3) In 1848, p. 184, Gray, in his ‘‘List of the Specimens of Mammalia in the Collection of the British Museum,” employed the Latin spelling, as appears from the following citation: The African Elephant. Loxodonta Africana. Klephas Africanus, Blumenb. Abbild. t. 19. f.c¢. TE. maximus, part, Linn. Mam. Lithog.t. Harris,W.A. Afric. t. 22. Cuvier, Oss. Fos. I. t. 2 to 8. “ee Again in 1869, p. 359, in his “Catalogue of Carnivorous, Pachydermatous, and Edentate Mammalia in the British Museum,’’ Gray employs the same spelling: 2. Loxodonta. Lamina of the teeth with lozenge-shaped crown. Skull subglobular, forehead shelving, crown rounded; front of lower jaw acute, produced. Trunk conical, thick at the base. Ears very large. Loxodonta, F. Cuvier, Dents Mamm. [1825,1827.] Loxodonta africana. (African Elephant.) B.M. Loxodonta africana, Gray, List Mamm. B.M. 1843; Gerrard, Cat. Bones B.M. Klephas africanus, Blumenb. Abbild. t. 19. f.c.; Kirk, P.Z.S. 1864, p. 654; Giebel, Sdugeth. p. 159; Blainv. Ostéogr. Gravigrades, t. 3 (skull), t. 7 & 9 (teeth). Elephas maximus (part.), Linn. (4) Falconer and Cautley (‘Fauna Antiqua Sivalensis,’’ 1847, Pl. xinm) and Falconer (Quart. Journ. Geol. Soc. London, 1857, pp. 315, 318) employed the shorter masculine term Loxodon [preoccupied], which Falconer (1857, p. 315) defined as follows: For this subgeneric group, the name of Loxodon (Footnote: ‘From Aogés obliquus, and ddovs dens, having reference to the rhomb-shaped dises of the worn molars; an adaptation of the term ‘Loxodonta’ proposed by Fred. Cuvier, Hist. Naturelle des Mammiferes, tom iii., “Article Eléphant d’Afrique.” 1835 [1825].’], first indicated by Frederick Cuvier, has been adopted. It comprises both extinct and living species. ‘Blumenbach’s original description (1799, p. 697) of Elephas primigenius cites as an example the Burgtonna (Gotha) skeleton, which certainly belongs to ‘Blephas antiquus,’ as fully explained in Chapter XVIII above; see also the present chapter (p. 1181 below). *Compare Chap. V, p. 119, Chap. VI, pp. 135, 165, and Chap. XXI, pp. 1363 and 1372, for the history of the term Mastodonte. THE LOXODONTIN#: CLASSIFICATION AND HISTORY OF DISCOVERY 1175 This most appropriate term, however, is preoccupied by Loxodon Miiller and Henle, 1841, for a genus of sharks, e.g., Loxodon macrorhinus; consequently Osborn adheres to the orthography Lozxodonta (F. Cuvier, 1825, 1827; Gray, 1843), although it involves the feminine termination of the species referable to this great genus. (5) Lydekker (Ency. Brit. 11th Ed.) never admitted the generic distinction of the genotypic and other species of African elephants, but adhered to Elephas africanus. (6) Heller and Roosevelt (1914, p. 739) erroneously em- ployed the trinomial Loxodonta africana africana, as listed under the existing African species below. ELASMODON Fatc. (PREOCCUPIED) AND EUELEPHAS Fautc. (NoMEN NUDUM).—(1) In 1846, Falconer and Cautley first named (p. 45), and figured (1847, Pls. x11.4, x11.B) a new Upper Siwalik species as ‘Elephas namadicus,’ the specific name referring to the Nerbudda [or Narbada] River, the Namadus River of Ptolemy. (2) A year later (Fauna Antiqua Sivalensis, 1847, caption to Pl. xin, figs. xrx—xxiv), they proposed the new subgeneric term Elasmodon [in allusion to the waving enamel ridges], to include the six species HE. antiquus, HE. hysudricus, E. meridionalis [not figured], H. namadicus, E. indicus (Dauntela var., Mukna var., and young), and EL. primigenius; these six heterogeneous species are the ‘genotypes’ of Elasmodon and of its substitute Huelephas. (3) In this same caption they erroneously applied the generic name Loxodon to E. planifrons, grouping it with EF. africanus. (4) In 1857, learning that the name Hlasmodus had been used for a series of fossil fish, Faleoner (p. 315) substituted the generic term Huelephas for Elasmodon: “For this subgeneric group we propose the term of Huelephas |Foot- note, ‘From ed bene, and édégas, having reference to the typical Elephants most familiarly known.’].”” (5) W. L. Selater (1900, p. 317) was in error in making Elephas planifrons the genotypic species of Huelephas, because this species was not included in the original definition of Elasmodon referred to above. (6) Palmer (1904, p. 275) Tutrp (INFERIOR) AND SECOND SUPERIOR MOLARS OF THE AFRICAN (LOXODONTA) AND Asiatic (ELEPHAS) ELEPHANTS. Fig. 88. Fig. 89. African Elephant. 4 nat. size. Asiatic Elephant. Fig. 1044. Crown view of: (A) second right superior molar of Loxodonta Fig. 1043. Crown view of the third inferior molar of the right side of: Fig. africana, with eight lozenge-shaped ridge-plates; (B) second right (?) superior 88, Lozodonta africana; Fig. 89, Elephas indicus, one-third natural size. molar of Hlephas indicus, exhibiting eleven worn ridge plates. After Lydekker, Compare Owen, “History of British Fossil Mammals, and Birds,” 1846, pp. ‘“‘Catalogue of the Ungulate Mammals in the British Museum (Natural 230-232, figs. 88, 89. History),’’ Vol. V, 1916, p. 80, fig. 22. Loxodonta africana (Jumbo) Amer. Mus. 3283 | 1/8 nat, size Cranta or Arrican (Lerr) AnD or Nerpuppa, [Np1A (Ricur), ELppHANTS Both one-eighth natural size LoxoponTA (MALE) PALAOLOXODON (I°EMALE) Vig. 1045. Cranium and jaws of the typical African elephant of the Fig. 1046. Cranium of the type of Palwolorodon namadicus. In the palate Sudanese or Abyssinian subspecies Loxodonta africana oxyolis. In the jaw — observe the closely parallel and relatively compressed molar ridge-plates char- observe the extremely lozenge-shaped or centrally expanding molar ridge- acteristic of all species of Palxoloxodon. Compare figure 1070, also figure 1072. plates characteristic of all species of the genus Loxodonta. Compare figure 1043. 1176 THE LOXODONTINA:: CLASSIFICATION AND HISTORY OF DISCOVERY 1177 simply quoted Sclater’s erroneous statement. (7) Falconer in his Memoir of 1857 (p. 318) defined in Latin the collective genus Hlephas (Linn.), on the basis of premolar and molar tooth structure, as embracing three subgenera, namely: Subgen. 1. Srecopon.—Dentium molarium 3 utrinque intermediorum coronis complicata colliculis hypisomeris (e.g. 7+7-+8), mammillatis, tectiformibus. Pramolares nondum observati. Subgen. 2. Loxopon.—Dent. molar. 3 utrinque intermedior. coronis lamellosa colliculis hypisomeris (e.g. 7-+-7-+8), cuneiformibus. Przemolares raro utrinque 2. Subgen. 3. HurLtepHas.—Dent. molar. 3 utrinque intermedior. coronis lamellosa colliculis deinceps numero auctis, anis- omeris (e.g. 12+ 14+ 18), attenuatis, compressis. Praemolares nulli. (8) In Falconer’s classification (1847) Elephas |Archidiskodon] meridionalis was erroneously grouped with Elasmodon (= Euelephas), while the closely related E. |Arch.| planifrons was grouped with Loxodon; this obvious discrepancy was corrected in 1857 (Synop. Tab. opp. p. 319), but Falconer fell into a fresh error by uniting EZ. (Loxod.) planifrons and HE. (Loxod.) meridionalis in the same group with the African elephant, Hlephas (Loxod.) africanus, because of the possession of ‘digitated’ ridge-plates. (9) From the above historic analysis it is clear that ‘Hlasmodon’ Fale. is preoccupied and that the substitute name ‘Huelephas’ Fale. is an invalid and unusable nomen nudum, because it was based on an artificial and hetero- geneous assemblage of six species, which, as shown in the present Memoir, belong to five or six entirely distinet generic phyla, namely: Elephas antiquus = Hesperoloxodon! Elephas namadicus = Palxoloxodon (Syn.: Sivalikia, Pilgrimia)? Elephas hysudricus = Hypselephas Elephas indicus = Hlephas Elephas meridionalis = Archidiskodon Elephas primigenius = Mammonteus® (10) These criticisms imply no disrespect of Falconer’s genius, but rather the insuperable difficulties under which he labored in disregarding cranial characters and in defining from ridge-plate characters only, as pointed out above in Chapter XV, ‘‘Classification of the Elephantoidea.’”’ As repeatedly shown in the present Memoir, the arrangement of the ridge-plates, whether separated (‘‘colliculi grossé digitati””) or compressed (‘‘colliculi approxi- mati”) is an unreliable and misleading basis of phyletic and generic distinction, because it is subject to parallelism or convergence; Cuvier (1806-1836) used both cranial and dental characters in separating the only three species recognized by him, viz., Hlephas africanus, E. indicus, and EF. primigenius, as shown above in Cuvier’s legends beneath figure 1042. Falconer’s reliance, for identification and definition of genera, on ridge-plate formule is shown in the entirely unnatural arrangement of genera in his Note Book for August 25, 1862 (Pal. Mem., 1868, II, p. 176) as follows: [Falconer, 1862,1868] Milk True Totals] [Present Memoir] Dinotherumy fase sea. Le eG rere a naeroEs e Sareea Ah costatoanot kai peeean ae Deinotherium giganteum Trilophodon Ohioticus —............... SOE SoBe oe oe eigec a RORe Ge Pea a ones = Oagh desi ks Mastodon americanus Tetralophodon Arvernensis ......... Dl Bate Ae Sr. are aye (peta) cobboawes ¢ =P GI faire Anancus arvernensis Pentalophodon ear eats ciao a. cage Oar Bap -G sesaagaces =28(26]....... Pentalophodon sivalensis Stegodon insignis =—S—~—ST Pe tay ee icles Gand e (ae tS PlOAM oo cso suaue AQ penis. rcs Stegodon insignis-ganesa Loxodon meridionalis —«................ Bt Oates 8 cvocseya evans Sap Gale sccaccasud =AG 6 a ae Archidiskodon meridionalis Huelephas antiquus =—S%=—............ Sot O- {all Opa LO=FI2AENG? vee sans tied aoa ae en Hesperoloxodon antiquus af PLUMS ens see ee 4+8+12.......... WAEPUIGS HEE gan o ob ober =76 .......Mammonteus primigenius ee WAOKOUE ~~ oeeaoavct 4+8412.......... PE SOY es ae = 67, Loree Elephas indicus See p. 1217 below for definition of Hesperoloxodon.—Kditor.| *[See explanation on p. 1179 below.—Kditor. } {For doubtful validity of the genus Mammonteus, see footnote on p. 1117 above.—Editor.] 1178 OSBORN: THE PROBOSCIDEA SEPARATION OF AnTIQUUS (III) AND ArricaNna (V) Lines.—Depéret and Mayet (1923, table, pp. 204 and 205) in their phyletic arrangement of the Proboscidea distinguished five absolutely separate phyla or lines of elephants, all of which, however, they included within the genus Elephas. I. Primigenius line: astensis, primigenius, trogontherti, subiricus. = Il. Meridionalis line: planifrons, hysudricus, meridionalis mut. cromerensis. GENUS ser aiterell arene sire sain anplitoners. GrCHOUS 7 - are III. Antiquus line: ausonius, antiquus: melitensis, creticus, cypriotes, atlanticus. HERES IV. Indicus line: indicus, namadicus. : V. Africanus line: africanus [= Loxodonta F. Cuvier]. In the following year Matsumoto (1924.2, Sept. 20) raised Depéret’s ‘III. Antiquus line’ into a subgenus of the genus Loxodonta, namely, Palzxoloxodon, cited below. Fatconer, 1857, OsBorn, 1918.—Falconer himself was the first to recognize the mutual resemblances of his British Pleistocene species Elephas antiquus to his Siwalik species HZ. namadicus (1857, Synop. Tab. opp. p. 319), because they both exhibited ‘‘colliculi approximati”; he did not perceive, however, their phyletic resemblance in cranial characters to the African elephant (Loxodonta africana), but only the difference in the ridge-plates (‘“‘col- liculi dilatati’’?) of the latter. It remained for several subsequent writers (Pohlig, Pilgrim) to perceive this very close resemblance, and Osborn (1918.468, table opp. p. 134) finally placed Hlephas africanus, E. antiquus, and E. namadicus in the generic phylum Loxodonta, subfamily Loxodontine. Osborn six years later (1924.633, p. 2, Dec. 20) selected Hlephas namadicus Fale. and Caut. as the geno- type of his new genus Sivalikia, to embrace also EH. antiquus, both distinguished from Loxodonta by more numer- ous and more compressed ridge-plates (i.e., ‘‘colliculi approximati’’). Sivalikia, new genus. Typified by Loxodonta namadica Falconer, type species, and distinguished by broad grinding teeth, numerous ridge-plates, and absence of ‘loxodont sinus.’ Unfortunately the name Falconeria is preoccupied for a genus of reptiles, consequently the name Szvalikia is proposed in honor of Dr. Hugh Falconer’s great work on the Siwalik fauna. In the same paper (1924) he proposed the genus Pilgrimia, typified by the narrow-toothed Hlephas falconeri of the Mediterranean Islands (Osborn, 1924.633, p. 2) as follows: Pilgrimia, new genus. Typified by Hlephas falconeri Busk, type species, H. melitensis Falconer, HL. mnaidre Adams, and L. antiquus Recki Dietrich; distinguished by narrow grinding teeth, numerous ridge-plates, ‘loxodont sinus’ vestigial or absent. The name Pilgrimia is given in honor of Dr. Guy E. Pilgrim of the Geological Survey of India, to whom paleontology is in- debted for the complete solution of the stratigraphy of the Siwaliks and of other mammaliferous horizons of India and Burma. PaLaoLoxopon Matsumoto, 1924.1—Independently of Osborn’s proposal of the genus Sivalikia, and at a somewhat earlier date (Sept. 20) in the same year (1924), Matsumoto (pp. 256, 260) selected a subspecies of Elephas namadicus, namely, EL. namadicus nawmanni Makiyama (1924), as the type of a new subgenus Palzxoloxo- don, a name which now proves applicable to the narrow- to broad-toothed? loxodonts of Japan and the East Indies as well as to similar loxodonts of Eurasia and Africa. An abstract of Matsumoto’s paper is as follows (pp. 257, 260): Palxoloxodon corresponds to Depéret’s group of #. antiquus, with molars consisting of a small number of ridge-plates, others with numerous ridge-plates. When worn the molars show clearly the rhombic form [loxodont sinus] and the enamel folds are almost parallel in the anterior and posterior portions of the unworn teeth. The molars vary in width, some being narrow, others wide. This subgenus is divided into two lines (a) E. melitensis-atlanticus and (b) E. antiquus-namadicus, according to the classification of Depéret. 2. antiquus is not extremely narrow, and the number of the ridges is rather numerous. The species of this group belong to a later period, and the ridge-plates of the molars are less rhombic in form; easily confused with ZH. trogontherti. E. ausonius equals the Calabrian in Europe, 2. antiquus the middle and lower Diluvian in Europe, 2. namadicus naumanni the lower Diluvian in Japan, #. namadicus the middle Diluvian in Japan and India. 1Unfortunately this article of 1924 did not reach the present author until much of the text of this Memoir had been prepared. The original was probably lost in transmission to America. “In all elephantine phyla the primitive grinders are relatively narrow; they become progressively broad as the cranium shortens, broadens, and deepens. Hence the narrow to broad or wide adaptation. THE LOXODONTIN: CLASSIFICATION AND HISTORY OF DISCOVERY 1179 OsBorn, 1924.—During these years Osborn was engaged in a very close comparison of the erania and denti- tion of the loxodonts of Asia, Europe, and Africa, the results of which are fully set forth in the present chapter, finally describing the entire structure of all the known Loxodontine. Unaware of Matsumoto’s publication of September 20, 1924, he independently separated the genus Sivalikia, based upon the genotypic species ‘Elephas namadicus’ Fale. and Caut., publishing the same on December 20 of the same year (1924), three months later than Matsumoto’s publication. In the present Memoir it is shown that the EZ. namadicus nawmanni of Makiyama (genotype’ of Palzoloxodon) is very close in character to the ‘Elephas namadicus’ of Falconer and Cautley (genotype of Swalikia). Although Matsumoto proposed the name Palzoloxodon as a subgenus of Lozxodonta, and Osborn proposed the name Sivalikia as a genus entirely distinct from Loxodonta, Dr. T. 8. Palmer (letter of March 12, 1929) rules that even if incorrectly conceived and defined, the subgeneric name Palxoloxodon (Sept. 20, 1924) preoccupies and anticipates the full generic name Sivalikia (Dec. 20, 1924). Osborn accordingly accepts Palmer’s ruling, although it appears to do injustice to his four years of research, in which his conclusions quite differ from those of Matsumoto. Meanwhile the generic name Sivalikia runs throughout the text and many of the illustrations; the reader, therefore, should make the following substitution: Palzoloxodon (syn. Sivalikia, Pilgrimia). I. Genus: Loxodonta F. Cuvier, 1825, 1827. renotypic species: LHlephas africanus Blumenbach. Relatively narrow grinding teeth with relatively few ridge- plates: Conservative in cranial and grinding tooth structure. Distinguished by very broad ‘joxodont sinus’; enamel non- plicate, or smooth. Descended from unknown an- cestors? to the modern Loxo- donta africana. Premaxillaries broad. Parietofrontal cranial vertex rounded, subacrocephalic. ORIGINAL DEFINITION OF GENERA (1825-1924) These four generic lines were originally separated and defined by the characters of the subgenera and species grouped within them, as follows: II. Subgenus: Palzoloxodon Matsumoto, September 20, 1924. Genotypic species: Hlephas namadicus naumanni Makiya- ma.! Ridge-plates expanded some- what mesially, ‘loxodont sinus’ rudimentary, vestigial, or ab- sent, enamel typically plicate; relatively narrow grinding teeth, with numerous ridge-plates: Max.: M 3 23. Large elephants of Japan. Premaxillaries broad. Cranium [of genotype] un- known. [Cf. P. namadicus fig- ured by Falconer and by Pil- grim (Fig. 1041).—Editor.| The genus Loxodonta is read- ily distinguished by its primi- tive cranium, primitive ridge- plate formula, and specialized lozenge-shaped or ‘loxodont sinus.’ The subgenus Palxoloxodon Matsumoto, 1924, founded on the species ‘Hlephas namadicus naumannv’’ Makiyama, 1924, is also characterized as relatively narrow toothed. [See footnote by Dr. E. H. Colbert on page 1247 below.—Kditor.] *[Compare Lozodonta prima and L. africana var. obliqua of Dart, 1929, on page 1287, below.—Editor.] 5[Sivalikia and Pilgrimia finally determined by Professor Osborn as synonyms of Palxoloxodon Matsumoto.—Editor.] III. Genus: Sivalikia? Osborn, December 20, 1924. Genotypic species: Elephas namadicus Falconer and Caut- ley. Relatively broad grinding teeth with numerous ridge- plates: roe 168-17 Max.: M:é 16-17 Progressive in cranial and grinding tooth structure. ‘Loxodont sinus’ absent or rudimentary; enamel plicate. Including animals of gigantic size. Premaxillaries very broad. Incisive tusks widely divergent. Parietofrontal cranial vertex broadly expanded, with promi- nent frontal crest. The genus Sivalikia, typified by ‘Elephas namadicus,’ is dis- tinguished by its relatively broad grinding teeth, numerous parallel ridge-plates, and the ab- sence of a ‘loxodont sinus.’ It proves to be the same as the subgenus Palzxoloxodon Mats. IV. Genus: Pilgrimia® Osborn December 20, 1924. Genotypic species: falconer’ Busk. Elephas Very long and narrow grind- ing teeth, with numerous and closely compressed ridge-plates: Max.: ?M 3 23. Dwarfed elephants of the Mediterranean Islands. ‘Loxodont sinus’ vestigial or absent. Including animals of diminu- tive size. Premaxillaries broad. Parietofrontal vertex expand- ed, with frontal crest. The genus Pilgrimia, founded on the narrow-toothed, dwarfed species ‘Hlephas falconeri,’ with less numerous ridge-plates, may prove to be ancestral to the subgenus Palgxoloxodon of Mat- sumoto, 1924.* 1180 OSBORN: THE PROBOSCIDEA DEFINITIONS AND DISTINCTIONS BY OSBORN OF THE Two GENERA There can be no question of the clear generic distinction of Palxoloxodon' (synonyms Sivalikia and Pilgrimia) from Loxodonta. Subsequent research may reveal more than one generic phylum within the genus Palzoloxodon, but at present it would appear, as explained in the following paragraphs of this chapter, that the phylum Palzolox- odon originated in Africa, migrated northward into Europe, possibly developing into the ‘Hlephas antiquus’ group, also northeastward through the Mediterranean Islands, leaving dwarfed descendants resembling either the ‘EB. antiquus’ group or the ‘HE. namadicus’ group, finally arriving in southern and southeastern Asia, developing into the ‘EHlephas namadicus’ group. Meanwhile the animals composing the great generic phyla Palxoloxodon and Hesperoloxodon, as in parallel species of Archidiskodon, increased in size and multiplied their molar ridge-plates from the minimum, M 3 {31} (Palzxoloxodon atlanticus), to the maximum, M 3 +? (P. namadicus naumannz), and from the minimum, M 3 {47-44 17 (Hesperoloxodon antiquus typicus), to M 3 \73° (H. antiquus germanicus), to M 3 +; (H. antiquus italicus). The species in these two great phyla are progressively distinguished by the successive addition of ridge-plates, by the increasing hypsodonty of the grinding teeth, by the absence (in the typical ‘H. |Hesperoloxodon| antiquus’ group) of the transverse bony ridge above the orbits, or by the presence (H. mnaidriensis, E. melitensis, EH. nama- dicus |= Palzxoloxodon}|) of this broad bony ridge above the orbits. Consequently according to this interpretation the subfamily Loxodontine contains three chief generic phyla, as follows: I. Genus: Loxodonta F. Cuvier, 1825, 1827. II. Genus (subgenus): Palzolorodon Mat- III.? Genus: Hesperolorodon Osborn, 1931. sumoto, 1924; Subgen. or syn.: Sivalikia Osborn, 1924. Palgrimia Osborn, 1924. Genotypic species: Elephas africanus Genotypic species Elephas namadicus Genotypic species Palewoloxodon an- Blumenbach. naumanni Makiyama. tiquus italicus Osborn.* Relatively conservative and primitive Relatively progressive in cranial and Cranium domelike with flattened in cranial and grinding tooth structure. grinding tooth structure. forehead, more hypsicephalie and bathycephaliec than that of Palxoloxo- don. Grinders distinguished by broadly Grinders distinguished by rudimen- Grinders hypsodont, ‘loxodont sinus’ open ‘loxodont sinus’ (lozenge shaped), tary, vestigial, or absent ‘loxodont vestigial or absent. on wear; enamel non-plicate or smooth. sinus’; enamel typically plicate. Grinders relatively narrow, low crown- Primitive grinders relatively narrow, : 3 : : AS es ‘ ; Ridge-plate formula: ed, with comparatively few ridge-plates: withincreasingly numerousridge-plates: 1: ‘ $-17 20 é é Min.: M 3 3% Max.: M 3 734 MaxesViioic aaa: Min.: M 3 33-44 Max.: M3 22. Tee a Premaxillaries broad with widely di- Premaxillaries extremely broad, in- Premaxillaries extremely broad, in- vergent incisive tusks, elongate, slightly cisive tusks widely divergent, straight cisive tusks widely divergent, slightly incurved. or slightly incurved. upeurved and incurved. Parietofrontal cranial vertex low, Parietofrontal cranial vertex some- Lacking prominent parietofrontal rounded, subacrocephalic, of Persistent what more progressive, acrocephalic, crest. so distinctive of Palzoloxodon primitive form. with smooth or with prominent frontal namadicus. ridge-crest for attachment of the muscles of the trunk. ‘(Subsequently (1931) Professor Osborn’s researches on the skull cf Palzxoloxodon antiquus italicus led him to the conclusion that this subspecies belonged in a quite distinct phylum from that containing ‘Elephas’ namadicus; consequently he provisionally assigned to it and to other members of the ‘H. antiquus’ group the new generic name Hesperoloxodon (see Osborn, 1931.846, p. 21), retaining the name Paloloxodon for members of the ‘EZ. namadicus' group.— Editor. | *{Compiled from statements in the text.—-Editor.] %{See footnote by Dr. E. H. Colbert on page 1247 below.—Editor.] THE LOXODONTINA: CLASSIFICATION AND HISTORY OF DISCOVERY 1181 2. HISTORY OF DISCOVERY AND SEPARATION OF EUROPEAN, INDIAN, MEDITERRANEAN, AFRICAN, JAPANESE, AND JAVANESE SPECIES Europe: ScELETO ELEpHANTINO TonN«#®, 1695 (TeNTzELIUs, 1698).—In 1695 occurred the discovery near Gotha, northern Germany, of the classic ‘Burg-Tonna skeleton,’ followed by the first scientific descriptions, as fully set forth in Chapter XVIII above, pp. 1118-1119; this famous skeleton, which aroused the interest of all the savants of western Europe, was referred to by Blumenbach (1799) in his type description of Elephas primi- genius (also cited in Chapter XVIII, p. 1141), but it now proves to belong to the ‘Hlephas antiquus’ phylum, judging by the few parts that remain of this originally fine specimen. As shown above in Chapter X VIII (footnote, p. 1119) of the present Memoir, parts of the Burgtonna skeleton were presented by Tentzelius to the Royal Society through Sir Hans Sloane and are now lost. Other parts are still preserved in the Gotha Museum. Dietrich writes (April 14, 1930) that there is no doubt that the classic Burgtonna skeleton belongs to Hlephas antiquus; its geological horizon is the travertine sands of the last Inter- glacial Period, that is, it belongs to the same deposits as those of Weimar-Taubach-Ehringsdorf, a more recent phase of the last Interglacial. Amthor of Gotha confirms the fact that the skeleton, incisors, tusks, ete., are present in the Gotha Museum, but illustrations are not available. Other European remains now known to belong to ‘EH. antiquus’ were for a long period confused with those of the southern mammoth, ‘Hlephas meridionalis.’ It is also a singular and generally unnoticed fact that the generically related ‘Elephas namadicus’ of India was described by Falconer and Cautley in 1846, a year before they figured (1847) the ‘Elephas antiquus’ of Europe, as shown in the following history of nomenclature. In the present Memoir are reproduced the type figures of all the species of Loxodonta, Palzoloxodon (syn.: Sivalikia, Pilgrimia), and Hesperoloxodon so far as published by the authors, also so far as the type specimens have been available. InprA: ELEPHAS NAMADICUS FaLc. AND Caut. (1846).—It was a long time before the relationship of the Indian species Hlephas namadicus to the African genus Loxodonta rather than to the Asiatic Elephas indicus was recognized. In 1846 Falconer and Cautley (1846, p. 45) observed: Another extinct Indian species #. Namadicus (to be described in the sequel), which is closely allied to the existing Indian form [i.e., Hlephas indicus], comes between it and HL. Hysudricus, together with a European fossil species [probably referring to the as yet unnamed Hlephas antiquus Falconer and Cautley, 1847], which we believe to be distinct from the Mammoth [i.e., Elephas primigenius Blum.]; and the gap between the existing African Elephant and #. planifrons is filled up by another well- marked European fossil species, 2. priscus (?), pl. 13, fig. 7, which is closely allied to the former. The type of ‘E. namadicus’ (Fig. 1070) was designated (Falconer, 1867, p. 15; 1868, Vol. I, p. 435) as from the “valley of the Nerbudda. Probably a female, from small size of tusks.’ Falconer writes (op. cit., pp. 15 and 435): ‘TI have named the species LH. Namadicus, after the Nerbudda river, the Namadus of Ptolemy.”’ FALCONER AND CAutTLey, 1847.—Falconer became impressed with the close resemblance between this Middle [to Upper] Pleistocene species of India, which he named H. Namadicus in 1846, and the British Lower Pleistocene proboscidean, which he named Elephas antiquus in 1847. GREAT BriTaIn: HLEPHAS ANTIQUUS FALCONER (1847).—Up to 1844 all the British fossil elephants had been referred to ‘Elephas primigenius.’ At that time Falconer was arranging and describing his rich collections from the sub-Himalaya and river deposits of central India, which had been aided by the contributions of Sir Proby T. Cautley, Mr. Fraser, and others. The superb plates of the “Fauna Antiqua Sivalensis,” beginning in 1845, dis- 1182 OSBORN: THE PROBOSCIDEA play Falconer’s confusion of the Elephas meridionalis of Nesti with the animal which he first erroneously named ‘Elephas priscus’ {after Goldfuss = Loxodonta africana] and finally in 1857 clearly distinguished in his mind as Elephas antiquus. The first figure of ‘Elephas antiquus,’ published in 1847, Pl. x11.p, fig. 4 (Brit. Mus. M.2006), may be taken as Osborn’s lectotype, although designated in the plate as ‘““Hlephas meridionalis,” but corrected by Falconer in a copy of the ““Fauna Antiqua Sivalensis” belonging to the British Museum. Falconer’s first! printed description of E. antiquus is that published by Murchison in 1867, p. 18, in the legend of figures 4, 4a, of Pl. x1r.p; this de- scription applies to a first true molar (M,, fide Lydekker, 1886, p. 130) with twelve to thirteen ridges. On a sub- sequent page (op. cit., 1867, p. 21) he described (PI. xtv, fig. 7), as ‘Hlephas priscus,’ an unmistakable specimen of E. antiquus from Gray’s Thurrock (Brit. Mus. 39,370), designating it as a “last lower molar, left side, containing eight ridges, heel inclusive’’ [].M., see Fig. 1076 below]. He adds: ‘‘Besides the great expansion,this tooth differs from all true FE. antiquus specimens in the lowness of the crown ridges. . . . If not a separate species, [it] is a very marked variety.” Leith Adams (1877-1881, p. 47) erroneously determined the ridge formula of the typical Elephas antiquus as: +2-3+ +5-7+ +9-10+ +9 -12+ +#12-13+ +15-20+ Dp 2 +3+-? Dp 3 +6-8+ Dp 4 +9-11+ M 1 +11-12+ M 2 +12-13+ MSc +16-19+° MEDITERRANEAN ISLANDS: Mata, SARDINIA, AND Sticrty (1862-1907) The existence of pigmy elephants on the island of Malta (Hlephas melitensis Falc.), as first announced at the Cambridge meeting of the British Association, October 6, 1862, astounded the paleontologists. Besides two dwarfed species (#. melitensis Fale., 1862, 1868, and E. falconeri Busk, 1867), both found in the Zebbug Cave of Malta, there were traces of a third elephant of nearly normal size referred to Elephas antiquus. In addition to reviewing this important discovery, Leith Adams (1870, p. 224) added a third species (EL. mnaidrz), a name amended by its author to H. mnazdriensis in 1874, p. 116). “This species [Lydekker, 1886, p. 138] is considered to have averaged between six and seven feet in height and to have been allied to the narrow- crowned race of H£. antiquus and also to E. africanus. The ridge-formula is given by Leith Adams [Footnote: ‘Trans. Zool. Soc. vol. ix. p. 112.’| (exclusive of talons) as’’: ‘Elephas mnaidrz’: Dp 2% Dp3% Dp4 &2 M12 M243 M3223 12-13° This formula is much lower than that of Hesperoloxodon antiquus, as cited from Leith Adams above; it is actually lower than that of H. antiquus as now determined, namely, M 3 1£%:42; but it is higher than that of Loxodonta africana, namely, M 3 --*° 1 Boa a Sicity.—The fossil elephants discovered from early times in Sicily included forms which Falconer considered identical with the full-sized ‘Hlephas antiquus’ and with ‘EH. melitensis,’ a dwarfed form. Pohlig (1898, p. 81) described the latter, from the Elephant Cave of Carini near Syracuse, as Elephas (antiquus) Melite Fale. Soergel (1912.2, p. 1) applies the term E/. antiquus var. insularis to the Sicilian stage from Carini. According to our record no new species have been described from Sicily, in which large island also the nearly full-sized elephant recorded as ‘Elephas antiquus’ prevailed at a certain period before it became dwarfed. ‘Compare Preface by F. A. Bather to Memoir by Andrews and Cooper “On a Specimen of Elephas antiquus from Upnor,” 1928, p. iii. THE LOXODONTIN/A: CLASSIFICATION AND HISTORY OF DISCOVERY 1183 Ramiro Fabiani (1928, p. 34) records the distribution of the dwarfed elephants of Sicily as follows: Elephas (antiquus) mnaidriensis Leith Adams.—Grotte di 8. Ciro, Olivella, Benfratelli, Billiemi, S. Elia, dei Puntali, Maccagnone?, 8. Teodoro. Corso Vittorio Emanuele, Teatro Massimo, Giardino Inglese, Cappuccini, Acqua dei Corsari, Fossa della Garofala (Palermo), Presso Case Capizzi (Monreale). Montelepre, Aleamo, Marianopoli, regione ‘La Romana’ presso Sciacca, Siracusa, T’abuna di Ragusa. Hlephas (antiquus) melitensis Falconer (incluso |’E. Falconeri Busk).—Grotte 8. Ciro, Addaura, Luparello e Stazione di Boccadifaleo, Cava dell’ Arena, Carburanceli, Cava Muletta (Capaci), S. Teodoro, della Santa (Siracusa). Montelepre, Tabuna di Ragusa. SARDINIA.—In 1883 Forsyth Major described the dwarfed species, Elephas Lamarmorae, from this island. EasteRN MEDITERRANEAN.—In 1903 Dorothea Bate described the diminutive form Elephas cypriotes from the island of Cyprus. In 1907 the same author described the diminutive form Hlephas creticus from the island of Crete. The dwarfed elephants of the Mediterranean Islands were long considered as subspecies of ‘Hlephas antiquus,’ but the cranium of ‘H. melitensis,’ as figured by Pohlig (Fig. 1121), more closely resembles that of ’. [Palzoloxodon| namadicus of India. In October of 1929 appeared Raymond Vaufrey’s admirable Memoir ‘‘Les Eléphants Nains des Iles Méditer- ranéennes”’ (Arch. Inst. Pal. Humaine, Mem. VI). This Memoir, with its invaluable observations and detailed studies, has been briefly abstracted and annotated in Section V of the present chapter. Osborn accepts many of Vaufrey’s conclusions but gives his reasons for hesitating or declining to accept others. Extinct AFRICAN SPECIES AND SUBSPECIES, DISCOVERY AND DESCRIPTION (1875-1929) About 1875 discoveries began in the Pleistocene of North Africa. The first fossil species described, Hlephas atlanticus Pomel, 1879, proves to be a primitive loxodont and in the present Memoir is treated as Palzxoloxodon atlanticus. ‘To the same species may be assigned Elephas africanus foss. of Thomas, named in 1884. In 1895 Pomel also described another form from Algeria, namely, Elephas jolensis, which is also a loxodont and is here _ treated as Palxoloxodon jolensis. Ancestral forms of the true ‘Elephas africanus’ (cf. Pomel, 1895, pp. 66, 67, Pls. 1-111) suddenly occur in the more recent Quaternary deposits of North Africa, as excellently described and figured by Pomel, who in his ‘‘Ré- sumé Général Stratigraphique”’ distinguishes the Quaternary Proboscidea of the North African littoral as follows: Most Recent Elephas africanus [= Loxodonta africana] not included in any of the prehistoric drawings of more recent age. Certainly living in Barbary at a prehistoric period, but not figured. Found in the geologic horizons of least antiquity. Ridge-plate and dental formule corresponding with Falconer’s formule of Loxodonta africana. ELEVATED Marine LiTroraAu, PLEISTOCENE Elephas [Palzoloxodon| atlanticus Pomel most characteristic and abundant of this period, affording a complete dental formula. More primitive than Elephas |Hesperoloxodon| antiquus, a contemporary of Bubalus antiquus in southern Oran; figured in wall drawings by contemporary prehistoric men, who showed the difference from the African species by the conformation of the ears (ef. Fig. 1047). Elephas [Palzoloxodon| melitensis(?) Fale. ref. Quaternary of Ternifine. Elephas |= Palzoloxodon|] jolensis Pomel, of Jol ou Julia casarcea, Cherchel, related to antiquus and to mnaidriensis of the island of Malta, but with less numerous ridge-plates. PLIOCENE SERIES Mastodon |= Zygolophodon|] borsoni ref. Y Elephas |= Archidiskodon] meridionalis ref., possibly referable to Archidiskodon planifrons. 1184 OSBORN: THE PROBOSCIDEA It is to these six species of proboscideans that Pomel’s Memoir of 1895, “Les Kléphants Quaternaires,”’ is devoted. oy PAL®OLOXODON AND HESPEROLOXODON AS DeEpicTED BY THE CAVE MEN oF NortH AFRICA AND SPAIN. WITH SMALL EARS The left-hand figure, after Pomel, probably relates to Palxoloxodon atlanticus; the right-hand and middle figures, after Breuil, probably relate to Hesperoloxodon antiquus platyrhynchus Fig. 1047. The left-hand figure is designated by Pomel (1895, Explanation of Pl. x1v, fig. 4) as follows: “Eléphant atlantique de Guebar-Rechim.” Algeria. The right-hand and middle figures are designated by Breuil (1911, p. 61, text fig. 57, and p. 239, text fig. 245) as follows: (Right figure) “Tléphant tracé en rouge (largeur o™ 44) et signes formés de lignes rouges verticales en faisceau.”” Cavern of Pindal, province of Asturias, northern Spain. Middle figure) “Figures d’Eléphants gravés dans le Sud Oranais ....d’aprés Flamand.” Algeria. Observe that these figures, although drawn by different artists, present a striking general agreement, that is, in the general body profile, the elevation of the limbs, the slight concavity of the forehead, the shortness of the inferior abdominal region, and especially, as noted by Pomel, in the contour of the rel- atively small and depressed ears, which entirely differ from those of the living African elephant or even from those of the pigmy variety (Loxodonta africana pumilio). Consequently these figures are used in the preparation of our restorations of the Upnor elephant (Fig. 1074) and other representatives of Hesperoloxodon antiquus. The contour of the ear resembles that cf the Indian elephant (Fig. 1120) rather than that of the African elephant (Fig. 1052). The first fossil elephant to be described from South Africa was Scott’s Elephas (Loxodon) zulu, 1907, which we refer to Loxodonta zulu (see Osborn, 1934.925, p. 2). This was followed in 1916 by Dietrich’s description of Elephas antiquus Recki of northern Tanganyika Territory, referred in the present Memoir to Palxoloxodon recki. The supposed Loxodonta griqua of Haughton, from South Africa, described in 1922, proves to belong to the more ancient genus Archidiskodon, namely, A. griqua.! YERMANY AND RUMANIA In 1924 Sabba Stefanescu described the two subspecies Elephas antiquus rumanus from Tulucesti, Rumania, and H. antiquus germanicus from Tanganu (Ilfov) Rumania, horizon of Taubach and Weimar, Germany, as well as a referred 2. antiquus ausonius and a referred EL. antiquus germanicus also from Ifov, Rumania. It is somewhat doubtful whether these moderately broad-plated types (Figs. 857 and 1089) are truly referable to the Hlephas |Hesperoloxodon| antiquus group or to the extremely broad-plated Archidiskodon planifrons. Judging from Stefanescu’s figures it would appear that: Blephas antiquus rumanus 8. Stef., of Rumania =Archidiskodon planifrons rumanus. Upper Pliocene of Rumania. Elephas antiquus germanicus 5. Stef., of Germany and Rumania = Hesperoloxodon antiquus germanicus. Upper Pleistocene of Germany and Rumania. Elephas antiquus ausonius ref., of Ilfov, Rumania = Hesperoloxodon antiquus ausonius. ?Upper Pliocene. [Professor Osborn in 1934 (1934.925, p. 12) made Loxodonta griqua the type of a new genus, namely, Metarchidiskodon (see Chap. XVI, p. 994).— Editor.] THE LOXODONTINA: CLASSIFICATION AND HISTORY OF DISCOVERY 1185 JAPAN AND JAVA (Compare abstract of Matsumoto’s articles, also letters of July 14 and November 20, 1924, pp. 901-908, Chap. XIV) JAPAN.—For a long period Japan constituted the extreme eastern portion of the continent of Asia, and, as described in recent papers and memoirs by Matsumoto and others, contained a stegodont and elephant fauna closely related to that of Burma and India. Beginning in the year 1924 Matsumoto has given a complete synopsis of this mastodont, stegodont, and elephantine fauna, which is summarized above (Chap. XIV, pp. 901-908). In the present chapter it is shown that all the elephants described by Matsumoto under different generic and specific names probably belong to the Loxodontine and are more or less closely related to Palxoloxodon (syn. Sivalikia) namadicus; none of them appears to be related to the genus Loxodonta or to the genus Parelephas as Matsumoto supposed. In the order of description, these Japanese species are as follows: OriainaL Name, Matsumoto, MAKIYAMA Speciric REFERENCE IN PresENT Memoir Elephas namadicus naumanni Makiyama (English description: June 30, 1924, p. 264) = Paleoloxodon namadicus naumanni Made the genotypic species of Palxolorodon Matsumoto (Sept. 20, 1924, p. 257) from an admirable type which is fully described and refigured on pp. 1294-1296 of the present Memoir. Elephas namadicus namadi Makiyama (English description: June 30, 1924, p. 264) = Palzxoloxodon namadicus namadi! Loxodonta Cuvier Type Elephas africanus. Subgenus. Palzolorodon, n. Matsumoto (Sept. 20, 1924, pp. 257, 260) =F. antiquus-namadicus group = Genus PALAMOLOXODON Euelephas protomammonteus, sp. n. Matsumoto (Sept. 20, 1924, p. 262) = Palzxoloxodon protomammonteus Loxodonta (Palxoloxodon) tokunagai, sp. n. Matsumoto (Sept. 20, 1924, p. 267) = Palzxoloxodon tokunagai Loxodonta (Palzxoloxodon) namadicus naumanni (Mak.), Matsumoto (Sept. 20, 1924, p. 268) = Palzxoloxodon namadicus naumanni Loxodonta (Palxoloxodon) namadicus Yale. and Caut. (typicus), Matsumoto (Sept. 20, 1924, p. 269) = Palxoloxodon namadicus Parelephas protomammonteus proximus Matsumoto (1926.2, pp. 48-50) = Palxoloxodon protomammonteus proximus Loxodonta (Palxoloxodon) namadica (Yaber) Matsumoto, 1929, p. 4 = Palxoloxodon namadicus yabei Loxodonta (Palxoloxodon) Tokunagai junior mut. Matsumoto, 1929, p. 10 = Palzxoloxodon tokunagai mut. junior Of the above species six type figures have been published which are available for reproduction in the present Memoir, namely, (1) Makiyama’s type figures of Hlephas namadicus Nawmanni (here reproduced as Fig. 1152 below) and (2) of Elephas namadicus namadi (Fig. 1153); (3) Matsumoto’s type figures of Huelephas (Parelephas) protomammonteus (Fig. 1154) and (4) of Parelephas protomammonteus proximus (Fig. 1155); also (5) of Loxodonta (Palxoloxodon) namadica (Yabei) (Fig. 1156), and (6) of Loxodonta (Palxoloxodon) Tokunagai junior mut. (Fig. 1157). In 1924 (Sept. 20) Matsumoto published in the Journal of the Geological Society of Tokyo, Vol. XX XI, his article ‘‘Preliminary Note on Fossil Elephants in Japan” (pp. 255-261), in which were listed the species named in the column above, with the exception of the subspecies mentioned in the preceding paragraph, namely, Nos. 2,4,5, and 6. This Japanese paper was not received by the present author until February, 1929; if sent, the first copy was lost in transmission. [See ‘1924 Elephas namadicus namadi Makiyama,” Chap. XXI, p. 1409, of the present Memoir.—Hditor.] 1186 OSBORN: THE PROBOSCIDEA In 1926 (Matsumoto, 1926.1) the same treatment was repeated in the English language and the same series of species was listed with the localities in which each was found, as reproduced in Chapter XIV, pp. 901-908, of the present Memoir. This list includes the following (ef. Chapter XIV, pp. 906-908, for full citations from Matsu- moto’s printed and written synopses) : 9. Euelephas protomammonteus Matsumoto, 1924, p. 262. Smaller than H. trogontherii. Of Calabrian age. 10. Euelephas trogontherii (Pohlig), 1924, p. 265. Of Cromerian age. 11. Hlephas indicus Linn., 1924, p. 266. Probably of Post-Monastirian and Pre-Neolithic age. 12. Loxodonta (Palzxoloxodon) tokunagai Matsumoto, 1924, p. 267. Possibly older Pleistocene. 13. Loxodonta (Palxoloxodon) namadica naumanni (Makiyama), 1924, p. 264. Of Cromerian [Lower Pleistocene] age. 14. Loxodonta (Palxoloxodon) namadica (Fale. and Caut.), 1924, p. 269. Of Milazzian-Tyrrhenian age. As stated above the specific and generic reference of these elephants awaits further study and comparison. With the possible exception of the specimen identified as Elephas indicus, Osborn is inclined to regard them all as belonging to the Palzoloxodon (syn. Sivalikia) phylum. Java.—In 1908 Dubois described from the Kendeng formation, Pithecanthropus zone, Lower or Middle Pleistocene of Java, Elephas hysudrindicus as standing near the Elephas hysudricus Falconer of the Siwaliks, but still more close to Hlephas indicus, accordingly naming it hysudrindicus. Stremme (1911) related this species more closely to ‘Elephas antiquus.’ Accompanying the publication of figure 1160, this species may confidently be placed with Palxoloxodon as P. hysudrindicus. Soe ~ PALAEOLOXODO Fig. 1048. Geographic distribution (according to the numbers in the following list, which represent the chronologic sequence of type description) of the principal species of the Loxodontine. See also figure 1055, map of distribution of existing African elephants. 28. THE LOXODONTINA:: CLASSIFICATION AND HISTORY OF DISCOVERY 1187 ORDER OF DISCOVERY AND DESCRIPTION OF THE FIFTY-THREE TYPE SPECIES 1797 1798 1821 1823 1846 1846, 1847 1847 1857 1857 1862, 1868 1867 1868 1870 1870 1875-1886, 1923 1879 1880 1883 1883 1883 1883 1884 [1888 [1889 [1891 1891, 1902 1895 1897 1903 1907 1907 1908 1912 1916 [1922 1924 1924 South Africa South Africa Germany Germany India England Europe Europe England Malta Malta England (Gray’s Thurrock) Malta Sicily (Catania) Italy North Africa Northern Italy England (Norwich) England (Norwich) England (Forest Bed, Norfolk) Sardinia North Africa Germany Germany England (lorest Bed, Norfolk) England (Forest Bed, Norfolk) Algeria Spain Cyprus Crete South Africa Java Sicily Tanganyika Terri- tory South Africa Japan Japan OF THE EXTINCT LOXODONTIN/A See Figure 1048 Elephas africanus Blumenbach. Probably Cape region; exact locality unknown Elephas capensis G. Cuvier, Cape of Good Hope Elephas priscus Goldfuss, near (?) Cologne Elephas antiquitatis Kriiger, Thiede, Thuringia Elephas namadicus Falconer and Cautley, Nerbud- da Valley E. [Elephas| priscus? Falconer and Cautley, Gray’s Thurrock—see E. (Loxod.) priscus (Goldf.) Falconer, 1857, below, this list. Elephas antiquus Falconer and Cautley, locality unrecorded KE. (Eueleph.) antiquus Falconer B. (Loxod.) priscus (Goldf.) Faleoner—see Elephas (Loxod.) priscus Falconer, 1868, below, this list Elephas Melitensis Falconer, Zebbug Cave Elephas falconeri Busk, Zebbug Cave E. (Loxod.) priscus Falconer Elephas mnaidrae Adams, Mnaidra Gap Elephas Cornaliae Aradas Elephas ausonius Major (MS., 1875), in Verri, 1886 (name only), Depéret and Mayet, 1923, San Romano, Val d’Arno inferior Elephas atlanticus Pomel, Ternifine, Algeria Elephas antiquus nana Acconci, near Monti Pisani, Cucigliana, Tuscany Leptodon minor Gunn Leptodon giganteus Gunn E. Gunnii Lartet Elephas Lamarmorae Major, near Gonnesa Elephas africanus fossilis Thomas Elephas (primigenius) Leith-Adamsi Pohlig Elephas (primigenius) Leith-Adamsi minor Pohlig Elephas antiquus Nestii Pohlig E. giganteus intermedius Gunn Elephas jolensis Pomel, Algerian seacoast, below Kromer-Roubia Elephas platyrhynchus Graells, San Isidro del Campo, near Madrid Elephas cypriotes Bate, Kerynia Hills Elephas creticus Bate, near Cape Maleka Elephas (Loxodon) zulu Seott, Zululand Elephas hysudrindicus Dubois, Kendeng-Schichten Elephas antiquus var. insularis Soergel (name only), Carini y Elephas antiquus Recki Dietrich, Oldoway-Tuffe, = Loxodonta africana = Loxodonta africana capensis = Loxodonta africana = Hesperoloxodon antiquus germanicus = Palxoloxodon namadicus = Hesperoloxodon antiquus = Hesperoloxodon antiquus = Palzxolorodon melitensis = Palzxolorodon falconeri = Hesperoloxodon antiquus = Palxoloxodon mnaidriensis = Loxodonta cornaliae = Hesperolorodon antiquus ausonius = Palzoloxodon atlanticus = Hesperoloxodon antiquus nanus = Hesperoloxodon antiquus = Hesperoloxodon antiquus = Hesperoloxodon antiquus = Palxoloxodon lamarmorae = Palexoloxodon atlanticus See Mammonteus? (Chap. XVIII)] See Mammonteus? (Chap. XVIII)] See Parelephas (Chap. XVII)] = Hesperoloxodon antiquus = Palxoloxodon jolensis = Hesperoloxodon antiquus platyrhynchus = Palxoloxodon cypriotes = Palxoloxodon creticus = Loxodonta zulu = Palxoloxodon hysudrindicus Nomen nudum Serengetisteppe, northern Tanganyika Territory =Palxoloxodon recki Loxodonta griqua Haughton Elephas namadicus Naumanni Makiyama, Sahamma, Tét6mi Province Elephas namadicus namadi Makiyama, island of Shédo, Sanuki Province See Metarchidiskodon (Chap. X VI)] = Palzxoloxodon namadicus naumanni = Palxolorodon namadicus namadi 1188 29. 30. ol. 32. 30. 40. 1927 1927 1929 1929 1929 1929 1929 1929 1929 1929 1929 1929 1931 1931 1931 1932 1932 1934 1935 1937 1939 Japan Rumania Rumania Japan Japan Japan Japan South Africa South Africa Japan Japan Japan South Africa South Africa South Africa South Africa South Africa South Africa South Africa Southern Italy England Japan South Africa South Africa Japan India Africa OSBORN: THE PROBOSCIDEA Loxodonta (Palxoloxodon) Tokunagai Matsumoto, Soyama, Gokayama, Hira-mura, Higashi- Tonami District, Province of Etcht Elephas antiquus rumanus 8. Stefiénescu Elephas antiquus germanicus S$. Stefinescu, Tanganu (Ilfov), horizon of Taubach and Weimar Euelephas protomammonteus Matsumoto. See be- low under Parelephas protomammonteus Matsu- moto, 1926 Parelephas protomammonteus (Matsumoto) typicus Matsumoto, Nagahama, Kimitsu District, Province of Kazusa Parelephas protomammonteus proximus mut. Mat- sumoto, Isone, Kokubo, Onuki-mura, Kimitsu District, Province of Kazusa Elephas indicus Buski Matsumoto, Ninohe Dis- trict, Province of Mutsu Archidiskodon transvaalensis Dart, near Bloemhof, Vaal River Archidiskodon sheppardi Dart, near Bloemhof, Vaal River Loxodonta (Palzoloxodon) namadica (Yaber) Mat- sumoto, Inland Sea Lox. (Pal.) Tokunagai junior, mut. Matsumoto. Exact locality unknown Elephas (Palxoloxodon) namadicus setoensis Maki- yama, Seto Archidiskodon andrewsi Dart, Gong-Gong, Vaal River Archidiskodon hanekomi Dart, Delport’s Hope, Vaal River Pilgrimia yorki Dart, below Christiana, Vaal River Pilgrimia wilmani Dart, below Christiana, Vaal River Pilgrimia kuhni Dart, Pniel Estate, Vaal River Loxodonta prima Dart, Pilandsberg, Limpopo River Loxodonta africana var. obliqua Dart, Kranzkloof farm, Steelport River Palexoloxodon antiquus italicus Osborn, Pignataro Interamna, near Naples Palzoloxodon antiquus (andrewsi?) Osborn, Upnor, Kent Parelephas protomammonteus (Matsumoto) mat- sumotoi Saheki, Mishima, Kimitsu District, Province of Kazusa Pilgrimia archidiskodontoides Haughton, Sydney- on-Vaal Breakwater Pilgrimia subantiqua Haughton, Delport’s Hope, Vaal River Palxoloxodon yokohamanus Tokunaga, Yokohama Palxoloxodon priscus var. bosei Chakravarti, Jammu, Siwaliks Palaeoloxodon darti Cooke and Clark, Victoria Falls, northern Rhodesia = Palxoloxodon tokunagai See Archidiskodon (Chap. XV1)] = Hesperoloxodon antiquus germanicus = Palexoloxodon protomammonteus =Palxoloxodon protomammonteus proxi- mus =?Palexoloxodon buski (See Chap. XX, p. 1833, for description and figure) = Palzxoloxodon transvaalensis =Palzxoloxodon sheppardi = Palzxoloxodon namadicus yabet =Palzxoloxodon (?Archidiskodon) tokun- agai mut. gunior {Not determined by the present author] =?Palxoloxodon andrewsi = Palzxoloxodon hanekomi = Palxoloxodon yorki = Palxoloxodon wilmani = Palzxoloxodon kuhni = Loxodonta prima = Loxodonta africana var. obliqua = Hesperoloxodon antiquus italicus = Hesperoloxodon antiquus {Not determined by the present author] = Palxoloxodon archidiskodontoides = Loxodonta subantiqua [| =Palzxoloxodon yokohamanus| {Not determined by the present author] [| Not determined by the present author] HABITAT OF THE AFRICAN ELEPHANT (LOXODONTA) I'OREST AND SAVANNA OF THE UASIN GisHu PLATEAU, Kenya CoLony, NEAR THE ’Nzor River. ArrER PHOTOGRAPHS BY Kermit RoosrvELtT (RIGHT) AND Cart E. AKELEY (LEFT) Vig. 1049. Calf of old female charging elephant. Open savanna country. Fig. 1050. Females and young bulls in forest. Photograph by Kermit Photograph by Carl E. Akeley. Roosevelt. LAST OF THE AFRICAN ELEPHANT (LOXODONTA) Vig. 1051. A small herd of elephants, containing two bulls, two cows with calves, and one young bull in the center, passing through the grassy meadows of an open savanna country in British East Africa. After a film photograph taken by Martin Johnson in the year 1923. 1189 AKELEY Group OF AFRICAN ELEPHANTS IN THE AMERICAN Musrum Fig. 1052. A special expedition of the American Museum of Natural History was sent out in 1909 under Carl E. Akeley to collect materials for this group of African elephants, male, female, and young, for the African Hall of the Museum. The cow elephant was shot by ex-President Theodore Roosevelt, and the calf by Kermit Roosevelt, in 1909, on the Uasin Gishu Plateau where the Roosevelt. and Akeley expeditions met. The large bull was shot by Mr. Akeley. The composition (A, B) shows the bull scenting danger, silently feeling fer scent with his trunk, ears fully extended to catch the least sound, for he does not see the source of disturbance. The attitude of the cow indicates that she has scen the intruder and has “frozen,” ears back, trunk pendant, prepared for any move she may decide on, whether attack or retreat. The calf, conscious of the alarm, is snuggling up to its mother for protection. The young bull to the right (C), startled, has started forward to swing around and face the danger, his trunk thrown back to catch the seent, and his ears forward to catch the sounds. MEASUREMENTS IN Minuimerers or ADULT AND YOUNG Amer. Mus. 32727 Amer. Mus. 32732 Amer. Mus. 32734 Amer. Mus. 54085 Calf l'emale Small Bull Large Bull Height of shoulder... .. ae ne ae 1358.9 2590.8 2711.45 3251.2 =10 ft. 8 in. Height to top of head...... Ents ae 1409.7 2679.7 2628.9 3403.6 =11 ft. 2% Circumference of forefoot. t ats : 654.05 1174.75 1352.55 1473.2= 4 ft. 10 Height above pelvis. . . eters « : 1346.2 2451.1 2489 .2 3048. =10 ft. Total spread of ears. . : 1066.8 2400.3 2870.2 3149.6 =10 ft. 21% Exposed length of tusks: (ayiletti msn. + sas > Othe ; 38.1 508 635. 1422.4= 4 ft. 8 (b) right. . eo: 44.45 508 660.4 1447.8= 4 ft. 8% GeroarapHic DATA oF THE FouR AFRICAN ELEPHANTS IN THE GROUP 1. The large bull (Loxodonta africana albertensis). 3. The old female (Loxodonta africana peeli). Locality: Budongo Forest, east of Lake Albert, Unyoro, northern Uganda. Locality: Near ’Nzoi River, Uasin Gishu Plateau, Kenya Colony. Amer. Mus. Dept. Mam. 54085. Shot by Colonel Theodore Roosevelt. (According to condition of molars about middle-age.) Amer. Mus. Dept. Mam. 32732. 2. The small bull (Loxodonta africana albertensis). (According to condition of molars approaches senile condition.) Locality: Budongo Forest, east of Lake Albert, Unyoro, northern Uganda. 4. The bull calf (Loxodonta africana peeli). Amer. Mus. Dept. Mam. 32734. Locality: Near ’Nzoi River, Uasin Gishu Plateau, Kenya Colony. Shot (According to condition of molars rather older, but certainly not younger by Captain Kermit Roosevelt. than the large bull No. 54085.) Amer. Mus. Dept. Mam. 32727. 1190 II. SYSTEMATIC REVISION OF THE LOXODONTINA SUPERFAMILY: KLEPHANTOIDEA Osborn, 1921 FAMILY: ELEPHANTID Gray, 1821 SuBFAMILY: LOX ODONTINZ@ Osborn, 1918 Original reference: Osborn, Bull. Geol. Soc. Amer., X XIX, table (opp. p. 134), p. 136 (Osborn, 1918.468); Amer. Mus. Novi- tates, No. 1, 1921, p. 15 (Osborn, 1921.515). Compare Pohlig, 1885-1888, Loxodonten, Loxo (disko) donten, Loxo (disko) don (Elephas africanus, E. priscus, E. antiquus). SUBFAMILY CHARACTERS.— United by the common dental and cranial characters of Loxodonta, Palzxoloxodon, and Hesperoloxodon. Cranium relatively primitive, platycephalic, brachycephalic (Loxodonta); somewhat more elevated or hypsicephalic in Palxoloxodon and Hesperoloxodon. Pre- maxillary rostrum broadened for insertion of widely divergent incisive tusks. Tusks relatively straight or slightly incurved, continuously serving in uprooting habits, chiefly in forests. Grinding teeth moder- ately hypsodont, typically narrow to broad; lozenge shaped (Loxoedonta) or with ‘loxodont sinus’ rudimentary or absent (Palzoloxodon, Hesperoloxodon). Habits chiefly browsing, crushing of coarse leaf- age, herbage, and wood fiber. Ridge formula progressive from M 3 |,"°5 (Loxodonta), to M 312 (Palzolox- odon), to M 3 38; (Hesperoloxodon). The grinding teeth of Palxoloxodon, as shown in P. namadicus, with ‘‘colliculi approximati”’ of Falconer (Figs. 1070 and 1189), certainly display no resemblance to the lozenge-shaped grinders of the existing species of Loxodonta, all of which are of the typical ‘losange’ form (Figs. 1043, 1058) first observed by F. Cuvier. Of somewhat intermediate ridge pattern are many of the species of Hesperoloxodon, such as are shown in figure 1076 in the ‘Elephas priscus’ of Falconer and Cautley, but less distinctly in the lectotype specimen Elephas [Hesperoloxodon| antiquus (Fig. 1075). This narrow ‘loxodont sinus,’ although but a rudiment, is characteristic of many of the species of Palzoloxodon in the Mediterranean Islands and in Africa; it is also observed in primitive Archidiskodon molars. From the resemblances and contrasts enumerated above in the comparative figures and definition of Loxo- donta, Palzoloxodon, and Hesperoloxodon, it is certain that we have to do with three distinct generic phyla, the descent lines of which begin to be known only in Upper Pliocene and Lower Pleistocene times. One of the strongest proofs of this profound phyletic cleavage is seen in the fact that all the modern living species of Loxodonta have a more primitive cranial structure and a more primitive ridge formula, namely, M 3 ,,*°, (Loxodonta afri- 11-12 cana), whereas the long extinct Palzoloxodon namadicus exhibits M 3 +2 and the typical Hesperoloxodon antiquus 1642-17 M 16%-17° In brief, in all cranial and dental characters the surviving species and subspecies of Loxodonta africana are more primitive than any of the known fossil species or subspecies of Palzoloxodon namadicus or of Hesperoloxodon antiquus. Genus: LOXODONTA F. Cuvier, 1825, 1827 Compare Loxodonte F. Cuvier, 1825; unsigned Loxodonta, 1827; Loxodonta Gray, 1843; Loxodon Falconer, 1847-1857. The South African form Elephas africanus Blumenbach, 1797, is probably the genotypic species. It is important to note that in 1798 G. Cuvier distinguished the extreme South African form as Elephas capensis. GENERIC CHARACTERS.— Ridge-plates of grinding teeth expanding into broad ‘losange’ or ‘lozenge- shaped’ median sinus. Relatively narrow superior and inferior grinding teeth, with relatively few ridge- plates; total permanent ridges Dp 4—M 3: 22. Enamel borders thick and simple without foldings or plications. Superior tusks widely divergent as they issue from the broadened premaxillaries; relatively straight, sightly upcurved (Fig. 1059) and incurved (Fig. 1063); marked sexual disparity in the tusks of females (Fig. 1063). Typical ridge formula: Dp 2%Dp3%Dp44M17M2,2, M3 2 11-12° 1191 1192 OSBORN: THE PROBOSCIDEA Premaxillaries broadened; parietofrontal cranial vertex rounded, platycephalic to subacrocephalic. Cranial profile and section much more primitive, less hypsicephalic and bathycephalic than in the Mam- montine (ef. pp. 921-925) or Elephantine (ef. p. 921). Vertebral formula: Cervicals 7, dorsals 20-21, lumbars 3, sacrals 4, caudals 26-31. Adaptation of the grinding teeth (cf. Falconer, p. 927 above). The grinders of the African elephant are relatively primitive in construction; the aggregate series of upper ridge-plates (Dp'—M_*) amounts to only 32 as compared to 48 in ‘Elephas antiquus’ and 64 in Elephas indicus. The molars are shorter, narrower, and of less elevation than in EL. [Hesperoloxodon] antiquus or E. indicus. In E. [H.] antiquus the numerous and closely set ridge-plates, without mesial expansion, indicate feeding habits similar to those of E. indicus. In Loxodonta africana the dises of wear exhibit the well-known rhomboidal expansion or ‘Josange’ characteristic of all the species. These low grinding teeth are best adapted to squeezing and crushing leaves and succulent stems or roots. Besides browsing on the foliage of the Mimosas and Acacias, these African elephants used their tusks like a crow-bar to tear up the trees of certain species by the roots. In Elephas |Hesperoloxodon| antiquus the numerous narrow transverse ridge-plates of Dp 4—M 3, totaling 48, form a perfect triturating mechanism like that of the Indian elephant. This food adaptation alone, in Falconer’s opinion of 1868, would entitle his Elephas [Palxoloxodon| namadicus to subgeneric distinction from his ‘Loxodon’; he accordingly grouped it under the unusable generic names of Elasmo- don (preoccupied) and EHuelephas (invalid). Generic Description, 1825-1827.—Cuvier, Frédéric, et Geoffroy Saint-Hilaire, Etienne, 1824-1829, “Histoire Naturelle des Mammiféres,” Vol. III, Livr. LI, LII, 1825, p. 2 (Loxodonte): Je proposerai pour nom générique de cette espece, le mot de Loxedonte, qui peut rappeler le caractere de ses dents, les losanges qu’on apercoit sur leur coupe. A review (unsigned) of this work appeared in the Zoological Journal, London, 1827, 1828, III, p. 140, noticing the: dismemberment of the genus Hlephas, for the purpose of establishing a new one under the name of LoxoponrTa. . . . For the Elephant of Asia he [Cuvier] retains the original generie name Elephas. The surfaces of its molar teeth present fasciee of enamel irregularly festooned; while in those of the African Elephant, the type of the new genus Loxoponra, the enamel is disposed in lozenges. In addition to this striking distinction derived from the dentary system, M. F. Cuvier also enumerates the other characters which have hitherto been regarded as specific. The smaller, more elongated, and less irregular head of the African animal when compared with the Asiatic; the rounded forehead of the former, strongly contrasted with the deep depres- sion in the middle of that of the latter; the ear of the former also twice the extent, while the tail is only half the length, &e. 1. ORDER OF DESCRIPTION OF EIGHTEEN LIVING AFRICAN SPECIES AND SUSBPECIES' Osborn does not express any opinion as to the validity of all these eighteen African geographic species and subspecies, as listed below with the aid of Mr. Herbert Lang and Dr. Paul Matschie, but furnishes this geographic- varietal list as a real contribution to paleontology and as a picture of past conditions, namely: Jn past time each extinct collective species, by local and continental adaptive radiation, doubtless embraced numerous geographic sub- species and varieties adapted to local conditions and as clearly distinguishable from each other in external and internal characters as are the eighteen living African species and subspecies. From Blumenbach’s first notice in 1797 to Roosevelt and Heller’s work? in 1914 and finally Frade’s articles of 1924 and 19282, eighteen species and subspecies have been proposed led by the Elephas capensis of G. Cuvier, a name which is probably a synonym of Loxodonta africana. This list is extremely valuable to the paleontologist as demonstrating the very wide range of geographic-varietal characters in existing loxodonts of a single continent. Undoubtedly a similar range of geographic-varietal characters distinguished the great herds of elephants belonging 1Prepared with the codperation of Mr. Herbert Lang of the American Museum and of Dr. Paul Matschie of the Berlin Museum. Chronological list of the eighteen names proposed for the living African elephants, with the name as the author originally described it (to the left) and the present proposed name (to the right), together with references to the type figures, locality of the type specimens, and the museums in which they are to be found. 2Roosevelt, Theodore, and Heller, Edmund. 1914. “‘Life-Histories of African Game Animals,” Vols. I, Il, New Yerk. Also Frade, Fernando, 1924, “Notes de Mammalogie Africaine,” Pt. 2—Les Eléphants du Jardin Zoologique de Lisbonne, Bull. Soc. Portugaise Sci. Nat., Tome IX, Fasc. 3, pp. 180-135, text figs. 1-6, and 1928, “‘Titulos e trabalhos cientificos (Curriculum vitae),’”’ pp. 15, 16, Lisboa. THE LOXODONTIN: CLASSIFICATION AND HISTORY OF DISCOVERY 1193 to the genera Archidiskodon, Parelephas, Mammonteus, and Elephas. A similar range of geographic-varietal characters is now observed in the four to five living species and subspecies of the true genus Hlephas, as described below (Chap. XX). According to different observers, the vertebral formule, as given in detail on pages 930 and 931 of the present Memoir, undoubtedly differ among these numerous subspecies and geographic varieties. The food of the African elephant, as observed by Falconer (p. 927), also probably differs in the various regions of Africa. Elephas africanus Blumenbach, 1797, No. 19, fig. C. Type loc.: South Africa; according to SUBSERCIEG Matschie (letter, 1921) probably from the Cape. [Location of type molar unknown.— Editor.] - = Loxodonta africana africana! Elephas capensis G. Cuvier, 1798, p. 149. Type loc.: ?Upper Orange River, eastern S. Africa. Type fig.: skull, in Cuvier, 1799, pp. 1-22. Type probably in Museum, Paris = Loxodonta africana capensis Elephas cyclotis Matschie, 1900, p. 194. Type loc.: Mwelle district northeast of Yaunde on the left bank of the Sanaga, east of the Nachtigall Falls, S. Cameroon. Type fig.: Heck, 1899, p. 116, Pl. exivi (live animal from the Berlin Zoological Garden). Type (skull) in Berlin Museum = Loxodonta africana cyclotis? Elephas (Loxodonta) oxyotis Matschie, 1900, p. 196. Type loc.: Upper Atbara, Sudan. Type not specified; species founded on numerous examples brought back from the Upper Atbara region, Sudan, by Casanova, Hagenbeck and Menges; according to Matschie (letter, 1921) askull, undoubtedly similar to oxyotis, is in the Berlin Museum. = Loxodonta africana oxyotis Elephas (Loxodonta) knochenhauert Matschie, 1900, p. 197. Type loc.: Barikiwa, Tanganyika Territory. Type (skull) in Berlin Museum = Loxodonta africana knochenhaueri Elephas africanus albertensis Lydekker, 1906, p. 1089. Type loc.: South end of Lake Albert. Type fig.: Lydekker, 1907.1, text fig. 121 (skull). Type in British Museum (Natural History) = Loxodonta africana albertensis Elephas africanus pumilio Noack, 1906, p.631. Typeloc.: French Congo. Type fig.: Horna- day, 1905, pp. 237, 238 (photographs of live animal). Type in the American Museum of Natural History (Amer. Mus. 35591) = Loxodonta africana pumilio Elephas africanus toxotis Lydekker, 1907, pp. 385, 388. Type loc.: Mossel Bay, western Cape Colony. Type fig.: 1907.1, text fig. 106 (head). Type (mounted female) in South African Museum at Capetown = Loxodonta africana toxotis Elephas africanus selousi Lydekker, 1907, pp. 387-389. Type loc.: Mashonaland, southern Rhodesia. Type fig.: 1907.1, text fig. 108 (head). Type in Imperial Institute, London = JLozxodonta africana selousi Elephas africanus peeli Lydekker, 1907, pp. 393, 394. Type loc.: Aberdare Mts., Kenya Colony. Type fig.: Lydekker, 1907.1, text fig. 114 (mounted head). Lectotype in private collection of Mr. C. V. A. Peel, 12 Woodstock Road, Oxford. Cotype in Mr. Roth- schild’s Museum at Tring = Loxodonta africana peeli Elephas africanus cavendishi Lydekker, 1907, p. 395. Type loc.: Lake Rudolf district. Type fig.: Lydekker, 1907.1, text fig. 115 (mounted head). Type in British Museum (Natural History) = Loxodonta africana cavendishi Elephas africanus orleansi Lydekker, 1907, p. 398. Type loc.: North Somaliland. Type fig.: Lydekker, 1907.1, text fig. 118 (dried right ear). Type in collection of Due d’Orléans at Wood Norton = Loxodonta africana orleansi Elephas africanus rothschildi Lydekker, 1907, p. 399. Type loc.: French Sudan (‘‘probably southward of Lake Chad’’). Type fig.: Lydekker, 1907.1, text fig. 119 (head from life). Type (skeleton) in the American Museum of Natural History. Lydekker states that he takes as type the statuette of “Jumbo” in the British Museum = Loxodonta africana rothschildi* Elephas africanus cottoni Lydekker, 1907 (1907.2), p. 783. Type loc.: Northeastern Congo. Type fig.: Lydekker, 1907.1, text fig. 111 (ear). Type probably in Powell-Cotton’s private collection = Loxodonta africana cottoni Elephas africanus Fransseni Schouteden, 1914, p. 396. Type loc.: M’Paa near Bongo, north- west of Lake Leopold II. Type fig.: Schouteden, 1914, Pl. x1, figs. 1, 2 (dead animal). Type (skin and skull) in Tervueren Museum = Loxodonta africana fransseni Elephas africanus mogambicus Frade, 1924, pp. 131, 133. Type loc.: Maputo, Mozambique. Type fig.: Frade, 1924, text fig. 5. Type: Female living at the time of description in the Jardin de Zoologique de Lisbonne ’ = Lozxodonta africana mo¢ambica Loxodonta africana Zukowskyi (Strand in Zukowsky, 1924), p. 68. Typeloe.: Kaoko District, southwest Africa. = Loxodonta africana zukowskyi Loxodonta africana angolensis Frade, 1928, p. 15. Type loc.: Region of Cunene, southern Angola; at the time of description, living in the Jardin de Zoologique de Lisbonne = Loxodonta africana angolensis 'Loxodonta africana africana Roosevelt and Heller, 1914, Vol. II, p. 739. Type loc.: near Albert Nyanza. The typical L. africana africana is believed to be Blumenbach’s type from the Cape Colony region. Consequently the Roosevelt and Heller name cannot be used for the subspecies from the Albert Nyanza = Loxodonta africana ?subsp. *[Compare, however, ‘‘Captive pigmy elephants in America,” Journ. Mam., 1934, Vol. XV, p. 248, by C. Emerson Brown.—Editor.] 5[Professor Osborn referred “Jumbo” to the subspecies Lozodonta africana oryotis (Osborn, 1931.846, p. 21), which would make rothschildi a synonym of oxyotis.—Editor.] 1194 OSBORN: THE PROBOSCIDEA Osborn, 1929: In the present Memoir the skull and skeleton of ‘‘Jumbo”’ (Amer. Mus. Dept. Mam. 3283) are referred to the subspecies Loxodonta africana oxyotis rather than to L. a. rothschildi Lydekker. The eighteen subspecies listed above are partly distinguished by size, partly by geographic distribution, and partly by the shape of the ears. The names adopted were listed by Matschie and Lang, with the exception of Loxodonta africana mocambica Frade, Loxodonta africana zukowskyi Strand, and Loxodonta africana angolensis Frade. Fig. 1053. Two growth stages in the Sudanese elephant ‘“Khartum”’ of the species Loxodonta africana oxyotis Matschie, from the Upper Atbara, Sudan, formerly living in the New York Zoo- logical Park. Both photographs taken by Elwin R. Sanborn and reproduced by courtesy of the New York Zoological Society. Reduced to a uniform one-fiftieth scale. ‘‘Khartum’ was born about 1903 and was captured on the Blue Nile in 1906; collected by Capt. Stanley 8. Flower, Director of the Government Zoological Gardens, Egypt. (Left) Young “Khartum”’ photographed in 1908 at the age of 5 years and measuring 5 ft. 1%4in. or 1567 mm. (Right) Young adult “Khartum”’ photographed in 1930 at the age of 27 years and measuring 10 ft. 84 in. or 3257 mm. The measurements of the successive heights of this elephant, abundantly fed on a varied diet, indicate a growth at the shoulder of 2007 mm. or 6 ft. 7 in., in twenty-four years elapsing between October, 1906, and January, 1930, namely, an increase from 1250 mm. (4 ft. 14 in.) to 3257 mm. (10 ft. 8}in.), or an annual growth of 83.6 mm. (34in.). The rate of growth is very rapid until the fifth year, then it slows down gradually to 2 in., and finally to in. a year, namely, from January, 1929 (10 ft. 74 in.) to January, 1930 (10 ft. 84 in.). Fig. 1054. Remarkable cave paintings recently discovered in South Africa representing still living species of the white rhinoceros and African elephant. Observe especially the extremely accurate profile of the small-tusked female elephant, herewith reproduced one thirty-seventh natural size; also the correct proportions of the long-headed white rhinoceros reproduced to the same one thirty-seventh scale; both by courtesy of the London Illustrated News. In the great area north of the dotted line (Fig. 1055), comprising the Atlas Mts., the desert of the Sahara, and the more or less mountainous and formerly forested regions bordering the Mediterranean Sea, other sub- species of the African elephant formerly ranged in large numbers, as recorded by Pomel and as summarized by THE LOXODONTIN:: CLASSIFICATION AND HISTORY OF DISCOVERY 1195 Sir Harry Johnston from various classical historians. Pomel (1895) also records ‘Hlephas africanus’ in the Upper Pleistocene of Oran, northern Africa. Fig. 1055. Distribution of the existing African elephant (Loxodonta); location of the proposed species and subspecies as indicated by various authors. The four known type localities are marked by a circle. The northern limit of distribution has been indicated by an interrupted line. 1. L. africana Blumenbach. Range unknown, probably South Africa. L. africana albertensis Lydekker. Lake Albert region. 2 3. L. capensis G. Cuvier. Upper Orange River District. 4. L. africana cavendishi Lydekker. Galla Country. 5 L. africana cottoni Lydekker. Congo. 6. L. capensis cyclotis Matschie. Southern Cameroon. 7. L. africana fransseni Schouteden. M’Paa near Bongo, region of Lake Leopold IT. 8. L. africana knochenhaueri Matschie. Barikiwa, Tanganyika Territory. \ 9. L. africana orleansi Lydekker. North Somaliland. a : y 10. L. africana oxyotis Matschie. Upper Atbara, Sudan. ran Got Bi f ll. L. africana peeli Lydekker. Aberdare Mts., Kenya Colony. 12. L. africana pumilio Noack. French Congo. 13. L. africana rothschildi Lydekker. French Sudan. 14. L. africana selousi Lydekker. Mashonaland, southern Rhodesia. 15. L. africana torotis Lydekker. Mossel Bay, western Cape Colony. 16. L. africana mocambica Frade. Maputo, Mozambique. 17. L. africana zukowskyi (Strand in Zukowsky), Kaoko District, southwest Africa. 18. L. africana angolensis Frade. Cunene, Angola. Meridian _0 of Greenwich From Sir Harry Johnston’s “A History of the Colonization of Africa by Alien Races,” 1905, pp.6, 7: “This Punic explorer [Hanno the Carthaginian] started from Carthage some time in the sixth century before Christ (perhaps about 520 B.C.) ... on a voyage of discovery mainly. . . . In the account given of the journey it is stated that after passing the Straits of Hercules, and stopping at the site of the modern Sebu, they rounded Cape Cantin and came to a marsh in which a large number of elephants were disporting themselves [Footnote: ‘This is an interesting observation. Not only does the statement repeatedly occur in the writings of ancient Greek and Roman geographers that the African elephant was found wild in Mauretania in these times, but this animal is pictured in the remarkable rock sculptures in the Sus country in the extreme south of Morocco, and in the Roman mosaics and frescos found in the interior of Tunis, and now to be seen at the Bardo Museum near Tunis. (See for this the travels of the Moroccan Jewish Rabbi, Mordokhai.)’] . .. The Carthaginians do not seem to have tamed the indigenous African elephant (which was certainly still found in Mauretania), but they introduced and used the Indian elephant.” From Sir Harry Johnston’s “The Opening Up of Africa,” 1911, pp. 103-105: “Jugurtha [King of Numidia (134 B.C.)] adopted a plan of fighting learnt from the Carthaginian armies. He had war elephants of the African species, which he placed in the van of his attack; but somehow they did not make much impression on the dogged Roman infantry. After the Roman conquest the African elephant disappears from the annals of North Africa and, no doubt, became extinct everywhere north of the Sahara except in Morocco, where—in the country near the High Atlas—it seems to have lingered till the arrival of the Arabs. The camel had been introduced into North Africa from Egypt about 200 B.C., and was rapidly adopted by the nomad races of Mauretania as an animal very useful in war... . About 45 A.D. another general of British fame, Suetonius Paulinus, marched up the valley of the Muluya river in Morocco and reached the High Atlas range. He ascended these mountains to the snows, and descended to the southern side of them into the valley of the Gir stream, and gave a vivid de- scription of the burnt-looking rocks of the desert and the swarms of elephants in the Atlas forests.” 1196 OSBORN: THE PROBOSCIDEA [The question having arisen as to the existence in the same forest of the South Cameroon elephant and the “pygmy” elephant, recourse was had to the article of Dr. Glover M. Allen on the “Zoological Results of the George Vanderbilt African Expedition of 1934. ... The Forest Elephant of Africa” (Proc. Acad. Nat. Sci. Phila., LXXXVIII, 1936, pp. 15-44), in which he comes to the following conclusion. The African elephants are repre- sented by two general types—the large, more or less typical Bush elephant of the eastern and southern plains, and the smaller Forest elephant of the Congo forests. The former is Loxodonta africana and its several (?)sub- species or (?)varieties; the latter is L. cyclotis or L. africana cyclotis, type from the South Cameroons (syn. L. pumilio, ete.). He also provisionally relegates both cottoni and fransseni of the Congo region to the synonymy of cyclotis, giving his reasons therefor on pages 21-24. Furthermore he regards the term “pygmy” for the Forest LOXODONTA AFRICANA PUMILIO 2032mMm.,6’8” W. AFRICA LOXODONTA AFRICANA OXYOTIS 3568MM..11/8y2/ CENTRAL AFRICA Fig. 1056. Restorations by Margret Flinsch Buba, under the direction of Henry Fairfield Osborn. One one-hundredth natural size. The tusks of Loxodonta africana oxyotis were restored after the longest (Fig. 1062) and heaviest (Fig. 1065) pairs of tusks recorded. See caption to figure 1093, this chapter. Elephant as a misnomer—it is a small variety or a small species if one wishes to so designate it, which may reach a shoulder height of 91; feet as compared with perhaps 11% feet in the largest Bush Elephants. Thus there is no tremendous disparity in size between the two. In the closing paragraph (p. 41) of his article, Doctor Allen remarks as follows: ‘Very probably the Forest Elephant represents more nearly the older original stock from which the larger Bush Elephants have been derived. The evolutionary development of the latter has doubtless taken place slowly and over a long period of time with intergradation and interbreeding of long duration before they became sufficiently differentiated to maintain complete segregation, in the absence of physical barriers. . . For this reason, and because of the slight but apparently constant differences in carriage, form of ears, and size, the Forest Klephant is given tentatively the rank of a separate species. Nor is there any ground as yet for believing that it breaks up further into geographical races within its rather limited range in the rain forest.’’—Editor.] THE LOXODONTIN#: LOXODONTA 1197 2. SYSTEMATIC DESCRIPTION OF SPECIES OF LOXODONTA Loxodonta africana Blumenbach, 1797 Figures 794, 805, 806, 810-814, 816, 893, 908, 912, 992, 995, 1013, 1031, 1040-1045, 1049-1054, 1056-1065, 1067, 1081, 1093, 1107-1109, 1112, 1120, 1167, 1190-1192, 1226, 1234, Pl. xxi Elephas africanus Blumenbach, 1797. Syn.: Hlephas capensis Cuvier, 1798: Elephas priscus Goldfuss, 1821. Type locality: Probably Cape Colony, South Africa. Eighteen living or recently extinct subspecies described from various parts of Africa south of the equator. Extinct forms of the true ‘Elephas africanus’ described by Pomel (1895) from the Upper Pleistocene of northern Africa. Range of the species, formerly abundant, the North African coast and Atlas Mountains in the time of Hannibal. See caption to figure 1055. Elephas africanus, 1797. Blumenbach’s paper (1797.1), in which the name Hlephas africanus was first used, was accessible to the present author by photostat only, through the courtesy of the British Museum. The chief references are as follows: 1. (Matschie, letter, 1921): “Not in the edition of the ‘Handbuch der Natur- geschichte’ of 1779 [First Edition] but in the Fifth Edition (1797); in ‘Abbild. Naturhist. Gegenstiinde,’ Heft 2, No. 19, fig. C, the name is also used. . . . The arrangement of the lamelle would indicate its Cape origin. It can only be a question of the Cape Colony and the Congo, perhaps the French Congo. The confluence of the anterior lamelle is peculiar.’ 2. (Andrews, C. W., and Bather, F. A., June 6, 1922): ‘Handbuch der Naturgeschichte.”’ Fifth Edition. Elephas asiaticus, p. 124. EHlephas africanus, p. 125. Mammontovaiakost, p. 708. 3. (Sherborn, ‘Index Animali- um,” 1902): Sherborn lists HZ. africanus in the Fifth Edition of the “Handbuch,” but reference is not made to a figure in this edition. Typr Mo.uar, R. Me.—Blumenbach’s type figure (Fig. 1057), probably a second right inferior molar, r.Ms, serves to characterize clearly the species with very prominent lozenge-shaped ridge-plates to which also the generic name Loxodonta applies so aptly; this was the feature which commanded the attention of F. Cuvier in applying the descriptive generic term ‘Loxodonte.’ TypPE Locauity.—Matschie (1900, p. 190) states that no exact locality is known for the type of H. africanus Blumenbach. It is probable that the type came from the Cape region of South Africa, be- cause at the time the Central African elephants were inaccessible to European naturalists, whereas the South African mammal fauna was comparatively well known. Consequently it is probable that the type of Hlephas africanus Blumenbach came from the same region as the type of Hlephas capensis Cuvier (see note under FE. capensis below). In 1797 Blumenbach clearly defined these teeth, as in the legend of his type (Fig. 1057). In 1846 Owen summed up the distinction between these teeth and those of the Indian elephant (Owen, 1846, pp. 230-232—see Fig. 1043 of the present Memoir) : “Thus in the African Elephant, (fig. 88,) in which the lozenge- shaped plates are always much fewer and thicker than the flattened ones in the Asiatic species, the variation which can be detected in any number of the grinders of the same size is very slight. . . . In the molars of the Asiatic Elephant, (fig. 89,) which, besides the difference in the shape of the plates, have always thinner and more numerous plates than those of the African species, a greater amount of variation in both these characters obtains; . and the like caution is still more requisite in the comparison of the molars of the Mammoth (Hlephas primigenius), which, having normally more numerous and thinner plates than in the existing Asiatic Elephant, present a much greater range of variety.” Elephas capensis Cuvier, 1798, “Tableau Elémentaire de VHistoire Naturelle des Animaux,” p. 149. TYPr.— (Matschie, letter, 1921): ‘‘Cuvier had at his disposal one skeleton from the Senegal and one skull from the Cape [Cap de Bonne- Espérance, Cuvier, Mém. Inst. (de France) National des Sciences et Arts, sometimes called the Académie des Sciences, Vol. II, année 7, (1799)].” Figure.—Cuvier, 1799, Pls. 1, tv, fig. 2. Typx Locatrry.—Cape of Good Hope, South Africa. Tur Expanpina ‘Loxopont Sinus’ or THE AFRICAN Mouar Crown (LoXopoNTA AFRICANA) TENE TI EL ee =e = =a Fig. 1057. Type r.Mz unworn crown of Elephas africanus eam Lie 1797. After Blumenbach, 1797.2, No.19: “C. vom Africanischen. . : Nur bilden jene beym Elephas asiaticus geschliingelte an beiden Ender en paar- weis zusammenlaufende Linien; hingegen beym africanus rautenformige Leisten. Diese Zihne der beiderlei Elephanten sind nach Originalen im hiesigen academischen Museum gezeichnet.”’ [According to Pohle (see Dr. Glover Allen, 1936, p. 16) the present location of the original tooth of Elephas africanus is unknown: “The molar tooth on which Blumenbach founded his Elephas africanus is not now known to be in existence, although Pohle (1926) searched for it in recent years among the collections of the Zoological Museum and of the Anatomical Institute of the University at Gottingen.””—Editor.] Fig. 1058. Type of Elephas priscus Goldfuss, 1821, Pl. xuiv, one-half natural size. Type of a worn molar tooth said to have been found in the neighborhood of Cologne described by the author as follows: (op. cit., p. 485) “Beschreibung eines fossilen Backenzahns vom afrikanischen Elephanten. . . . Daher verdient ein unbezweifelt fossiler Backenzahn, dessen Rhomben auf der Kaufliche denen des afrikanischen Elephanten entsprechen, welchen das Museum der hiesigen Universitit kiirzlich als ein schitzbares Geschenk emp- fing, die Aufmerksamkeit der Naturforscher . . . und es ist daher zu vermuthen, dass auch dieses seltene Stiick in der Gegend von Céln gefunden worden sey.” 1198 (1) According to the above note, the type of Blephas capensis is probably a skull from the Cape of Good Hope. Lydekker (1907.1, p. 384) states that Hlephas capensis may “‘really be in- separable from Blumenbach’s HE. africanus typicus |= Loxodonta africana africana], based on teeth from a locality unknown.” (2) Matschie (1900, p. 190) states that if it were possible to prove that the elephant in different parts of Africa shows certain dis- tinctive characteristics, i.e., occurs as different subspecies, one must not readily synonymize EF. capensis Cuvier with EH. africanus Blumenbach; it would then be necessary to distinguish EH. capensis as the elephant of the Orange River District and E. africanus as some other subspecies.”” (3) Osborn, 1924: In the list, as fully cited above, Elephas capensis appears as follows: “Hlephas capensis G. Cuvier, 1798, p. 149. Type loc.: ?Upper Orange River, eastern S. Africa. Type fig.: skull, in Cuvier, 1799, pp. 1-22. Type probably in Museum, Paris = Loxodonta africana capensis.” Elephas priscus Goldfuss, 1821, 1823. The same lozenge- shaped ridge-plates are observed in the type of Elephas priscus Goldfuss, 1821. This type undoubtedly belongs to the living species of Loxodonta africana, but it misled Goldfuss into the im- pression that he was describing a fossil tooth, as shown in the legend to figure 1058. Cuvier considered the Goldfuss type as belonging to a recent African elephant. Falconer remarks (1868, Vol. I, p. 95): “In the autumn of 1847 I had an opportunity of examining the specimens above referred to, in company with Dr. Goldfuss. . . the fracture and texture of the ivory yielded the .. . appearance characteristic of recent teeth, and conveyed to my mind a corresponding impression that the molar was probably of modern origin.’ Nevertheless Falconer employed the name Elephas (Loxod.) priscus for undoubted Pleistocene fossil teeth from Gray’s Thurrock and elsewhere. Subsequently (cf. Leith Adams, 1877-1881, pp. 1, 2) Faleoner abandoned the name Elephas priscus Goldfuss, and it is now considered (fide Pohlig) as a synonym of Loxodonta africana. Pomel, 1895, p. 20, revives the name, after a very careful re- view: Elephas africanus priscus, Goldfuss. He says that the teeth of the africanus priscus type are found in Europe and certainly in northern Africa. In Plate 1, figure 1, he refers the latter, from the mountains of Oued Bourkika, to 2. africanus. CONSTANT RIDGE FORMULA OF LOXODONTA AFRICANA AS OBSERVED BY VARIOUS AUTHORS De Blainville (1839-1864, fide Falconer, 1865, p. 264): Dp 2 4 Dp3+ Dp4+M1¢M2"—2M3>,-33- Owen (“Odontography,” 1840-1845, p. 638): Dp 2 + Dp 3 3 Dp42M12M2,,M3 TEE Faleoner (1865, p. 265) on De Blainville’s and Owen’s materials, “excluding the two talons’: Dp 2 3 Dp 3 § Dp 4? M1? M2355 M3. Falconer (1865, p. 265) observes that a left M*, eleven inches in length, from Cape Colony [region typical of Loxodonta africana capensis Cuyv.], exhibits thirteen plates, ‘‘7.e., eleven principal ridges, besides front and back talons,” namely, M 3 2-*~*. FaALconer, 1857—1868.—Although Falconer did not adopt the term Loxodon as of full generic value, he referred to these animals as Loxodon, a subgenus of EHlephas. In his ‘Paleontological OSBORN: THE PROBOSCIDEA , Memoirs” of 1868, and in the plates accompanying that work, after most precise study, he gave a description of the lower ridge formula, including the half ridge-crests or “talons,” as follows: (Faleoner, ‘Paleontological Memoirs,” Vol. I, 1868, pp. 422, 440, 441, Plates 1, x11.a, XIv): Elephas africanus. Plate n, fig. 4a, Me with 944 ridges; fig. 4b, Me with 9 ridges. Plate xr, fig. 8, lower Jaw with Dpu, M,; My, with %~-7-s ridges. Plate xrv, fig. 4, lower jaw, r.Dps with 6% ridges; figs. 5, 5a, r.Me2 with 8 ridges. From the above may be deduced Falconer’s lower ridge-plate formula of referred Elephas africanus, namely: Dp 3 Mi as M 2 Bi-9-0'% [M 3 qworalk 6% Lerrx Apams, 1879-1881, p. 48.—Leith Adams gives the ridge formula of Loxodonta africana as follows: Dp 2 2=2 Dp 3 2=* Dp 4 42 M12M 2 = M3 yous: Feurx, 1912, p. 17.—Felix gives the following ridge formula of Loxodonta africana: Dp2= Dp3. Dp4— Ma Nb2) = Ne Fig. 1059. Photograph of the dome-shaped skull and of the tusks (Amer. Mus. Dept. Mam. 21889) of adult male Lozodonta africana peeli ref., reduced to about one-twentieth natural size. Represents the cranium characteristic of elephants of eastern equatorial Africa. Specimen collected in 1911 by Mrs. Carl E. Akeley on southern slopes of Mt. Kenya. Tusks measure 8 ft. 54 in. and 8 ft. 9% in. respectively and weigh 112 and 115 lbs. each. Now exhibited in the Age of Man Hall, American Museum, beside the skull of Hesperolozodon antiquus italicus. Compare figure 1107 of the same crania. THE LOXODONTINA: LOXODONTA Constant Ripce Formuia.—According to the above, the ridge formula of Loxodonta africana is constant and very conserva- tive, since the M 8 ridge formula of the living African elephant closely corresponds with that of the primitive Upper Pliocene Archidiskodon planifrons. The constancy of the ridge formula in the two existing species of elephants, Loxodonta africana and Elephas indicus, is a very im- portant fact in its bearing on the probable constancy of ridge formule in extinct species of elephants of a single geologic time period, for example, in Mammonteus primigenius of Upper Pleisto- cene time, as observed both by Falconer and Osborn. CRANIAL CHARACTERS AND TUSKS OF LOXODONTA AFRICANA (Continued from Chapter XV, pp. 915-927) CraniAu Axts.—The profound characters of the platycephalic cranial axis of Loxodonta africana in comparison with the hypsi- cephalic EHlephas indicus and Archidiskodon imperator are clearly shown in figure 805, also in figures 806, 812, 811,810, and 813. In Chapter XV, the introductory section on the cranium of the Elephantoidea, these profound differences between the cranium of Loxodonta and the crania of other genera of elephants are much more apparent than the superficial differences which are displayed in the accompanying figures (Figs. 1041, 1059, 1060, 1061, 1062). The summary of the sectional or axial characters of the Loxo- donta africana skull is as follows: (1) Occipital plane, perpendicular to basis cranii, extremely long; (2) frontal plane extremely short, convex anteroposteriorly and transversely; (3) nasals broad and rounded; (4) occipitohorizontal section extremely broad with deep pit for ligamentum nuche; (5) cranial dome, i. e., occipito- horizontal contour, uniformly rounded; (6) cranium more platy- cephalic and brachycephalic and less hypsicephalic and bathy- cephalicthan that of Mammonteus, of Archidiskodon, or of Parelephas. These primitive characters of low, rounded cranial dome and of relatively mesocephalic profile contour are best displayed in figure 1059 and in figure 1060. The latter illustrates the classic mid- dle-aged skull of ‘Jumbo,’ a Sudanese subspecies. We observe: (1) That in the fully adult skull the dome is continuously rounded from the occipital condyles to the broad extremities of the nasals, presenting the widest contrast to the profiles of Archidiskodon, of Parelephas, of Mammonteus, and of Elephas; (2) that while actually brachycephalic, the cranium of Loxodonta is also compara- tively mesocephalic or elongate (shown in the relatively long mandi- bular ramus, Fig. 1060), as compared with the deeply depressed mandibular ramus of Elephas indicus, or with the extremely bathy- cephalic and abbreviate mandibular ramus of Mammonteus primi- genius; (3) that since both superior and inferior molars, M*, Ms, are shorter and less hypsodont, the maxillary and mandibular dental cavities are much less deep, thus accounting for the less hypsicephalic proportions of this part of the skull; (4) the superior aspect of the mandibular rami (Fig. 1060) also displays the relative prominence and horizontal distinction of the rostrum, again presenting a very wide contrast to the deep, hypsicephalic rostrum of the Mammontine and of the Elephantine; (5) the superior or frontal aspect of the cranium is beautifully displayed in figure 1061A, showing (a) the short frontal bones, (b) the massive orbital prominences, (¢) the broad narial openings, (d) the widely separate maxillo-premaxillary sockets for the enormous incisive tusks, 1199 similar to those in Palzxoloxodon namadicus and Hesperoloxodon antiquus; (6) the palatal aspect of the Loxodonta cranium should be examined side by side with the palatal aspect of the Hlephas M\, Loxodonta africana (Jumbo) ’ Amer. Mus. 3283 | 1/8 nat, size AFRICAN ELEPHANT CRANIUM AT THE AGE OF TWENTY-FOUR, M?, Mo 1n UsE Fig. 1060. Middle-aged skull of the Sudanese or Abyssinian subspecies, Loxodonta africana oxyotis (Amer. Mus. Dept. Mam. 3283) with nine-plated M2 in use and an unworn M3 still embedded in the jaw. One-eighth natural Observe the angular vertex. For full details, see legend to figure 1061. This is the famous individual, named ‘‘Jumbo,” purchased in 1883 by P. T. Barnum, the American showman, from the London Zoological Society. “Jumbo” died Sept. 15, 1885; skin at Tufts College, Medford, Massachusetts; skeleton in the American Museum. size. 1200 OSBORN: THE PROBOSCIDEA indicus cranium (Fig. 800). It will be observed that the Lorodonta skull (1061 B) is relatively broader, more brachycephalic in all its dimensions. Compare the measurements from the occipital condyles to the extremities of the premaxillaries in Loxrodonta africana (Fig. 1061 B) with the measurements in Elephas indicus (Fig. 800); compare also the zygomatic breadth of Loxodonta W. K. Grecory on E. arricanus (JUMBO SKULL).—The re- markable foreshortening (cyrtocephaly) and deepening (bathy- cephaly) of the African elephant cranium, while less extreme than that of other elephants, is nevertheless far more advanced than in any species of mastodont. (1) Note extreme compression of the hinder part of the palate, correlated with the very backward exten- Loxodonta africana (Jumbo) Amer, Mus. 3283 1/8 nat. size AFRICAN ELEPHANT CRANIUM AT THE AGE OF TWENTY-FOUR. SAME CRANIUM AS IN F'raurE 1060 Fig. 1061. Superior and palatal views of cranium of “Jumbo’’ (Loxodonta africana oxyotis), Amer. Mus. Dept. Mam. 3283; same skull as that shown in figure 1045. One-eighth natural size. Compare B with similar views of Elephas indicus (Fig. 800). A middle-aged male (twenty-four years) taken in 1861 at the Setit River, northwestern Abyssinia; famous in history as having lived three years in the Jardin des Plantes, Paris, and then transferred to the London Zoological Society, which, doubtful of his temper, accepted in 1882 an offer of $10,000 made by the celebrated circus proprietor, P. T. Barnum, of the United States. Jumbo is supposed to have been twenty-four years of age at the time of his death. africana with the zygomatic breadth of Elephas indicus. ‘The comparison shows that what the Loxodonta skull loses in length it gains in breadth, i.e., in brachycephaly, and in platycephaly. It also shows that while relatively narrow and long, the skull of Elephas indicus gains greatly in vertical depth; it is more hyp- sicephalic, more acrocephalic. sion and great vertical depth [bathycephaly] of the maxillary alveoli containing M?, M*. (2) Note anterior position of suture separating malar from maxillary portion of zygoma. In the primi- tive mastodont Trilophodon the malar extends forward over the maxillary to form the postorbital process; very primitive. (3) Note extremely robust [brachycephaly| buccal portion of zygoma. THE LOXODONTINA: LOXODONTA 1201 (4) Note confluence of anterior palatine canals and wide divergence of incisive alveoli. (5) Note reduction in size of auditory bulla which is squeezed down close against the basis cranii; the anterior prolongation under the eustachian tube is less prominent. (6) Note reduction in size and in-sinking of foramen for carotid artery in the bulla. EXTERNAL CHARACTERS AND TUSKS OF LOXODONTA AFRICANA (Fies. 1062, 1052 A, B, C).—The external characters of the male The different views of the head of this full-grown male ele- phant, as displayed in figures 1063, 1052 A,B,C, all show the same depressed, relatively flattened or platycephalic profile, correspond- ing with the short space between the eyes and the top of the head, due to the lesser development of the frontal sinuses, as compared with Hlephas indicus or with the extreme acrocephaly and hypsi- cephaly of Archidiskodon and of Mammonteus. Mate AND FrMALE Cranta.—The wide disparity between Vig. 1062. (Left). Tusks of Loxodonta africana oxyotis in the New York Zoological Society collection, one twenty-fourth natural size. ComMPARISON OF TUSKS OF LOXODONTA AND OF MAMMONTEUS Right tusk 11 ft. 5!%sin. on outer curve; left tusk 11 ft. Circumference of right tusk 18}¢in.; of left tusk 18in. Weight of the pair 293 lbs. The locality record of these tusks is East Africa; it is reported that they were recently owned by King Menelek of Abyssinia who presented them to a European political officer, eventually they were offered for sale in London, were purchased by Rowland Ward, and finally were presented to the New York Zoological Society by Mr. Charles T. Barney. (Right). shaped arrangement in Lozxodonta. and female and young African elephant are beautifully displayed in the great group (Fig. 1052 A) which includes two subspecies of Loxodonta africana (peeli and albertensis) collected and mounted by Carl E. Akeley between the years 1909 and 1922 for the African Hall of the American Museum of Natural History. The bull is a typical example of the male African elephant, as carefully ob- served and measured by Mr. Akeley in the field; it attains a shoulder height of 10 ft. 8 in. [as mounted, height of head 11 ft. 2/4 in.]; the expanse of the ears is 10 ft. 24 in. Tusks of Mammonteus primigenius from Alaska, showing the circular curvature and crossing in Mammonteus as compared with the lyre- From original photograph brought from Alaska. One twenty-fourth natural size. male and female crania and tusks in species of Loxodonta is shown in the accompanying figure (Fig. 1063) of two mounted heads from east central Africa. The male is the great bull collected and mounted by Carl I. Akeley; the geographic locality is northern Uganda; the specific reference is Loxodonta africana albertensis Lydekker. The female is a specimen collected by Paul Rainey north of Mt. Kenya; the species is Lorodonta africana peeli. These two heads display the following generic and sexual char- acters: (1) Uniform elongation or mesocephaly of the cranium, as 1202 compared with that of Elephas indicus; (2) the male and female crania are substantially of the same elongate, mesocephalic propor- tions; (3) the gigantic ears are relatively of the same size; (4) there is a great disparity in size of the female as compared with the male cranium; (5) the female tusks are extremely slender as compared with the male tusks, yet they constitute formidable weapons for the defense of the young; (6) this sexual disparity of size is equally apparent in extinct species (e.g., Hesperoloxodon antiquus, ef. Fig. 1090), and it is important to note that there should be observed a very marked difference in the development of the Vig. 1063. OSBORN: THE PROBOSCIDEA fronto-occipital crest in the male as compared with the female cranium. (7) Similar disparity, between male and female tusks, to that shown in figures 1063, 1090, and 1106, is displayed in all the known members of the Elephantide or true elephants. The disparity in the known members of the Mastodontide is somewhat less extreme between the male and female tusks. PGTAL SKELETON OF LOXODONTA AFRICANA (?)COTTONI Ears, 1926-1929.—The foetal cranium, jaw, and milk denti- tion of Loxodonta africana (?)cottoni (Fig. 1064), as fully described Female Heap or ApuLT MALE AND FEMALE LOXODONTA AFRICANA Compare figure 1090, male and female tusks of Blephas |Hesperoloxodon| antiquus [germanicus?], after Pohlig, 1888, Taf. 1, figs. 1, la, 1b, 2, 2a. Full-grown male head of Loxodonta africana albertensis (Amer. Mus. Dept. Mam. 54085), from the Budongo Forest, east of Lake Albert, northern Uganda, as mounted by Carl E. Akeley in the American Museum, 1922. Full-grown female head of Loxodonta africana peeli collected north of Mt. Kenya by Paul Rainey and mounted under the direction of Carl E. Akeley. It is now in the private collection of Mr. Perey Madcira of Philadelphia. THE LOXODONTINA: LOXODONTA in Miss Nellie B. Hales’s Memoir of 1926-1929, should be compared with the juvenile stages of the cranium of Mastodon americanus (Chap. VI, Fig. 131B, Vol. I), also with the juvenile cranium of Elephas indicus (Chap. XV, pp. 915-918, Figs. 796, 797). The cranial contours and the elongated jaw, the occipital condyle on the same plane as the dentition, the budding incisors with delicate tips, the primitively plated crowns of the lower milk molars, Dp2s, contain no forecast of the tremendous transformation which this foetal cranium is destined to undergo into the typical African 1203 elephant skull, such as appears in figures 1060 and 1061. No trace of the permanent premolars is found, as might be expected in so young a specimen; the ridge formula, as shown in figure 1064, is: Dp 22 Dp 37 Dp 4% the posterior plates in Dp 4 not being calcified; this formula agrees with that of Owen, Flower, and Lydekker. AS -- finf.o. ¢ EE fen. rot.} b.ex.o. PRIMITIVE FORM OF THE LOXODONTINE CRANIUM AND DENTITION, AFTER HALES Pig. 1064. Foetal cranium, jaw, and milk dentition of Loxodonta africana (?)cottoni of the Belgian Congo, locality not given. After photographs kindly sent the present author by Miss Nellie B. Eales. Compare Eales on “The Anatomy of a Foetal African Elephant, Elephas africanus (Loxodonta africana),” Trans. Roy. Soc. Edinburgh, Vol. LIV, Pt. III (No. 11), 1926, PI. rx, fig. 29, Pl. x, figs. 32 and 34 on the “Anatomy of the Head,” etc.; see also Vol. LV, Pt. III (No. 25), 1928, on the ‘“Body Muscles,” and Vol. LVI, Pt. 1 (No. 11), 1929, on the ‘“‘Contents of the Thorax and Abdomen and the Skeleton.” In Vol. LVI, Pt. 1 (No. 11), 1929, of this invaluable Memoir, the author’s conclusions as to the phylogenetic relationships of Lorodonta are fully set forth. 1204 OSBORN: THE PROBOSCIDEA : bid | a Fig. 1065. Tusks of the African elephant, believed to be the heaviest (combined weight 461 Ibs.) in the world, shown in front of a typical Arab door at Zanzibar. From Kilimanjaro, East Africa, purchased in Zanzibar in 1900 and exhibited for some time by Tiffany & Company of New York. The length of the larger tusk is given as 10 ft. 1 in. on outer curve, circumference 23% in. at hollow end. One of these tusks is now in the British Museum (weight 226 lbs.). Reproduced after “Ivory and the Elephant in Art, in Archaeology, and in Science,” 1916, p. 411, by Dr. George I’. Kunz, through the courtesy of the Executors of his Estate. Loxodonta cornaliae Aradas, 1870 Figure 1066 From Catania, near the monastery of Santa Chiara, Sicily. Quaternary or Post-Pliocene. Elephas Cornaliae Aradas, 1870. “Sopra un Molare Elefantino Fossile Riferibile a Specie Distinta dalle Cono- sciute.” Atti Accad. Gioenia sci. nat. Catania, (3), IV, op. 233-235. Typr.—sSuperior molar of the right g side. Horizon AND Locatiry.—From Catania, near the monastery of Santa Chiara, Sicily. Quaternary or Post-Pliocene. TyprE Figure.—Op. cit., figs. 1, 2. Typrr Description.—(Aradas, op. cit., pp. 233-235): “Hsso 6 un molare vero, superiore, destro; ¢@ rotto in mezzo quasi e per lungo, talehé manca di poco meno della meta; manea pure del tallone anteriore e del posteri- ore, e conseguentemente della estrema lamina anteriore, non che della posteriore; le radici si conservano in ottimo stato, meno quelle che dovevano spettare alle lamine mancanti. Sulla sua superficie triturante, qualora stato fosse intiero, si avrebbero potuto contare otto a nove lamine. I dischi di logoramento sono romboidali, in forma di losanghe; le lamine leggermente increspate, non molto avvicinate. Le digitazioni della corona dal lato esteriore son quasi perpendicolari alla superficie triturante e rego- larmente arrotondite.”’ “Or, se fossero questi soltanto 1 caratteri che il molare in esame presenta, certamente potrebbe per la configu- razione dei dischi di logoramento, come si @ detto, rom- boidali, riferirsi all’ Elephas africanus di Blumenbach. Ma é da considerare, che gli angoli mediani delle losanghe son poco approssimati a quelli delle contigue, e che se la specie africana offre da 9 a 10 lamine in esercizio sopra 0,20 o 0, 24 di superficie triturante in lunghezza secondo Lartet, nel nostro molare, come si disse, non pit di 7 0 8 se ne possono contare sopra 0, ‘24 di superficie triturante.”” “QOltre a cid @ da notarsi, che la corona del molare, che deseriviamo, calcolandola nella sua totalitd, cioe, comprendendovi cid che manca, presenta tale proporzi- one tra la lunghezza e la larghezza, da non trovarsi in altri uguale, essendo, a meglio spiegarci, assai pil largo proporzionalmente degli altri molari conosciuti.” “Pure si potrebbe, non ostante le premesse consid- erazioni, e qualora non si volesse tener conto di altri saratteri del descritto molare, riconoscere in esso una certa affinita coll’Elephas africanus. Perd, confrontandolo coi molari riferiti a questa specie, ben descritti e figurati nella eccellente monografia degli Elefanti siciliani del prof. Gemmellaro e del Barone Anca, saremmo forzati a dire, o che il nostro spetti veramente all’africano e quelli dei sullodati autori appartenenti a specie distinta, oppure tutto al contrario, essendo che la configurazione dei loro THE LOXODONTIN/A: dischi di logoramento e le altre proporzioni fra loro apertamente differiscono.” “Ma tutte queste riflessioni debbon cedere e cadere a fronte di un carattere singolare, che il nostro molare appresenta, e che in alcun altro non si osserva, ed il quale basterebbe solo a far riguar- dare come una specie distinta l’Elefante cui esso appartenne. Le lamine sono tra loro separate da solehi lati e profondi, molto levigati, 1 quali a mo di semi-coniche escavazioni, vanno elevandosi man mano dagli orli esterni, laterali e ad archi interrotti della corona sino ai punti nei quali si pongono in avvicinamento gli angoli mediani delle lamine. assai meglio mostrare di qualsiasi descrizione, non sono da attribuirsi Questi solchi, che le figure possono a cause accidentali, per esser tutti regolarmente architettati, senza che si possa rilevare alcuna alterazione nella materia che la corona esternamente compone, né a dislocamento delle lamine, perché, oltre che questo sarebbe un fatto senza aleuno stento riconoscibile, gli abbassamenti sarebbero solo di un lato ed in modo di dare alle lamine una disposizione scalariforme. abbiamo detto, formano un carattere nuovo e molto singolare, a nostro modo di vedere, potrebbero significare il passaggio degli Questi solchi, che, come LOXODONTA 1205 Elefanti ai Mastodonti. Per noi il descritto molare dee appartenere ad una specie nuova, che, ove fosse per tale riconosciuta dai dotti che saranno per giudicare la nostra opinione, vorremmo portasse il nome dell’egregio Prof. Cay. Emilio Cornalia, che sui vertebrati ha fatto studii positivi ed utili alla scienza, chiamandolo Elephas Cornaliae.”’ Tyre or LoxopDONTA CORNALIAE Fig. 1066. Type superior molar of the right side of Hlephas Cornaliae Aradas, 1870, fig. 2 of plate, two-thirds natural size. From Catania, Sicily. Fig. 1067. Colony. three feet. Young Addobush Elephant (Loxodonta) from Cape Probably six months old, with a height at shoulder of about Reproduced through the courtesy of Mr. Henry C. Raven, who took the photograph in 1919. PALAEOLOXODON NAMADICUS: 3736MM., 12’3%e INDIA J “ d/ x HESPEROLOXODON ANTIQUUS ITALICUS: 4068MM., 13’ 4% e P MNAIDRIENSIS: I900MM., 62% e P. MELITENSIS: 1400Mm., 4746 ROME P. FALCONERI: 900MM.,2/11%%e = \ ( ) | [i " ? a 3 HESPEROLOXODON HESPEROLOXODON ANTIQUUS ITALICUS: 3905MM., 12’934”e ANTIQUUS PLATYRHYNCHUS: 3828MM., 12'6%"e PIGNATARO PINDAL N. SPAIN / ee es (4 f re pa - \ oN ~ 4 Lae ay \\ 7) A (| ’ \A / 4 (> vf & ! Y \ ae f a0 soe 4 “Qa 3. HESPEROLOXODON ANTIQUUS: 3934INM., 12°107% HESPEROLOXODON ANTIQUUS PLATYRHYNCHUS: 3828mM,12/6%4e UPNOR N. SPAIN Fic. 1068. Restorations sy Mareret liinscu Bursa (1931), UNDER THE DIRECTION OF HENRY FAIRFIELD OSBORN, OF SPECIES OF H&rsPEROLOXODON OF Europe AND OF PAL-ZOLOXODON OF INDIA AND OF THE MEDITERRANBAN ISLANDS, IN COMPARISON WITH A DRAWING OF HESPEROLOXODON BY THE Cave MEN oF NORTHERN SPAIN. ONE ONE-HUNDREDTH NATURAL SIZE. 1206 III. EURASIATIC SPECIES OF PALZOLOXODON AND HESPEROLOXODON SUPERFAMILY: KLEPHANTOIDEA Osborn, 1921 FAMILY: ELEPHANTID& Gray, 1821 SUBFAMILY: LOxoDONTIN® Osborn, 1918 Genus: PALZOLOXODON Matsumoto, 1924 Original reference: Matsumoto, Journ. Geol. Soc. Tokyo, 1924, XXXI, No. 371, pp. 257, 260 (Matsumoto, 1924.2). Compare Elephas namadicus Fale. and Caut., 1846. Genotypic species Elephas namadicus naumanni Makiyama, 1924. Syn.: Hlasmodon (preoc.) Falconer, 1847 (in part); Huelephas Fale., 1857 (in part). Sivalikia Osborn, 1924, and Pilgrimia Osborn, 1924 (in part). Subgenus Palzxolorodon =‘E. antiquus-namadicus group’ of Matsumoto (1924-1926). GENERIC CHARACTERS.—Ridge-plates of grinding teeth parallel, closely compressed, waving or plicate. ‘Loxodont sinus’ rudimentary or absent. Progressively narrow to broad superior grinding teeth, with numerous ridge-plates composed of thin, plicate enamel foldings; twelve ridge-plates in 10 em.; total number of ridge-plates in lower grinding series (Dp 4—M 38) fifty-one. Ridge-plate formula progressive from: 15 19 P. melitensis, 7,3, to P.atlanticus, M3 +5-,;4, to P. namadicus +2, to P. namadicus naumanni, +. Correlated with more hypsodont and numerous ridge-plated grinding teeth, cranium more hypsi- cephalic and bathycephalic than that of Loxodonta; broad, rugose parietofrontal crest (P. namadicus) overlapping the forehead, adapted to muscular insertion of the very broad proboscis. Premaxillaries (Fig. 1069) broadening inferiorly for the insertion of widely divergent superior incisive tusks. Tusks relatively straight, shightly upcurved and incurved toward the extremities. The genotype of Palzoloxodon Matsumoto is ‘Elephas namadicus nawmanni’ Makiyama, 1924. The geno- typic species of Szvalikia Osborn is ‘Elephas namadicus’ Falconer and Cautley, as fully explained in the historical introduction to the present chapter; it is also shown that the generic name Sivalikia Osborn (Dee. 20, 1924) is technically preoccupied by the subgeneric name Palxolorodon Matsumoto (Sept. 20, 1924). In 1924 (MS8.) Osborn observed: “The gigantic elephants typified by the Hlephas namadicus of Falconer are believed by Osborn to constitute a distinct generic phylum, to which the name S7valzkia is applied in honor of Dr. Hugh Falconer’s great work on the Siwalik fauna. They agree with Loxodonta in certain characters of the cranium, especially in the low occiput and the distally broadened premaxillaries; they differ from Loxodonta in the com- pressed ridge-plates lacking the ‘loxodont sinus,’ also in the breadth of the grinding teeth. This generic phylum may spring from the giant species of North Africa, e.g., Loxodonta atlantica |Palzoloxodon atlanticus], but it is certainly distinct from the typical Loxodonta africana group.” Historic Periop.—It is not impossible that some elephants of the Palxoloxodon type survived into early historic times, but it is an open question whether the elephants described from Mesopotamia were of the ancient ‘loxodontine’ or of the modern ‘elephantine’ type, probably the latter; drawings and inscriptions will probably be found some day which will determine these relations. (Letter, A. H. Godbey, July 19, 1927): Assyrian Royal Inscriptions mention elephant hunting in northern Syria, Meso- potamia, etc. But no reliefs, so far, portray the elephant. Have you any information as to the species that once existed there?— any skeletal evidence? . . . Also, Josephus mentions monster bones exhibited as ‘giant’s bones’ in Palestine. Benjamin of Tudela, A.D. 1173, was shown a ‘giant’s rib’ at Damascus, 9 cubits long. (Letter, Godbey, April 5, 1928): ‘Hunting elephants on upper Euphrates is a notable achievement of Assyrian kings. The only portrayal so far is on Black Obelisk Shalmaneser, small ears like Indian type, tusks project up from lower jaw; like a boar’s, or ‘Babyroussa.’ It may be an artist’s blunder.— The ideogram for elephant is ‘Big-Horned Bison’ (‘Mountain Ox’), compare Bos Lucas, from Lucania, the ‘boot-toe’ of Italy. 1207 Cindi) \ Bi S. NAMADICA Ref. Falc., 1847, Pi, XXIV A, Fig. 4¢ S&S. NAMADICA Type S. NAMADICA Type Felc., 1847, Pi. XI1A Falc., 1847, Pl. XII B, Fig. 1 (rev.) S. NAMADICA Ret. Feic., 1847, Pil. XXIVA, Fig 4 a Ss. NAMADICA Ref S. NAMADICA Ref. Felc. 1847, Pi. XLII, Fig. XXI Felc. 1847, Pt. XLIV, Fig. XXII (rev. {A We at int SS '' Ny) j yf \ id a Myo ’ v S. ANTIQUA (NAMADICA) Ref. Pilgrim, 1905, Vol. XXX, Pl 1 Piigrim, 1905, Vol. XXXII, PI. 12 S. ANTIQUA (NAMADICA) Ref. S. ANTIQUA, Ref. Pignataro, Itely. From photo. S. ANTIQUA PLATYRHYNCHA, Type Greells, 1897, Pi. XVIII, Fig © 98 S. ANTIQUA NESTII Type Pohlig, 1891, p. 350, Fig. 109 S. ANTIQUA Ref. Weithofer, 1890, Taf. Il, Fig. 2 S. ANTIQUA NESTII Type Pohlig, 1891, p. 350, Fig. 109 All 1/20 nat. size S. (ANTIQUA) MELITAE Ret S. (ANTIQUA) MELITAE Ref, Pohlig, 1893, Tef. |, Fig. 1 Ponlig, 1893, Taf. |, Fig. 1« Fig. 1069. See explanatory legend on opposite page. 1208 THE LOXODONTIN#E: PALHOLOXODON 1209 As shown below (Chap. XX) in comments of Murray on the etymology of the word Elephas, signifying ivory, it would seem that the animal first became known from trade in its tusks originating in Africa, as mentioned by Homer, Hesiod, and Heroditus, whereas Aristotle treats only of Elephas indicus. From this it does not appear that the Mesopotamian elephants were known to the Greeks. CRANIAL CHARACTERS OF PALAOLOXODON NAMADICUS AND HESPEROLOXODON ANTIQUUS COMPARED WITH LOXODONTA AFRICANA.—In figure 1041 above the cranial characters of the recent Loxodonta africana are shown in comparison with those of the fossil Indian species Palxoloxodon namadicus and those of the European species Hesperoloxodon antiquus. The relatively low, brachycephalic, platycephalic, and mesocephalic form of the Loxodonta cranium, as compared with the hypsicephalic crania of Hlephas, of Parelephas, and of Mammonteus, is more clearly shown in the preceding figures (Chap. XV, pp. 915-926) relating to the comparative structure of the crania in these genera. In the present comparison (Figs. 1041, 1069) of the crania of Loxodonta africana, Palzoloxodon namadicus, and Hesperoloxodon antiquus, we are especially struck by the short and extremely broad rostrum and by other charac- ters, as follows: (1) Rostrum short and extremely broad, the premaxillaries diverging to the point where the tusks issue from the skull; (2) thus the bases of the tusks are very far apart instead of being close together as in Mam- monteus primigenius or relatively close as in HL. indicus; (3) whereas the premaxillary sockets are relatively of the same length in all three species, the divergence of the sockets in P. namadicus is about the same as in L. africana; (4) the premanillary sockets are relatively longer and diverge still more widely in H. antiquus; (5) the crania of both P. namadicus and H. antiquus are distingushed from the cranium of L. africana by the greater development of the fronto-occipital crest which in P. namadicus engulfs the frontal bones so that there is a very short space between the lower border of this crest and the extremities of the nasals and the narial openings; (6) the narial openings are extremely broad and shallow, they exhibit approximately the same hour-glass-shaped form in L. africana, P. namadicus, and H. antiquus; (7) the skull of P. namadicus, moreover, is relatively broader and more flattened or platycephalic than the skull of L. africana; (8) this broadening and flattening of the summit of the cranium reaches an extreme in the gigantic P. namadicus cranium. The phyletic affinities of these three types of crania, namely, Loxodonta africana, Palxoloxodon namadicus, and Hesperoloxodon antiquus, to each other are obvious, while there is a wide separation from the cranial type of Elephas and a still wider separation from the cranial type of Archidiskodon and of Mammonteus, which represent the opposite extreme of hypsicephaly. Fig. 1069. Comparison oF CRANIA OF PALHOLOXODON NAMADICUS, HESPEROLOXODON ANTIQUUS ITALICUS, H. ANT, AUSONIUS, AND H. ANT, PLATYRHYNCHUS All to the same one-twentieth scale. Compare figures 1041, 1121, 1096, 1105, 1106 [At the time this figure was prepared Professor Osborn was not aware that Matsumoto had preceded him by three months in the description of Palzoloxo- don, thereby rendering his own genus Sivalikia a synonym. Consequently the reader should disregard the letter “‘S” in this figure (=Sivalikia) and substitute Matsumoto’s genus Palzoloxodon and Osborn’s Hesperoloxodon (a subsequent determination for the antiquus group), as indicated in the following caption.— Editor.] (Upper row) Palzxoloxodon namadicus: One juvenile and two adult crania, front and side views. After Falconer and Cautley, 1846 [1847]. J (Second row) Hesperoloxodon antiquus italicus (left), excavated at Pignataro Interamna, Italy, about fifty miles north of Naples. Drawn after photograph of specimen as it lay in the quarry (Fig. 1096); subject to modification after restoration of the cranium (Fig. 1098 and especially Fig. 1106, an orthogonal drawing from the mount). (Right) Aged Palzoloxodon namadicus, after Pilgrim, 1905, front and side views. (Third row) Hesperoloxodon antiquus ausonius? (left), from the Val d’Arno of Italy, after Weithofer, 1890; H. antiquus platyrhynchus (middle), from San Isidro, Spain, after Graells, 1897; H. antiquus ausonius (right), from the Val d’Arno of Italy, after Pohlig, 1891, erroneously referred to Elephas (antiquus) Nestit. (Lower row) Palzoloxodon melitensis (left), from the island of Sicily, front and side views, after Pohlig, 1893; Hesperoloxodon antiquus ausonius (right), from the Val d’Arno of Italy, after Pohlig, 1891, erroneously referred to Elephas (antiquus) Nestii. 1210 OSBORN: THE PROBOSCIDEA GEOLOGIC AND GEOGRAPHIC DISTRIBUTION OF THE LOXODONTIN®.—Originally described by Falconer from deposits now determined as of the Upper Pleistocene of India (Palzoloxodon namadicus) and the Lower(?) Pleis- tocene of England (Hesperoloxodon antiquus), the geologic range has been extended from the relatively small Upper Pliocene! H. ausonius to the great ascending Upper Pleistocene mutations (i.e., H. antiquus in Germany = H. antiquus germanicus), also throughout western Europe, to the progressive Palwoloxodon namadicus of the Middle to Upper Pleistocene of India, to smaller subspecies or varieties of Palzoloxodon in Japan, described by Matsumoto, and in Java, described by Dubois, to the P. atlanticus of northern Africa and a long series of probably antecedent species in central and southern Africa, as well as to larger and smaller varieties in the Mediterranean Islands. The straight-tusked elephants of western and southern Europe have been treated, in succession to Falconer, by Leith Adams in Italy, by Pohlig, Weithofer, Soergel, and Berckhemer in Germany and Italy, by Forsyth Major and Depéret and Mayet in the Pliocene of Italy and of England, by Andrews and Forster Cooper in the description of the Upnor elephant (Hesperoloxodon antiquus) of England, and finally by Osborn in his description of Palzoloxodon |Hesperoloxodon| antiquus italicus of Italy. As the whole Pleistocene period is now estimated at approximately 1,000,000 years, the straight-tusked elephant Hesperoloxodon antiquus lived in western Europe for an enormously long period of time, it survived many climatic phases, it doubtless passed through many ascending mutations which will be determined by final mono- graphic comparison and description. First appearing in Upper Pliocene time, it survived three successive glaci- ations of northern Europe, but perished toward the close of the 3d Interglacial. Throughout this long period its companions in southwestern Europe were the Parelephas trogontherii, the hippopotami and the rhinoceroses (Rhinoceros etruscus and R. merckii). The provisional geologic succession and companionship of these four types may be presented in the table after Osborn and Reeds, 1922-1929 (Pl. xxrv). Osborn, 1929: Pilgrim (1905) first notes that Hlephas namadicus is entirely absent from all the Pliocene Siwalik strata, in which there is no ancestral type from which it might arise; this suggests the probability that the genus Palxoloxodon originated in Africa, migrated north into Europe, thence to India and the Oriental regions. As soon as we begin to examine the Pleistocene deposits of the Godavari, the Nerbudda, and the Ganges, Palzoloxodon namadicus occurs in great abundance; it is also found sparingly in Burma, China, Java, and Japan, as described by Owen, Martin, Koken, Schlosser, Naumann, Makiyama, and Matsumoto. SYSTEMATIC DESCRIPTION OF SPECIES OF PALAXOLOXODON AND HESPEROLOXODON The present revision by Osborn makes no attempt at finality, but is devoted to the establishment of the characters and geologic age of the principal generic and specific types, awaiting fuller monographic revision based only upon close comparison and measurement of the rich materials which the museums of western Europe and of India afford. To the writer’s knowledge, no complete cranium of Hesperoloxodon antiquus has been figured, and it is accordingly of interest to reproduce the photograph of a cranium? recently discovered at Pignataro Interamna, in Valle del Liri, near Cassino, Italy, between Naples and Rome, which appears to be exceptionally complete (see Figs. 1096, 1098, and 1101 below). See footnote 1 on p. 1049 regarding the possible Lower Pleistocene age of the Villafranchian.—Editor.] *This cranium was presented (May, 1929) by the author to the American Museum of Natural History and bears the number Amer. Mus. 22634. When discovered it was in perfect condition (as shown in Figs. 1096 and 1069), but it was seriously damaged in removal and in attempts to repair it by the owner. Full description and figures are given below in the present chapter. THE LOXODONTIN/: Paleoloxodon namadicus [lI alconer and Cautley, 1846, 1847 Figures 1041, 1046, 1068-1070, 1072, 1073, 1108, 1110 Type locality: Valley of the Nerbudda (or Narbada), Goddvari formation Vig. 729). Narbada Alluvium horizon, compare ‘Elephas antiquus (namadicus)’ Pilgrim, 1905, containing also referred Stegodon insignis, S. ganesa, and Rhinoceros unicornis. (see Upper Pleistocene. Speciric CHARACTHERS.—Transverse occipitofrontal rugosity very prominent in males and females. Typical ridge formula as below (p. 1212). Third superior molars with an estimated total of fifteen ridge-plates; 8e ridge-plates in 10 em.; maximum breadth Fig. 1070. skull, showing portions of the third true molar of either side; ten and a half ridge-plates remain (there were possibly three or four anterior to these, making fourteen or fifteen). One-sixth natural size. (Fig. 729). 101mm. Breadth of Mz 84mm.; estimated length of M2 264 mm. Ridge-plates broad, close set, entirely lacking ‘loxodont sinus’; enamel borders thin. Skeleton of gigantic size, height estimated at 12-18 feet (cf. Fig. 1068). In 1846, Falconer and Cautley in the ‘Fauna Antiqua Sival- ensis,”’ p. 45, first named the Upper Pleistocene species of the “Valley of the Nerbudda,”’ India, Elephas Namadicus, but did not figure the species until 1847 (Pls. xm.a and B of the “Fauna Antiqua Sivalensis’’). Subsequently, in the year 1847 (op. cit., Pl. x11.p), they figured and named the Lower Pleistocene species of western Europe Elephas antiquus. Consequently the name EHlephas namadicus Fale. and Caut. antedates the name Elephas antiquus Fale. and Caut. by nearly a year. Later Falconer clearly Brit. Mus. M.3092; cast Amer. Mus. Warren Coll. 10381. PALMHOLOXODON 1211 recognized and pointed out that his species #. namadicus of India most closely resembled his species H. antiquus of western Europe. Meanwhile he gave his usual thorough description and character- ization of this very important Indian species. references are as follows: E. |Elephas| Namadicus Faleoner and Cautley, 1846, 1847. “Fauna Antiqua Sivalensis,’” 1846, p. 45, and Atlas, 1847. (Op. cit., p. 45): “Another extinct Indian species H. Namadicus (to be described in the sequel).”’ Typr.—Skull showing portions of the third true molar of either side (Brit. Mus. M.3092). Hort- ZON AND Locauiry.—Valley of the Nerbudda, Upper Pleistocene, India. Typr Figure.—Op. cit., Pls. x1t.aA, X11.B, figs. 1 and 3. The systematic fig 3 TyPE SKULL OF PALMOLOXODON NAMADICUS (?)Female type cranium of Elephas Namadicus Falconer and Cautley, 1846 [1847, Pl. x1r.s, figs. 1 and 3]; lateral and palatal views of the Valley of the Nerbudda (or Narbada), India (see Tyrer Descrretion.—(Falconer in Murchison, 1867, p. 15, Pls. x1r.4 and x11.B: “From the valley of the Nerbudda. Probably a female, from small size of tusks. . . . It was chiselled out by Dr. “aleoner, and determined by him to be a new species. In a letter to Lieut.-Colonel Ousely, Dr. F. writes thus: ‘It is probably the most perfect specimen of a fossil elephant’s cranium in Europe. The species is especially interesting from the form of the cranium, which is so grotesquely constructed that it looks the caricature of an elephant’s head in a periwig. I have named the species EH. Namadicus, after the Nerbudda river, the Namadus of Ptolemy.’ There is a very similar specimen in the Museum of the Asiatic Society of Bengal. . . . Length of remaining portion left molar, 7.5 in. [12e ridge-plates in 7.5 in. or 191 mm.]. Width of remaining 1212 portion left molar, 3.7 in. N.B.—Twelve plates in this extent. Width of palate in front (between molars), 2.8 in. Width behind, 4.1 in.” Falconer gives full measurements (op. cit., pp. 15 and 16) and comparison with EF. indicus, BH. hysudricus, and E. primigenius. FaLcoNer (1868) MarTerIALS AND RipGeE FormuLA.—The ridge formula of Palxoloxodon namadicus may be derived by Fal- coner’s careful examination and comparison of the type skull (Fig. 1070) containing M*, an imperfect tooth, also of referred upper jaws (Pl. x1) and of several referred lower jaws (PI. x11.c, x1.D), as annotated herewith: Falconer, ‘Paleontological Memoirs,” 1868, Vol. I, pp. 435— 438. Typeskull: Plates x.a,B,c,p and x11, ridge-plates M 3**-7°. REFERRED Upper JAws.—Plate xu, figs. 1, la, 1b, upper jaw, M$, ridge-plates 11+. REFERRED LoweER JAws.—Plate x11.c, figs. 3, 3a, lower jaw, Dp, ridge-plates 10%; figs. 4, 4a, lower jaw, Ms, ridge-plates 2045; Plate x1r.p, figs. 1, la, lower jaw, M,, ridge-plates }s-13-'4, 15 in all; figs. 2, 2a, lower jaw, Mi, ridge- plates 13; figs. 3, 3a, lower jaw, Me, about 15 ridge-plates. Falconer’s type and referred ridge formula of Hlephas [= Palzxoloxodon| namadicus is: Dp 4 10% M 1 &-33-% M 2715 M3 i555. LyDEKKER (1886.2).—Lydekker does not give the complete ridge formula of Palxoloxodon namadicus but states (p. 169) that the third lower molar in specimens referred to Hlephas namadicus from Japan has but sixteen ridge-plates. He characterizes the grinding teeth of 2. namadicus as follows (p. 167): ‘‘The cheek- teeth of this species appear frequently almost or quite indistin- guishable from those of the broad-toothed variety of HZ. antiquus, although the ridge-formula is, on the whole, rather higher and the ridges themselves are somewhat taller; some teeth, however, especially those from Burma, China, and Japan, show excessive plication of the enamel, and thereby approximate to EL. indicus, although with a lower ridge-formula. . . . The adult cranium is characterized by the presence of a bold, overlapping, transverse ridge on the frontals. . . . which appears to be wanting in FE. antiquus. . . . In India the species occurs in the Pleistocene of the Narbada valley, and it is probable that the other specimens are from strata of equivalent age.” Cuina.—From China is recorded a 94+ ridge-plated third superior molar (Brit. Mus. 29007); length 166 mm., breadth 101 mm., height 180 mm.; laminar frequency 6 in 100 mm. (fide Hopwood, letter, August 9, 1928). Osborn, 1928: It is important to observe: (1) That the type ridge formula of Palxoloredon namadicus, as deduced above from Falconer’s observations, greatly exceeds the constant ridge formula of Loxodonta africana; (2) while Falconer also includes an M, with twenty ridge-plates, it is probably an erroneous generic reference, the true M 3 formula appears to be M 3 }-42; (3) in P. namadicus the ridge-plates are relatively broader, more numer- ous, more closely compressed, and entirely lacking the loxodont sinus character. (4) In 1868 (Vol. II, p. 261) Falconer observes: ‘In the fossil £. antiquus of Europe, the dentition of which I have been OSBORN: THE PROBOSCIDEA able to determine with precision, the formula for the three inter- mediate molars, and the last true molar, above and below, is 10: 10, 12, 16, being nearly intermediate between the Indian and African Elephants.” COMPARISON OF PALAHOLOXODON NAMADICUS AND HESPEROLOXODON ANTIQUUS Piterm™ (1905) ON THE GODAVARI SPECIMENS OF PALO- LOXODON NAMADICUS.—The fullest and most recent treatment of the species Palxolorodon namadicus is that of Pilgrim (1905): (1) He erroneously adopts Pohlig’s opinion that Llephas namadicus and F#. antiquus are closely related if not identical species.! (2) Consequently he describes the Nerbudda species as “Hlephas antiquus (namadicus),’’ giving namadicus the rank of a subspecies; inasmuch, however, as Falconer named the species 2. namadicus prior to naming the species Z. antiquus, this usage (Pilgrim) cannot be adopted; nor are these species identical. (3) He regards the Godavari river gravels (Lat. 20° 1’, Long. 74° 11’) as of the same Lower [Upper] Pleistocene age as the typical Nerbudda deposits in which the type of #. namadicus was discovered, containing alike EL. namadicus, Stegodon insignis ref., S. ganesa ref., Equus namadicus, Hippopotamus palxindicus, H. tetraprotodon, also Rhinoceros unicornis. He regards FE. namadicus as abundant also in the Pliocene of Burma, of China, of Java, and of Japan. Pilgrim’s description of the skull and skeleton of Hlephas namadicus may be freely cited as follows (Pilgrim, 1905, pp. 203- 206): ‘““The cranium and bones, which I am describing, and which represent the species Hlephas antiquus (namadicus) Fale. et Cautl., belonged to an individual of remarkable size. It cannot have stood much less than 16 feet at the shoulder. The cranium, as found, is larger than any hitherto recorded. . . . The cranium either on one or both sides possesses all the essential features of the portion above the maxillaries and the foramen magnum... . The accompanying plates exhibit all the more important characters of the present cranium, and the most casual inspection of it can leave no doubt as to its identity with the crania from the Narbada beds, figured by Falconer in the Antiqua Fauna Sivalensis, Plates 12A, 12B, figs. 1-3, and Plate 24A, figs. 4, 4a, as Hlephas namadicus. These are the two most complete crania which have been known up to now, and are preserved in the British Museum. One of them has small tusks and probably belonged to an adult female. .. . The present skull is that of a fully grown male. It seems that the supra-orbital ridge grew forward with age, so that in young skulls there is a considerable interval between its margin and the extreme tip of the nasal process; in the large female skull in the British Museum this interval is sensibly diminished, while in this latest specimen, which represents the largest and presumably the most aged type with which we are acquainted, the supra-orbital ridge almost overhangs the nasal fossa, and the interval is reduced to its smallest dimensions. . . . Considering only the teeth and mandible of E. antiquus and of H. namadicus, Leith Adams . . . remarked that they seemed to him to be indistinguishable. This opinion of Leith Adams has been endorsed by many subsequent writers, among whom I need only mention Naumann, Weithofer, Pohlig, [Professor Osborn (1931.846, p. 21) provisionally made his subspecies Palzoloxodon antiquus italicus the genotype of a new genus Hesperolozodon, as distinct from Palzoloxodon, to include also other members of the antiquus group (see Fig. 1068 of the present Memoir), which he finally adopted (Osborn, 1934. 926, p. 285, 1935.937, fig. 2, p. 407, and Pl. x1, Vol. I of the present Memoir).—Hditor.] E1GI “XI [Td ‘V061 Wag 103j8 poonpoidoy “punoy svn (SOTT 3 u : 0 ae (13]I SBM (8 PUB OLLT ‘TFOT ‘84g ‘trowopy quosoid jo FI ZT ‘ZIZT “dd 99s) ¢ y QLosop snoippupu Uoporojox2)Dq JO [[NYS pediojod oyy UOTsoI YOupM ur ‘BIpup “IBMYSOUIPYY InpuBNy Ie Z eo age eee ied pled I g D INGLES UST TE DO i : O TLOT “4t 1214 OSBORN: THE PROBOSCIDEA and Lydekker. ... So far is this from being the case, however, that all the skulls of the dwarf forms which Pohlig . . . has figured from the Grotto di Pontale von Carini in Sicily bear a striking resemblance to Elephas namadicus, and leave us no excuse for separating the two forms specifically. The accompanying text figure, taken from one of Pohlig’s plates [Fig. 1041], brings out these resemblances in a remarkable degree. There is no doubt that future discoveries will prove that the original #. antiquus of Europe possesses the same yy craniological peculiarities as its Indian variety. Pilgrim (op. cit., pp. 208, 209) gives detailed comparative Tig. 1072. Comparison OF H&SPEROLOXODON AND PAL®OLOXODON (syn. SIVALIKIA) SUPERIOR AND INFERIOR MoLars Drawn to the same one-sixth scale. Compare figures 1073 and 1152 A, Hesperoloxodon antiquus after Forster Cooper, 1924, Pls. rx, x. Left M*, 17 ridge-plates; left M*, 12 ridge-plates; left M3, 15 ridge- ’ plates. Lower Pleistocene. From Barrington, England. B, Hesperoloxodon ausonius after Depéret and Mayet, 1923, Pls. x, x1. Left M*, 15 ridge-plates; left Me, 11 ridge-plates; left M3, type, 2). 19 ridge-plates. Upper Pliocene [Lower Pleistocene San Romano. Val d’Arno inf., Italy. C, Hesperoloxodon antiquus Fale. and Caut., 1847, Pls. xi1.p, Xiv.A, and Leith Adams, 1877-1881, Pl. 1v. Left M%, 18 ridge-plates; left M?, 14 ridge-plates; left Ms, type, 12+ ridge-plates; left M3, 17 ridge- plates. Lower Pleistocene. From England. D, Palzxoloxodon namadicus Fale. and Caut., 1847, Pls. x11.B, D, c. Left Mg, type, 14+ ridge-plates; left Mo, 14 ridge-plates; left M3, 19% ridge-plates. Upper Pleistocene. Nerbudda Valley, India. Observe the progressive increase in breadth both of the superior and inferior molars and the corresponding decrease in length. The superior and inferior molars of Hesperoloxodon ausonius are relatively the longest and the narrowest; the grinders of H. antiquus (A, C) are intermediate in width and length; the grinders of Palxoloxodon namadicus are relatively the shortest and the broadest, also the most elevated or hypso- dont. This change of proportions from the extremely long and narrow type, H. ausonius, to the relatively short and broad type, P. namadicus, is correlated with the progressive hypsicephaly, bathycephaly, and brachycephaly of the cranium. The same principle of the relative shortening and broadening of the grinders is displayed in figure 1073. measurements of the crania of Elephas antiquus (namadicus), E. antiquus, and E. antiquus melitensis in the museums of Calcutta, London, Florence, and Palermo, and concludes (p. 210): “2. EH. africanus |Loxodonta africana] in the flattened shape of the vertex, antiquus |Hesperolorodon antiquus| approaches nearest to in the shortness and breadth of brow, in the form of the temporal All 146 Nalural A Superior molars Infervor molars Stze LM3 ANTIQUA P, ANTIQUA P. NAMADICA fossa and of the interjugal space, and in the obtusely angled june- tion between the frontal and occipital surfaces. It differs in the slope of the occipital surface, in the greater length and shallowness of the intermaxillary median fossa, in the convexity of the occiput, in the higher position of the maxillary zygomatic process, and in the shorter length and greater width of the sub-orbital foramen.” Professor Osborn regarded E. antiquus as referable to his genus Hesperoloxodon (see footnote 1 on p. 1180 above).—Editor.] THE LOXODONTINA: PALHOLOXODON GENERIC CHARACTERISTICS OF PALAXOLOXODON NAMADICUS CRANIUM (Compare Figs. 1069 and 1041) (Pilgrim, 1905, pp. 206, 207): ‘‘The crania of FH. [= Hespero- loxodon] antiquus (stem. sp.), imperfect as they are, show the following points in common with the Indian variety [namadicus] and the pygmy types which serve to distinguish them from all other R M3 (rev) Ref. wil 124 SIVALIKIA SIVALIKIA NAMADICA ANTIQUA 1215 Narbada elephants exhibit in addition the following points of likeness:— 4. The shortness and breadth of the brow and the widening out of the cranium from below upward. 5. The rhom- boidal outline of the temporal fossa and its sharply-cut, acute- angled upper margin. 6. The presence of protuberances on either side of the occipital fossa. 7. The almost rectangular bend by which the occipital passes into the parietal, and the ob- tusely-angled junction between the parietal and frontal surfaces. After Falconer After Falconer AUSONIA After Deperet, Sayet “/ Roman Fig. 1073. THREE PROGRESSIVE BROADENING STAGES IN THE PALMOLOXODON (SYN. SIVALIKIA) AND HrSPEROLOXODON SUPERIOR GRINDING TEETH. ALL TO THE SAME ONE-FOURTH SCALE. Compare Figures 1072, 1152, 1075, 1078 (Upper) Palzoloxodon (syn. Sivalikia) namadicus. Right M*, 12+ ridge-plates; left M3, 19} ridge-plates, after Falconer and Cautley, 1847, Pl. xm, fig. la, and PI. x11.c, fig. 4, respectively. (Middle) Hesperolozodon antiquus. Right M*%, 19+ ridge-plates, after Falconer and Cautley, 1847, Pl. xu.p, fig. 5, Kent, England (Canterbury Mus.); left M3, 17 ridge-plates, Pl. x1v.a, fig. 11, Saffron Walden, England. Of intermediate length and width. (Lower) Hesperoloxodon ausonius. Left M%, 15e ridge-plates, after Depéret and Mayet, 1923, Pl. x, fig.3, Malafrasca (Val d’Arno sup.), Italy; left M3, type, 19 ridge-plates; Pl. x, fig. 1, San Romano (Val d’Arno inf.). Both in the Inst. Géol., Florence. Relatively long and narrow grinders. Observe as in figure 1072 the progressive broadening and shortening of the third superior and inferior grinders. Also observe as in figure 871 that the superior ridge-plates are concave posteriorly, the inferior ridge-plates concave anteriorly. Cement areas dotted. elephants:— 1. The extreme divergence of the incisive alveoli and the broad shallow depression which occupies their centre. 2. The great distance of the occipital fossa from the foramen Magnum and the basal breadth and extreme depth of the fossa. 3. The strong convexity of the occiput in a horizontal direction, which pushes the zygomatic process of the temporal to the front in an unusual degree. The crania of the Sicilian and of the 8. The well-marked frontal projection [namadicus] of the crown, which must have given an exceedingly beetling aspect to the living animal. 9. The approximately transversely oval contour of the cranium, when viewed in a direction at right angles to the plane of the occiput. It is much broader than high. {[10.] Z. africanus also approaches them to some extent in regard to the Ist, 4th, 5th, and 9th of the above characters.”’ STRAIGHT-TUSKED ELEPHANT OF UPNOR (HESPEROLOXODON ANTIQUUS) IN THE BritisH MuskuM. RESTORATION, UNDER THE DIRECTION OF HENRy Farr- FIELD OsBorN, BY Mararet Fiinscu Bupa (1935) To A ONE-FIFTIETH SCALE (BULLS) AND A ONE-SIXTIETH SCALE (CALF) Skeleton and growth stage based upon the Upnor elephant (Fig. 1079); head or cranium based upon the Pignataro Interamna elephant (ig. 1098); the one fragmentary tusk found at Upnor furnishes the length of the ivories in the restoration; ears based upon the cave drawings of Africa and Spain (Fig. 1047). Several months’ research and all the studies involved in the preparation of figures 1079, 1080, 1084, 1083, and 1081 were also preliminary to the execution of this restoration. Fig. 1074. This giant straight-tusked elephant, which formerly ranged over western Europe, is here shown in its relatively early Pleistocene stage of evolution and in a stage of growth which corresponds to the twenty-fifth year of living African and Indian elephants, namely, in which the second superior and inferior molars are still in use, while the third superior and inferior molars have not ccme into use (ef. Fig. 1061 of Loxodonta africana). In other words, the Upnor straight-tusked elephant in the British Museum was about 25 to 30 years of age; it had not attained the full height character- istic of this species nor the maximum length of the tusks; yet it was a giant in life, measuring from the summit of the scapula to the ground, as originally estimated by Andrews and Forster Cooper, 12 ft. 146 in. (3700 mm.), and, as estimated by Osborn, 12 ft. 10% in. (3934 mm.) in the flesh. The views of the skull and tusks, drawn directly from the Pignataro skull (Amer. Mus. 22634), are extremely accurate and characteristic, with great breadth between the tusks and correspondingly broad proboscis. The relatively level vertebral contour lacks the marked mid-depression and elevation above the pelvis which are characteristic of the African elephant (Fig. 1084). The relatively small, low-set ears are restored from the admirable cave drawings re- produced from Pomel (1895) and Breuil (1911) in figure 1047 above. As observed in the legend of that figure, the ears resemble those of the Indian (Fig. 1120) rather than the enormously enlarged and elevated ears of the African elephant (Fig. 1052). 1216 Genus: HESPEROLOXODON Osborn, 1931 Original reference: Amer. Mus. Novitates, No. 460, 1931, p. 21. Genotypic species: Palzoloxodon antiquus italicus Osborn, 1931.846, p. 21.1 [The genus Hesperoloxodon was provisionally proposed by Professor Osborn in his article entitled, ‘‘Palxoloxo- don antiquus italicus Sp. Nov., Final Stage in the ‘Elephas antiquus’ Phylum” (see Osborn, 1931.846, p. 21), to include Palxoloxodon antiquus italicus from Pignataro Interamna, Italy. Subsequently in various ways he indicated his opinion that the typical ‘Hlephas antiquus’ and other members of the antiquus group should be referred to his genus Hesperoloxodon, and finally in 1934 (Osborn, 1934.926, p. 285) the name appears as follows: ‘‘Hesperoloxo- don Osborn, a Loxodontine of western Eurasia and Africa, never reaching America.’ Also the name appears on a chart between Loxodonta and Palzxoloxodon (see Osborn, 1935.937, p. 407, fig. 2), and on page 12 as well as on Plate XI of Volume I of the present Memoir. The accompanying definition, therefore, has been compiled from several sources and embodies, as far as can be determined, the distinctive characters on which Professor Osborn separated Hesperoloxodon from Palxoloxodon Matsumoto. GENERIC DeFinition.—(Osborn, 1931.846, p. 21): ‘‘Comparison [of the cranium of Palzxoloxodon antiquus italicus] with the cranium of ‘Elephas namadicus’ shows a strong resemblance in the breadth of the premaxillary rostrum but an extreme difference in the summit of the cranium, which in ‘EZ. namadicus’ is relatively low and reinforced by the overhanging parieto-frontal crest. This points to Pal. ant. italicus as a member of a phylum quite distinct from that of the Siwalik ‘#. namadicus,’ a phylum which if sup- ported by other cranial and skeletal differences might well constitute a new genus to which the name HESPEROLOXODON, or ‘loxodont of the west,’ might be applied. This name is provisionally proposed, as I would not like to be forestalled a second time, as in the case of Palxoloxodon, a generic name assigned to ‘EH. namadicus naumanni’ by Matsumoto but a few weeks prior to my description of Sivalikia.”’ Supposed diagnostic characters of Hesperoloxodon compiled from statements in the present Memoir: Cranium domelike with flattened forehead, more hypsicephalic and bathycephalic than that of Palxoloxodon; the promi- nent frontoparietal crest or “bold overlapping transverse ridge on the frontals”’ as mentioned by Lydekker in his description of H. [Palxoloxodon| namadicus (1886.2, p. 167) is lacking. Occiput relatively narrow and high (broad and low in namadicus). Grinders nypsedent; ‘loxodont sinus’ vestigial or absent; ridge formule: M 318#-17 (typical), to M 3 42-12 (germanicus), to M 3 3. (ztalicus). divergent, slightly upeurved and incurved.—Editor. | Premaxillaries extremely broad, incisive tusks widely Bed, Norfolk). All these beds probably belong to the Lower Pleistocene, /st Interglacial stage, and are geologically older than the 2d Interglacial beds of Mosbach, ete., Germany, which contain larger and more progressive stages of Hesperoloxodon antiquus, as published by Pohlig. They are much older than the beds of Tau- bach and Weimar containing H. antiquus germanicus. Hesperoloxodon antiquus Falconer and Cautley, 1847, 1857 Figures 794, 871, 1068, 1072-1084, 1088, 1109, Pl. xxu1 Type locality: Not recorded, probably Lower Pleistocene. numerous referred specimens of Lower and Middle Pleistocene age (see Falconer, ‘Paleontological Memoirs,” Vol. II, pp. 176-188). Syn.: ‘Elephas priscus’ Falconer and Cautley, 1847; Hlephas (Loxod.) priscus Falconer, 1857. Also very History.—As noted above, the ‘‘Sceleto Elephantino Tonnze Grotocic Acr.—While the locality of Falconer’s type speci- men (Brit. Mus. M.2006) is not recorded, Falconer’s referred specimen, Hlephas (Loxod.) priscus, is from Gray’s Thurrock (Essex); the same locality yields six of the British Museum speci- mens described by Leith Adams (1877-1881), while one specimen comes from Clacton (Essex) and another from Cromer (Forest [See footnote on p. 1247 below.—Editor. | of Tentzelius (1698) is the first scientific description of the ‘Hlephas antiquus’ of Europe; Blumenbach confused this skeleton with that of ‘#. primigenius’ of Europe. The species ‘H. antiquus’ was then confused even by Falconer until 1847 with ‘2. meridionalis,’ owing to the wide separation of the ridge-plates. 1217 1218 Spreciric CHARACTERS.—Prominent transverse fronto-occipi- tal crest known to be lacking. Typical ridge formula of Hesperoloxo- don antiquus somewhat greater than that of Palzolorodon namadi- cus, namely; Dp22Dp3£ Dp 473 M12232M222M3i8". 10 Grinding teeth relatively longer and narrower than in P. namadi- cus; height more than double the width of the crown (Fale.). ‘Loxodont sinus’ absent in the type (Fig. 1075); absent or vestigial in other specimens (Fig. 1076); ‘‘mesial rhomboidal expansion of the dises of wear” (Fale., 1868, II, p. 176). Inferior ridge-plates concave anteriorly; superior ridge-plates concave posteriorly; crowns with thicker cement and rela- tively thicker and less plicate enamel than in P. namadicus; “great crimping of the enamel-plates”’ (op. cit., Faleoner, 1868, p. 176). Extremely narrow ‘loxodont sinus’ indicated in worn ridge-plates of Hesperoloxodon antiquus and P. namadicus (Fig. 1072). See also characters noted under H. antiquus italicus and H. antiquus germanicus below (pp. 1238- 1256). The above specific characters, including Fal- coner’s final definition (“Paleontological Memoirs,”’ 1868, Vol. II, p.176) doubtless apply to a collective species embracing many ascending mutations from Upper Pliocene to Middle Pleistocene time, but the typical Hesperoloxodon antiquus is probably of Lower Pleistocene age. Falconer and Cautley named this species of straight-tusked elephant in 1847, a year after naming the Indian species Elephas namadicus in 1846; consequently if the two species are identical, as alleged by Pohlig and by Pilgrim, the name Elephas namadicus has the technical priority. There are many reasons, however, for treating the great straight-tusked elephant of southern Europe as a species distinct from, although nearly allied to, its Indian relative Elephas namadicus. All authors agree as to the relationship of these two animals and there is little doubt as to their affinity, but it is important to observe that in Falconer’s type of Elephas namadicus (¥ig. 1070) the grinding teeth are much broader than in Faleoner’s type of Elephas natural size. antiquus (Fig. 1075), which are relatively narrow. Consequently #. antiquus cannot be the same species as E£. namadicus, however much these animals re- semble each other in cranial characters.! CONFUSED HISTORY OF NAME AND TYPE Blephas antiquus Faleoner and Cautley, 1847. ‘Fauna Anti- qua Sivalensis,” 1847, Atlas, figs. 4, 4a of Pl. xi1.p, figured as “PB. meridionalis” but corrected by Dr. Faleoner in copy of “Fauna OSBORN: THE PROBOSCIDEA Antiqua Sivalensis” belonging to British Museum; also Pl. xrv.B (the first time the name was published).? E. (Eueleph.) antiquus ‘aleoner. “On the Species of Mastodon and Elephant occurring in the fossil state in Great Britain,” Quart. Journ. Geol. Soc. London, 1857, Vol. XIII, table opposite page 319. Typr.—Lower jaw with second lower molar, Me, of the left side (Brit. Mus. M.2006). Tyre Locauiry.—Unnkown, undoubtedly Eng- LecroryrE Seconp Lerr INFERIOR Mouar OF HESPEROLOXODON (SYN. SIVALIKIA) ANTIQUUS FALCONER AND CAUTLEY One-third natural size Fig. 1075. Lectotype, l.Me, of Elephas antiquus, first figured as “H. meridionalis” (Falconer and Cautley, 1847, Pl. xu.p, figs. 4, 4a); Brit. Mus. M. 2006, locality unrecorded. One-third Named E. (Eueleph.) antiquus (Falconer, 1857, Synop. Tab. opp. p. 319), de- fined by Falconer, 1867, p. 18, and 1868, Vol. I, p. 488, as follows: “Hlephas antiquus. . . . jaw, left side, with first [?] true molar. thirteen ridges, with front ridge and heel. It narrows excessively in front and behind, like fig. 3 of EZ. Namadicus! The crimping, &c, are also exactly alike.—B. M. [Brit. Mus. M. 2006]. Length of molar, 8.in. Width at middle, 2.6in. Width in front, 1.3 in.” Lower This tooth is a beautiful specimen; shows twelve to land, probably of Lower Pleistocene age. Typr F'icurn.— ‘aleoner and Cautley, 1847, Pl. x1r.p, figs. 4, 4a. FaLconer’s TypeE.—There can be no question that this second inferior molar (ig. 1075) should be regarded as Falconer’s type. '{In 1931 Professor Osborn proposed the genus Hesperoloxodon to embrace members of the ‘Elephas’ antiquus group, retaining Matsumoto’s genus Palzxoloxo- don, 1924, for members of the ‘HZ.’ namadicus group (see pp. 1212 and 1217 of this chapter).—Editor.] *(Compare Bather, in Andrews and Cooper, 1928.1, p. iii; also for descriptive legend, see Falconer, 1867.1, pp. 18 and 23, and 1868.1, II, pp. 438, 441, and 443.—Editor.]} THE LOXODONTIN«: (Osborn, 1924): This twelve-plated Mo, of the left jaw, may be regarded as the type, since it was the first specimen figured and named (1847), and first described in 1867 by Falconer. Lydekker (1886.2, p. 130) designated it as follows: ‘‘M.2006. Part of the left ramus of the mandible, containing the half-worn ,-s; locality unknown. Figured by Falconer and Cautley in the ‘Fauna Antiqua Sivalensis,’ pl. xii.p, fig. 4, 4a. No history.” At the time, unfortunately, Falconer was misled by the spurious fossil type of Elephas priscus Goldfuss, for in the same table (Falconer, 1857, opp. p. 319) he cites “‘E. (Loxod.) priscus (Goldf.) . . . Pliocene ... England; Lombardy ... Imperfectly known. Fossil remains rare.’ This citation agrees entirely with his previous reference (Falconer, 1846, p. 15), in which he recognizes EF. priscus Goldfuss, 1821, as a valid fossil type, although disputed by Cuvier. We must therefore regard the tooth described as FH. (Loxod.) priscus, and reproduced in this Memoir (Fig. 1076), as referable to the true Elephas |Hesperoloxodon| antiquus of Falconer. E. (Loxop.) priscus (Goupr.) of FALconrR, 1857 = HESPEROLOXODON ANTIQUUS E. (Loxod.) priscus (Goldf.) Faleoner, 1857. ‘On the Species of Mastodon and Elephant occurring in the fossil state in Great Britain,’’ Quart. Journ. Geol. Soc., London, Vol. XIII, pp. 345, 346, and table opposite page 319. Typr.—A second lower molar of the left side. Horizon AND Locauiry.—Gray’s Thurrock, England, Pleistocene. Fraure.—Faleoner and Cautley, 1847, Pl. xiv, figs. 7, 7a, 7b, under the name £. priscus? (Brit. Mus. 39370). Falconer’s type tooth of E. antiquus (Fig. 1075) and the tooth designated as EH. (Loxod.) priscus (Fig. 1076) supplement each other in the fact that the lorodont character is more evident in the priscus variety. This is the eight-crested grinding tooth from the brick earth of Gray’s Thurrock, Pleistocene, to which Falconer applied (1865, pp. 276, 277) the preoccupied name Hlephas (Loxod.) priscus, in allusion to the lozenge-shaped crests, which remotely suggest those of the type of Elephas priscus Goldfuss (= Loxodonta africana). Falconer described this tooth (1865, pp. 270, and 1868, p. 96) as a “last molar [an error], left side, of the lower jaw.” Lydekker described the tooth (1886.2, p. 133) as a “second left lower true molar.’’ In the same publication, Lydekker (p. 122) treats it under Elephas antiquus as follows: “Syn. Elephas (Loxo- don) priscus, Falconer and Cautley.” ELEPHAS ANTIQUUS AND E. NAMADICUS, FALCONER’S NOTES OF 1867 AND 1868 RELATING TO SPECIMENS FIGURED IN THE PLATES OF THE “FAUNA ANTIQUA SIVALENSIS,” 1845-1847 Osborn, 1923: Future research will probably distinguish between the lectotype ridge formula of Elephas antiquus from Essex, Norfolk, ete., and the collective ridge formula of specimens from other localities. Couuective Ripcr FormutA.—Faleoner’s conception of the dental characters of Elephas antiquus and their points of similarity to the molars of Hlephas namadicus are to be found in the beautiful plates and legends of the “Fauna Antiqua Sivalensis’”’ and in the “Paleontological Memoirs” of 1868, as follows: HESPEROLOXODON 1219 Falconer, ‘Paleontological Memoirs,” Vol. I, 1868, pp. 438— 440, 442, 443, 447, legends to Plates x11.p, XIII.A, XIV, XIV.A, XIV.B. Elephas antiquus. Lxecroryen. Plate x11.p, figs. 4, 4a, M, [Mg] with 12-13 ridge-plates, and front ridge and heel, closely similar to E. namadicus, crimping, ete., exactly alike. Upper JaAws.— Plate x1r.p, figs. 5, 5a, M%?, with 16% ridge-plates [19+]. Plate XIV, figs. 2, 2a, 2b, Dp’, ridge-plates 6). Plate xtv.a, figs. 1, la, Dp’, ridge-plates 5; figs. 2, 2a, Dp*, ridge-plates 10; figs. 3, 3a, Dp’, ridge-plates 10, figs, 4, 4a, M!, ridge-plates 8; figs. 5, 5a, r.M§, ridge-plates 14% [164]. Plate xrv.s, figs. 16, 16a, entire upper molar, M°, ridge-plates 16-17, length 11 in.=280 mm. LOWER Jaws.—Plate xu.a, fig. 4, lower jaw, Mo, ridge-plates 6+, Ms, ridge-plates 17; fig. 5, lower jaw, My, ridge-plates 124%. Plate xtv, figs. 1, la, 1b, lower jaw, Dps, ridge-plates }s-6—4, from Grays, Essex. Plate xiv.a, figs. 8, 8a, r.M,, ridge-plates 12; figs. 10, 10a, lower jaw, r.Mz, ridge-plates 124; figs. 11, lla, 1.Ms, ridge-plates 15-1635. REFERRED HESPEROLOXODON ANTIQUUS Vig. 1076. Aged second molar of the left side, 1.Me, from Gray’s Thurrock (Essex), figured as E. [Hlephas] priscus? by Falconer and Cautley, 1847, Pl. xIv, figs. 7, 7a, and as Hlephas (Loxod.) priscus by Falconer, 1868, Vol. II, Pl. vir, fig. 1. One-third natural size. Brit. Mus. 39370. Falconer’s type and referred ridge formula of EH. antiquus: 5-68 ¥s-6-18 Dp 3 Dp 4 *#2*°%4*M155%455y The above observations of Falconer (1867, 1868) and on Faleoner’s plates (by Osborn) are consistent with Faleoner’s less detailed formula of 1863. We may regard the following as his collective ridge formula of Hlephas [= Hesperolorodon| antiquus: M3 164-17 Dp 3 : Dp 4 10 164-17" 1220 OSBORN: FALCONER’S OBSERVATIONS SUMMARIZED BY MURCHISON “Paleontological Memoirs,” 1868, Vol. II, pp. 176-188 We owe to Murchison (“‘Palzontological Memoirs,” 1868, Vol. II, pp. 176-188) the complete enumeration of the numerous speci- mens regarded by Falconer as belonging to the above species, as observed in the museums of England, Italy, and Sicily; they doubtless belong to many successive geologic horizons. Falconer himself observed many variations in the ‘loxodont sinus’ or loop, the thickness of the enamel, the width and length of the crown, and the greater or less crimping or undulation of the ridge-plates. The typical ‘Elephas antiquus’ of Falconer appears to have been of Lower Pleistocene, or Cromer Forest Bed age, as shown by the fact that the collective ridge formula cited above agrees precisely with that assigned to Cromer Forest Bed superior and inferior molars as cited below by Osborn. Tuirp Riaut Surerior Mouar or HESPEROLOXODON ANTIQUUS Compare diagrammatic figure 1088 Fig. 1077. Referred 16) ridge-plated molar, r.M*, of Elephas antiquus, after Falconer and Cautley, 1846 [1847, Pl. xiv.a, figs. 5, 5a). designated on Falconer’s plate (xtv.a) as ‘Elephas meridionalis,’ but corrected in his handwriting in copy of the “Fauna Antiqua Sivalensis” belonging to the British Museum. Falconer defined this specimen as follows (1867, p. 22, 1868, Vol. I, p. 442, Pl. x1v.a): “Figs. 5 and 5a.- side. Erroneously Last true molar, upper jaw, right Has [+] fourteen plates and a heel, well crimped. Ostend, Norfolk. Green collection.—No. 16,229 B.M. Length, 10. in. Width, 3.4 in. Height, 6.5 in.” One-third natural size. Elephas |Hesperolorodon| antiquus (Pig. 1075). rom forest. bed, Compare with the type of THE PROBOSCIDEA Vig. 1078. Upnor ELmpHant (HESPEROLOXODON ANTIQUUS). SECOND Superior AND INFERIOR MOLARS OF SKELETON DESCRIBED BELOW ONE-THIRD NATURAL SIZE A, Crown view of twelve ridge-plated second right superior molar, r.M?’. B, External view of twelve ridge-plated second left superior molar, 1.M?; anterior ridges partly worn. C, Lateral view of second right inferior molar, r.Me, retaining ridge-plates 2-11. D, Crown view of second right inferior molar, r.Me, retaining ridge-plates 2-114 estimated. After retouched original photographs by C. Forster Cooper. Observe that these molars were identified as second superior and inferior molars by Andrews (1915, p. 11), as third superior and inferior molars by Forster Cooper (1928, pp. 23, 24). The two upper molars, r.M?, 1.M’, show twelve remaining ridge-plates with a posterior talon; they measure +221 mm. in length, 83 mm. in breadth, 150 mm. in maximum depth; laminar frequency 5 ridge-plates in 10cm. The lower molar, r.Me, measures +231 mm. in length, 70 mm. in breadth, 123 mm. in depth; laminar frequency 5 in 10 em. Loxo- dont sinus rudimentary. This r.Me compares closely with Falconer’s type 1.Me (Fig. 1075). THE LOXODONTINA: Falconer found no trace of successional premolars such as are observed in Archidiskodon planifrons nor of the first lower milk molar. The other priceless observations contained in Falconer’s Note-book of 1862 are summarized by Murchison (op. cit., Pal. Mem., 1868, p. 176) as follows: “4, Elephas (Huelephas) antiquus. . . . compiled from entries in Dr. Falconer’s Note-books.—|Ep.] The distinctive characters of the teeth of Elephas antiquus may be expressed in the following terms.— if height. 2. Great height of the plates. double the width of the crown. Narrowness of the tooth in proportion to its length and The height is more than 3. Mesial rhomboidal expansion of the discs of wear. 4. Great crimping of the enamel-plates. The dental formula of H#. antiquus is as follows: Milk Molars. 3+6+10 3+6+10 True Molars. 10+12+16 9) 10+12+16 The comparative enumeration of the ridge-plate totals and of the characteristic formule was also commented upon in Fal- coner’s posthumous Memoir of 1865. Observe throughout. his notes that the: First upper milk molar = Dp? of the present Memoir. Second upper milk molar = Dp‘ of the present Memoir. Third upper milk molar = Dp? of the present Memoir. HESPEROLOXODON 1221 CONFUSION WITH PARELEPHAS MOLARS, BY ADAMS, LYDEKKER, AND MATSUMOTO Leith Adams (1877-1881, p. 47), after careful consideration of Falconer’s work and ridge formule, concludes as follows: ‘From the foregoing details it seems to me that the ridge formula of Elephas antiquus, as far as British specimens in particular demon- strate, is, . . . [without talons], in upper and lower jaws, as follows: Dp 23-7 Dp 355 =t Dp 4 8" Mf 1 222 Mi 222-48 Mf 3 18-20 11-12 12-13 16-19° Leith Adams’ error arose from his confusing the ‘HZ. antiquus’ molars with those of the contemporary Parelephas. DISTINCTION FROM PARELEPHAS TROGONTHERII.—Lydekker (1886.2, p. 122), although making use of the same materials as Falconer, erroneously assigns to H. antiquus a higher ridge formula; because, like Leith Adams, he includes within the narrow-plated E. antiquus the ‘broad-plated’ molars belonging to Parelephas trongontherti. He observes: ‘The ridge-formula [Footnote: ‘Slightly modified from Leith-Adams, “British Fossil Elephants’’ (Mon. Pal. Soec.), p. 176. If the talons be included the formula will be higher, see Leith-Adams, op. cit., p. 231.’], excluding talons, may be represented as: (5-7) Mm. [Dp 2] a [Dp 3] (6-8) (M 2] os [Dp 4] (aan [M 3] M.[M 1] (33 -12) (15-20) 9 (16-21) 3) 3) The higher numerals in this Leith Adams and Lydekker formula probably pertain to Parelephas trogontheriz, namely: M122 M2'3 M322, Similarly Matsumoto (1924-1926) erroneously considered that the Palzxoloxrodon phylum represented by his ‘Parelephas proto- mammonteus’ was ancestral through the Parelephas trogontherii phylum to the Hlephas [= Mammonteus] primigenius phylum, as indicated by his choice of the specific name ‘protomammonteus’ (ef. p. 1297 below). PRIMITIVE AND PROGRESSIVE RIDGE-PLATE FORMULAi Research to the end of the year 1929, aided by the cranium from Pignataro Interamna, Italy (the type of Palxoloxodon [= Hesperoloxo- don| antiquus italicus), reveals (Fig. 1088) a progressive hypsodonty and ridge-plate addition from the Upper Pliocene to the 3d Inter- glacial stage, when H. antiquus germanicus and H. antiquus italicus mark the extinction of this phylum in western Europe. ProGressive Ripcr Formut&#.—The progression from the typical Lower Pleistocene M 3 42 cited above, to the progressive ridge formule given by Zuffardi, Depéret, Soergel, and others, namely M 3 2° cited below, does not represent contemporary variations; it represents rather progressive or ascending mutations. As shown in the Osborn-Reeds diagram (Pl. xxiv), Hesperoloxodon antiquus lived for an enormously long period of time, perhaps hundreds of thousands of years, during which ridge-plates were constantly being added. During the same long geologic period, as shown in figures 1072 and 1073, the ridge-plates were constantly broadening. Forster Cooper cites the primitive and progressive ridge formule (1924, p. 117), as given by more recent authorities, as follows: Zuffardi, 1913: M 212/13 M3 72/29 | Hlephas antiquus Depéret, 1923: M273775 M3 42/29 } [=germanicus], M2 5242 M3121 | Hlephas antiquus mut. ausonius Soergel, 1912: M2913 M3 pate | mid-Pleistocene | 1222 OSBORN: THE PROBOSCIDIEA He adds the following characters: Blephas antiquus: ‘Thick, much folded plates regular rhombic or rectangular [S]. Fairly thick, always folded. Loxodont sinuses always well marked [D]. Folding not particularly noticeable, lozenges marked, a considerable difference in size between upper and lower molars [Z]. Long straight high crowns [Z].”’ Elephas antiquus As antiquus but more deeply folded, and with folds more numerous, a very regular feature. mut. ausonius: Loxodont sinuses variable but on the whole stronger and more prominent [D]. Elongated straight crowns about }s smaller than antiquus from Clacton, Saffron Walden, ete. [D].”’ Osborn, 1930: The ridge formule cited by Forster Cooper (1924, p. 117) and the specific references and identifications are super- seded by the more precise ridge formule and specific identifications in the present Memoir; the unique ‘EHlephas ausonius’ (M 3 48) does not occur in the Forest Bed, the mut. ausonius is probably ‘Parelephas trogontherti nestii’; the higher ridge formule attributed to Zuffardi, Depéret, and Soergel belong only with the mid-Pleistocene [Upper Pleistocene] progressive H. |Hesperoloxodan| antiquus germanicus, as shown in figure 1088. Primitive RipGe-PLATE FoRMULa AND MEASUREMENTS Ridge-plates Length Breadth Height Cromer Forest Bed R. M3 16-17 280 mm. =11 in. 90 mm.=3.50 in. 175 mm.=6.80 in. Happisburgh (Forest Bed) R. M; spud 267 mm. = 10.5 in. 88 mm.=3.40 in. 146 mm. =5.70 in. Upnor Elephant (after Forster Cooper) R. M2 12% 22 emmi—s sees 83 mm. =3.25 in. 150 mm. = 5.90 in. R. Me 12% est. 2olemms— 9 inne 70 mm.=2.75 in. 123 mm.=4.76 in. E. antiquus type L. Me 12 204 mm.= 8 in. 66 mm. = 2.60 in. THE HESPEROLOXODON ANTIQUUS SKELETON OF UPNOR, KENT, ENGLAND! Illustrated by figures 1078-1084, 1074 One of the most fortunate events in the recent history of mammalian paleontology is the discovery and restoration of a nearly complete skeleton of an elephant (lacking only the skull). This skeleton was discovered in 1911-1912 in a trench near Upnor on the banks of the Medway, opposite Chatham Dockyard, in Kent, England. >. Thus the type of Hespero- loxodon antiquus ausonius exhibits an exceptionally high ridge formula (M 3 ;,—s5) for an Upper Pliocene! stage. Hisrory.—In 1875, Forsyth Major (following Falconer, 1868) discovered in the Upper Pliocene! of Italy extremely long molar teeth, supposedly related to H. [Hesperoloxodon] antiquus but specifically distinct, to which he gave the manuscript name of Elephas ausonius. Similarly in 1891, Pohlig observed teeth (in the Upper Pliocene! of Italy) which he erroneously referred to his Forest Bed species Elephas (antiquus) Nestiv. This renders it certain that in the Upper Pliocene! of Italy there was a relatively small narrow-toothed variety of the larger Hesperoloxodon antiquus of the Forest Bed level, clearly distinguish- ed by the name of Elephas [= Hesperoloxodon| ausonius; with the greater ridge formula: ‘Elephas ausonius’: M 3 y3s6 Grotocic Levet.—Depéret and Mayet (1928, p. 162) and Falconer (1868, Vol. II, p. 187) also describe Hlephas antiquus as found in the marine Pliocene of Rignano and in the lacustrine TyPE OF HESPEROLOXODON ANTIQUUS AUSONIUS Fig. 1087. Type of Hlephas ausonius Major. After Depéret and Mayet, 1923, Pl. x, figs. 1 and 2, p. 220: “Fig. 1 et 2.—Hlephas ausonius, de San Romano (Val d’Arno inférieur), piéces types de l’espéce. Msg droite et gauche. (Voir p. 166.) Institut géologique de Florence. Photographie du professeur Stefanini.” Third lower molars of the right and left side. About one-fourth natural size. Pliocene of Astésan, at San Paolo de Villafranca: “‘. . . two last upper and two last lower molars, also from St. Paolo: . .. Each of the upper teeth consists of nineteen plates, the rear part being broken off. . . . Of the lower molars, the right shows twenty plates, and is very narrow for its height.” Forsyth Major (1875) first observed in the upper layers of the Val d’Arno inférieur at San Romano teeth related to EH. antiquus but sufficiently distinct to be separated under the manuscript name of “Elephas ausonius.” Depéret and Mayet note that F. Major’s manuscript name, cited by Weithofer (1891 [1890], p. 194), quoted by Verri (1886) from labels in different museums, deserves to be re- vived and retained. Pohlig (1891, pp. 303, 350), while also clearly discerning the differences between the Upper Pliocene form and the [See footnote 1 on page 1049 above regarding the possible Lower Pleistocene age of the Villafranchian.—Kditor.] THE LOXODONTINA: true E. antiquus, erroneously refers an Upper Pliocene! skull (Fig. 1041 of the present Memoir) to his “‘special race,’ namely, Elephas (antiquus) Nestiz; this skull is probably a referred Hesperoloxodon antiquus ausonius. Consequently the type and referred specimens of this true Upper Pliocene H. ausonius appear in the previous literature under various names as follows: 1868 Vol. II, p. 187 EHlephas antiquus ref., Falconer. 1868 Vol. II, p. 250 Hlephas armeniacus ref., Falconer. 1875 Elephas ausonius F. Major MS. in collection labels. 1886 p. 453 Hlephas ausonius F. Major, in Verri. 1890 p. 194 Hlephas antiquus ref., Weithofer. 1891 p. 303 Elephas (ant.) Nesti? ref., Pohlig. 1923 pp. 162-166 Llephas ausonius type, Depéret and Mayet. Elephas ausonius ¥. Major, MS. in collection labels, 1875; in Verri (1886, Soc. Geol. Boll. [Ital.], V, p. 453); in Depéret and Mayet “Les Eléphants Pliocénes,’”’ 1923, p. 162. TyPE.— Third lower molars of the right and left side. Horizon AND Locatity.—San Romano, Upper Pliocene! of Italy. TYPE Freurr.—Not figured by Forsyth Major; Depéret and Mayet chose the same types as those described by Major (Depéret and Mayet, 1923, Pl. x, figs. 1 and 2). Originals in the Geological Institute of Florence. Typr Drscription.—(Depéret and Mayet, 1923, p. 166): “Le Musée de Florence contient, de cette localité, deux belles M3 du méme sujet, qui ont été déterminées par F. Major et doivent étre regardées comme les préces types de l’espéce; nous les figurons pl. x, fig. 1 et 2. Leur couronne est remarquablement étroite et allongée. La dent gauche, la mieux conservée, comprend 13 lames en action, plus 7 autres lames intactes 4 l’état de colonnettes isolées, soit en tout 20 lames, plus les talons. Du cété droit, on voit 13 lames déja entamées, plus un nombre indéterminé de lames non encore dégagées de l’alvéole. La fréquence laminaire est 4 peu pres de 5, comme dans le crane de Florence. L’émail est finement et assez réguliérement plissé; il existe de forts sinus loxodontes en arriére dans les lames moyennement usées; ces sinus disparaissent en avant par l’usure. Longueur de la couronne, 240 millimétres, avec une largeur maximum de 74.” CHARACTERS OF HESPEROLOXODON ANTIQUUS AUSONIUS.—ACc- cording to Depéret and Mayet, the two type inferior molars, selected by F. Major for his manuscript name Elephas ausonius and revived by them, exhibit the following ridge formula and size; M 3 35; length of crown 240 mm.; width of same 74 mm. Con- sequently Depéret and Mayet describe and figure the same type lower molars as those named Elephas ausonius in manuscript by Forsyth Major. These type molars are shown in the accompanying figure 1087 after photographs furnished the authors, Depéret and Mayet, by Professor Stefanini. Pohlig (1891, p. 303) erroneously applied the name FL. Nestii to the rostrum of the cranium described by Nesti as E. meridionalis, Crane B; this piece is shown in our figure 1041, lower row, after Pohlig, 1891, p. 350, fig. 109. It is the same ros- trum as that figured by Weithofer, 1890, Taf. 1, fig. 2, as L. HESPEROLOXODON 1233 antiquus. Consequently it is now demonstrated clearly that the type of the species #. ausonius Major-Depéret-Mayet does not belong to the same race as the type of the subspecies Elephas (antiquus) nestii [= Parelephas? trogontherii nestii] Pohlig, from the Forest Bed of Norfolk. Rewation To E. antiquus.—Depéret and Mayet (1923, pp. 163-170) give a detailed description of similar Upper Pliocene! stages from specimens in the museums of Florence, Rome, Bologna, and Norwich, derived from various Upper Pliocene beds of Villa- franchian age, and conclude that the species E. ausonius is without doubt a direct ancestral form or Pliocene mutation of the Pleistocene species Elephas antiquus. In free translation: Hlephas ausonius is a Pliocene mutation of the typical Quaternary Elephas antiquus. (1) Its molars are one-third smaller than those of the typical Lower Pleistocene stage of Grays Thurrock (Essex). (2) The crowns of the molars, especially of Ms, are relatively longer than those of the Quaternary H. antiquus and equally narrow. (3) The ridge-crests are less elevated, the height ranges from 75 mm. to 160 mm. (in the type M; of San Romano), whereas in F. antiquus the ridge-plates range from 175 mm. to 240 mm. (in the tooth from Taubach). (4) The ridge formula does not vary greatly from that of HE. antiquus but in the type M; of San Romano there are twenty ridge-plates, while in general the ridge-plate formula is from fifteen to sixteen. (5) The laminar frequency is from five to six lamine in 100 mm. (6) In EZ. ausonius the enamel thickness is equal to that of #. antiquus; it is more largely and more numerously crimped. (7) The loxodont sinus is generally strong and more striking in #. ausonius, but there are notable individual variations. (8) The most constant and most important character is the inferior dimensions of the molars of H. ausonius, the length of M 3 not exceeding 220 mm. in the superior teeth and 240 mm. in the inferior teeth; whereas Pohlig assigns to Z. antiquus of the Middle [Upper] Pleistocene of Germany 380 mm. for M* and 480 mm. for M3. Osborn, 1929: Depéret’s conclusion that ‘Elephas’ ausonius is directly ancestral to ‘E.’ antiquus is contrary to our present knowl- edge of the ridge-plate formule in these two species, namely: Typical Lower Pleistocene Hesperoloxodon antiquus: M 348-42 Typical Upper Pliocene! Hesperolorodon ausonius: M 374-5 - Despite its smaller size and narrower molars, Hesperoloxodon ausonius is More progressive in the addition of three posterior ridge-plates. While very primitive in its long, narrow proportions, it has a higher ridge-plate formula than either the Mediterranean dwarfed species or the North African species [of Palzoloxodon|], as shown in the comparative ridge-plate formule below; it exceeds the typical Palzoloxodon namadicus (M 3 73) and equals the progressive P. namadicus illustrated in figure 1073; the typical : 6 1 = Hesperoloxodon ausonius exceeds that of H. antiquus (M3 ie 12 BE (A Hesperoloxodon antiquus germanicus 8. Stefadnescu, 1924 Figures 1068, 1088-1091, 1114-1116, 1152, Pl. xx Type level: Geologic horizon of Weimar and Taubach, Germany. Type from Tanganu (Ilfov), Rumania. Upper Pleistocene. Now amplified by crania discovered at Steinheim (see p. 1253 below). Osborn, 1930: For the progressive Upper Pleistocene stage of ‘Elephas antiquus’ we may adopt the subspecific name germanicus Stefanescu, 1924, and amplify the characters of this stage from '[See footnote 1 on p. 1049 above, regarding the possible Lower Pleistocene age of the Villafranchian.—Editor.] R. M3 yey outer vie, Bril-/us. 16229 C R.M3 rex outer view R. MS yey outer view 1277 wear SSSEEF 10 tn wear 16 in wear ORD J¢ 1% wear as, R. Mz trrer vrew Jena Mus. WSS # Brit Mus. 39964 See HESPEROLOXODON ANTIQUUS (TYPICUS) - HESPEROLOXODON ANTIQUUS GERMANICUS After Faiconer After Fohlig /89/ HESPEROLOxoDON ANTIQUUS Amer: us. 22654 Jyoe ITALICUS Neotypes Fig. 1088. Progressive STAGES IN THE Evowurtion or HEsPEROLOXODON ANTIQUUS GRINDING TEETH All figures to the same seale, one-fifth natural size REFERRED Neroryprs Yreee Hesperoloxodon antiquus (typicus) third superior and inferior molars, r.M®, ].Mg. R.M®, from the Forest Bed, Ostend (Norfolk), England. Brit. Mus. 16229, Green Collection. Length 10 in. =254 mm., breadth 3.4 in. =85 mm., height 6.5 in. =166 mm. [Tallest ridge-plate 174 mm., 544 ridge- plates in 10 em., total ridge-plates 16). After Falconer, 1846 [1847, PI, XIV.A, fig. 5). Compare figure 1077, which shows another anterior plate, i.e., 1644 ridge- plates. L.M3, from Saffron Walden (Essex), England. Brit. Mus. 39464. Bowerbank Collection. Length 12.3 in. =315 mm., breadth 3 in. =77 mm., height 5 in. =126 mm. [Tallest ridge-plate 126 mm., 5}g ridge-plates in 10 cm., total ridge-plates 17.) After Falconer, 1846 (1847, Pl. XIV.A, fig. 11a]. Hesperoloxodon antiquus germanicus. Upper Pleistocene, 3d ] nterglacial stage of Taubach and Wei- mar, R.M3, external view (reversed), with 17+ ridge- plates, of which 10 are worn. Length 295 mm. (max. 315 mm.), breadth 80 mm., height 210 mm. (max. 235 mm.), [height of tallest ridge-plate 190 mm.], 6 ridge- plates (i.e., 7- 12) in 10 em. After Pohlig, 1891, Taf. 1 bis, fig. 7a, Original in Weimar Museum. Molar from Taubach. R.Ms3, external view, diagrammatic drawing, with 1835 ridge-plates, 20+ est. Total length of crown (actual 425 mam., estimated 480 mm.), breadth 85 mm., height 170 mm. [Height of tallest ridge-plate 128-130 mm., 4% ridge-plates in 10 cm.] After Pohlig, 1888, Taf. vi, fig. 1, Original in Jena Museum. great Taubach skeleton. From the 1234 TYPE Hesperoloxodon antiquus ttalicus, progressive stage from southern Italy. Right Second and third superior and inferior molars, original diagrammatic drawing after Amer. Mus. 22634, type. R.M? with 20 ridge-plates, height. of tallest ridge- plate 190 mm., 6 ridge-plates (1.e., 7-12) in 10 cm., 8 ridge-plates in greatly worn r.M2. Length 295 mm., breadth 82 mm. R.M3 with 18+ ridge-plates, height of tallest ridge-plate 128 mm., 414 (.e., 9-13) ridge-plates in 10em. In front is the greatly worn r.Mo, with 6 ridge- plates. Total ridge-plates in use above (M2, M3) 1243; total ridge-plates in use below (Mo, M 3) 13. These molars are placed in their natural position as found in the Jaw and skull of the type of Hespero- loxodon antiquus italicus of Pignataro Interamna, Italy. THE LOXODONTINA:: Pohlig’s monographic descriptions of Taubach grinding teeth, selecting the most characteristic specimens he has described and figured (Fig. 1088) as neotypes of Hlephas [= Hesperoloxodon| antiquus germanicus. In brief, Stefanescu designated as ‘Hlephas antiquus german- icus’ the Upper Pleistocene mutation of Weimar and Taubach. He designated Weimar and Taubach (Allemagne) as corresponding with the geological horizon of the type. The type itself, however, comes from Tanganu (Ilfov), Rumania (Fig. 1089). In the same notice Stefanescu characterized this subspecies as follows: Elephas antiquus germanicus S. Stefanescu, 1924. “Sur la présence de |’Hlephas planifrons et de trois mutations de |’Hlephas antiquus dans les couches géologiques de Roumanie.’”’ Compt. Rend. Acad. Sci., Paris, Tome 179, p. 1418, December 15, 1924. Typn.—Fractured crown of a second inferior molar of the right side, r.Mo, in the Laboratory of Geology, University of Bucharest. Horizon anpD Locauity.—Type from Tanganu (Ilfov), Rumania. Upper Pleistocene horizon of Weimar and Taubach, Germany. Typre Figure.—s. Stefanescu, 1927. Typr Derscription.—(Sabba Stefanescu, 1924, p. 1418, also 1927): The brief description of the subspecies germanicus is quoted with the context under Archidiskodon planifrons rumanus in Type oF HESPEROLOXODON ANTIQUUS GERMANICUS Vig. 1089. Type of Elephas antiqu- us germanicus 8S. Stefanescu, 1924. After Stefainescu, 1927, Figs. A and B of plate. (A) Posterior half of a second in- ferior molar of the right side, r.Ms, exposing 11 posterior ridge-plates (L. 1-11); 7-8 anterior ridge-plates wanting. (B) Six intermediate ridge-plates of the same inferior molar, r.Mo: “Face trituratrice de la couronne de M? [Mg] d’Elephas antiquus germanicus (A) et de ses six lames antérieures (Ly-Ls) détachées (B): talon postérieur (Tp), deux lames (Ly-Ly0) depouillées de cément, deux lames (Lo-Ls) en- veloppées de cément, sept lames (L7-L1) entamées par l|’érosion.” Chapter XVI, pp. 968, 969; it is also fully embodied in the follow- ing sentence: ‘Je n’insiste pas pour le moment sur les caractéres de cette mutation qui, 4 mon avis, est la plus rapprochée de l’origine mastodontide de Vespéce antiquus. J’ajoute seulement que la mutation ausonius de l’Italie lui suecéde et que la mutation de Weimar et Taubach (Allemagne), que je désigne sous le nom germanicus, est la plus récente.”’ HESPEROLOXODON SpecrFic CHARACTERS.—Comparing this type specimen with third superior and other molars of ‘Hlephas antiquus’ of Taubach and Weimar figured by Pohlig (1888-1891, Taf. 1 bis, fig. 7a (explanation of plate, p. 272), and Taf. v1, fig. 1'—Fig. 1088) we record the following dimensions, after Pohlig and Osborn, of the neotypes of germanicus. Length Breadth Height Ridge-plates Right M* 295-315 80 190-235 17+—18}5 Right M; 375-425 85 128-170 18+ EXPLANATION OF Figure 1088.—As shown in the partly dia- grammatic figure: (1) The progressive height of the upper ridge- plates (174-190 mm.) is measured on the enameled portions only and does not include the roots lacking enamel; (2) the progressive length of the superior molars (254-295, 315 mm. max.) is measured directly across the ridge-plates, as indicated by the oblique arrow; (3) the progressive inferior molars (815-425 mm.) are measured horizontally directly across the ridge-plates; (4) the height of the lower ridge-plates (126-128 mm.) represents the enameled portion of the tallest ridge-plate. Neroryee CHARACTERS OF HiSPEROLOXODON ANT. GERMAN- 1cus (Ponuic, 1888-1891, pp. 183-185, 298).—In Pohlig’s great Memoir he described a number of third superior and inferior grinding teeth from Taubach and Weimar, of which two especially typical examples from Taubach, as shown in figure 1088, are selected as neotypes. (1) Of the giant specimens from Taubach (“Theile des grossen Taubacher Skelettes”) are figured and de- scribed the superior and inferior molars of one individual (Taf. v1, figs. 1-6); the inferior molar (left), with 18+ ridge-plates, has a total length of 375 mm., the inferior molar (right) has an actual count of 18+ ridge-plates, total length 425 mm., estimated 20+ ridge-plates, total estimated length 480 mm. (or nearly 20 inches), breadth 85 mm., height 170 mm. (see Taf. vi, figs. 1, 1’. (2) Of the same Taubach skeleton are gigantic maxillary grinders in the Jena Museum (Pohlig, op. c7t., Taf. v1, figs. 2, 2a), namely, an r.M*%, with 18% ridge-plates (length 340 mm., breadth 98 mm., height 240 mm.), and an |.M’, with 17+ ridge-plates (length 320 mm., greatest breadth 98 mm., according to Pohlig, probably due to pressure). (38) Also from Taubach, in the Weimar Museum, an r.M’, with 17+ ridge-plates (Pohlig, op. cit., Taf. mr bis, fig. 7a); length 295 mm., (max. 315 mm.), breadth 80 mm., height 210 mm. (4) Another highly typical example of gigantic length is an ].M; from Weimar, in the Halle Museum (Pohlig, op. cit., Taf. v1, figs. 11, lla), with ?16+ ridge-plates in 330 mm.; length 410 mm., breadth 90 mm., height 180 mm. ERRONEOUS REFERENCE.—In 1924 the veteran palzontolo- gist of Hungary, Dr. Sabba Stefanescu, also described ‘Hlephas antiquus rumanus,’ a mutation from the Pliocene of Tulucesti, as an ancestral mutation of ‘Hlephas antiquus’; this specimen is referred by Osborn (p. 968) to Archidiskodon planifrons rumanus and is not regarded as ancestral to ‘H#. antiquus.’ Stefanescu adds that all three mutations are found in Rumania, namely, rwmanus, i.e., Archidiskodon planifrons rumanus, from Tulucesti (Covurlui), ausonius, i.e., Hesperolorodon antiquus ausonius, from Colintina (Ilfov), and germanicus, 1.e., H. antiquus germanicus, from Tan- ganu (Ilfov). 1236 OSBORN: THE PROBOSCIDEA TUSK OF HESPEROLOXODON ANTIQUUS GERMANICUS IN FIELD MUSEUM OF NATURAL HISTORY, CHICAGO [A deeade ago Dr. Henry Field acquired by purchase for Field Museum of Natural History an ‘Elephas antiquus’ tusk | = Hesperoloxodon antiquus germanicus], found at Steinheim on the Murr, Germany, which at that time was supposed to be the largest of the antiquus species in existence. While this tusk measures 2925 mm. or 9 ft. 7% in. in length, it is slightly smaller in 2950 mm. Fig. 1090. MaLe AND FEMALE TusKS OF HESPEROLOXODON ANTIQUUS GERMANICUS All figures one-twentieth natural size. Compare figure 1106, Hesperoloxodon antiquus italicus, one twenty-fourth natural size (Left) Right tusk (Field Mus. Nat. Hist. 201616) excavated at Steinheim on the Murr, Germany. Length on inner curve 2770 mm. or 9 ft. 1 in., on outer curve 2925 mm. or 9 ft. 7gin. Diagrammatic sketch furnished through the courtesy of Dr. Henry Field of Field Museum of Natural History, Chicago. (Center) Male and female tusks unearthed at Tonna, Germany, preserved in the Museum of Gotha. After Pohlig, 1888, Taf. 1, figs. la, 1b, 2,2a. Compare figure 1063 (Lozodonta africana). The male tusk (Taf. 1, fig. 1) measures 2950 mm. or 9 ft. 84in.; the female tusk (ig. 2) 1820 mm. or 5 ft. 11} in. 1820 mm. size than a male tusk from Tonna, Germany, preserved in the Gotha Museum, which, according to Dr. Hans Pohlig (1888, p. 49, Taf. 1—Fig. 1090 of the present Memoir), measures 2950 mm. in length or 9 ft. 8}s in. Furthermore, Dr. Fritz Berckhemer mentions (1930.1, Abb. 4) an isolated tusk in the Stuttgart Museum of even larger provortions, namely, 3390 mm. or 11 ft. 3500 mm. (Right) Right male tusk (Stuttgart Mus. 16274) found in 1929 at Steinheim on the Murr, Germany, measuring 3500 mm. or 11 ft. 594 in. Photograph kindly sent by Dr. Fritz Berckhemer. Doctor Berckhemer writes (letter, May 15, 1937) that the largest tusk of ‘Blephas antiquus’ in the Stuttgart Museum collections measures about 3750 mm. or 12 ft. 35 in.; the longest tusk recorded by Pohlig (op. cit., p. 51), excavated from the ‘‘schottern” (rubble) of the ancient river Arno at the Porta S. Lorenzo, is in the University of Rome and measures nearly 3900 mm. or 12 ft. 9% in. [=Hesperolorodon antiquus italicus of Rome—scee Fig. 1068 above]. The male tusks of Hesperolorodon antiquus italicus (Vig. 1106) measure 3070e mm. or 10 ft. /% in., within the alveolus 840-845 mm., free length beyond alveolus 2230- 2270 mm., a total of about 10 ft. THE LOXODONTINA:: HESPEROLOXODON 5% in., and in a letter of May 15, 1937, he states that the largest tusk in the collection found at Steinheim measures about 3750 mm. or 12 ft. 3% in. Doctor Field very generously offered Professor Osborn the privilege of first description in the present Memoir of the Field Museum specimen, an opportunity of which he intended to avail himself, but, as in other instances, his regrettable death prevented the consummation of his plan. The following description has been prepared with the kind assistance of Doctor Field, supplemented by the diagrammatic sketch of the tusk (Fig. 1091).—Editor.] Rieut Tusk oF HESPEROLOXODON ANTIQUUS GERMANICUS IN THE Couuections oF Firtp Museum or Naturau History, CHICAGO F.M.N.H. specimen No. 201616. Right tusk excavated at Steinheim on the Murr, Wiirttemberg, by the late Professor EH. Fraas, Stuttgart, from interglacial river-sands associated with on (A) sib oy dong we lr tore 292.5 ems (B) L850 Cms 1237 Rhinoceros merckii and Bison priscus. This specimen was pur- chased by Dr. Henry Field, leader of the Marshall Field Archeo- logical Expedition to Western Europe (1927-1928), from Dr. F. Krantz, Herwarthstrasse 36, Bonn, Germany, who very kindly supplied the above information. The length of the tusk on the inner curve is 2770 mm. or 9 ft. 1 in., on the outer curve 2925 mm. or 9 ft. 7% in., along the pro- jection of the base 2850 mm. or 9 ft. 4); in., and the maximum circumference is 520 mm. or 1 ft. 8% in. The curvature of the tusk is clearly shown in the accompanying sketch, which is reproduced herewith one-twentieth natural size; “the dotted lines indicate the probable terminal extension and the depth of the pulp cavity.” The alveolar portion cannot be determined. The ivory is in fair condition, although the tusk was broken into four sections. - The tusk has been repaired by Mr. L. L. Pray (who also made the diagrammatic sketch, reversed in this figure) and is now on exhibition in the Hall of the Stone Age of the Old World (Hall C) in Field Museum of Natural History. S16 Cen$ tre fren oboe bo thou curuatine | Fig. 1091. Wiirttemberg, by the late Professor E. Fraas, Stuttgart, from interglacial river sands, associated with Rhinoceros merckii and Bison priscus. Lateral and vertical views of right tusk of Elephas antiquus [=Hesperoloxodon antiquus germanicus] exacavated at Steinheim on the Murr, One-twentieth natural size. Compare figure 1090 (left). Sketch by Mr. L. L. Pray, reproduced through the courtesy of Dr. Henry Field. 1238 OSBORN: THE PROBOSCIDEA HESPEROLOXODON ANTIQUUS ITALICUS OF SOUTHERN ITALY AND HESPEROLOXODON ANTIQUUS GERMANICUS OF NORTH CENTRAL GERMANY A flood of light on the cranial structure of the ‘ancient’ or ‘straight-tusked’ elephant of Europe has resulted from the discovery in 1911-1912 at Pignataro Interamna, near Cassino, Italy, of the skull and jaws herein de- scribed as Palxoloxodon |= Hesperoloxodon| antiquus italicus, a subspecies of the classic ‘Hlephas antiquus’ of “aleoner. During the years 1926 and 1928 excavation in the region of Steinheim a.d. Murr, Germany, yielded to the Stuttgart Museum one exceptionally perfect cranium and another less perfect cranium which were finally restored RESTORATION OF HESPEROLOXODON ANTIQUUS ITALICUS Adult but not full grown. Shoulder height (fully adult) estimated at 13 ft. 6 in. (ef. p. 1250 below). One-fiftieth natural size. Vig. 1092. This restoration of Paloloxodon | =Hesperoloxodon] antiquus italicus is based entirely on the type cranium (Amer. Mus. 22634) combined with the skeletal characters of the Upnor ‘straight-tusked’ elephant Hlephas [Hesperoloxodon] antiquus. Compare figures 1079, 1080, 1096, 1098, 1105, 1099, 1100, and 1083 of the present Memoir, with accompanying skeletal measurements and comparisons (pp. 1245-1252). The relatively small ears are drawn from outlines by Paleolithic artists of North Africa and of Spain represented in figure 1047. Both in frontal and lateral aspects this ‘straight-tusked’ elephant is widely different from the recent African elephant. After crayon drawings by Margret Flinsch, 1931. and reconstructed by Dr. Fritz Berekhemer in 1929-1930, and herein referred to Hesperoloxodon antiquus german- icus of Stefanescu and Pohlig. Thus within a period of twenty-years (1911-1931) there came a most welcome mass of new cranial knowledge with a very important bearing on the evolutionary history and relationships of Falconer’s species ‘EHlephas antiquus.’ As shown in the preceding pages of this chapter, the ‘ancient’ or ‘straight-tusked’ elephant has previ- ously been known by isolated portions of the cranium, jaws, grinding teeth, and tusks, which can now be con- Sidered in their mutual mechanical relations, especially in the Italian specimen. THE LOXODONTINA:: HESPEROLOXODON 1239 LOXODONTA AFRICANA ALBERTENSIS, AFTER DRAWING BY MARGRET FLINSCH, UNDER DIRECTION OF AUTHOR. ONE-FIFTIETH NATURAL SIZE Fig. 1093. This combination drawing of a Central African elephant is based upon the bull elephant in the Carl E. Akeley group (Amer. Mus. Dept. Mam. 54085—see Fig. 1052 above) collected in 1909 by Mr. Akeley in the Budongo Forest, east of Lake Albert, Unyoro, Northern Uganda, also upon the Sudanese elephant ““Khartum” (Lozodonta africana oxryotis—Fig. 1053), formerly living in the New York Zoological Park, and upon a photograph by Marius Maxwell showing an elephant drinking at the river border. The tusks (measuring 8 ft. 5!4in. and 8 ft. 9% in. respectively and weighing 112 and 115 lbs. each) of a bull shot by Mrs. Akeley on the slopes of Mt. Kenya (Loxodonta africana peeli—Fig. 1059) furnished the basis for those in the present restoration. The longest tusks of the African elephant on record are the lyre- shaped pair in the Heads and Horns Collection of the New York Zoological Park (L. a. oxyotis—Fig. 1062), which measure 11 ft. 5} in., circumference 18}4 in. (right), 18% in. (left), with a combined weight of 293 lbs. The heaviest pair (although not the longest) recorded may be seen in our figure 1065, with a com— bined weight of 461 lbs. The tusks of L. a. oxyotis (Fig. 1056) were restored after these two record pairs. The large ears are drawn in extended position for contrast with the relatively small, low-set ears of Hesperoloxodon antiquus italicus (Fig. 1092). The height at the shoulder in the flesh, namely, 11 ft. 6% in. or 3520 mm., is after Ward’s skeletal record (1928) of 10 ft. 9% in. or 3290 mm. (ef. caption to Fig. 912, Chap. XVI, above). HESPEROLOXODON ANTIQUUS ITALICUS OF SOUTHERN ITALY Twelve localities have been recorded in which discoveries by Italian paleontologists of more or less perfect remains have been made in this region since the first note by Oronzio Costa in June, 1864. In July, 1926, the cranium and jaws of the ‘Pignataro In- teramna’ elephant were discovered and exposed by a farmer, Saverio Tiseo, while excavating for building purposes, in the extra- ordinarily perfect condition shown in figure 1096 and most for- tunately reported to Professor Giuseppe De Lorenzo, Director of the Institute of Geology of the University of Naples and a member of the R. Accademia Nazionale dei Lincei. Professor De Lorenzo promptly made a preliminary communication (1926) on this most important discovery, and in the following year (1927) published, with the codperation of Professor Geremia D’Erasmo, also of the University of Naples, a superb memoir entitled ““L’Elephas anti- quus nell’Italia Meridionale”’; this memoir (pp. 1-35) affords a valuable review of the discoveries previously made in the valley of the river Liri (see Fig. 1094) in the following localities: Castelliri Casalvieri Isoletta Arpino Roceasecca Pontecorvo Fregellae Cassino (grotto) PiGNaTaRo INTERAMNA Ceprano Aquino Caianello Gxrotocic Acu.—According to recent estimates of Pleistocene time, a 500,000-year interval elapsed between the typical Lower = : CASTELUR' ARPINO CEPR ISOLETTA PONTECOR' Upper PLEISTOCENE HorIzON OF HESPEROLOXODON ANTIQUUS ITALICUS Compare Osborn, 1931.846, p. 4, fig. 3, also figure 1097 of the present Memoir. Fig. 1094. Valley of the Liri River displaying the principal exposures along the eastern and western banks and slopes of the bordering hills, varying from 60 to 70 meters above the present sea level, of a thickness of about 50 meters, where the re- mains of Hesperolorodon antiquus italicus, of Hippopotamus, of Cervus, and of other Pleistocene animals have been found. After De Lorenzo and D’Erasmo, 1927, p. 7, fig. 1. Upper portion only. Scale 1:3,000,000. 1240 OSBORN: THE Pleistocene ‘Blephas antiquus’ Falconer and the new subspecies herein described, which is even somewhat more progressive than the ‘Elephas germanicus’ Pohlig of Taubach-Weimar. The more or less fragmentary mammalian remains found in proximity to the type include the following (see Fig. 1095): (1) Cervus elaphus Linn.: (a) Three antlers from the left side, (b) two antlers from the right side, and (c) right astragalus. Stag. (2) Bos primigenius Boj.: (a) Portion of left My and (b) frag- ment of mandible. Primitive ox. (3) Palzoloxodon [Hesperoloxodon| antiquus italicus: (a) Frag- ment of r.Ms, (b) symphysis of mandible, (c) a nearly complete female tusk, (d) right scapula (probably belonging to the type), and (e) central portion of a right humerus. Ancient elephant. (4) Hippopotamus amphibius Linn. (Pleist.=H. Cuvier): (a) Complete mandible, (b) cervical 5, (c) left meta- tarsal IV, and (d) left femur (juvenile). African hippopotamus. major (5) Rhinoceros merckii Kaup: (a) Right radius and (b) cervi- cal 7. Merck’s rhinoceros. (6) Associated with the mammal fauna was the imperfect shell of a freshwater mollusc, referable to the genus Unio, of the family Unionide. PROBOSCIDEA DESCRIPTION BY DE LORENZO AND D’ERASMO Dr Lorenzo AND D’ERAsMo (1926, 1927).—On pages 35 to 39 (see also Tav. 1) of the De Lorenzo and D’Erasmo Memoir of 1927 is given a complete description of this superb cranium in its original state (as cited in full from De Lorenzo’s original contribu- tion of 1926, pp. 185-188), of which the following is a literal translation in part?: “Tn the past month of July [1926], the farmer Saverio Tiseo, of Pignataro Interamna near Cassino, excavating, for building purposes, a piece of ground on his farm situated on the southern slope of the hill which borders the village and is really in Fon- tanarosa, found, at a depth of about 8 metres, a large cranium of a mammifer.”’ “The locality in which the fossil in question was found forms, with the hill of Pignataro Interamna, part of this whole system of gently rolling hills, which extend from Aquino and Pontecorvo along the left bank of the Liri as far as the river Rapido below Cassino, and which are made up of large alluvial deposits of the early quaternary, deposited first among the chain of the Aurunci and that of the Mainarde, later moulded by backwaters, defluents in lesser volume of the present and more confined course of the river Liri, which still today, as in the day of Orazio, continues with its calm waters the taciturn corrosion of its plains.” MAMMALIAN FossiLs ASSOCIATED WITH THE TyPpE CRANIUM OF HESPEROLOXODON ANTIQUUS ITALICUS About one-thirteenth natural size After Osborn, 1931.846, p. 14, fig. 11 Fig. 1095. Type right scapula (3d) of Palzoloxodon [Hesperoloxodon) antiquus italicus. (3a, b, c, e) Fragment of right second inferior molar (r.M»), symphysis of mandible, nearly complete female tusk, and central portion of right humerus of Palxoloxodon {| =Hesperoloxodon| antiquus italicus, (3d) right scapula (probably belonging to type). (4a, b, c, d) Mandible, 5th cervical, Mts. IV, juvenile femur of Hippopotamus amphibius major ref. (5a, b) Right radius and 7th cervical of Rhinoceros merckii ref. (2a, b) Portion of left inferior molar (M;) and fragment of mandible of Bos primigenius. (la, b, c) Portions of three antlers from the left side, two antlers from the right side, also right astragalus of the stag Cervus elaphus. ‘Kindly prepared by Miss Francesca LaMonte of the Department of Ichthyology of the American Museum of Natural History. THE LOXODONTINE: HESPEROLOXODON 1241 PIGNATARO INTERAMNA CRANIUM (TYPE) BEFORE REMOVAL, SHOWING (LEFT) THE ORIGINAL OWNER SAVERIO TISEO After Osborn 1931.846, p. 2, fig. 1 Fig. 1096. Type cranium in situ of Palzoloxodon [Hesperoloxodon] antiquus italicus (Amer. Mus. 2 Interamna, near Cassino, Italy, and measured and described by Giuseppe De Lorenzo in 1926 and 1927. Compare Tav. 1, 1927 by De Lorenzo and D’Erasmo. “The present plains, formed in recent times by the Liri, extend below Pignataro Interamna to about twenty metres above sea level; while the hills mentioned above, formed by the early quaternary diluvium, vary between 60 and 70 metres, thus giving a thickness of about 50 metres to the uncovered Pleistocene ground. This territory, prevalently clayish in the lower parts, becomes on top sandy,—yellowish sand and gravel interspersed with layers of clay and mud, and, on top, voleanic cinders.” “Tn this early quaternary district some time ago there were already found remains of fossil mammifers, especially elephants. These have been preserved in part in the Museum of Geology and Paleontology of the University of Naples. Oronzio Costa first noted them in the Rendiconti della Reale Accademia di Scienze fis. e mat. di Napoli for June, 1864. They were fully described by Giustiniano Nicolucci in his memoir Sw gli elefanti fossili della Valle del Liri [Concerning the fossil elephants of the Valle del Liri] (Memorie della Soc. ital. delle Scienze, detta dei XL, vol. IV, 1882). Cacciamali contributes further to this in the Bollettino della Societa geologica italiana, 1890, describing some molars of the elephants of Val di Comino and of Aquino. Finally, they are mentioned by Antonio Weithofer in his memoir on the fossil Proboscideans of Valdarno (a memoir which serves as the descrip- tive matter for a geologic atlas of Italy, vol. IV, part 2, Firenze, 1893). Hans Pohlig also speaks of this in his big monograph on Elephas antiquus, published in the Nova Acta Academiae Caes. 2634) as found and exposed by Saverio Tiseo at Pignataro figs. 1 and 2, of the Memoir of °Pignataro Cajazeo® i 1,18605¢ apua Veter ye Upper PLEISTOCENE Horizon OF HESPEROLOXODON ANTIQUUS ITALICUS Compare Osborn, 1931.846, p. 3, fig. 2, also figure 1094 for details Fig. 1097. Pignataro Interamna is near Cassino (circle), southwestern Italy, about fifty miles north of Naples. Region of the Valley of the Liri (Liris) occupied in Pleistocene time by large herds of the ‘ancient’ or ‘straight- tusked’ elephant now known as Hesperoloxodon antiquus italicus, also by Hippopotamus, Cervus, Bos, Dicerorhinus, and other species of 3d Interglacial time. After Pl. 94 of the Century Atlas, edition of 1913. 1242 OSBORN: THE Leopold. Carol. Germanicae naturae curiosorum, vol. 53, Halle, 1889, and vol. 57, Halle, 1892.” “My own and the observations of others are gathered together in my ‘Geologia e Geografia fisica dell’Italia meridionale,’ Bari (Laterza), 1904. On page 157 I spoke of the certain existence of Elephas (Euelephas) antiquus Fale. in the early quaternary deposits of the Valle del Liri.”’ “The fact that remains of fossil elephants already existed in the Valle del Liri does not diminish the importance of the present discoveries at Pignataro Interamna; of really exceptional impor- tance because of the completeness of the exhumed cranium and because of its position, a position which leads to the deduction that it was found in its original posture, not a secondary one caused by transportation, and this leads to the hope that it may be con- nected with the rest of the animal’s skeleton.” Dr Lorenzo's ORIGINAL SKETCH AND MEASUREMENTS OF THE TYPE CRANIUM or HESPEROLOXODON ANTIQUUS ITALICUS Compare Osborn, 1931.846, p. 8, fig. 7 Fig. 1098.—Diagrammatic sketch, prepared in the American Museum to aid in the eighteen months’ process of reconstruction. Cranium of Palzolozo- don [Hesperoloxodon| antiquus italicus (Amer. Mus. 22634), based upon original photographs (Fig. 1096), showing the exact measurements recorded in De Lorenzo’s contribution of 1926 and fully quoted in his Memoir of 1927. The American Museum reconstruction, completed November, 1930, in front view accords exactly with the 1926-1927 measurements of De Lorenzo, except as to width of rostrum [857 mm., 520 mm.], as follows: Apex of right tusk to vertex of cranium 3500 mm. 11 ft. 5%4in. Vertex of right tusk to border of premaxillary socket 2200 7 254 Lower border of premaxillary socket to vertex of cranium 1400 4 7% Diameter of incisive tusk at exit from socket 150 5% Transverse breadth across premaxillary sockets 900 [857] 2 11°%[9%4] Space between inner sides of incisive tusks 500 1 7% Transverse space across rostrum just below orbits 420 [520] 1 446 [814] Midline of premaxillary rostrum to midline of nasal opening 750 2 56 Transverse across narrowest portion of frontals 800 2 7% PROBOSCIDEA THE CRANIUM AND TUSKS “The enormous head rests with its longitudinal axis, which measures not less than 3.50 metres from the frontal protuberance to the apex of the tusks, in an almost perfect horizontal position (PI. 1, figs. 1 and 2) in such a way as to lead one to suppose that the animal, descending to bathe in a muddy and richly vegetated brook, sank in the sand and mud, and, unable to swim, tried to keep its head and proboscis above in order to breathe as long as possible, until it sank altogether, and a lower alluvial deposit than that of its prey covered and surrounded the creature. This hypothesis is strengthened by the fact that in close proximity to the head and interspersed among the yellow gravel is a soft black layer, muddy and agreeing with the herbaceous vegetation of the bottom of the swamp in which the elephant probably sank. For these reasons it is anticipated that probably behind the head, still in situ, there is to be discoveed all the skeleton of the elephant, which can therefore be dug out, given the opportunity.” “But the excavation is not easy, not so much because of the depth as because of the easily shattered nature of the skeleton itself. The bones excavated up to now, that is, those of the head, as well as being petrified are, as it were, decalcified, in such a way that to isolate and gather them, there is need of great delicacy and accuracy.” “Tn this formation and in such conditions now lies the cranium of the elephant, truly imposing in its silent grandeur. From the peak of the cranial protuberance [vertex of cranium] to the distal border of the intermaxillaries it is 1.40 metres long, its width on the frontal line between the parietals is 0.80 metres. Characteristic is the fan-shape of the two intermaxillaries, which together, below the nasal cavity, measure 0.42 metres [520 mm.]wide, while on the anterior border, concave and circular, they widen to 0.90 m. [857 mm.], keeping in the free internal space between the two inci- sive tusks a width of 0.50 m. and having a total length, along the median suture, of 0.75 m. The two incisive tusks, 0.15 m. in diameter at the exit of the sockets and 2.20 m. long, very beauti- fully and perfectly formed and gently and elegantly curved both on the inner and outer curves, diverge widely, following the external line of the intermaxillaries in such a way that 0.45 m. from the exit of the sockets they are already a metre apart, and the two apices more than two metres from each other.” “The mandible is so closely adherent to the upper maxillary that its symphysis appears to be almost soldered to the internal surface of the intermaxillaries. The symphysis is wide, near the condyles, about 0.60 m., with as much again in measurement on the bisettrice [midline?]. On detaching the cranium, the plates of the upper molars, devoid, as we have said, of cement, and deprived of support, fell at once. The lower molars of the mandible stayed in place; especially on the right are of the mandible one can see plainly the traces of the abrasion of the first (or second) molar, 0.10 m. long, 0.07 m. wide, with six residual plates, and the second (or third) molar, 0.17 m. long, 0.07 m. wide, with 10 plates.” ‘All these characters, and especially the length of the cranium, the prominence of the protuberance and of the occipital fossa (see Pl. 1, fig. 1), the enormous divergence of the intermaxillaries and of the incisive tusks (PI. 1, fig. 2 and interpolated figs. 10 and 11), the frontal depression, the narrowness of the plates of the molars in proportion to their height and to the length of the molars them- THE LOXODONTINA: HESPEROLOXODON 1243 selves, the form of the difese [framework?], show clearly that the cranium from Pignataro Interamna belongs to the species Hlephas (Huelephas) antiquus Falconer, characteristic of the interglacial phases of the early quaternary and the largest of the few species of elephants which have inhabited the earth. The fortunate dis- covery of this complete cranium further shows that, contrary to what Pohlig wrote, Falconer was right in maintaining that his Elephas antiquus was closely allied to the living Indian elephant, Euelephas indicus.” The above citation from G. De Lorenzo and G. D’Erasmo (1927), pages 35-39, figures 10 and 11, and plate 1, figures 1 and 2, affords invaluable information as to the Pignataro Interamna cranium in its original undisturbed condition partly buried in the matrix. These drawings and photographs demonstrate the superb condition of the cranium and tusks when first exposed and our knowledge is fortunately amplified by two photographs (repro- duced in our Fig. 1096) subsequently taken by Saverio Tiseo before he attempted to remove this priceless fossil from its original bed in the matrix. The precise knowledge of the specimen in its original condition afforded by these four photographs, also by the measure- ments and outline sketches by De Lorenzo reproduced in our figures 1099 and 1100, is in close accord with the measurements in De Lorenzo’s original paper of 1926, pages 187 and 188, as re- $1,000.00 by Henry Fairfield Osborn, mounted step by step to $4,375.34, almost nine times the amount originally set aside for the purpose. On opening the boxes containing the fossil, the extremely painful discovery was made that between 1927, when the negotia- tions of Tiseo with De Lorenzo were concluded, and 1929, the owner Saverio Tiseo had irretrievably damaged the entire upper Der LoRENzO’s ORIGINAL SKETCHES OF THE TYPE CRANIUM OF HESPEROLOXODON ITALICUS One-fortieth natural size Fig. 1099. Pignataro Interamna cranium (Amer. Mus. 22634) in situ. After De Lorenzo and D’Erasmo, 1927, p. 37, fig. 11: ‘“Cranio dell ‘21. antiquus di Pignataro Interamna, visto di fianco, ancora parzialmente immerso nella sabbia (Mo della grand. nat.).’’ Reproduced same size. Compare Osborn, 1931.846, p. 5, fig. 5. produced in our diagram (Fig. 1098). Had it not been for these priceless measurements, sketches, and figures, we should find ourselves obliged to record one of the most tragic losses in the history of vertebrate paleontology, namely, the characters of the cranium, jaws, and tusks of an adult Hlephas antiquus in a perfect condition of preservation. ACQUISITION BY THE AMERICAN MUSEUM IN THE YEAR 1929 In the hope that the excavations would be continued and the precious remains would become a part of the collection of the State, Professor De Lorenzo, as Director of the Institute of Geology of the University of Naples, made every effort (1927, p. 39) to secure the specimen for the Naples Museum, but without success. During the following year (September 4, 1928), the American Museum of Natural History was informed of the desire of Saverio Tiseo to dispose of the specimen and began negotiations (No- vember, 1928) on condition that no step would be taken without due permission from the authorities of the Italian Government. On December 3, 1928, these terms were formulated in detail, and in May, 1929, the specimen was received in the American Museum. The total cost to the American Museum, including the donation of Fig. 1100. Front view of the Pignataro Interamna cranium (Amer. Mus- 22634) in situ. After De Lorenzo and D’Erasmo, 1927, p. 36, fig. 10: “Cranio dell’. antiquus di Pignataro Interamna, visto di fronte(}4o della grand. nat.).’” Reproduced same size. Compare Osborn, 1931.846, p. 5, fig. 4. portion of the cranium by attempting to remove it for purposes of exhibition. Thus the superb and unique cranium shown in Profes- sor De Lorenzo’s description and in the figures and photographs above mentioned and reproduced in the present text no longer existed. The remaining parts of the specimen, namely, the rostrum, tusks, palate, jaws, and the lower portion of the occiput were also seriously damaged, while the entire upper portion was irrevocably lost to science with the exception of three small pieces extricated with great difficulty from the hard cement in which Tiseo had attempted to repair the terrible injury which he had inflicted on this priceless specimen. At first the reconstruction of the skull appeared hopeless, and the present author, who had donated it to the American Museum collection on the basis of the excellent photographs showing the specimen in its original condition, was not even allowed to see it in the laboratory. After eighteen months of arduous labor on the part of Mr. Jeremiah Walsh, under the direction of preparator Charles Lang, and of Curator Barnum Brown, and finally of Honorary Curator-in-Chief Osborn, the reconstruction entered its final stages in which the precise measure- ments, figures, and photographs secured by Professor De Lorenzo of the cranium in its original unfractured condition were of in- 1244 Fig. 1101. (Amer. Mus. 22634). One-fourth natural size. After original photographs, retouched and numbered, displaying ridge-plates 1 to 20 in r.M®, ridge-plates 4to 12inr.M?. Compare Osborn, 1931.846, p. 10, fig. 8. (Upper) Second and third right superior molars, r.M?, r.M®. (Lower) Palate exhibiting right and left superior molars, M?, M®. Type superior grinders of Hesperoloxodon antiquus italicus calculable value and importance. Thus, after almost continuous and very expensive labor between May, 1929, and November, 1930, the specimen was ready for complete description and exhibi- tion, although not open to the public until January 1, 1931. Despite all these drawbacks and scientific disappointments, the restored cranium and tusks, jaws, and scapula still afford a wealth of new knowledge regarding the relationships of the classic ‘Elephas antiquus,’ which, added to the equally priceless Upnor skeleton, also the Steinheim crania (Hlephas [Hesperolorodon| OSBORN: THE PROBOSCIDEA antiquus germanicus), remove this great branch of the family Elephantide from its previous obscurity and uncertainty and enable us to rank it as among the best known of the fossil elephants of Eurasia. Whereas Hlephas |Hesperoloxodon| antiquus belongs in the Lower Pleistocene, or 1st Interglacial, the present specimen belongs in the upper Middle Pleistocene,! or 3d Interglacial; it is somewhat more progressive than the H. antiquus germanicus of Weimar. Hesperoloxodon antiquus italicus is far superior in size to Palxoloxodon namadicus and greatly surpasses H. antiquus typicus both in the number and height of the ridge-plates (see Figs. 1102 and 1088). It is a very progressive ascending mutation, equaling in size, but exceeding in the number of its superior ridge-plates, the most progressive H. antiquus germanicus of Weimar. How- ever, from close comparison with all the numerous specimens de- scribed from Lower Pleistocene deposits in England (by Falconer and others) to 3d Interglacial deposits in Weimar (by Pohlig and Soergel), H. antiquus italicus appears to be the largest and most progressive member of the ‘Hlephas antiquus’ phylum thus far discovered. C N \\, ANY HESPEROLOXODON ANTIQUUS ITALICUS Amer: us. 226354 Type- Fig. 1102.—Type right superior and inferior grinders, M 2, M 8, of Hespe- roloxodon antiquus italicus (Amer. Mus. 22634). Diagrammatie key to the superior and inferior ridge-plates: M 2 12% Mss The principal measurements of r.M® are: Length of 20 ridge-plates 295 mm.; height of 5th ridge-plate 190 mm.; 6 ridge-plates in 10 cm. The principal measurements of r.M3 are: Length 306 mm., height of 8th ridge-plate 128 mm., 44 ridge-plates in 10 em. After Osborn, 1931.846, p. 13, fig. 10. 1(Careful consideration of the several statements by Professor Osborn in the present Memoir, as well as the general opinion today of geologists and paleon - tologists, has led to the conclusion that Professor Osborn would have placed the 3rd Interglacial in the Upper Pleistocene; hence throughout it will be observed gists, PI & that this determination has been adopted.—Kditor.] THE LOXODONTINA:: HESPEROLOXODON i Fig. 1103.—Type mandible, right lateral and superior aspects, of Hespero- Joxodon antiquus italicus, one-sixth natural size. A, Right lateral view, with second and third superior teeth, M?, M°, super- posed on corresponding inferior teeth, Mo, Ms. After Osborn, 1931.846, p. 16, fig. 12. B, Superior view, with second inferior molar, Mo, exhibiting ridge-plates 7-12, third inferior mclar, Mg, exhibiting ridge-plates 1-8. After original photograph. [See footnote on opposite page.—Kditor.] 1245 Fortunately the second and third superior and inferior molars of both sides were preserved in situ and their characters are very clearly displayed in the accompanying type figures, in which all the ridge-plates are shown both in crown and lateral view, with clear enumeration of the ridge-plate numbers in figures 1101, 1103, and 1104, and in the diagrammatic figure 1102. Very important is figure 1011 (lower) in which 15 ridge-plates are shown in simul- taneous use, namely, r.M?, ridge-plates 4 to 12', plus r.M?°, ridge- plates }s-1 to 6. male, corresponding with the attrition of Hlephas indicus estimated This stage of attrition represents a young adult to be about forty years of age. To the 20 ridge-plates actually observed in r.Mé (Fig. 1101) there may possibly be added ridge- plates 21 and 22; whereas in a much older individual of the typical Hesperolorodon antiquus only 16%-17 ridge-plates are shown. SUBSPECIFIC DESCRIPTION Compare Osborn, 1931.846, pp. 17-24 Hesperoloxodon antiquus italicus Osborn, 1931 Figures 1068, 1069, 1088, 1092, 1095, 1096, 1098-1108, 1111, 1112, Pl. xxi Pignataro Interamna, near Cassino, Italy. gravels of 3d Interglacial time. Upper Pleistocene river Palxoloxodon antiquus italicus Osborn, 1931. ‘‘Palxoloxodon antiquus italicus sp. nov., Final Stage in the ‘Hlephas antiquus’ Phylum.” Amer. Mus. Novitates, No. 460, pp. 1-24. TyPr.— Cranium and jaws with superior and inferior dentition (M 2—M 3) of a young adult male; also right scapula (Amer. Mus. 22634). Locauiry AND Horizon.—‘‘Pignataro Interamna, near Cassino, Italy. Found in upper Middle! Pleistocene river gravels, of 3d Interglacial time, equivalent to, or slightly more recent in geologic age than, the 3d Interglacial stage of Taubach- Weimar of the Ilm River valley, Saxe-Weimar, Thuringia, northern Germany, 40 kilometers east and a little south of Burgtonna, north of Gotha, where the first skeleton of ‘Hlephas antiquus’ (cited by Blumenbach as Elephas primigenius) was discovered in 1695.” Tyrer Figure.—Op. cit., figs. 1, 4-16. AA WNalural Szze Fig. 1104.—Type of Hesperoloxodon antiquus italicus (Amer. Mus. 22634). -Inner view of third right inferior molar, r.M3, exhibiting +-18+ ridge-plates. One-fourth natural size. After Osborn, 1931.846, p. 12, fig. 9. Amer. Mus. 22634 Ying. Nal. size PALAEOLOXODON ANTIQUUS\. ITALICUS Zyoe 4 Fig. 1105. HesprROLOXODON ANTIQUUS ITALICUS TYPE, OF PIGNATARO INTERAMNA, ITALY, VALLEY OF THE LirgI, IN THE AMERICAN MusEeuM or NATURAL History (Amer. Mus. 22634) After photographs and indicated measurements (March 10, 1931) by the present author. Reduced to a uniform scale of one-sixteenth natural size. See also type figure 1106. A, Left lateral view of cranium. Al, Anterior view of cranium, exhibiting upturned portions of tusks in slight perspective. A2, Posterior view of cranium and jaws, exhibiting exposed lower grinding teeth of the right side, tusks in strong receding perspective. Observe the relatively high, narrow occiput exceeding in height that of Hesperolorodon antiquus germanicus. The adult cranium (A) is about equal in depth or bathycephaly (990e mm.) to that of H. ant. germanicus (980 mm.—Fig. 1114, B); this is owing to the greater depression of the tritural surface of M® below the occipital condyle. 1246 THE LOXODONTINA: In the following description (Osborn, 1931.846, pp. 17-24) references to the figures in the present Memoir have been substi- tuted for the original figure numbers. “SUBSPECIFIC CHARACTERS.— Displayed in the measurements, ridge-plate formule, and height of the ridge-plates, enumerated below; also illustrated in the type figures 1101, 1102, and 1103. Progressive evolution indicated as follows: Pal. ant. typicus;"! M3 16 M$, length 254 mm., height of tallest ridge-plate 174 mm.; M3, length 315 mm., height of tallest ridge-plate 126 mm. 11+ Pal. ant. germanicus; M 3 74%. M?’, length 295 mm., height of tallest ridge-plate 190 mm.; M;, length 425 mm., height of tallest ridge-plate 120 mm. Pal. ant. italicus (type); M 3 725. Mj, length 295 mm., height of tallest ridge-plate 190 mm. ; Mz, length 306 mm., height of tallest ridge-plate 128 mm.” “TncIsIvE Tusks.—The incisive tusks of the type are not fully grown [see Figs. 1100, 1099, 1106, 1107, 1108]. Total estimated length 3030-3070 mm., that is, 800e mm. within the alveolus plus 2230- 2270 mm. beyond the alveolar border. The longest tusk recorded by Pohlig in the University of Rome collection measures 3900 mm., or 12 ft. 9% in., in comparison with 10 ft., length of the present specimen. This indicates that a full-grown adult male of Pal. ant. italicus attained gigantic size.”’ “Lowmr Jaws [Fias. 1103, 1106].—The type inferior mandible is entirely complete, as represented in figure [1103], requiring little or no restoration. As compared with the more or less complete mandibles figured by Falconer, from the Lower Pleistocene of England, it closely resembles in profile aspect the typical ‘Hlephas antiquus’ jaw but is very much larger and more massive; it differs widely in every aspect from the mandibles belonging to any species of Archidiskodon, Parelephas, or Mammonteus; the rostrum is abruptly truncated but less abbreviate than in ‘Hlephas primi- genius,’ which is more of the extreme bathycephalic type.” “SKULL.—The extremely broad rostrum, characteristic of all stages of the ‘Hlephas antiquus’ phylum, measuring 857 mm. or 2 ft. 9%4 in., is exactly the same width as that of the ‘Hlephas platyrhynchus’ of Graells, which measures 860e mm. or about 2 ft. 10 in. transversely; this animal is from the Pleistocene, at San Isidro, near Madrid, Spain.” “Most novel and surprising is the dome-like, highly arched occipito-parietofrontal contour [Figs. 1099, 1100, 1106, 1107, 1108] which superficially resembles that of the Indian elephant more HESPEROLOXODON 1247 » PALAEOLOXODON ANTIQUUS ITALICUS Va Ail 24 Nat- size HESPEROLOXODON ANTIQUUS ITALICUS, TYPE One twenty-fourth natural size. See figure 1105, one-sixteent h natural size Fig. 1106. Type cranium of Palxoloxodon antiquus italicus (Amer. Mus. 22634) as reconstructed and mounted in the American Museum during 1929 and 1930. One twenty-fourth natural size. Restored parts (oblique lines); parts preserved (shading), namely, occipital condyles, portions of premaxil- laries and maxillaries, and complete mandible. After Osborn, 1931.846, p. 7, fig. 6. The measurements, as seen from the front, with but two exceptions accord with those given by De Lorenzo. The few original fragments seen in frontal aspect lie at the back of the narial chamber (c); along the border of the left temporal fossa (¢), and along the temporal arch (¢'); in lateral aspect of the left side, as shown in the fragments in the left temporal fossa (¢’, t*) just above the fragment of the right temporal arch (), transferred to the left side for purposes of restoration. In posterior aspect, the occipital parts preserved are portion of the superior border (o'), and parts of the posterior occipital plate (o?, o*) and the very broad condyles (0%). 11On page 17 of American Museum Novitates, No. 460, Osborn presents the descriptions of three subspecies of Palzoloxrodon antiquus, designated as P. antiquus typicus, P. antiquus germanicus, and P. antiquus italicus. According to current rules the name typicus, designating the type of the species antiquus, must be regarded as a still-born synonym of antiquus, since the name of the typical subspecies should be the same as the name of the species. Consequently the three subspecies would be: Palzxoloxodon antiquus antiquus (Falconer) Palxoloxodon antiquus germanicus (Stefinescu) On page 21 of this same publication, Osborn proposes the new genus Hesperolorodon, with Palxolorodon antiquus italicus as the generic type. a genus is based upon a species, so that the type of Hesperolorodon should be antiquus and all of its subspecies. between two or more genera on the basis of its subspecies. (E. H. Cotpert.) Palxoloxodon antiquus italicus Osborn But In other words, a species cannot be split Nevertheless the author’s plain intention was to recognize the distinction of a “new loxodont phylum of the west’”’ (as opposed to the Indian namadicus) ; as a “type” of the “phylum” or “genus’’ (terms often used interchangeably by him) he designated Palzoloxodon antiquus ttalicus. Since this was his intention the subspecies italicus should have been raised to the rank of a species to serve as a type of the genus Hesperoloxodon. (W. K. Greaory.)] 1248 OSBORN: THE closely than that of either Lorodonta africana or Palzoloxodon namadicus. ‘This lofty profile and corresponding bathycephaly are represented correctly in figures [1106 and 1107 of the present Memoir], because they accord . . . [Fig. 1098] with the measurements and photographs taken by De Lorenzo and Saverio Tiseo before this cranium was damaged. Moreover, beside the well preserved and extraordinarily broad occipital condyles (280 mm.), there are portions of the occiput ([Fig. 1106], 01, 0?, 0%, 0+) PROBOSCIDEA 12 ft. 1% in.; thus it appears from the cranial proportions only that the skeleton of Pal. ant. italicus was about fifteen per cent. taller at the shoulder than that of the African elephant and closely similar in height to that of the Upnor elephant.” COMPARISON WITH PALAOLOXODON NAMADICUS.—‘‘Examined closely, the above measurements prove that the cranium of Pal. ant. italicus is profoundly different from that of ‘#. namadicus’ (the genotypic species [!] of Palxoloxodon) as well as from that of COMPARISON WITH AFRICAN ELEPHANT (1) From front of orbit to back of occipital condyle Loxodonta africana peelt [Percentages plus Hesperoloxodon (+) and minus (—)] antiquus italicus (2) From summit of parieto-occipital crest to attritional surface of M? (bathycephaly) Occipital condyles, transverse Temporal arches, transverse width across Premaxillary rostrum, maximum width across Mandibular condyle, height above angle of jaw Mandibular length, condyle to apex of rostrum Incisive tusks: total length of free length beyond rostrum maximum diameter at exit from rostrum total circumference of (3) (4) (5) (6) (7) (8) which prove that the occiput is forwardly inclined. The chief comparative measurements between this restored cranium and the cranium of the adult African bull elephant (Lovodonta africana peeli—see Fig. 1107) are as [above].”’ Pohlig (letter May 5, 1929) gave a very high estimate of the length of the incisive tusks of ‘Hlephas antiquus’ of Germany, namely, an average length of 5000 mm., or 16 ft. 4% in., as compared with the incisive tusks of a Siberian ‘Elephas primigenius’ in the St. Petersburg Museum (4300 mm., or 14 ft. 14 in.). He adds: “The average incisor length of a Siberian Mammoth at Peters- burg, measured and figured by me (1891, p. 321, Pl. B, fig. 3) is 4 m. 30,—so that the average length of the Hlephas antiquus incisor (op. cit., p. 51) must have been more than 5 meters.” COMPARISON WITH LOXODONTA AFRICANA PEELI.—‘“‘In (4) width across temporal arches, L. africana [peeli| exceeds Pal. ant. italicus by six per cent. In percentages the cranium of Pal. ant. italicus in all other measurements is from seven to twenty- nine per cent. larger than that of L. africana, as follows: (1) Orbit to occipital condyles 10% (2) Cranial apex to grinding surface of M* 24% (3) Transverse occipital condyles 13% (5) Width across premaxillary rostrum 29% (7) Mandibular length Weiou “On the average of the five principal measurements, the cranium of Pal. ant. italicus is about fifteen per cent. larger than that of L. africana. If this fifteen per cent. obtains throughout the entire skeleton, the skeletal height of Pal. ant. italicus should be about 3673 mm. or 12 ft., as compared with the height zm the flesh of a large adult bull of L. africana, namely, 3450 mm. or 11 ft. 4 in., or with the skeletal height of the Palxolorodon antiquus (andrewsi?) [= Hesperoloxodon antiquus| of Upnor, from the top of the scapula to the ground, namely, 3700 mm. increase or 746 mm. +10% = 820e mm. 800 +24% =990e 248 +13% = 280 797 — 6% =750e 665 +29% = 857 489 — 2% =478 749 + 7% = 800 L R L R 1960e 2010e 3030e 3070e 1905 1960 2230 2270 156 160 145 139 460 467 433 432 Ee ANTIQUUS AFRICANA ITALICUS 7yge PEELI A. 17. 22634 A. 17. 2/889 Vig. 1107. CRaANIA or HeESPEROLOXODON AND LOXODONTA IN LATERAL VIEW After Osborn, 1931.846, p. 18, fig. 13, one twenty-fourth scale. Compare figures 1106 and 1108, also measurements above. (Left) Cranium and tusks of type of Palxoloxodon [Hesperoloxodon| antiquus italicus (Amer. Mus. 22634), right lateral aspect. (Right) Cranium and tusks of Loxodonta africana peeli (Amer. Mus. 21889), adult male, from Mt. Kenya district. Same as in figure 1059. ‘ADescribed by Matsumoto (1924) under the subspecific name Palzoloxrodon namadicus naumanni”’ [genotype Elephas namadicus naumanni Makiyama, 1924. See, however, footnote on p. 1247 above regarding genotypic usage.—Hditor. | THE LOXODONTINA: HESPEROLOXODON Loxodonta africana, which is a relatively primitive cranium. Coordinated with its very tall grinding teeth, the cranium of the Italian specimen is much more bathycephalic (depth 990e mm. as compared with 800 mm. in L. africana); it is correspondingly less brachycephalic (750e mm. as compared with 797 mm.); this is in accord with the cranial proportions which are much nearer those of A 8 G ANTIQUUS ITALICUS A./1 22654 .NAMADICUS Fale. /847 PEXLY AFRICANA AM. 2/889 L. AFRICANA A./1. 2/889 P. NAMADICUS Pilgrim /905 PI P. ANT. ITALICUS A./1.22634 CoMPARATIVE BATHYCEPHALY OF THE LOXODONTIN.® Fig. 1108. Frontal and lateral views of three adult males, one-fortieth natural size. After Osborn, 1931.846, p. 20, fig. 14. A, Al, Loxodonta africana. Bathycephaly, 800: 710 mm. Male. B, Palzoloxodon namadicus. Bathycephaly, 728: 592 mm. Female. B1, Palzoloxodon namadicus. Male cranium of the Godvari Alluvium. After Pilgrim, 1905. C, Cl. Palxoloxodon [Hesperoloxodon| antiquus italicus type. Bathy- cephaly, 990e.: 820e mm. Observe that Pal. [Hesperoloxodon| antiquus tlalicus is much more bathy- cephalic (990e: 820e mm.) than Loxodonta africana (800: 710 mm.), which is approximately the same as Pal. namadicus (728: 592 mm.). The female cranium (B) is much smaller than the large male cranium of the Godsvari Alluvium (B1); compare the more accurate diagrammatic figure (ig. 1110) of the same cranium. SCAPULA Height, superior border (restored) to center of glenoid border Width, median, across pre- and post-scapular borders Anteroposterior diameter of neck of scapula Anteroposterior diameter of glenoid border Equals Hesperolorodon antiquus—see footnote on p. 1222 above.—Editor.] PALALOLOXODON ANTIQUUS ITALICUS ye AM 22634 1249 the Indian elephant than of the African elephant. This bathy- cephaly is, however, a parallelism rather than a point of affinity, because the very broad rostrum of Pal. ant. italicus presents an extreme difference from the very narrow rostrum of Hlephas indicus. Comparison with the cranium of ‘Elephas namadicus’ shows a strong resemblance in the breadth of the premaxillary rostrum but an extreme difference in the summit of the cranium, which in ‘EZ. namadicus’ is relatively low and reinforced by the overhanging parieto-frontal crest.” “This points to Pal. ant. ztalicus as a member of a phylum quite distinct from that of the Siwalik ‘#. namadicus,’ a phylum which if supported by other cranial and skeletal differences might well constitute a new genus to which the name HrspEROLOXODON, or ‘loxodont of the west,’ might be applied. This name is provision- ally proposed, as I would not like to be forestalled a second time, as in the case of Palxolorodon, a generic name assigned to ‘EF. namadicus naumanni’ by Matsumoto but a few weeks prior to my description of Sivalikia. “ScapuLta.—The right scapula (1109A, Al) was found near the type and may be considered as belonging to the same individual (Amer. Mus. 22634); it greatly exceeds in size that of the African and Indian elephants of the same age [Fig. 1109C, D, E]. On the 28 Natural size CoMPARISON OF SCAPULS One twenty-eighth natural size Fig. 1109. [ =Hesperoloxodon| ant. italicus (Amer. Mus. 22634) drawn to the same scale as the corresponding scapula of (B) Pal. ant. (andrewsi?),! of (C) Lox. africana, and of (D) Elephas indicus, juvenile (both after Andrews and Cooper, 1928, fig. 2), also of (E) Elephas indicus, adult (Amer. Mus. 54453), Vernay’s middle- aged male, of which the entire forelimb is shown in figure 1194 of the present Memoir. Right scapula (A, Al in reversed outline) of the type of Pal. other hand, it is slightly exceeded in size by the left scapula [Fig. 1109B] preserved in the Upnor skeleton. The measurements of the scapule of Pal. ant. italicus and Pal. ant. (andrewsi?)! are taken with the restored border indicated in dotted lines: Pal. ant. italicus Pal. ant. (andrewsi?)! Pignataro Upnor [Amer. Mus. 22634] [Brit. Mus. | +15% =1065mm. +10% =1170 mm. Lox. africana oxyotis (Jumbo) {[Amer. Mus. 3283] 925 mm. 594 +29% = 770 +12% = 868e 240 +28% = 307 E697 324 187 +35% = 253 +15% = 290” PALAEOLOXODON ANTIQUUS NAMADICUS Pilgrim, /905, Vol. XXXII, Pls 10. 1/. 12 OSBORN: THE PROBOSCIDEA Larce Mate CraNniuM OF PALMOLOXODON NAMADICUS OF THE GODAVARI ALLUVIUM One-twentieth natural size Fig. 1110. Palzxolorodon namadicus, large male cranium discovered in the Godavari Alluvium, at Nandtir Madméshwar, India (cf. Pilgrim, 1905, Pls. 10, 11, 12). Measurements after Pilgrim and Osborn. Fronto-vertical length Transverse premaxillary rostrum Occiput; transverseswidthyacrossia5 6. sitet « scrac anja cei ameter © cele P. namadicus H. ant. italicus sects .sie/erd aileyeccvalerevereve eisue afaverelevenehe che 1270 mm. 1400 mm. andes vakeranelie leas) orev ener Stare LaPebed sv spaNet ORG fos Stes 920 857 v's iatayates qyatavero Weval ar cususcepextelacctepare nee 1075 875 This comparison shows that Palxolorodon namadicus exceeds in breadth, while Hesperoloxodon antiquus italicus exceeds in depth. “RSTIMATES OF SKELETAL AND FiesH Heicut.—It is im- portant to compare the estimates of the shoulder height derivable from the scapula as well as from the cranium; they are found to agree exactly, as shown below. Neither the Upnor nor the Italian specimen is full-grown, yet combined they afford a priceless means of estimating the height of the full-grown ‘EHlephas antiquus.’ ”’ “Tn each of the four above dimensions the scapula of Pal. ant. italicus is from fifteen to thirty-five per cent. larger than that of Lox. africana oxyotis (‘Juambo,’ Amer. Mus. Dept. Mam. 3283); the actual skeletal height of “Jumbo” is 10 ft. 5% in. or 3194 mm.; consequently if we add fifteen per cent. (the difference in scapular height) to the skeletal height of “Jumbo” we obtain 3673 mm. or 12 ft. % in. as the estimated skeletal height of the Pal. ant. italicus type; this agrees with the height estimated from the proportions of the cranium, namely, about 3673 mm. or 12 ft. % in.” “Tn height the scapula of Pal. ant. ctalicus is about ten per cent. less than that of Pal. ant. (andrewsi?)! of Upnor, the skeletal height of which is 3700 mm. or 12 ft. 19¢ in.” Pal. ant. italicus: Total skeletal height estimated from proportions of cranium 3673 mm. = 12 ft. % in. ea. Total skeletal height estimated from proportions of scapula Pal. ant. (andrewsi?)': Total skeletal height estimated from proportions of entire fore limb 3673 mm. = 12 ft. % in. ca. 3700 mm. = 12 ft. 1°s in.” “To this estimated skeletal height should be added about six and one-third per cent. to obtain the height in the flesh, giving us an estimated height at the shoulder of 3905 mm. or 12 ft. 934 in.” “Pal. ant. italicus of Pignataro Interamna, adult: Estimated height in the flesh 3905 mm.=12 ft. 9%4 in. Pal. ant. (andrewsi?)' of Upnor, young adult: Estimated height in the flesh 3934 mm.=12 ft. 10% in. Lox. africana, adult: Height in the flesh 3450 mm.=11 ft. 3% in.” “The Pignataro Interamna specimen is several years older than the Upnor specimen, as indicated by the fact that the posteri- or ridge-plates of the second molar and the anterior ridge-plates of the third molar [Fig. 1101] are in use, while in the Upnor specimen the ridge-plates of the second molar only are in use. Comparison with the growth of the large African elephant ‘“Khartum” in the New York Zoological Park shows that in captive conditions at the age of twenty-seven years the animal grows three-quarters of an inch a year. By such an estimate the Pignataro Interamna adult is about five years older than the Upnor young adult; had it con- tinued to increase in height, the fully adult bulls would measure about 13 ft. 6 in. in height, or two feet above the shoulder height of a large fully adult African bull elephant.”’ THE LOXODONTINA: HESPEROLOXODON 1251 FEMORAL LENGTHS AFTER POHLIG The humerus bears a more constant ratio to the height of an elephant than the femur, because the relative height of the fore and hind quarters differs in each phylum. Yet the femoral lengths where obtainable are very important as a key to pelvic heights. Consequently the following notes as to femoral lengths recently communicated by letter from Dr. Hans Pohlig, the leading his- torian of the fossil proboscideans of western Europe (whose portrait beside the femur of Rome appears in figure 1111 of the present Memoir) are of interest. Pou.ic, 1929-1931.—(Letter, ‘‘Bonn, 5.III, 29’): “The... antiquus femur (from Rome) is now at Lumbres, P. Calais (Dr. Pontier) [see Fig. 1111]. As my own height is 1 m. 70, the femur will be about 1 m. 50, but it is not quite entire and not quite adult; that of the large Taubach skeleton (the huge grinders of which are figured in my Mon., plate v1, fig. 2a, 1b, and the premaxillaries (average breadth 1 meter! ibid. fig. 3), is 1 m. 60 at least... . The largest [Archidiskodon| meridionalis femur at Florence is 1 m. 38. The [Parelephas] trogonthervi femur at Budapest 1 m. 48, according to my Stegodon Osteologie.” The length measurements of femora thus kindly recorded by Doctor Pohlig for comparison with those recorded by others are as follows: Hesperoloxodon antiquus italicus ref., femur from Rome now at Lumbres, Pas de Calais (Dr. Pontier), partly restored Hesperoloxodon antiquus germanicus ref., femur of large Taubach skele- ton (ef. huge molars, Pohlig, 1889, Pl. vi, figs. 2a, 1b) Archidiskodon meridionalis Florence Museum Parelephas trongontherii ref., femur, Budapest 1500e mm. 4 ft. 11 in. 1600e mm. 5 ft. 3 in. ref., 1380 mm. 4 ft. 64 in. 1430 mm. 4 ft. 8% in. COMPARATIVE BRAIN CHARACTERS OF HESPEROLOXODON Although the entire upper portion of the type cranium of Hesperoloxodon antiquus italicus was practically destroyed, the solid walls of the brain case fortunately were preserved, revealing for the first time the brain characters of a member of the ‘Hlephas antiquus’ phylum. For immediate comparison intracranial casts were taken from the relatively young crania of the African and Indian elephants in the American Museum, affording the following brain cube comparison, as displayed in figure 1112: LOXODONTA AND Brain and Volume Weight cubie centimeters grams C, Elephas indicus (Amer. Mus. 54261), young adult male 6686 —20% = 5349 B, Loxodonta africana (Amer. Mus. 51939), young adult male 6651 —20% = §321 A, Hesperoloxodon antiquus italicus (Amer. Mus. 22634), adult male not full grown 6807 —20% = 5446 APPROXIMATE EXsTIMATES.—Comparing the figures given by Weber (1896, p. 115) and by Gregory and Colbert (letters, May 26, 1931), the actual brain weight is about twenty per cent. less than the cubic centimeter volume of the intracranial cast (which in- cludes the space occupied by the dura mater and the intracranial fluids). Consequently by deducting twenty per cent. from the above intracranial casts shown in figure 1112, we obtain a means of estimating the actual brain weight of Hesperoloxodon antiquus italicus, in comparison with the brain weights of Hlephas indicus and Loxodonta africana, as in table below (p. 1252). It will be observed that the large male EHlephas indicus ratio (1:560) is selected for the male Hesperoloxodon antiquus italicus, in which the estimated brain weight of gr. 5446 and estimated body weight of gr. 3,049,760 exceed the corresponding weights of the Fig. 1111. Left femur, largely restored, referable to Hesperoloxodon antiquus italicus. After photograph bearing the following inscription: “H. Pohlig mit dem Femur des Elefas antiquus von Rom. September 1918.” This femur is estimated at 4 ft. 11 in. or 1500 mm. large male Indian elephant. If we select the Loxodonta africana ratio (1:375), the body weight of H. ant. italicus falls to gr. 2,042,250, probably very much under the actual weight of an animal 12 ft. in height at the shoulder. These roughly approxi- mate estimates of body weight are given in comparison with the maximum actual body weights in pounds and grams below. From these actual figures of the ratio between body weight and shoulder height we may allow an average of sixty-five pounds for one inch in height; with such allowance we deduce the following: Hesperoloxodon antiquus italicus, age 50 yrs. height 12 ft., estimated body weight 9360 Ibs. or 4,250,000 grs. 1252 OSBORN: THE PROBOSCIDEA Brain weight Weber’s ratio of Body weight esti- | Body weight esti- (in grams) brain weight to mated in grams mated from brain Max Weber (1896) body weight weight in pounds Elephas indicus, large male 5430 1: 560 3,048,000 6705 Elephas indicus female (20 yrs. old) 4660 1: 439 2,047,000 4503 Loxodonta africana, female 4370 ils S75) 1,642,000 3612 Amer. Mus. specimens C, Elephas indicus, male, not fully adult 5349 1: 560 2,995,440 6590 B, Loxodonta africana, male, about half grown 5321 1: 375 1,995,375 4390 A, Hesperoloxodon ant. italicus, male, three- ets fourths adult 5446 1: 560 (Indian ratio) 3,049,760 6709 AcruaL Bopy Wertcuts (NEw York ZoonogicaL Park) Age Height Estimated Weight Loxodonta africana oxyotis (male) 28 yrs. 10 ft. 8% in. 8,500 lbs. 3,863,636 gr Elephas indicus (female) 35 8 ft. 6 in. 6,200 2,818, 182 cy Loxodonta africana pumilio (female) 13 6 ft. 2,450 1,113,636 Elephas indicus (male) 4 4 ft. 5 in. 1,025 465,909 Elephas indicus average weight from measurements of eight individuals: Average age: 30 yrs. Average height: 7 ft. 94. in. Average body weight 6074 Ibs. or 2,750,000 grs. SUMMARY OF THE PIGNATARO INTERAMNA ELEPHANT, PALASOLOXODON [HESPERO- LOXODON] ANTIQUUS ITALICUS, NOVEMBER, 1931 Taken altogether, the Pignataro Interamna straight- PATE US tusked elephant, genotypic species' of Hesperoloxodon, 6666 cc marks a great step forward in our knowledge of the ‘Elephas antiquus’ phylum of Falconer, 1847, 1857. (1) The lofty cranial dome resembling that of the Elephas indicus of India is in close agreement with the ‘‘Eléphant tracé en rouge” in the cavern of Pindal, also with the anterior dorsal hump of the Spanish and Algerian ele- phants (Fig. 1047). The entire cranial and dorsal hump silhouette (Fig. 1092) is quite different from that of the African elephant (Fig. 1093). (2) The cranial profile of LOXODONTA AFRICANA Hesperoloxodon italicus is also entirely different from that 665/ c.¢ of Palzxoloxodon namadicus (Fig. 1070) and of P. melitensis? (Fig. 1121), both of which are characterized by a very prominent transverse frontal ridge for the attachment of the gigantic proboscis. (8) In both cranial and dental characters H. italicus differs profoundly from the African elephant, which is extremely conservative in its structure, in fact, much more conservative than any of the known fossil Pleistocene elephants of Eurasia. (4) H. ctalicus affords additional and positive evidence that the dwarfed if elephants of the Mediterranean Islands (Palxoloxodon BL OKD INN Hila el hale Spee falconeri, P. melitensis, and P. mnaidriensis) were not 6507 « derived from the ‘Elephas antiquus’ phylum of Falconer, as hitherto universally believed, but sprang from some INTRACRANIAL Casts or PaLawoLoxopon [=HESPEROLOXxODON], LOXODONTA, AND ELEPHAS undiscovered phylum of elephants of African origin, which ene pn ieee me racratna ly bralit cash Ob aie Rigi ntasostnteraimns. type, Specimen gave rise on the one hand to the dwarfed elephants of the (Amer. Mus. 22634) compared with casts of the African elephant and the Indian elephant, as exhibited in the American Museum. After Osborn, 1931.846, p. 22, fig. 16. Mediterranean Islands and on the other to the gigantic Brain Cube P. namadicus phylum of India and the Far East extending os é ae to Japan. C, Elephas indicus (Amer. Mus. 54261) 6686 = B, Loxodonta africana (Amer. Mus. Dept. Mam. 51939) 6651 See footnote on page 1247 above.—Editor.] A, Palzoloxodon [Hesperoloxodon) antiquus italicus (Amer. Mus. 22634) 6807 *{In Fig. 1121 referred to Palzoloxodon mnaidriensis.—Editor.] THE LOXODONTINA: HESPEROLOXODON 1253 HESPEROLOXODON ANTIQUUS GERMANICUS OF STEINHEIM Many years ago (Feb. 26, 1901) Dr. Eberhard Fraas presented to the American Museum from his Steinheim collection a finely preserved superior molar, r.M* (Amer. Mus. 10655—Fig. 1115) which agrees approximately in its ridge formula (M 3**) with progressive molars (M 3 +2“) from the Taubach-Weimar region of Thuringia, described by Pohlig (1888-1891), as set forth in pages 1233-1235 of the present Memoir, and by Soergel (1912, Tab. 1, VI, VII). During the years 1926 and 1928 the region of Steinheim on the river Murr yielded to the Stuttgart Museum a number of invalu- able remains of ‘‘Der Waldelefant” or ‘forest elephant,’ a pre- liminary account of which has been given by Dr. Fritz Berckhemer, Konservator, Wiirttemberg Naturaliensammlung, Stuttgart. Following his two articles of 1929 and 1930, Doctor Berckhemer kindly furnished (April, 1931) excellent photographs (Fig. 1114), with complete measurements, for inclusion in the present Memoir in comparison with Hesperoloxodon antiquus italicus (Fig. 1105). Gro Logic Acr.—Doctor Bereckhemer describes (1929, p. 188) the related geologic levels and fauna as of 2d Interglacial age. The present author regards them rather as of 3d Interglacial age. “(T. 1926] In Steinheim wurde zuerst die Kiesgrube von Sigrist aufgesucht, die 1926 einen ersten Schidel von Elephas antiquus geliefert hat, und in der vor zwei Jahren ein prichtiger Riesenhirsch . . . zutage kam. Im SO-Teil der Grube sind die Schotter zurzeit in einer Miachtigkeit von rund 10 m freigelegt; der anstehende Untergrund soll nach Aussage des Besitzers noch 3 m tiefer leigen. Uber dem Schotter folgt hier eine Decke von rund 4 m Léss und Lésslehm. Der antiquus-Schadel [1926] lag in diesem Teil der Grube 8 m tief unter der Grenze Schotter/Léss- lehm. Kurz vor der Tagung war noch ein Stirnschadel vom Wisent gemeldet worden, dessen Bergung in Gegenwart der Gesellschaft vorgenommen wurde. Das eindrucksvolle Stiick fand sich im tieferen Teil der tiber dem antiquus-Lager folgenden Mammut- schotter. Auf der gegeniiberliegenden Seite der Strasse befindet sich die neue Grube von Bauer, die durch den Fund des Buffelus murrensis Berckh. . . . bekannt geworden ist.” “{TI. 1928] Sie lieferte im vergangenen Jahr einen zweiten Schadel von Elephas antiquus, der soweit erhalten war, dass er zur vollen Gestalt ergiinzt werden konnte. Er erméglichte zu- sammen mit einem im Juni d.J. im selben Lager gefundenen antiquus-Stosszahn (einen Rest der bei der Bergung entstandenen H6hlung konnten die Teilnehmer in der Schotterwand noch be- merken) erstmals eine Rekonstruktion des Hlephas antiquus durch den Stuttgarter Oberpraparator Bock.” In his article of 1930 (pp. 332, 333) Doctor Berckhemer adds the fauna: “e” In this layer were discovered remains of (1) Rhinoceros merckii, (2) of the Murr water-buffalo (Buffelus murrensis), (3) a nearly perfect skull of the diluvial urus or aurochs (Bos primigenius), (4) very numerous remains of the stag (Cervus elaphus) and of the royal stag (Megaceros). Obviously during this period of the deposition of the lower white sands “‘e’’, the surround- ing country was richly forested as the home of the forest elephant, ‘Elephas antiquus,’ of the urus and of Megaceros; the layer ‘“e’’ containing the cranium and tusks of Elephas antiquus of Steinheim accordingly belongs toward the close of the 2d Interglacial period [3d (Osborn)]; in the same layer was found (5) the remains of Leo sp., of Ursus sp., and of Meles taxus (the badger). 5 Fig. 1113. W. Bauer Quarry, STEINHEIM A. D. MURR, SHOWING THE SITE OF THE 1928 DiscoveRY After original photograph kindly furnished by Dr. Berckhemer (ef. Berckhemer 19380, pp. 331, 332) a. ‘Zu oberst haben wir noch Lésslehm und Loss.” c. “Es folgen nach oben rotlichbraune Schotter (c), die links im Bild auch an die Stelle der hier durch eine abtragende wasserstromung zuvor entfernten Lettenbank treten (Abtragungszone durch weisse Linie bezeich- net).”’ [Hlephas primigenius fraasi.| d. “Daruber liegt eine feinsandige Lettenbank von graugriiner bis dunkelbrauner Farbe.” e. [The layer consists of clear white sand of variable fineness, which toward the bottom is mingled with more or less gravel and sand.]} x. Elephas antiquus [Hesperoloxodon antiquus germanicus| cranium of 1928. ‘“‘d” The fine-sanded loam bank is a still water formation in which are found remains of Bos, Megaceros, and the bear (Ursus arctos). “c” The reddish brown gravels and sands mark a decided change of climate, the Hlephas primigenius fraasi replacing the ‘forest elephant’; other remains, including the wisent (Bison priscus) and the wild horse (Hquus sp.) which is quite frequent, indicate a drier and colder climate of the Glacial Period, with restricted forests. Osborn: The Elephas antiquus horizon of Steinheim yields a fauna distinctive of the 3d Interglacial stage, consequently these Steinheim remains may be provisionally referred to Hesperoloxrodon antiquus germanicus. ; AZ A Stuttgart Mus./5930 (1928) B Stuttgart Mus 15344 (1926) (/926) (reyersed) PALAEOLOXODON ANTIQUUS GERMANICUS “ef HESPEROLOXODON ANTIQUUS GERMANICUS REF. OF STEINHEIM ON THE Morr, IN THE STUTTGART MusrEuM Fig. 1114. After photographs and indicated measurements (March 16, 1931) by Dr. F. Berekhemer. Reduced to a uniform scale of one-sixteenth natural size A, Right lateral view of cranium of 1928 (Stutt. Mus. 15930). A3, Posterior or parieto-oecipital view of cranium. Al, Superior view of cranium A2, Palatal view of cranium, with third superior grinding teeth. B, Cranium of 1926 (Stutt. Mus. 15344, reversed for comparison). exceeding in breadth that of Hesperolorodon antiquus italicus (Fig. 1105), correspondingly broad in the palate. See comparative measurements (p. 1255). Observe also 16+ ridge- Observe the relatively broad, low occiput, The adult cranium (B) is about equal in depth (bathycephaly) to that of H. ant. italicus. plates exposed out of a probable total of 19 ridge-plates (ef. Pig. 1115). 1254 THE LOXODONTIN#:: PAL. ANT. GERMANICUS REFERRED Mo.uar OF HESPEROLOXODON ANTIQUUS GERMANICUS Fig. 1115. Third superior molar of the right side, r.M* (Amer. Mus. 10655) found at Steinheim a. d. Murr and presented to the American Museum by Dr. Eberhard Fraas, February 26, 1901. Observe 19 ridge-plates, 11 anterior worn; length 280 mm., height of 11th ridge-plate 195 mm. Compare figure 1088B, Weimar molar, length 295 mm., height 190 mm.; also figure 1114 A2, Steinheim specimen. STEINHEIM SKULLS OF 1928 (FIG. 1114A) AND OF 1926 (FIG. 1114 B) Doctor Berckhemer’s descriptions refer to an exceptionally perfect cranium (Fig. 1114, A) of 1928 (Stutt. Mus. 15930) and to a less perfect cranium (Fig. 1114, B) of 1926 (Stutt. Mus. 15344). These priceless new materials, with associated faunal remains, Summit of occiput to tip of right premaxillary Transverse across widest portion of occiput Mid-length of cranium, occipital condyles to tip of premaxil- lary junction Breadth across zygomatic arches Summit of parieto-occipital crest to attritional surface of r. M$ (bathycephaly) Summit of parieto-occipital crest (partly restored) to attri- tional surface of 1.M?’ Width of occipital condyles Width of anterior nares Width across narrowest portion of maxillaries Width across broadest portion of premaxillaries Occipital condyle to front border of orbit (left) Occipital condyle to front border of orbit (right) 1928 (Stutt. Mus. HESPEROLOXODON 1255 apparently enable us to establish the cranial characters of Hespero- lorodon antiquus germanicus as quite distinct from those of H. ant. italicus above described. In Cranium A of 1928 (Stutt. Mus. 15930) sixteen ridge- plates are exposed on the worn surface of the left superior molar, 1.M*; the right molar, r.M%’, may show 15-16 ridge-plates as com- pared with the perfeet superior molar (Fig. 1115) from Steinheim exhibiting 19 ridge-plates as compared with M 3 78% in the Taubach-Weimar horizons described by Pohlig (1888-1891) and Soergel (1912) mentioned above. The diameters of the crowns of M 3 are: Amer. Mus. 10655 A. Stutt. Mus. 15930 R. M* Length of 16 exposed ridge- plates 245 mm. Breadth 88 Length of 11 exposed ridge-plates Maximum height of 11th ridge- plate L. M® Length of 16 exposed ridge- plates (total ?+17) 245 Breadth 89 In the Stuttgart Museum |.M?* (No. 15344) exhibits a total of 19 ridge-plates; an r.M* (Stutt. Mus. 16515), length 300 mm., exhibits +18-+ ridge-plates (total 20 ridge-plates). In the right maxillary alveolus of A (Stutt. Mus. 15930) are found the remains of a tusk (Berckhemer, 1930, Abb. 5) which had been broken off during the lifetime of the animal, consequently the right alveolus had been diminished by the formation of new bony parts and the right premaxillary had correspondingly di- minished in breadth as compared with the normal width of the left premaxillary (Fig. 1114, A2). The remaining left superior incisive tusk was not preserved with the skull. [The isolated right tusk, 3500 mm. in length, does not belong to this skull; Doctor Berck- hemer, however, does not hesitate to ascribe it to the same species. | 168 mm. 195 Total width of premaxillaries as seen from above 850 mm. Width of diminished right premaxillary 400 Width of full-grown left premaxillary 450 The cranial and dental characters of this remarkably well- preserved individual are beautifully shown in the accompanying photographs (Fig. 1114, Al-3). The principal measurements, in addition to those given above, are the following: Hesperoloxodon antiquus italicus (Amer. Mus. 22634) Hesperoloxodon antiquus germanicus Cranium of B: Cranium of 1926 (Stutt. Mus. 15930) 15344) 1375 mm. 1400 mm. 1095 875e 1150 1130 880 865 mm. 750e 860 980+ 990e 885 295 307 280 400 385e 545 480 530 850 890e 857 775 850 765 820e 1256 The B Cranium of 1926 (Stutt. Mus. 15344), from the above measurements, appears to belong to a somewhat older individual than the A Cranium of 1928 (Stutt. Mus. 15930). More of the dental ridge-plates in M? are exposed; the bathycephaly or depth (from the summit of the occiput to triturating surface of M?®) is 980+ mm. as compared with 860 mm. in the A Cranium of 1928; the horizontal measurement from the occipital condyle to the front border of the orbit in the B Cranium of 1926 is 850 mm. as com- pared with 775 mm. in the A Cranium of 1928; the occipital condyles as well as the orbit are more prominent and pediculated; the premaxillaries are more elongated. In brief, the B Cranium of 1926 (Stutt. Mus. 15344) belongs to a fully mature and adult, but not aged, bull of Hesperolorodon antiquus germanicus. HESPEROLOXODON ANTIQUUS GERMANICUS FROM THURINGIA ELEPHAS ANTIQUITATIS KruGer, 1823 [= HEspEROLOXODON ANTIQUUS GERMANICUS].—(Kriiger, 1823, p. 832): ‘Eleph. anti- quitatis. Europiischer Elephant. An einigen Orten in Europa werden einzelne Spuren von einer Elephantenart angetroffen, welche weniger dem asiatischen, und mehr dem afrikanischen Elephanten sich nahert. Sind auch die bei Thiede*) und bei Teschen am Boberfluss im Jahre 1795 entdeckten Zaihne**) achte Mammuthsziihne mit abgenutzter Kauflache gewesen, und gehoren sie nicht einer davon abweichenden Art an, so zeigen sich doch bisweilen andere, welche den Zihnen des afrikanischen Elephanten sehr dhnlich sind. Selbst Cuvier, der nur eine Art, die asiatischen Mammuths, annimmt, zugestehen, dass sich bei Eichstedt ein dem afrikanischen Elephantenzahn gefunden habe***). So sind wahrscheinlich mehrere Ueberreste in den Sammlungen vorhanden, nur hat man sie bis jetzt ibersehen, und aus Vorliebe fiir den nordischen Elephanten auf Abweichungen nicht geachtet.” Osborn, 1931: It is apparent from the above description, and figure from Breislak reproduced herewith, that Kriiger had in mind the ‘ancient’ or ‘straight-tusked’ elephant as distinct in tusk structure from the modern Asiatic elephant and resembling rather the African elephant, since the first locality mentioned (Thiede) is in the Thuringian region, northern Germany. The tooth which we may select as Kriiger’s type (after Breislak, Bd. II, s. 428) is not improbably of the same 3d Interglacial geologic age as Pohlig’s and Stefanescu’s ‘Hlephas germanicus’ of the Taubach-Weimar horizon. It is therefore for German paleontologists to determine whether the specific name ‘Hlephas antiquitatis,’ 1823, has priority over ‘Blephas germanicus’ Pohlig—Stefanescu. muss doch ahnlicher *Abgebildet in Breislak’s Lehrb. d. Geologie. 2ter Bd. 8 428. **Arch. d. Urw. III., 2. S 396. OSBORN: THE PROBOSCIDEA Paid Seite 428 Bhai tiche CLV2CS fegailen Gi lephunten. Cahias ron She HESPEROLOXODON ANTIQUUS GERMANICUS REF. Fig. 1116. the present author to Hesperoloxodon antiquus germanicus. 1820, p. 428. Type molar of Hlephas antiquitatis Kriiger, 1823, referred by After Breislak, «**J’ai vu une molaire donnée comme d’Aichstedt, dans la collection d. M. Ebel 4 Brémen; quoique d’apparence bien fossile, elle étoit remarquable par sa ressemblance avec les molaires d’Afrique. Rech. s. 1. Oss. foss. Th. I. S. 127. IV. EXTINCT DWARFED SPECIES OF THE MEDITERRANEAN ISLANDS (Continued from pp. 1182, 1183 of the present chapter, modified and extended; see also observations of Raymond Vaufrey, 1929, below, pp. 1268 to 1272.) Of the greatest interest are the full-sized and dwarfed species of elephants found in the cavern deposits of the different islands of the Mediterranean, successively described by Busk (1867), Falconer (1862, 1868), Leith Adams (1870, 1874), Forsyth Major (1883), and Bate (1903 and 1907). To the latter author we are especially indebted for the most recent discoveries and descriptions of these insular proboscidean species and subspecies. Falconer and the older authorities related these dwarfed species to the ‘Elephas antiquus’ of the European continent, but subse- quent discovery has shown that they are more probably derived from certain of the extinct ancestral African species described below as Palzoloxodon (synonym Pilgrimia).' BaTHYMETRIC Map OF THE MEDITERRANEAN ISLANDS Fig. 1117. Dwarfed Mediterranean species of elephants are found east to west on the islands of Cyprus, Crete, Malta, Sicily, and Sardinia. The white areas are within the 100 fathom line (600 feet); the maximum depression of the sea or elevation of the land (in certain limited areas) was to the 216 fathom line (about 1300 feet) or 400 m. Reproduced by permission of Longmans, Green and Company, publishers, from map edited by Chisholm and Leete. From a recent hydrographic chart (No. 4300 U. S. Navy, January, 1930), it is shown that the submerged land connections are now at the following depths: (1) Cyprus to Turkish mainland (N.E. to Gulf of Alexandria near Adana, Turkey), shallowest depth 320 fathoms, 1920 feet; (2) Crete to Turkish main- land (N.E. viaisland of Rhodes), shallowest depth 250 fathoms, 1500 feet; (3) Malta to Sicily, shallowest depth 74 fathoms, 444 feet; (4) Sicily to Italy across strait of Messina, 109 fathoms, 657 feet; (5) Tunisia to Sicily, shallowest depth, 168 fathoms, 1008 feet; (6) Sardinia to Corsica, 46 fathoms, 276 feet; Corsica (northern end) to Italy, 140 fathoms, 840 feet. Before discussing these questions of affinity, and the conclusions reached by Pohlig (1888, 1893), we may follow the chronological order of systematic description. 1862 Malta, Zebbug Cave. Type molar tooth of Hlephas (Loxodon) Melitensis Falconer, described in 1862. 1867 Malta, Zebbug Cave. Type skeleton of EHlephas falconeri Busk, 1867. 1870 Malta, rock-fissure, Mnaidra Gap. Type molar of Elephas mnaidre Adams, 1870. 1883 Sardinia, sands of Morimentu. Elephas Lamarmorae Major, 1883. Carpal and tarsal bones. 1903 Cyprus, Kerynia Hills. Elephas cypriotes Bate, 1903. Cotype molars. 1907 Crete, cave near Cape Maleka. Elephas creticus Bate, 1907. Nine imperfect molars, portion of an incisor, and dorsal half of a vertebra. 1912 Sicily, Carini. Elephas antiquus var. insularis Soergel (name only). Falconer was the first to describe before the British Association, October 6, 1862 (see ‘“The Parthenon” for October 18, 1862, p. 780), one of the pygmy elephants of Malta found in the Zebbug Cave, namely, Hlephas In a footnote on page 1 of Professor Osborn’s Novitates article on the “Primitive Archidiskodon and Palaeolorodon of South Africa” (Osborn, 1934.925) appears the following statement: “Pilgrimia Osborn (December 20, 1924) is antedated by Palxolorodon Matsumoto (September 20, 1924).”’ Hence Pilgrimia becomes a synonym of Palzolorodon.—Kditor. | 1257 1258 OSBORN: THE PROBOSCIDEA Melitensis, which is supplemented by more detailed description in the ‘“‘Palzeontological Memoirs” of 1868 (Fal- coner, 1868, Vol. I, pp. 292-308). According to Leith Adams (1870, p. 223), Hlephas falconeri, although found in the same cave, represents a more diminutive animal than the Hlephas melitensis of Falconer. € ELEPHAS MNAIDRIENSIS’ ‘ELEPHAS FALCONERI, ‘ELEPHAS MELITENSIS’ = 1900 m™m = 900mm = 1400 mm: DwarFED ELEPHANTS [PALHOLOXODON (SYN. PruGRIMIA)| OF THE MEDITERRANEAN ISLANDS One twenty-ninth natural size. Compare Figure 1119 Vig. 1118. Diagrammatic representation of the manner in which the three restorations of figure 1119 were calculated. As a rule the humerus affords the most reliable estimate of the height of the shoulders. Comparison of the relative heights of these animals may best be made with the Upnor elephant (Hespero- lozodon antiquus). Length of Humerus Estimated Shoulder Height H. antiquus of Upnor 1290 mm. =4 ft. 2% in. 3700 mm. =12 ft. 15 in. P. mnaidriensis of Malta 573 mm.=1 ft. 10% in. 1900 mm.= 6 ft. 2%4 in. P. melitensis of Malta 478 mm.=1 ft. 6°%4 in. 1400 mm.= 4 ft. 7% in. P. falconeri of Malta 270 mm. = 10% in. 900 mm.= 2 ft. 11° in. Broadly speaking, Palzoloxodon mnaidriensis is one-half the height of Hesperoloxodon antiquus of Upnor, P. melitensis is about two-fifths the height of H. antiquus, while P. falconeri is only about one-fourth the height of the Upnor animal. As fully explained in the text, these dwarfed elephants appear to be related to P. namadicus and to be descended from North African ancestral stages of Palzxoloxodon. The cranium (Fig. 1121)' and jaw (Fig. 1124) of Palxoloxodon melitensis, with broadly overhanging fronto- parietal crest, more closely resemble the P. namadicus of India than the Hesperoloxodon antiquus of western Europe. OriGcin.—Comparison of the type grinding teeth of the dwarfed Mediterranean species with the type grinding teeth of the extinct African species described below reveals a striking general resemblance in the narrow proportions and in the rudiment or absence of the ‘loxodont sinus,’ characters which appear to relate these teeth to the phylum Palxoloxodon of Africa rather than to the typical Loxodonta africana. In Europe some of the narrow-toothed varieties of Hesperoloxodon antiquus may be related to the dwarfed insular elephants.” The characteristic body height and progressive ridge formul of these species were estimated by the authors as follows: 'Referred by Professor Osborn to Palxoloxodon mnaidriensis.—Editor.] *[See the opinion expressed by Professor Osborn on page 1252 above regarding the origin of the dwarfed elephants of the Mediterranean Islands.— Edtior.] THE LOXODONTINA:: DWARFED SPECIES OF THE MEDITERRANEAN ISLANDS 1259 Height Ridge Formula Malta Elephas melitensis Fale., 1862, 1868 Height of Indian Tapir, i.e., 5 feet (1525 mm.) M 3 == Elephas falconeri Busk, 1867 2 ft. 6 in. (760 mm.) to 3 feet (915 mm.) M 3 — Elephas mnaidre Adams, 1870 6 to 7 feet (1830 mm. to 2135 mm.) Misia Sardinia Hlephas Lamarmore Major, 1883 2 to 3 feet (610 mm. to 915 mm.) M 3 — Cyprus Elephas cypriotes Bate, 1903 22 to 3 feet (610 mm. to 915 mm.) M343 Crete Elephas creticus Bate, 1907 Slightly larger than H. cypriotes, max. 5 feet (1525 mm.) MES} Sear “pe hA ROI ne mem ee ma FE LomMseh 193 DwarFrED ELEPHANTS, PALAOLOXODON (SYN. PILGRIMIA), OF THE MEDITERRANEAN ISLANDS One twenty-fourth natural size. Compare Figure 1118 Vig. 1119. In descending scale of size from ‘Elephas mnaidriensis’ (left), to ‘#. melitensis’ (right), to the diminutive ‘H. falconeri’ (center), the relative heights are very carefully estimated from the respective length of the limb bones (indicated in solid black in Fig. 1118). Drawn under the direction of the author by Margret Flinsch, 1930. j The larger stage (EH. mnaidriensis) is that of which the complete cranium is known [=the #. (antiquus) Melite of Pohlig, 1893], probably resembling that of ‘E. namadicus’; the same form of cranium is attributed to H. melitensis and E. falconert. The tusks are drawn from the recent comparative figures of Vaufrey. This restoration is entirely different from that of Adams (Fig. 1127), which is based solely on the theory that these dwarfed elephants were related to Loxodonta africana. Young ArricAN Pyemy! ELEPHANT PASSING BENEATH ADULT INDIAN FEMALE ELEPHANT Fig. 1120. The existing pygmy elephant Hlephas africanus pumilio Noack, 1906, from the French Congo, attaining a height of 2 ft. 6 in. at the age of two and a half years and corresponding at this age in size with the most diminutive fossil species Palzoloxodon falconert, P. lamarmorx, and P. cypriotes. Photograph (1922) by Mr. Elwin R. Sanborn through the courtesy of the New York Zoological Park. The intimate companionship and friendship of these two elephants has been described (1928) in William T. Hornaday’s volume “Wild Animal Interviews.” At the age of four and a half years the same animal at- tained a height of 4 ft. 444 in., corresponding more nearly with Palzoloxodon melitensis and P. creticus. The full-grown Loxo- donta africana pumilio (Amer. Mus. Dept. Mam. 35591) attains a height of 6 ft. 8 in., equaling that of the larger fossil Medi- terranean species. 'See editorial note on p. 1196 above.] 1260 OSBORN: THE PROBOSCIDEA In comparison, the dwarfed or small forest! elephant of the Congo, Loxodonta africana pumilio (Fig. 1120), measured 2 ft. 6 in. in height (=760 mm.) at the age of two and a half years; consequently at this age it agreed with Palxoloxodon falconeri. Thesame animal at the age of four and a half years measured 4 ft. 44 inches in height at the shoulders, and therefore attained the approximate height (=5 ft. or 1525 mm.) of P. melitensis or of P. creticus. RipcEe Formuta.—In the following type and subsequent descriptions the ridge formule of the species are very carefully cited from the earlier and later publications by the authors who have successively treated the denti- tion of these interesting animals, namely, Falconer, Leith Adams, Pohlig, Busk, and Bate. These type ridge formule constitute an important part of the type descriptions; they differ widely from the collective ridge formule cited from Raymond Vaufrey (1929) below; compare, for example, the ridge formula of ‘HZ’. mnaidriensis’ (M 375-78), Fig. 1121 REFERRED PALAOLOXODON MNAIDRIENSIS. PALERMO COLLECTION, Cranium I One-eighth natural size. After Pohlig, 1893, who referred it to Elephas (antiquus) Melitx” Fig. 1121. Fully adult cranium of ‘‘Zlephas (antiquus) Melite’’ Falec., from the Interglacial layers of the Grotta di Pontale, Carini, Sicily, the most complete cranium in the collection of the Palermo Mu- seum, exhibiting the characteristic prominent transverse frontal crest, comparable to that of ‘Hlephas [ =Palxolozodon| namadicus’ of India. After Pohlig, 1898, Taf. 1, figs. 1, la, reduced to one-twelfth natural size. 1122 For the proportion of this insular form to the full-grown Palzoloxodon namadicus, see figure 1069, also the restoration (Fig. 1119). Although these dwarf insular species of Malta usually have been referred to ‘Elephas antiquus,’ the present specimen is obviously more closely related to P. namadicus, and, while adult, belongs to an individual inferior in size to the juvenile P. namadicus (Fig. 1069), after Malconer and Cautley, 1847, Pl. xxiv.a, fig. 4a. REFERRED PALZOLOXODON MNAIDRIENSIS. PALERMO CoLLECTION, Cranium VI One-eighth natural size. After Pohlig, 1893, who referred it to Hlephas (antiquus) Melite Fig. 1122. Palate of juvenile cranium (Palermo Coll. VI), after Pohlig, 1898, Taf. u, fig. 2, one-eighth natural size. This juvenile cranium (Palermo Coll. VI), retaining the 8+ ridge-plated r.M2, also the partly erupted r.M?® displaying only 9 plates, 3 partly worn, is in a larger but younger stage than the adult cranium (Palermo Coll. I—Fig. 1121 herewith). The measurement from condyle to extremity of maxillaries is approximately 744 mm., while in the adult cranium the same measurement is approximately 675 mm. This ontogenetic short- ening and deepening of the cranium as it grows older conforms with what we observe in the crania of all species of elephants. (Pohlig, op. cit., p. 90) “An Cranium VI, das, in basaler Ansicht, auf Tafel II, in Fig. 2, abgebildet ist, ist das Suborbitalforamen kaum 0 m 02 lang, obwohl der Schiidel, der grésste sicilische, mehr als 0 m 7 maximaler Liinge misst. . . . Die M. II. sind stark abradirt, M. III. noch fast intact; die maximale Breite der Schnauze, welche extrem divergente Lateralriinder hat, betrigt ca. 0 m 48... . die Condylen sind an diesem Exemplar, im Gegensatze zu den kleineren Cranien, nach Form, Stellung und relativer Grosse sehr iihnlich wie bei dem typischen und dem namadischen £. antiquus, und folglich auch bei £. africanus.” (See editorial note on p. 1196 above.] “Referred to ‘Elephas mnaidriensis’ by Leith Adams and Raymond Vaufrey. THE LOXODONTINA: DWARFED SPECIES OF THE MEDITERRANEAN ISLANDS 1261 given by Leith Adams, 1874, p. 112, with that (M 3 *14*) of Vaufrey (1929, pp. 124, 128). It appears that Vaufrey (pp. 113-132, figs. 28-37) has intermingled the ridge formule of the grinding teeth of more than one species or mutation. PoHLIG ON THE Monospeciric ‘ELEPHAS (ANTIQUUS) MELIT&’ OF SICILY Pohlig (1893, pp. 75-108) described and figured fully the referred ‘Elephas (antiquus) Melite’ Fale. from the famous elephant grotto of Carini, Sicily, where were found in great numbers the skeletal parts of this dwarfed species with numerous remains of the degenerate stag Cervus (elaphus) Siciliz and less abundant remains of both Bos and Bison, Bos (taurus) primigenii and Bison priscus; also traces of Hyena spelza and of partly worked flint implements. Pohlig (op. cit., p. 100) maintains that the cranial structure of the six best preserved skulls from Carini closely unites these insular pony-elephants with the European LH. antiquus. The extraordinary transverse frontal crest (Fig. 1121) observed in the adult Carini specimens Pohlig regards as the compelling ground for the specific union of the three forms “EH. Namadi, E. Melite und E. antiquus typus.” Pohlig adheres (p. 101) to the conclusions reached in his earlier (1891) monograph on E. antiquus of the specific identity of LZ. namadicus and E. antiquus. From these remains his geological inferences are as follows (op. cit., p. 82): 2. Landverbindung zwischen Sicilien und Italien einerseits, Afrika andererseits [error], und Einwanderung der grossen Saugethiere, gegen das Ende der ersten diluvialen Glacialperiode. 3. Erneutes siculares Steigen des Meeresspiegels zu Beginn der diluvialen Interglacialzeit, erneute Isolation Siciliens, Aus- bildung von diminutiven Formen grosser Siiugethierarten, Anhiiufung von Skelettheilen solcher auch in der Grotta di Pontale, einem Zufluchtsort namentlich fiir Elephanten, Edelhirsche und Rinder. Pohlig’s palzontological observations are (op. cit., p. 83): In meiner angefiihrten Monographie des Elephas antiquus habe ich zuerst nachgewiesen, dass die Angaben de Anca’s und Gemmellaro’s von Hlephas armeniacus und EF. africanus aus Sicilien, ebenso die Artbezeichnungen EH. Falconeri von Busk [Footnote: ‘Transactions of the zoolog. soc. London 1868, VI., p. 5, No. 10.’] und #. mnaidriensis von L. Adams [Footnote: ‘Tbid. vol. IX, p. 1. 1877.’] aus Malta auf Irrthum beruhen, und dass der ‘Hlephas melitensis’ von Falconer nichts anderes ist, als eine insulare Diminutivrasse des Urelephanten, Elephas antiquus, fiir welche ich daher die Bezeichnung E. (antiquus) Melite Fale. vorschlug. Zugleich erbrachte ich die ersten Nachweise der Thatsache, dass die gleiche Zwergelephantenrasse, wenn auch nicht bis zu gleich extremer Gréssenreduction, wie auf dem kleinen Malta, auch auf Sicilien und in anderen Mit- telmeergegenden gelebt hat. From this Carini grotto Pohlig describes and figures the most completely preserved crania in the Palermo Museum (op. cit., pp. 84-98) concluding as follows (p. 98): Die Vereinigung der drei Speciesnamen EHlephas melitensis Fale., E. Falconeri Busk und EF. mnaidriensis L. Adams, die lediglich auf den Dimensions verhiltnissen des Malteser Materiales beruhten, unter der Rassenbezeichnung Z. antiquus Melite Fale. wurde in meiner Elephantenmonographie vorzugsweise begriindet auf die wichtigste bis dahin bekannte Eigenthiimlich- keit jener Diminutivformen, deren Dentition,—wobei der iiberraschende Nachweis in der Gestaltung der friihesten, in gleicher Weise sonst bisher von keiner Species bekannten Milchdentition der Malteser Elephanten, auch fiir den typischen Taubacher E. antiquus, ausschlaggebend sein musste. According to my observations, Pohlig continues (op. cit., p. 99), neither in North Africa nor in lower Italy (in the west) does any fossil species occur excepting ‘HZ. antiquus,’ while in the Mediterranean islands not a single trace has been found of a specimen of the normal continental size. In Sicily occur only the larger types of dwarfed E. antiquus from one-half to two-thirds the size of the normal continental forms, corresponding in dimensions 1262 OSBORN: THE PROBOSCIDEA with the larger H. mnaidriensis Leith Adams of Malta. This larger subspecies obviously lived in Malta at a time when it was united with Sicily by a land bridge. There followed more severe living conditions on the smaller rocky islands which brought about further degeneration and reduction in size. According to the dimensions of the numerous limb bones from the Grotta di Pontale of Carini, in comparison with those of the typical EH. antiquus the Sicilian Hlephas (antiquus) Melite in its full growth or adult condition was about the size of a middle-aged Indian elephant; whereas on the continent the intermediate stages between the full-sized EH. antiquus and the smaller forms of the known E. antiquus remains from the islands are without exception diminutive and all the specimens from the very rich material of Carini belong to the diminutive race. On the other hand, the single Sicilian molar found elsewhere corresponds with a small example of the true E. antiquus. Osborn (1928) entirely dissents from Pohlig’s opinion as to the specific union of Hlephas antiquus Fale., 1847, with the earlier described species H. namadicus Fale., 1846. It would appear that Pilgrim (1905) was influenced by Pohlig in the same erroneous confusion of these two very distinct species of western Europe and the Siwaliks. Osborn, 1930: The cranium of EL. [Palxoloxodon| mnaidriensis (Fig. 1121) closely resembles that of H. [Palzxo- loxodon| namadicus of the Nerbudda, India, in the transverse crest; it differs widely from the cranium of Palzxoloxodon |Hesperoloxodon| antiquus italicus (Fig. 1106). SYSTEMATIC DESCRIPTION OF MEDITERRANEAN ISLAND SPECIES Compare the specific revision by Vaufrey (1929) Palwoloxodon melitensis Falconer, 1862 292, 299, 307, 308. Typr.—Third upper molar of the left Figures 1041, 1068, 1069, 1118, 1119, 1123, 1124, 1127, 1131, 1133, Pl. xxmr side, 1.M* (Brit. Mus. 44312). Horizon AND LocaLity.— Pleistocene, Zebbug Cave, island of Malta. From the lower cave de- Pleistocene, Zebbug Cave, island of Malta. Tyee FigurE.— posits or levels. Op. cit., 1868, Vol. II, Pl. x1, figs. 1, la. Elephas melitensis Falconer, 1862, ‘The Parthenon,” October, OricInaL Derscription.—This elephant was found in the 1862, p. 780; also ‘“Palzontological Memoirs,” 1868, Vol. II, pp. Zebbug Cave, and is the first described of the ‘‘pigmy” elephants of Malta. Falconer (‘‘Parthenon,” Oct. 18, 1862, p. 780—see reprint in “Paleontological Memoirs,’ Vol. II, p. 308) states: ‘The pigmy Elephant was an animal of remarkably small proportions; an adult individual could not have exceeded the Indian Tapir in height and bulk, a creature not much larger than a full-grown Hog. Contrasted with the bones and teeth of an adult African Elephant the difference in size of these portions of its frame exhibited were most striking. ... But though so small, the skeleton agreed in every particular with the one of greatest bulk. A series of harmonies ran through the two skeletons, one bone answering to another truly, and without ordinal or generic difference. The author could refer it unhesitatingly to his subgenus Lozxodon, in the African group of Elephants.” SuPpPLEMENTARY DescriptTion.—Falconer described the teeth of this species at the British Association, October 6, 1862 (an ab- stract of which appeared in “The Parthenon,” as stated above) Typr OF PAL®OLOXODON MELITENSIS and in 1868, Vol. II, pp. 292, 299, he designated the type as fol- Tig. 1123. Elephas Melitensis Falconer, 1862, 1868, type molar tooth lows: “One of the most characteristic of these [fossils] is an upper (Brit. Mus. 44312), one-half natural size. After Falconer, 1868, Vol. II, Pl. molar of the left side... . As this specimen is about to be returned x1, figs. 1, la: “Views in plan and profile of last upper true molar, left side. to Malta, at Captain Spratt’s request, it is necessary to make an Described at page 292.” Lydekker (1886, p. 154) catalogues this tooth (Brit. accurate description of it, to accompany the figures drawn by Mr. Mus. 44312) as follows: “The third left upper true molar; from Zebbug Cave. Dinkel. (PI foal anda? Heoaleactates 283. 284 Described and figured in ‘Falconer’s Paleontological Memoirs,’ vol. ii. p. 292, inkel. ( x1, figs. 1 and La.) ‘ elise sta SEO OE 35) 2 J pl. xi. figs. 1, la; and noticed by Busk, op. cit. [1867.1], p. 296, and by Leith- that Captain Spratt had “lately discovered in Malta numerous re- Adams, op. cit. [1874.1], x. p. 28. Leith-Adams Collection. Purchased, 1873.” mains of a surprisingly small fossil Elephant, of the sub-genus THE LOXODONTINAZ: DWARFED SPECIES OF THE MEDITERRANEAN ISLANDS Loxodon, which I have named EF. Melitensis,” and on the ‘‘Descrip- tion of Plate x1’ he refers to this species as “‘Hlephas (Loxodon) Melitensis.” Falconer (op. cit., 1868, p. 298) gives the ridge formula of E. melitensis as follows: [Dp 2] $ [Dp 3] [Dp 4] ¢ [M 1] [M 2] $ [M 3] ts. Bate (ef. 1904, p. 357) observes that the ridge formula of EF. cypriotes is slightly lower than that which Leith Adams, later than Falconer and after examining a further large amount of material, gives for HL. melitensis: Dp 2% Dp3 = Dp 4 22 M1 £2 M 222 M3 22. 9 9 REFERRED MANDIBLE OF PALAOLOXODON MELITENSIS, FROM THE GROTTA DI PoNTALE, SICILY One-fourth natural size Fig. 1124. Juvenile jaw containing right and left second inferior molars, r.Mg, 1.M2 (Amer. Mus. 2011), acquired by the Museum in exchange with the Padua University, Italy. Observe the sharply truncated vertical rostrum and the broad forwardly pitched coronoid process. The grinders exhibit nine complete elevated ridge-plates and two posterior half ridge-plates, or a total of eleven ridge-plates, namely, M 2 ga15- 1263 RESUME BY DEPRHRET AND MAYET, 1923, PAGE 175 Nous résumons dans le tableau ci-dessous les données numériques qui caractérisent les espéces ou mutations du rameau de |’E. meli- tensis ; Dimensions des Mut en millimetres OO SS Nombre de lames Longueur Largeur Hauteur aux Mur totale maxima au milieu E. cypriotes BER Ohege sesing ey Bee te 1I—12 114-125 26-29 sauoyee 8—9 105 33 84 E. melitensis . . . = ae a 7 E. iolensis ; ae ne So! 72) : hake 12 140 3 70 . 6 12—13 177 E. mnatdrensis . a5 7790" = a : 12 324 78 E. atlanticus . a Jah 3as =r ae E. priscus Falconer. care a3 a er Palwoloxodon falconeri Busk, 1867 Figures 1068, 1118, 1119, 1127, 1131, Pl. xxim Pleistocene, Zebbug Cave, island of Malta. From the higher cave deposits or levels. According to Leith Adams, this ‘lesser pigmy elephant” is the most diminutive insular proboscidean discovered, in which the average height could not have exceeded 2 ft. 6 in. to 3 ft. It is of more recent age than H. melitensis. Elephas falconeri Busk, 1867. ‘Description of the Remains of three extinct Species of Elephant, collected by Capt. Spratt, C.B., R.N., in the Ossiferous Cavern of Zebbug, in the Island of Malta,” etc., Trans. Zool. Soc. London, VI, Pt. V, p. 251. Typr.— Fragments of skeleton. Horizon anp Locaurry.—Pleisto- cene, Zebbug Cave, island of Malta. Typr Ficurr.—Op. cit., Pls. xtrx, L, LI. It is not practicable to reproduce these type fragments. Typr Drescription.—Busk’s designation of the name and the type is as follows (op. cit., p. 251): ‘With regard to the points of difference between it [femur of melitensis] and that of H. falconeri, I will reserve what I have to say until I come to that bone... . Of the remains referred to this second diminutive species, the follow- ing have been selected for the purpose of conveying some notion of the characteristics of the mature animal.” The author cites portions of the skeleton, namely, atlas, several vertebrae, second rib, scapula, right humerus, left humerus, knee, phalanx, pelvis, femur, astragalus, left metatarsal. He then proceeds (op. cit., pp. 251-283) to give a detailed comparison of the skeleton referred to E. melitensis with portions of the type skeleton of H. falconeri. This is followed by a very elaborate series of comparative measurements with H. indicus, E. africanus, EH. melitensis, and E. falconert. 1264 OSBORN: Of this animal, Adams (1870, p. 223) remarks: ‘‘The Lesser Pigmy Elephant. Hlephas falconeri, Busk. As be- fore remarked, the presence of this species in the exuvie of the Zebbug cave was determined by Mr. Busk after Dr. Falconer’s death, and seemingly more or less from fragments of bones, which on comparison with similar portions of the skeleton of Hlephas melitensis and a footbone (astragalus) discovered by me in Mnaidra gap, there appeared such differences both in size and configuration as to warrant the assumption that they represented a still more diminutive proboscidian, the average height of which at the withers could not have exceeded two feet six to three feet.”’ Osborn, 1930: As Busk’s type (1867) is a diminutive skeleton without teeth, it remained for Bate (1903, 1904) to give the ridge formula of ‘Hlephas cypriotes’ cited below, without reference to the ridge formula of ‘F#. falconert.’ So far as we can find, Vaufrey was the first to suggest that ‘EB. cypriotes’ is synonymous with ‘F. falconeri’ and to give (1929, pp. 98, 99) the high collective ridge formula of E. falconeri—E. cypriotes as follows: M 2 29-32 M 3 7834. This is a higher ridge-plate formula than that (M 3 73) finally assigned to EH. cypriotes by Bate, it equals that (M 3 42%) of E. mnaidriensis, fide Adams. This discrepancy partly depends upon the reckoning of the half crests or talons omitted above. Type OF PALAOLOXODON MNAIDRIENSIS Fig. 1125. Type of Hlephas mnaidre Adams, 1870, Pl. u, figs. 2, 2a [= EHlephas mnaidriensis Adams, 1874, p. 116], one-half natural size. Adams (1874, p. 33) describes the same tooth and gives a much more accurate figure (Fig. 1126) of the type (Pl. vu, figs. 2, 2a). Lydek- ker (1886, p. 142) designated this molar (Brit. Mus. 44304) as follows: “The third right lower true molar, about one-third worn; from Mnaidra gap. Described and figured, op. cit. p. 33, pl. vii, figs. 2, 2a. The crown is remarkable for its excessive lateral curvature.’’ See also figure 1126. THE PROBOSCIDEA NAT SIZE DENTITION Warden & Valen WD Wesley aft tah Tyre AND Paratype Mouars OF PALAOLOXODON MNAIDRIENSIS Reproduced herewith one-half natural size Vig. 1126. Hlephas mnaidriensis Adams, from Mnaidra Gap, Malta. After Leith Adams, 1874, Pl. vu, figs. 1, 2 and 2a. Original figures (natural size) reproduced herewith one-half natural size. (Upper figure, 1) Side view of a 14 ridge-plated third superior molar [paratype] of the right side, r.M®, described by Leith Adams (op. cit., p. 33) as follows: “2. Two beautiful and highly suggestive examples of what must be considered last true molars, are represented by the entire specimens Nos. 64 & 59 (Pl. vu. figs. 1&2 &2a). The former, an upper tooth, shows fourteen ridges, including the pygmy digitated posterior talon a, in a space of 7 inches [178 mm.]. Attached in front, although not shown in the figure, are two plates of the penultimate molar. As the crown is just being invaded, of course its pattern is not developed; the macherides are therefore well crimped, and the plates and enamel thick.” Brit. Mus. 44306. (Lower figures, 2, 2a) Crown and internal views of a right third inferior molar, r.M3. This is the type molar; the same tooth as that shown in Adams’ original figure (Fig. 1125 of the present Memoir). This beautifully preserved molar is de- scribed by Adams (op. cit., 1874, p. 33) as follows: The next, No. 59 (figs. 2 & 2a), is a much arcuated lower molar; the last ridge, although rounded and finger-like, rises like the others from the common base to the same level as the penultimate. There is a slight flattening on its base internally, but no trace of what could be called a pressure-mark. The crown is broad in front, tapering steadily posteriorly. The anterior talon is large and semilunar; and the anterior fang seems to support it and the succeeding plate only. Here we have four- teen ridges in 6.5 inches [166 mm.|.”’ Brit. Mus, 44804. THE LOXODONTINA: DWARFED SPECIES OF THE MEDITERRANEAN ISLANDS Palwoloxodon mnaidriensis Adams, 1870 Figures 1068, 1118, 1119, 1121, 1122, 1125-1127, 1131, 1132, Pl. xxur Pleistocene, Mnaidra Gap, island of Malta. This is the third species to be described from the island of Malta. Adams (1870, p. 223) calls it the “Large Elephant of Malta,” estimating its stature as six to seven feet, as compared with the Elephas melitensis of intermediate stature, and the Hlephas falconeri of extremely dwarfed stature, namely, two to three feet. Elephas mnaidre Adams, 1870. “Notes of a Naturalist in the Nile Valley and Malta,” etc., 1870, p. 223. Typr.—A last lower molar of the right side, r.M3 (Brit. Mus. 44304). Para- typE.—A third superior molar of the right side, r.M* (Brit. Mus. 44306). Horizon AND Locaurry.—Pleistocene, Mnaidra Gap, island of Malta. Type Fraurr.—QOp. cit., Pl. 1, figs. 2, 2a; also 1874, Pl. vn, figs. 2,2a. Paratype, op. cit., 1874, Pl. vm, fig. 1. Typr Description.—Adams (1870, p. 223) distinguishes this as “The Large Elephant of Malta,” estimating its average stature (op. cit., p. 228) as six feet ten inches to seven feet as compared with the average height of F. falconeri of two feet six inches to three feet (op. cit., p. 223). As EH. falconeri is distinguished by its extremely dwarfed stature, 2. mnaidrex is distinguished by its relatively large stature, and EH. melitensis by its intermediate stature. Adams (op. cit., p. 228) concludes: ‘Thus, to all present appearances, we have represented by the remains hitherto col- lected in the Maltese islands no less than three distinct species of elephants of about the proportions indicated, and represented in the spirited and well-executed drawing, page 161 [Fig. 1127 of the present Memoir], for which I am indebted to the able pencil of Mrs. Blackburn, whose admirable representations of animals have obtained for her a high position as a delineator of natural objects.” Subsequently Adams (1874, p. 116) added: ‘I believe that the bones of the Maltese fossil elephants are divisible into three varieties and two well-marked species, viz. a large and a small Elephant, the latter showing two forms represented by the Elephas melitensis of Faleoner and Busk, which may have seldom attained a height of 5 feet, and a diminutive or pygmy form named by Mr. Busk Hlephas falconeri, the smallest bones of which indicate an elephant about 3 feet in height. But there are intermediate- sized bones which easily bridge over the differences between the latter and the Hlephas melitensis; nevertheless Mr. Busk has pointed out characters appertaining to the two, and is of opinion that they are distinct species. . phant Llephas mnaidriensis, in consideration of the circumstance that the gap, or rock-rent, from which I obtained the most perfect specimens of its bony structure is situated close to the ruins of the Mnaidra temple, a prehistoric and megalithic structure bearing evidences of the earliest human occupation of the Island of Malta.” Leith Adams in this second contribution (1874, p. 116) changed the original name mnaidrx to mnaidriensis, the latter being the form cited by Lydekker (1886, p. 138), and gave an excellent new figure of the type molar (PI. vu, figs. 2, 2a), also the following ridge formula (p. 112 and Pl. vir): .. I have named the largest Ele- Dp 2% Dp 3 ¢ Dp 4 2-2 M1 22 8-9 M 2 io M3 224g -13-([14]" \ WigZ i Tn as 9: NY y DzENIZENS OF ANCIENT Matta.—l, Large fossil Elephant (Elephas mnaidre), p. 223. 2. Pigmy Maltese Elephant (Elephas melitensis), p. 216. 3, Smallest Dwarf Elephant (Elephas falconeri), p. 223. 4. Fossil Hippopotamus (H. pentlandi), p. 212. 5. Great Dormouse (Myoxus melitensis), p. 234. 5. Great extinct Swan (Cygnus falconeri), p. 237. 7. Large extinct fresh-water Turtle, p. 237. ApAMs’ RESTORATION AS DWARFED AFRICAN ELEPHANTS Fig. 1127. Reproduced directly from Adams’ figure in his book “Notes of a Naturalist in the Nile Valley and Malta,” 1870, opposite page 161. In this very picturesque drawing we observe that Adams has restored the external ears of these three species in direct imitation of the very large ears of Loxodonta africana, whereas it is probable that these dwarfed elephants had the smaller ears which we have shown in figures 1047 and 1074 to be doubtless characteristic of Hesperoloxodon antiquus and all its descendants. It should be remembered, however, that the external ears of Palxoloxodon namadicus, to which the type of P. melitensis is more nearly related, are still unknown. Compare the new restoration (I'ig. 1119), under the direction of the present author, based on the obvious affinity of these dwarfed elephants to Palxoloxro- don namadicus of India rather than to Hesperoloxodon antiquus of western Europe, also with small ears. 1266 Palwoloxodon lamarmorae [’orsyth Major, 1883 Sardinia, Quaternary sands of Morimentu near Gonnesa. Elephas Lamarmorae Major, 1883. “Die Tyrrhenis,” etc., Kosmos, 1883, VII Jahrg., Bd. XIII, p. 6. Typr.—The author in describing this species designates as the types “die Ex- tremitiiten-Knochen eines kleinen, aber vollstindig ausgewach- senen Elephanten zum Vorschein.” Horizon AND LocaLiry.— (Op. cit., p. 6): “Aus quatern. Sanden von Morimentu b. Gonnesa (Sardinien).”’ Type Ficurn.—No figures published. Typr Description.—The animal is said to agree in size with EB. mnaidre |E. mnaidriensis| from Malta, but differs pretty sharply in the carpal and tarsal bones which constitute the type. The author’s full description (Major, 1883, p. 7) is as follows: “Im vergangenen Jahre kamen in dem von LaMarmora beschriebenen quaterniren ‘Grés’ von Morimentu bei Gonnesa (Sardinien) die Extremititen-Knochen eines kleinen, aber vollstandig ausge- wachsenen Elephanten zum Vorschein. Die von Malta bekannten Zwerg-elephanten, mit deren grésstem, H. mnaidriensis, der sar- dische in den Dimensionen ziemlich gut iibereinstimmt, luden in erster Linie zur Vergleichung ein; es stellte sich aber heraus, dass die Carpal- und Tarsalknochen ziemlich bedeutend abweichen.” Summary.—Agrees in size with FE. mnaidriensis but differs pretty sharply in the carpal and tarsal bones which constitute the type. Palwoloxodon cypriotes Bate, 1903 Figure 1128 Pleistocene, Kerynia Hills, island of Cyprus. Elephas cypriotes Bate, 1903. ‘Preliminary Note on the Dis- covery of a Pigmy Elephant in the Pleistocene of Cyprus.” Proc. Roy. Soe., London, 1903, Vol. 71, No. 475, pp. 498-500; also “Further Note on the Remains of Elephas Cypriotes from a Cave- Deposit in Cyprus.” Phil. Trans. Roy. Soc. London, 1904, Vol. 197 (B), pp. 347-360. Cotyprs.—The folowing have been selected by the present author from the many cotypes: a second true molar of the right side (Brit. Mus. M.8588), also a last lower true molar (Brit. Mus. M.8591). Horizon AND LocALiry.— Pleistocene, a single cave deposit situated on the southern side of the Kerynia Range, island of Cyprus. Coryre FicurEs.— Op. cit., 1904, Pls. xx1, Xx. Bare (1903, pv. 500).—In the original description the author observes: ‘Taking into consideration the several characters in which the teeth of the Cyprus elephant differ from those of all the hitherto described dwarf species (putting on one side 2. lamarmorae ... from the Pleistocene of Sardinia, the teeth of which are un- known to science) as well as the distinct habitat of the animal, I have come to the conclusion that it is specifically distinct from these other small forms, though possibly they were derived from a common ancestor, and I, therefore, propose to name it Hlephas cypriotes.”’ Bate (1904, p. 357): In a supplementary description the author states: “Undoubtedly there is a strong resemblance be- tween the teeth of H. cypriotes and those of the Maltese and Sicilian forms, more especially 2. melitensis, but this likeness is OSBORN: THE PROBOSCIDEA very apt to be over-estimated owing to the respective dwarf proportions of these island races. The Maltese pigmy species have been considered to be most closely allied to H. antiquus and E. africanus. ... On the other hand it seems more probable that £. cypriotes, which so far as available material is concerned shows no close affinity to the African species, is rather connected with #. antiquus and FE. meridionalis, agreeing with this last in the lowness of its ridge formula, .. . though differing in wanting the persistence of the strongly marked digitation of the plates which is usually found in the molars of that elephant. CotypEs OF PALMOLOXODON CYPRIOTES Fig. 1128. Elephas cypriotes Bate, 1903, Kerynia Hills, island of Cyprus. Selection from the many cotype molars fully figured by Bate (1904, Pls. XXI, XXII). (Upper) Pl. xx1, fig. 3, “Crown view of second true molar in right man- dibular ramus. B.M., M.8588, p. 355.’ Reduced to one-half natural size. (Lower) Pl. xxn, fig. 6, “Crown view of last lower true molar. B.M., M.8591, p. 355’’; fig. Ga, “Side view of ditto.’’ Both reduced to one-half natural size. In this connection it is interesting to note that from the char- acters of the carpal and tarsal bones (the only remains known) of E. lamarmorae, the dwarf elephant of Sardinia, Dr. C. I. Forsyth Major considers it to have been most closely allied to HE. meri- dionalis. . . . Since writing the first notice of 2. cypriotes I have come to the conclusion that the average number of plates in the true molars is somewhat less than was at first supposed, therefore the corrected ridge-formula, exclusive of talons, would stand as follows: [E. cypriotes]: [Dp 2] +, [Dp 3] $, [Dp 4] 728, [M 1] 733, [M 2) Se), Mis) are This is slightly lower than that of 2. melitensis which Dr. THE LOXODONTINA: Leith Adams, later than Dr. Falconer and after examining a further large amount of material, gives as: [E. melitensis]: [Dp 2] 3, [Dp 3] 3, [Dp 4] 3-2, [M 1] 2:2, [M 2] 50, [M 3] +5. The marked lateral compression of the tusks of EH. cypriotes (Plate 22, fig. 9), which is a constant character in all the speci- mens so far obtained, would in itself be almost sufficient to distin- guish this from the other pigmy elephants of the Mediterranean region. .. . Presupposing that the dwarf elephants of the Mediter- ranean region were all derived from a common ancestor, the simpler construction of the molars of 2. cypriotes might be explained on the assumption that this species was isolated and subsequently differ- entiated from the parent stock, prior to a similar fate overtaking the Maltese and Sicilian races. This is borne out by the fact that E. cypriotes lived contemporaneously with H. [Hippopotamus] minutus, a more generalized and primitive form... than either H. pentlandi or H. melitensis, the associates of the pigmy elephants of Malta and Sicily. The geological evidence, as distinguished from the paleo-zoological, also supports this theory, for it appears prob- able that Cyprus became an island at an early period.” Palwoloxodon creticus Bate, 1907 Figure 1129 Pleistocene, cave deposit near Cape Maleka, island of Crete. Elephas creticus Bate, 1907. ‘On Elephant Remains from Crete, with Description of Hlephas creticus, sp. n.’”’ Proce. Zool. Soe., London, 1907, pp. 238-250. Coryprs.—Nine imperfect cotype molars. Two are figured in the present Memoir (Fig. 1129), namely, r.M; (Brit .Mus. M.9381) and an M2 (Brit. Mus. Horizon AND Locauiry.—Near Cape Maleka, Figurres.—Op. cit., Pls. x11, figs. 1-3, and M.9378). island of Crete. xin, figs. 1 and 2. DWARFED SPECIES OF THE MEDITERRANEAN ISLANDS 1267 Bate (1907, pe. 239).—The author described the type as fol- lows: ‘The remains of the smallest of the Cretan Elephants were all obtained from a much damaged and weathered cave-deposit in the limestone cliffs near Cape Maleka in the west of the island, which has already been described [Footnote: ‘Geol. Mag. n.s. dee. v. vol. ii. (1905) p. 195.’], and where only some teeth and limb- bones of small rodents were found besides those under discussion. These latter include nine imperfect molars and a few fragments, among which are a portion of an incisor and the dorsal half of a vertebra. As this small race differs from those of other Mediter- ranean islands, and its minute proportions being seemingly the result of specialisation due to isolation in Crete, it is suggested that it may be known by the above specific name denoting its island habitat.” Of the nine molars two belong to the upper series, namely, M! consisting of seven or seven and a half plates, length 54 mm., height about 832 mm. A referred M? includes eight plus plates. Of the cotype lower molars, one regarded as Mz includes nine plates; other lower second molars include eight to nine plates. Mz, in- cludes eleven plus plates. (Op. cit., p. 243): “From this brief account of the remains procured of F. creticus, it will be seen that this pigmy Elephant must have been of slightly larger proportions than FH. cypriotes and approached in size more closely to H. melitensis; that is to say, it would have attained as a maximum a height of five feet. . . All the molars obtained differ from those of the two last-named dwarf species in being much lower in the crown; this is perhaps the most noticeable feature of the series. At the same time the teeth are wide, the cement-areas broad, and the enamel simple, though at times broken up into a number of rings. So far as can be ascer- tained from the scanty amount of material the ridge-formula must have been low.” E. creticus: (M 1 ** M 2 8+ M3 w4G]. CoryPEs OF PALAOLOXODON CRETICUS Fig. 1129. Elephas creticus Bate, 1907, Pls. xu, fig. 3, x1u, fig. 1. Maleka, Crete. Both molars two-thirds natural size. Nine imperfect cotype molars from a cave deposit near Cape (Left) Pl. x1, fig. 3: “Crown view of right lower third molar of Z. creticus. ([Brit. Mus.] M.9381.)”’ +13 ridge-crests. (Right) Pl. x1, fig. 1: Crown view of second lower molar of E. creticus, Brit. Mus. M.9378. +7 ridge-crests. V. LES ELEPHANTS NAINS DES ILES MEDITERRANEENNES ET LA QUESTION DES ISTHMES PLEISTOCENES (VAUFREY, 1929) _ Enfin, par plusieurs beaux spécimens de Palerme, nous connaissons le crane de la plus grande, Elephas mnaidriensis, crane qui présente tous les caracteres propres al Elephas namadicus, c’est-d-dire al Eléphant antique, notamment le bourrelet suprafrontal si spécial. Il n’y a donc pas de doute que les formes naines des tiles appartiennent toutes trois au phylum de V Elephas antiquus et doivent étre considérées comme des races de cette espéce”’ (Vaufrey, 1929, p. 209). The valuable Memoir of Dr. Raymond Vaufrey, ‘Les Eléphants Nains des Iles Méditerranéennes” (October, 1929), reached the Osborn Library in December, 1929. The present abstract of the observations and conclusions of the author should be compared with the previous history and abstracts of the extensive literature cited more or less fully above. The author, under the guidance of Dr. Marcellin Boule, had the advantage of visiting the various caves and rock-fissures in Sicily and Malta, in which these fossils occur, and of examining and comparing all the collections. Among the many geologic observations of value are those in the Grotto of Luparello near Palermo (Fig. 1130) which show that the lower geologic level of ‘Elephas melitensis’ is much more ancient than the higher level containing ‘Elephas falconeri,’ from which Vaufrey rightly infers that the extremely dwarfed ‘E. falconerv’ is geologically more recent than the middle-sized ‘EH. melitensis.’ Vaufrey throughout regards the species ‘Elephas namadicus’ and ‘E. antiquus’ as synonymous. On the con- trary, it is shown above in the present Memoir, that ‘EH. namadicus’ Falc. (1846) antedates and is widely different specifically, if not generically, from ‘H. antiquus’ Fale. (1847) of western Europe. We need only abstract those portions of this Memoir which bear directly on the following questions: (1) The time of insulation or separation of these islands from the mainland; (2) the respective evidence of the rela- tionship of the dwarfed elephants to (a) the typical ‘Elephas antiquus’ of western Europe, or to (b) the ‘Elephas Couches a Elephas Falconer Principal niveau stalagmitique Couches a Elephas melitensis Mur de soutenement 3 0 5 10m Voie ferrée /(136750) eee fe PALA&OLOXODON FALCONERI MORE RECENT THAN P. MELITENSIS Fig. 1130. melitensis” at the bottom of the cavern and separated above by a broad stalagmitic layer from the “Couches 4 Elephas Falconeri.” After Vaufrey, 1929, fig. 7, p. 51. The underlying marine phase (“Plage marine’’), 136™, 40, is shown in the darkly tinted horizon at the bottom of the cavern. This “Plage marine’ may be geologically correlated with the Monastirian stage (Depéret, 1918-1921, named after Monastir, Tunis), elevation of the sea level, or insular depression. Section of Grotto of Luparello, Palermo, Sicily, showing below the ‘Couches 4 Elephas 1268 namadicus’ of India, or to (c) the north- wardly migrating elephants of Africa; (3) the time of entry of these dwarfed elephants into the Mediterranean Is- lands; (4) the geographic region from which they entered the islands. GEOLOGIC AND GEOGRAPHIC OBSERVATIONS The elevations and depressions of the Medi- terranean sea level and the consequent con- nections of its present islands with the sur- rounding mainland of Africa and Eurasia are fully treated in the Introduction (pp. 1-45), dis- cussed throughout the descriptions of the islands and in the formal text (pp. 45-202), and summarized in the Conclusions Générales (pp. 203-216). 1°». GroGRAPHIC AND FAUNAL RELATIONS OF THE Istanps.—Vaufrey (p. 2) adopts Boule’s (1906-1919) theory that in closing Miocene and early Pliocene time (Pontian) the western basin of the Mediterranean was by elevation reduced to large lakes between which, over great isthmuses, freely migrated the Hipparion THE LOXODONTINAZ: DWARFED SPECIES OF THE MEDITERRANEAN ISLANDS fauna of Léberon, Eppelsheim, Pikermi, Samos, Maragha, etc. After this early Miocene-Pliocene Pontian stage of elevation there followed a great depression of the land, several hundred meters below the present level, or a rise of the seashore lines, which completely isolated Europe from the African continent as well as from parts of Asia. In closing Pliocene time [Villafranchian! stage of northern Italy, containing Archidiskodon meridionalis, Hespero- loxodon ausonius, Parelephas trogontherioides| there is evidence of renewed African connections and migrations, but certainly no evidence of a Tunisian-Sicilian land bridge (p. 203), for from the latest hydrographic charts (1916) the Mediterranean Sea, while 400 m. below its present level, was too high to form an isthmus for intermigrations between Africa and Europe by way of Tunis and Sicily. Moreover, the absence of a Tunisian-Sicilian land bridge, or, in fact, of any land connection between Sicily and north Africa during late Pliocene and Pleistocene times, is proved by the contrast (Pomel, 1895, Boule, 1899) between the Pleistocene fauna 1269 ment apparentées: elles présentent toutes trois les mémes varia- tions de la morphologie dentaire, déja relevées par les auteurs dans les molaires d’Elephas antiquus. Enfin, par plusieurs beaux spécimens de Palerme, nous connaissons le crane de la plus grande, Elephas mnaidriensis, crane qui présente tous les caractéres propres A l’Elephas namadicus, ¢’est-a-dire 4 |’Eléphant antique, notam- ment le bourrelet suprafrontal si spécial. Il n’y a done pas de doute que les formes naines des iles appartiennent toutes trois au phylum de l’Elephas antiquus et doivent étre considérées comme des races de cette espéce.”’ Synonymy (op. cit., p. 143).—‘‘Les notions ainsi acquises nous ont permis, en méme temps, de vérifier l’identité des espéces nouvelles créées en Sardaigne par Forsyth Major, en Créte et a Chypre par Bate. II devient évident que le squelette de La Mari- menta se compare exactement par la taille au squelette de Luparello et se rapporte donc a l’Elephas melitensis, que |’Elephas creticus par la morphologie et les dimensions des molaires s’assimile par- SICILY MALta SARDINIA Cyprus CRETE 1B CREOLE VEER CSOT Ode tend OFS cho fsl HARES RETO AAI RE EE aE aa eee ET ect rs IN Is ne. ec cies Gan chs RSE HMInaLOmenszs meles (Crania TetCs) aa a MMNAlanLensts YPC. ac. ya a. esse sass ones ce eee E. mnaidriensis ref. IEINELULENSTS TEL.|. 1 0: fo oo gin wees E. melitensis type..... =H. lamarmoraetype..................... =F. creticus type Hpyalconeny Tele. cece aes 2s oes E. falconert type......... of north Africa, of Algeria, and of Tunis and that of the Mediter- ranean Islands, summarized by Vaufrey (pp. 204-206) as follows: (a) The Pleistocene fauna of north Africa is totally different from that of Europe or of the Mediterranean Islands; (b) the late Pleistocene entry of European forms into north Africa was not via a Mediterranean land bridge between Sicily and Tunis (p. 207) but by way of land connections along the eastern Mediterranean shore of Syria and the Suez; (c) whereas we find certain European and other Holarctic forms in the late Pleistocene of north Africa, we do not find a single African form in the Mediterranean Islands, more especially in the island of Sicily; (d) the fossil remains formerly attributed to the African elephant (2. africanus) belong rather to aberrant specimens of ‘Elephas mnaidriensis’; (e) thus rejecting (p. 208) all previous suggestions of north African relation- ship, Vaufrey summarizes his conclusions as to the geologic age, characters, and relationships of the dwarfed elephants as below. [THREE DWARFED SPECIES OR RAckES ONLY] “Cette démonstration, toutefois, ne pouvait se faire qu’en reprenant sur de nouvelles bases la question des Eléphants nains; elle n’efit done pas été possible si le hasard ne m’avait fait décou- vrir 4 Luparello de nouveaux documents. II ressort de cette étude que les Eléphants nains appartiennent A trois formes de tailles différentes: Hlephas mnaidriensis (hauteur au garrot: environ 1", 90), E. melitensis (hauteur au garrot: 1”, 40) et HE. Falconeri (hauteur au garrot: environ 0™, 90), caractérisées par la présence 4 la mandibule d’une premiére molaire [Dp2] 4 racine unique, particularité qui n’existe que chez |’Eléphant antique, 4l’exclusion dé toutes les autres espéces d’Eléphants. Par ce caractére comme par la morphologie des molaires, ainsi que par leur présence dans les mémes gisements, ces trois formes apparaissent comme étroite- faitement aussi 4 cette méme race, enfin que les diagnoses dentaires de |’Elephas cypriotes et de l’E. Falconeri sont si analogues qu’il n’y a aucune raison de les désigner sous deux noms différents. I] convient done de parler d’Elephas melitensis de Sardaigne et de Créte et non d’E. Lamarmorex et d’E. creticus, d’Elephas Falconeri de Chypre et non d’E. cypriotes. Exception faite, sans doute, pour quelques individus de taille normale, auxquels est dé l’introduction du phylum, les Eléphants ne sont donc représentés dans les les que par trois races de taille décroissante: EH. mnaidriensis, E. melitensis et E. Falconerz. D’une part, la morphologie dentaire, notamment en ce qui concerne les caractéres des figures d’abrasion, nous interdit de considérer les races naines comme appartenant a plus d’une espéce; d’autre part, les caractéres craniens d’Hlephas mnaidriensis, ainsi que la présence dans les trois races d’une premiére molaire inférieure 4 racine unique, désignent cette espéce comme étant indubitablement Elephas antiquus. INSULATION (oP. cIT., p. 181).—‘De méme qu’en Sicile, la faune A Eléphants nains nous apparait donc, d’une manieére générale, comme contemporaine du dernier Interglaciaire, et son extinction comme s’étant produite pendant la derniére période glaciaire au moment de la formation des couches qui la renferment. L’absence de toute trace de l’Homme dans ces couches confirme ce que nous ont appris déja les grottes siciliennes; Sicile et Malte nous apparaissent une fois de plus comme des finisterres.’”’ 1.. As to geologic age, Vaufrey concludes as follows (p. 209) : ‘“Rappelons enfin que les couches 4 Eléphants nains de Sicile et de Malte datent vraisemblablement de l’époque wurmienne, époque a laquelle |’Eléphant antique avait, sauf en Italie méri- dionale (Romanelli), disparu d’ Europe occidentale et que, dans le gisement de Luparello, la plus petite des trois races (Hlephas Falconeri) était superposée a celle de taille intermédiaire (2. melitensis).”’ [See footnote 1 on page 1049 above regarding the possible Lower Pleistocene age of the Villafranchian.—Hditor.] 1270 2. ReLaTionsHips.—The traditional opinion, seemingly shared by Vaufrey, is that the dwarfed elephants of the Mediter- ranean Islands should be regarded as the stranded descendants of the well-known early and late Pleistocene European species ‘Ele- phas antiquus,’ which roamed over the Mediterranean lands before they were broken up into islands. Vaufrey does not distinguish between ‘#. namadicus’ Falc., 1846, of India, and the typical ‘E. antiquus’ Fale., 1847, of western Europe; his comparisons and measurements are chiefly with those of the late Pleistocene [3d Interglacial| stages of Weimar and Taubach (see Fig. 1088 of the present Memoir), which he designates as ‘Elephas antiquus normal,’ selecting mean measurements between the larger and the smaller dimensions given by Pohlig (ef. Vaufrey, op. cit., p. 138—see Tables below). OSBORN: THE PROBOSCIDEA 108) 13-14, laminar frequence 3) (type pachyganal) to 6 (type endioganal). General specific characters (p. 110): Except for a difference in dimensions, (a) the general characters are similar to those of the typical ‘Elephas antiquus,’ with more feeble develop- ment of the enamel foldings correlated with reduction in the size of the molars; (b) asin ‘#. falconeri,’ there are two divergent tenden- cies in the grinding teeth, one of the fine enamel type (émazl mince) with compression of the lamine, the other of the thick enamel type (émail épais) with spreading of the lamin, corresponding respectively with the broad and short molars and the long and narrow molars described by Leith Adams. (5) Skeleton: Portions of mandibles, vertebre, femur, ulna, radius, attributable to ‘Elephas melitensis’; discovered also in Cyprus by Bate, in the cavern of Mnaidra by Adams, and in La Marimenta, Sardinia— COMPARATIVE MEASUREMENTS (cr. VAUFREY, 1929, p. 138) Measurements in Millimeters I. [Dp 2] II. [Dp 3] IIT. [Dp 4] IV. [M 1] V. [M 2] VI. [M 3] sey nh ne 3,60 5445 6,80 9,00 8 E. Falconert 1, 10 3,20 5,60 7,25 11,50 14,30? 7 chy es Qe 7,10 8,75. 12,70 14.50 Y. melitensis 1, 30 SPLIG - ADPASIE ERT Soe " Se pon 6,95 11,65 16,65 17,25 23,00 E. mnaidriensis 1, 60 Se _ ae seat Sane a 5 : ae 2 6,90 11,60 15,60 21,70 30,50 E. antiquus normal 1, 82 cee ERE = RRO aS ComParATIVE RipGE ForMUL® (cr. VAUFREY, 1929, pp. 91-128) . X4X-X5X X6X-X 8X 9 X10X-X12X 13-14 E. Falconer 3- 3X XBXTX = ee 10-X10X XXa Ti x14 E litensis ee end eM 8X-X8X 9X-10X 12X x12X 4. MelUens?s 2 6X-7X 8 10-11X 13)? 13-14 Pinion es ria X5X-7X 9x 11X 11X-13 16 (X14%) 4. MNALATLENSIS 3=19X x6ex K9X 12x X12X-13X 2X17 CHARACTERS OF THE SPECIES, AFTER VAUFREY In all the species the ‘premiére molaire’ (Dpz) is said to have a single root, a unique character of ‘E. antiquus’ (p. 58). Elephas antiquus Falconeri Busk (p. 91), represented by the rich dental series from Luparello, is regarded as identical with ‘Elephas cypriotes’ Bate of Cyprus. Distinguished by small tusks of sharp upward curvature, with a maximum diameter of 0,045 m. The teeth from Luparello are said to agree with the types of ‘E. falconer’ from Malta and of ‘E#. cypriotes’ from Cyprus. The superior and inferior molar series (pp. 91-101), examined in great. detail, give the following conclusions (p. 100): (1) Laminar fre- quence 45 to 6 in 10 em.; (2) marked reduction from ‘2. antiquus’ in the laminar ridge formula, accompanied by a relative thickening of the lamine: ‘2. falconeri,’ max. M 3 {+ [‘E. antiquus’ typicus M 3 {eictel; (3) incisive tusks with regular upward curvature [in contrast to the relatively straight tusks of ‘H. antiquus’]; (4) posterior grinders long and narrow, and elevated as in ‘EF. antiquus,’ with similar median sinus or expansion; (5) molars of two types, with thick (émail épais) and with thin (émail mince) enamel respectively (Pls. mr and tv); skeleton (p. 102), parts of vertebral column, humerus, ulna, radius, femur, tibia, carpus, and tarsus. Elephas antiquus melitensis Fale. (pp. 103-112), of inter- mediate size, from lower or more ancient (Luparello) cavern deposits than ‘#. falconeri,’ less completely known. (1) Incisive tusks relatively less robust and upeurved than those of ‘#. fal- known only in Luparello. (2) Grinding teeth (lower Luparello level) similar to those of the type found in the grotto of Mnaidra, Malta; laminar frequence 4 to 5 to 63 in 10 em. (Luparello and Malta). (3) Ridge-plates in third inferior molar (p. coneri,’ the latter described by Forsyth Major as ‘Elephas lamarmore,’ which Vaufrey regards as specifically identical with ‘Hlephas meli- tensis.’ Elephas antiquus mnaidriensis Leith Adams (pp. 113-139). Remains of this larger race not discovered in the cavern of Lu- parello, but an ulna found in the cavern of San Teodoro; richly represented by molars in the Museum of Palermo from other localities in Sicily and from Malta, including especially from the cavern of Puntali, Sicily, several crania and the greater part of the bones of the limbs, to which Pohlig (1893) devoted his short Memoir; also a complete series of grinders of ‘Hlephas mnaidrien- sis’ (Pl. v1) from Sicily and Malta, mostly of the pachyganal (émail épais) character. (1) The molars (p. 114) of ‘2. mnaidrien- sis’ almost reach the lower limits of size of the typical ‘Hlephas antiquus’ from which they are separated only by the constant absence of large individuals; nevertheless the bones from Puntali and the ulna from San Teodoro are invariably of inferior size to those of ‘2. antiquus’ of the continent. (2) Tusks (lig. 27, p. 115—Fig. 1131 of the present Memoir) longer and more slender, approaching those of ‘HF. antiquus’ of Tilloux (Fig. 11381,4) [which in turn resemble those of the Pignataro Interamna specimen (Fig. 1096)]; the length of the tusks is closely concordant with the length of the ulne (p. 113, figs. 26) as restored in figure 1134 of the present Memoir. (3) The grinders (pp. 116-132) include a nearly complete series (Dp 2-M 3) in which M 3 %,'4-" exibits a laminar frequence of 44 to 74 in 10 em.; the ridge-plates when worn show more or less median expansion [or ‘loxodont sinus’] at the base (igs. 28, 30) but none approaches the lozenge-shaped Loxodonta africana type; the enamel also varies in thickness from the émail mince with relatively close compression (Fig. 32, 4,6) to the THE LOXODONTINA: DWARFED SPECIES OF THE émail épais with wide compression (Fig. 37,5—Fig. 1132 of present Memoir). (4) The summary of dental characters (p. 131) is as follows: (a) A certain number of ‘#. mnaidriensis’ molars from the Sicilian horizons approaches the classic structure of the continental ‘BH. antiquus,’ namely, “couronne élancée et étroite, lames d’épais- seur modérée, émail plissé, souvent avec une expansion losangique médiane modérée”’; (b) from this central type, however, arose “nw BAe =? oo” nF MEDITERRANEAN ISLANDS 1271 region, a considerable period after the disappearance, during Inter- glacial Epoch 3 or Riss-Wiirm interval of the Alps, of the north German Hesperoloxcodon antiquus germanicus, and probably after the disappearance of the southerly H. antiquus italicus; (5) the ridge formule which Vaufrey assigns to ‘H. falconeri’ and to ‘E. meliten- sis’ are collective; these ‘collective ridge formule’ may belong to one or more specific stages, whereas in a certain stage of evolution the ridge formula is constant; thus Vaufrey may not be right in regarding as the same species the typical ‘EZ. falconeri’ of Malta and the typical ‘#. cypriotes’ Bate of Cyprus, which has a lower ridge formula; again Vaufrey’s ridge for- ny ! mula of ‘H. melitensis’ differs widely from that which Leith Adams finally 2 assigned to the same species; the ridge formula which Vaufrey assigns 3 to ‘EH. mnaidriensis’ namely, ++, is much below that of the Upper Pleistocene species of Taubach and 4 Weimar (Fig. 1088), i.e., germanicus, Fig. 27. — Schémas montrant les grandeurs relatives des défenses des races naines insulaires et de 1'Eléphant M3 ee italicus, M 3 Te it is also antique normal. 1, Elephas antiquus Falconeri de Wuparello; 2. E. a. melitensis de Luparello ; 3, E. a. mnaidriensis des Puntali; 4. E. antiquus de Tilloux (d’aprés M. Boule). — 1/20 de la grandeur naturelle. Fig. 1131. Relative size, thickness, and curvature of the incisive tusks in the three dwarfed subspecies of After Vaufrey, 1929, fig. 27, p. the Mediterranean Islands, one-twentieth natural size. antiquus’ of Tilloux (4) is found with some individuals of this species. variations, some of which accent the median sinus, broaden the ridge-plates, thicken the enamel, while others narrow the ridge- plates, diminish the median sinus, and thin the enamel. The latter reach an extreme degree of thinness and plate compression which remind us of the ‘Hlephas armeniacus’ Fale. [= Parelephas armeniacus of the present Memoir]. In brief, there exist in the cavern of Puntali all intermediates between these extreme types, but none approaches the loxodont type of ‘Elephas africanus’ or of the ‘Elephas priscus’ of Falconer. OSBORN (1930) SUBSTITUTES A THEORY OF AFRICAN DESCENT BY WAY OF EASTERLY RATHER THAN NORTHERLY LAND CONNECTIONS While recognizing the valuable and important observations of this Memoir, we are unable to accept many of Vaufrey’s conclu- sions, for the following reasons: (1) The cranium of the dwarfed elephants cannot be derived from that of the typical ‘Hlephas antiquus’ of western Europe, from which it differs very widely; (2) all the known Upper Pliocene and Pleistocene stages of ‘H. anti- quus’ described in detail in the present Memoir, namely, ‘ausonius,’ ‘antiquus’ (typicus), ‘germanicus,’ and ‘italicus,’ are too highly specialized in their ridge formule and other characters to give rise to these dwarfed Mediterranean species, which are primitive in ridge formule, and related to Palxolorodon namadicus in cranial structure; (3) the traditional opinion of ‘#. antiquus’ origin 7s not sustained by our present knowledge of the highly diverse characters of the African, west European, and south Asiatic members of the Loxodontine; (4) the dwarfed elephants date from the closing stage of Glacial time, namely, [IV GractaL= Wiirm of the Alpine below that of the Lower Pleistocene ‘E. antiquus’ typical of the Forest Bed, M 3 427232; (6) these facts seem to render highly improbable the descent of the dwarfed elephants from the large and specialized west European ‘EF. antiquus,’ or from the lower Upper Pliocene or the progressive Pleistocene stages, and render more probable the descent of these dwarfed elephants from east African forms, such as the ‘Elephas antiquus reckv’ Dietrich, 1916, M 3 ;s4@u, or the Algerian ‘H. jolensis’ Pomel, 1895, M3 or the Algerian ‘H. atlanticus’ Pomel, 1879, M 3,357. Finally, it appears probable that at least two separate lines of phyletic descent are represented in these dwarfed elephants of 115. The ‘Elephas 13) Fig. 1132. Two types of molars (A) and (B) figured by Vaufrey, 1929, fig. 37, Nos. 4, 5, 6, as belonging to ‘Hlephas mnatdriensis’ : (A) Nos. 4 and 6, from Shantiiin and Puntali respectively, of the ‘type endioganal’ (émail mince); (B) No. 5, from Puntali, of the ‘type pachyganal’ (émail épais). In Osborn’s opinion the ‘type endioganal’ (émazl mince) belongs to a different species from the ‘type pachyganal’ (émazl épais). One- sixth natural size. OSBORN: the Mediterranean Islands, as illustrated by Vaufrey (p. 109, fig. 23), namely: (A) with coarse enamel and ridge-plates widely separated, that is, with a low laminar frequence of 3% in 10 em., corresponding with Vaufrey’s ‘type pachyganal’ (émail épais), 6 Fig. 1133. Molars referred by Vaufrey to Elephas {| =Paleoloxodon| melitensis from Luparello, Sicily, (No. 1), and Benghisa, Malta, (No.6), showing respectively widely separated ridge-plates, i.e., low laminar frequence, with coarse enamel (‘type pachyganal’) and close laminar frequence, with fine enamel (‘type endioganal’). After Vaufrey, 1929, fig. 23, Nos. 1 and 6, one-half natural size. See also Fig. 1132 of the present Memoir. e.g., figure 23,1, and (B) with fine enamel and close laminar fre- quence, corresponding with Vaufrey’s ‘type endioganal’ (émail mince), e.g., figure 23,6. This distinetion of (A) and (B) is in accordance with the failure of all earlier authors clearly to dis- tinguish the phylum Parelephas and the phylum Archidiskodon from the phylum Loxodonta, which in the present Memoir are shown to be absolutely distinct from each other even at the close of Pliocene time. Thus we cannot support the author’s union (pp. 109, 110) of these (A) and (B) types in the single species ‘FZ. melitensis,’ in which (p. 110) he is erroneously supported by Leith Adams (1874, p. 35) as follows: ‘Comme chez I’Eléphant antique, il y a deux tendances divergentes, une vers la réalisation d’un type 4 émail mince avee resserrement des lames, l’autre vers celle d’un type a émail épais avee écartement des lames, aboutissant THE PROBOSCIDIEA dans le premier cas & des molaires larges et courtes (variété A de Leith Adams), dans le second 4 des molaires longues et étroites (variété B de Leith Adams).”’ Osporn’s Conciusions (1930).—(1) Whereas the progessive known west European stages neither of Hesperoloxodon antiquus nor of the Indian Palxoloxodon namadicus appear to be ancestral to the dwarfed Mediterranean species, we seem to find on the con- tinent of Africa a number of more primitive stages of Palxoloxodon, with more primitive ridge formule and long narrow grinding teeth, which further knowledge may prove to be ancestral to the dwarfed Mediterranean types. This will depend on the discovery of a cranium ancestral to that of ‘Hlephas melitensis,’ which, in turn, resembles that of ‘#. namadicus’ of India. (2) The true ‘mnaidriensis,’ ‘melitensis,’ and ‘falconerv’? seem to constitute a single phylum of regressive degeneration or dwarf- ing of the ‘endioganal’ or émail mince type. But intermingled with the most ancient and larger ‘mnazdriensis’ stage in certain of the caverns seem to be members of other phyla of the ‘pachyganal,’ or émail épais type. Fig. 26. — Figure montrant les grandeurs relatives des cubitus des trois races naines d’Elephas antiquus. De gauche a droite: E. a. mnaidriensis, E. a. meliten- sis (cubitus gauches), E. a. Falconcri (cubitus droit, avec un fragment du radius). — 1/6 de la grandeur naturelle. Fig. 1134. Ulne of the three species. After Vaufrey, 1929, fig. 26, p. 118. VI. ANCESTRAL STAGES OF PALAEOLOXODON IN AFRICA Whereas most previous writers, including Vaufrey (1929), derive the Mediterranean dwarfed insular species from Elephas antiquus of western Europe, we would be inclined to regard them as dwarfed insular derivatives of those extinct African species of Palzoloxodon, with which the genus Pilgrimia is synonymous. On a large scale ‘Elephas zulu’ Scott! resembles the types of Elephas antiquus Recki Dietrich, 1916, of Elephas jolensis Pomel, 1895, and of Elephas atlanticus Pomel, 1879, also of the dwarfed species of Malta, Hlephas melitensis Falconer, 1862, 1868, ‘Elephas falconeri’ Busk, 1867, and Elephas mnaidriensis Adams, 1870. From the subjoined table of ridge formule it appears that the dwarfed Mediterranean species differ from the west European species of ‘Hlephas antiquus’ |= Hesperoloxodon antiquus| but are closely similar to the African species of Palzoloxodon. Firsriy.—Since the first studies for this Memoir were begun (1900) evidence has rapidly accumulated not only to demonstrate that the Order Proboscidea originated in Africa, as recited in previous chapters of the present Memoir, but to render it probable that most of the separate genera and subfamilies also originated in Africa instead of Asia as formerly supposed. This apparently is true of the Archidiskodon phylum and now begins to be apparent in the Palzoloxodon phylum, because Africa and the Mediterranean Islands reveal stages of Palxoloxodon more primitive in molar ridge formule at least than any hitherto found in Eurasia, as displayed in the following comparative table com- piled from previous records and type formule: Middle to Upper Pleistocene of India Elephas |Palxoloxodon| namadicus Fale. and Caut., 1846, 1847 M 25, M3 Lower Pleistocene of England Elephas {Hesperoloxodon] antiquus Fale. and Caut., 1847, 1857 M 2 +5 M 3 164-17 Malta Elephas {|Palxoloxodon| mnaidriensis Adams, 1870 M2 M334 Elephas |Palzoloxodon| melitensis Falconer, 1862 M223 M33 Elephas |Palxoloxodon|] falconeri Busk, 1867 [ridge formula after Vaufrey, 1929} M 2 *%* M3 73:34 East Africa Elephas |Palxoloxodon| recki Dietrich, 1916 M2, MEG} aa Algeria Elephas |Palxoloxodon| jolensis Pomel, 1895 Me2 i M375 Elephas |Palzoloxodon| atlanticus Pomel, 1879 M2595 M3is5n Zululand Elephas (Loxodon) |Loxodonta] zulu Scott, 1907 M 2 — MD: =a East Central Africa Elephas |Loxodonta| zulu ref. from Kaiso Bone Beds M 2 — IMU) ss Recent Loxodonta africana M2 M3 SrconpLy.—The above ridge formule demonstrate that the recent Loxodonta africana is more primitive in dental ridge structure than any of the known Upper Pliocene or the Pleistocene species of Palzoloxodon; conse- quently L. africana cannot be descended from any known typical species of Palzoloxodon but may have sprung from a more primitive ancestral form still to be discovered.” Turrpiy.— It is a striking fact that the broad conspicuous ‘loxodont sinus’ of the recent Loxodonta africana is rudimentary or absent in all [see footnotes below, also pp. 1286-1288, this chapter] the extinct species of elephants thus far described from Africa or Eurasia. This absence or rudimentary condition of the ‘loxodont sinus’ removes certain extinct Pliocene and Pleistocene species from close relationship to the typical recent Loxodonta africana and relates them rather to the dwarfed species of the Mediterranean Islands which we also have grouped under the name Palxoloxodon (syn. Pilgrimia), as distinguished from the typical Palzxoloxodon namadicus and Hespero- loxodon antiquus. \(Elephas zulu Scott referred to Loxodonta by Professor Osborn (see Osborn, 1934.925, p. 2).—Editor.] 21See Lozodonta prima Dart, 1929, and L. africana var. obliqua Dart, 1929, below, this chapter, pp. 1287, 1288.—Editor-.| 1273 OSBORN: THE PROBOSCIDEA SYSTEMATIC DESCRIPTION OF AFRICAN SPECIES Palwoloxodon atlanticus Pomel, 1879 Figures 1047, 1135, 1136 Pleistocene. North Africa, Ternifine near Mascara, Algeria. SPECIFIC CHARACTERS (OSBORN, 1928).—The cotype second inferior molar (Fig. 1135), length 240 mm., breadth 60 mm., broadly resembles Falconer’s type of Elephas antiquus (Fig. 1075) except that the median ‘loxodont sinus’ is more widely expanded, re- sembling the conditions shown in Falconer’s aged referred ‘Elephas priscus,’ (Fig. 1076); the ridge formula, M 2 +2, agrees with his identification of the cotype molar (Fig. 1135) as a ‘‘Pénultiéme (5°) molaire inférieure’’; counting the two ‘talons’ as ridges, it agrees with the ridge formula at present known of Hesperoloxodon antiquus (typicus), namely, M 2 += M 3 122-47. The following is Pomel’s 1638-17" main ridge formula, omitting talons (1895, p. 51), of: E. atlanticus: Dp 2? Dp 3 $ Dp4?#M12M222M3 2. 10 History.—Pomel first named this species in 1879 but did not figure it until 1895. ONE-THIRD NATURAL SIZE Fig. 1135. Elephas atlanticus Pomel, 1879. One of the cotypes of Pomel, 1895, Pl. vin, fig. 1: ‘“Pénultiéme (5°) molaire inférieure, vue par la couronne, 4, et montrant le canal symphysaire, trés obtus; Ternifine;” identified as a right inferior molar, r.M». (Op. cit., p. 44): “‘J’ai fait figurer, Pl. vin, fig. 1 et 2, une magnifique molaire trés bien conservée, en partie encore contenue dans son os mandibulaire; elle a 240™™ de longueur avec une largeur de 60" au milieu; elle compte dix lames avec deux talons.” Observe: (1) That the central evaginations of the ‘loxodont sinus’ resemble those of the Hesperoloxodon antiquus, figured as ‘Elephas priscus’ by Falconer in 1868 and reproduced in our figure 1076; (2) that the ridge formula is 410-4 or 12, instead of 10 as given in Pomel’s description of 1895, p. 51; (3) that the measurements are, length 240 mm., breadth 60 mm. Cotyre R.Me oF PALZOLOXODON ATLANTICUS. Elephas atlanticus Pomel, 1879. ‘“Ossements d’fléphants et d’Hippopotames découverts dans une station préhistorique de la plaine d’Eghis (province d’Oran).” Bull soc. géol. France, Vol. VII, Ser. 3, 1879, p. 51. Corypre.—Second inferior molar, r.Me. Musée d’Oran, Algeria. Horizon AND LOocALITy.— Ternifine, Mascara, Algeria. Corypr FigurrE.—Pomel, 1895, Pl. vin, figs. 1, 2. Pomel figured this specimen as above in 1895, but he named it in 1879 (1879, p. 51): “Je me crois dés lors autorisé 4 donner A cette forme particuliére, actuellement disparue de la région atlantique, mais seulement depuis une époque trés-rapprochée des temps historiques, le nom d’Elephas atlanticus, rappelant le nom de la . superior molar from Ternifine M 3 2+. race d’hommes primitifs placés par la mythologie dans le massif du Nord de l’Afrique.”’ He designated the locality in 1895 (op. cit., p. 42): ‘J’ai désigné sous ce nom, dans diverses publications, une espéce trés particuliére d’éléphant découverte dans des stations de la pierre éclatée & Ternifine, pres de Mascara, retrouvée prés du village de la Sénia, par M. Féningre, dans des fouilles pour fondations et, plus réceemment, dans la caverne aux hippopotames de Pointe-Pescade.’”’ In the same publication (op. cit., p. 51) he defines the animal: “L’elephas atlanticus était armé de défenses robustes.” Osborn, 1930: Pomel’s detailed description (1895, pp. 42-51, Pls. vi-x) indicates a ridge formula (including ‘les talons’) for this species of : M 2 gyoce=a (length 240 mm., breadth 60 mm.). See page 44 of Pomel, also figure 1135 of the present Memoir. M 3 232-4214 (length $24 mm., breadth £4 mm., height of middle plate ;> mm.). See pages 43 and 47 of Pomel, also figure 1136 of the present Memoir. Thus Palzoloxodon atlanticus, while having a relatively low ridge-plate formula, namely, M 2 y;—s-% M 3 2232-3, is a very large elephant, equaling in size the typical Hesperoloxodon antiquus of the Forest Bed (Fig. 1088), namely, M 3 +5, length of inferior molar 315 mm., height of middle ridge-plate 126 mm. Counting the large anterior and posterior talons (Pl. vi), the ridge-plate count of the cotype is M 2 jz5, of the referred Consequently the ridge- plate count may be actually higher than that given by Pomel. Fig. 1136. atlanticus Pomel. Referred third superior molar of the left side, 1. M%, of Elephas After Pomel, 1895, Pl. vin, fig. 3: ‘“Derniére molaire supérieure, vue par la couronne; Ternifine; 44. Musée d’Oran.”’ This very large superior molar (op. cit., 1895, p. 47), length 324 mm., breadth 84 mm., exhibits a ridge formula of M 3 +4; the anterior and posterior ridge-plates rise to the surface of the crown and are described by Pomel as “talons.” Palwoloxodon jolensis Pomel, 1895 Figure 1137 Pleistocene. Algerian seacoast, North Africa. This is the second North African species described by Pomel, in the same year (1895) in which he published his type figure of Elephas {= Palzoloxodon| atlanticus. Osborn, 1924: Pomel’s type figure of Ms; of the left side ex- hibits the ridge formula M 3 js, a formula inferior to that of Hesperoloxodon antiquus, but exceeding that of Palzoloxodon THE LOXODONTINA:: ANCESTRAL STAGES OF PALASOLOXODON IN AFRICA atlanticus, laminar frequency 14 ridge-plates in 30 em.; it may be an ascending progression above P. atlanticus. Elephas jolensis Pomel, 1895. ‘‘Paléontologie Monographies. Les Eléphants Quaternaires.”’ Carte Géol. de l’Algérie, 1895, p. 32. Typr.—Third left inferior molar, 1.M;. Keole des Sciences, Algiers. Horizon aNpd Locauiry.—(Op. cit., ex- planation of Pl. v, fig. 3): “°. . . trouvée 4 la ferme Beauséjour, dans l’ancienne plage émergée,”’ Algeria, North Africa. TYPE Ficgure.—Op. cit., Pl. v, figs. 3 and 4. Type Description.—(Op. cit., p. 32): “La piece la plus im- portante pour la caractéristique de cette forme est représentée PI. v, fig. 3 et 4. Elle a été trouvée a la ferme de Beauséjour, en aval du Krober-Roumia, dans la plage marine soulevée et a été donnée 4 Ecole des Sciences par M. Maupas, conservateur de la Biblio- théque nationale d’Alger. Elle est remarquable par sa longueur, par son étroitesse et par son incurvation. Elle compte treize lames dont la derniére, trés étalée, peut 4 la rigueur passer pour un talon Tyee Lerr Turrp INrer1ioR MOLAR oF PALAOLOXODON JOLENSIS Fig. 1137. Type 1.M; of Elephas jolensis Pomel, 1895, Pl. v, figs. 3 and 4, one-half natural size: 3. “Derniére molaire inférieure, vue de profil, yy trouvée A la ferme Beausdéjour, dans l’ancienne plage émergée.”’ 4. ‘‘La méme dent, vue par la couronne.”’ (Pomel, 1895, p. 38): “Le fossile algérien qui a treize lames 4 son arriére- molaire inférieure, a cette dent longue de 140" avee une largeur de couronne de 36 et une hauteur, i la septiéme lame, de 70"™.” ambigu. ... Cette dent est assez particuliére par sa forme en long ruban, dont la partie postérieure est comme flanquée par une double rangée de disques de détrition séparés de la partie moyenne; ceux du coté extérieur se dédoublant plus habituellement que les intérieurs. Cette disposition est assez semblable a celle qu’offre la derniégre molaire inférieure de Velephas antiquus; mais dans celui-ci la dent est beaucoup plus grande; elle présente quatorze lames pour 30°"de longueur (ou méme quinze lames pour une longueur de 27°") au lieu de treize lames pour 14°".”’ Osborn, 1928: This narrow thirteen ridge-plated left third 1275 inferior molar, l.Mz2, clearly distinguishes this species from Pomel’s ‘Hlephas atlanticus.’ It is of very small size and very primitive, as shown by the comparative measurements given by Pomel (ef. 1895, p. 38): Length Breadth Ridge-plates Height 1.M; M; Max. Antiquus [Elephas antiquus (typicus) | 300 73 16 123 Mnaidriensis 170 50 13 70 Jolensis [Type] 140 36 13 70 Thus Palzoloxodon jo'ensis of Algeria is inferior in size to P. mnaidriensis of Malta; it is closely similar in size and in the number of ridge-plates to the P. melitensis of Malta. Palwoloxodon recki Dietrich, 1916 Figure 1138 Oldoway-Tuffe, Serengetisteppe, northern Tanganyika Territory. [Pleis- tocene.| Specrric CHARACTERS.—The relatively broad eleven to twelve ridge-plated second inferior molar suggests comparison with Loxo- donta zulu Scott, except that the ridge-plates appear to be more closely compressed and the dentinal exposures between the ridge- plates more narrow. This molar [lectotype], length 216 mm., breadth 62 mm. (Dietrich, 1916, p. 15), is relatively broader than Pomel’s cotype of ‘Elephas atlanticus’ (Fig. 1135). Mus. No. XVII 1384, 1.M. with 12 ridge-plates, length 216 mm., breadth 62 mm., height 120-130 mm. (ef. Taf. 1, fig. 2). Mus. No. XIII 711, 1.M:, with /-16- ridge-plates, length 330 mm., breadth 85 mm., height 150 mm. (ef. Taf. 1, fig. 3, and explanation of Taf. 1, also see Tab. 1, p. 15). Elephas antiquus Recki Dietrich, 1916. ‘‘Elephas antiquus Recki n.f. aus dem Diluvium Deutsch-Ostafrikas. .. .”’ Arch. Biontol., Bd. IV, Heft I, p. 22. Original in the Geologisch- Paliiontologisches Institut der Universitat, Berlin, XVII 1384. LecrotyPE.—Second inferior molar of the left side, 1.M.. Horizon anp Locatiry.—Oldoway-Tuffe, Serengetisteppe, northern Tanganyika Territory. Lecrorypr FiGuRE.—Op. cit., Taf. 1, fig. 2; cotypes, Taf. 1-vin (in part). Tyrr Description.—The lectotype, Me, is described by the author (op. cit., p. 22) as follows: ‘Wir kénnen aus den vorliegen- den Funden eine neue Lokalrasse oder geographische Abart des E. antiquus ableiten, die ich Herrn Dr. H. Reck zu Ehren Elephas antiquus Recki benennen méchte; sie verhalt sich in den Unterkieferzihnen folgendermassen: Die Kaufliche der Mz. ist wie bei H. antiquus typus schmal und bandférmig, die der M3 dagegen entschieden breiter als bei jenem und von ovalem Umriss oder mit geraden von vorn nach hinten zusammenlaufenden Riindern. . .. Die Zahl der Lamellen der hintersten Backenziihne ist trotz der Verlingerung dieser Zihne geringer geblieben als bei den geologisch jiingsten Formen des E. antiquus Da die Zihne keine Mittel- oder Ubergangsstellung einnehmen, d. h. keine neue Mutation, sondern eine neue Variante des Antiquustyps vor- liegt, so méchte ich die neue Bezeichnung E. antiquus Recki nicht als Namen fiir eine bestimmte Entwicklungsstufe im Antiquusstamm, sondern im zoologischen Sinn als Bezeichnung einer neuen geo- graphischen Rasse ... verstanden wissen.” Dietrich adds (op. cit., p. 75): ‘Nach den vorhandenen Lang- knochen diirfen wir fiir die Oldowayer Elefantenrasse Riesenmasse annehmen, so die Héhe im Widerrist mit 4m, die in der Kruppe nicht weniger, sondern eher mehr, denn die Riickenlinie des EL. antiquus senkt sich nicht wie beim Mammut von der Schulter nach hinten ab, sondern steigt wie beim afrikanischen Elefanten in der Kruppe wieder an. Der Riesenwuchs des E. antiquus Recki bestitigt die Tatsache, dass #. antiquus neben (nicht hinter) 2. meridionalis zu den gréssten Elefanten, ja zu den gréssten Land- siiugern iiberhaupt, zu rechnen ist.” Reck (1914, pp. 306 and 307) makes the following observa- tions: “Uber die Merkmale der Unterkiefermolaren (M» und Ms) macht Dr. Dietrich zur vorliufigen Charakterisierung der neuen Plefanten folgende Angaben: 1. Lamelle plattig mit gebogenen seitlichen Umrissen. 2. Schmale Seitenpfeiler, breiter Mittelpfeiler der Lamelle, ausgesprochener Mamillenbau. 3. Miassig dicker, missig gekriiuselter Schmelz. 4. Weite Distanzierung der Lamellen durch breite Zementin- tervalle; doch ist die Tialerweite ein schwankendes Merkmal. 5. Verhiltnismissig geringe Lamellenzahl (14 bis 16 bei Ms; 11 bei Mg). 6. Breite Zahnkronen der letzten Molaren (M;).”’ “ic ... Die Unterschiede zum Elefas Zulu Scott endlich legen hauptsiichlich in den Punkten 1, 3 und 4 der Charakteristik des Oldowayelefanten. Einige der Zihne, namlich die vorletzten Molaren erinnern stark an 2. antiquus Fale. aus dem europiischen Diluvium; doch entfernt die Gesamtheit aller Merkmale, besond- OSBORN: THE PROBOSCIDEA ers der M; den Elefanten von Oldoway vom Urelefanten, wihrend die Stosszihne beide Elefanten wiederum einander nihern (s. unten).” Dietrich’s ridge formula of Klephas antiquus recki is as follows: M 273 M3 iene Over-EstmateD Heicuts (Drerricn, 1916, p. 76, 1924, Pp. 24).—Dietrich concludes (1916, p. 76): ‘Als wichtigstes Ergebnis hebe ich hervor, dass die gréssten Hoéhen aller fossilen Elefanten um 4 m herum liegen, dass es keine 5 m hohen Elefanten gegeben hat, dass nicht einmal Dinotherium diese Hohe erreicht hat und LECTOTYPE OF PALMOLOXODON RECKI Fig. 1138. Lectotype 1.Me of Elephas antiquus Recki Dietrich, 1916, Taf. 1, fig. 2, one-third natural size. Geologisch- Paliontologisches Institut der Universitit Berlin, XVII 1384. From the Oldoway- Tuffe, Serengetisteppe, northern Tangan- yika Territory. dass schliesslich die stérksten Elefanten der Gegenwart hinter denen der Vergangenheit an Mass und Gewicht zuriickstehen.” According to Dietrich’s measurements and estimates of height, the ‘Hlephas antiquus recki’ is something over 4 m., namely, 4030 mm., in skeletal height, or 13 ft. 25, in., exceeding the tallest living African elephant by 2 ft., or610mm. This estimated height of ‘EF. antiquus recki’ is inferior to that of the ‘EH. antiquus german- icus’ of Taubach, in which the humerus is about 1300 mm. in length. According to this estimate both these animals exceed the height of the tallest of the known imperial mammoths, namely, Archidiskodon marbeni (3826 mm. or 12 ft. 65, in.), as shown in the comparative shoulder heights of living and extinct elephants (Chap. XVI, fig. 912). Dietrich’s shoulder height of the ‘2. antiquus germanicus’ of Taubach, estimated from the humeral length (1300 mm.), is still greater, namely, 4000+ mm. He assigns the same height (4000+mm.), estimated from the humeral length of 1300 mm., to the ‘Hlephas antiquus’ (typicus) of the Lower Pleistocene (Mauer) of western Europe. The length of the humerus, namely, 1290 mm., of Hesperoloxo- don antiquus of Upnor, as described above, agrees very closely with Dietrich’s estimates and measurements, giving an estimated shoulder height of 3700 mm. or 12 ft. 1% in.; this accords well with the fact that the Upnor skeleton is not full grown, because the second inferior and superior molars, M 2, are still in full use. By the practical agreement of the Dietrich humeral measure- ments with those of Forster Cooper (1928), we may reach a nearly THE LOXODONTINA: ANCESTRAL STAGES OF PALASOLOXODON IN AFRICA 1277 correct estimate of the maximum height of ‘Hlephas antiquus In brief, in comparison with the estimates of Archidiskodon germanicus’ of Taubach, namely: (tmperator) maibeni, as given in the legend of figure 912, and taking Humeral length 1300 mm. advantage of Dietrich’s excellent table of comparative measure- Skeletal height at shoulder 3729 ments (1916, p. 76), we may estimate the shoulder height of the Height in the flesh at shoulder 3965 skeleton (in millimeters) of the following species as below. COMPARATIVE SKELETAL MEASUREMENTS AND HEIGHTS WITH THOSE OF ‘HLEPHAS ANTIQUUS RECKI’ (Dierricn, 1916; Ossorn, 1930) The length of the humerus affords one standard means of estimating the skeletal height at the shoulder. In Archidiskodon maibeni the humerus is relatively shorter and the estimated shoulder height is made from the entire forelimb. ee Length Length Length Length Length Estimated | Millimeters Ms of humerus | of Mte. III of femur | of tibia skeletal height | at shoulder | E. antiquus recki Serengetisteppe, Tanganyika Territory 330 1235 240 1500! 900 3600 (H.F.O.) E. antiquus germanicus, Taubach 330-350 1300 ca. 250 1500 900 3729 (H.F.O.)| E. antiquus (typicus), western Europe 243-310 1300 1400+ 840 | Hesperoloxodon antiquus of Upnor 1290 246 21520e 1020 3700 Parelephas jeffersonii, Indiana 298e 1090 1250 3200 Archidiskodon maibeni, Nebraska 244 1251 3826 VII. PALAZ.OLOXODON AND LOXODONTA OF SOUTH AFRICA [The following section on Palzoloxodon and Loxodonta of South Africa has been compiled in accordance with Professor Osborn’s expressed views in his article in Novitates, No. 741, ‘Primitive Archidiskodon and Palaeoloxodon of South Africa’ (Osborn, 1934.925), that is, the type descriptions of the various species have been extracted in whole or in part from the original publications, under the generic reference of the present author, and reproductions have been made of the type figures. The Elephas |Loxodon] zulu Scott, 1907, at first regarded by Professor Osborn as referable to the genus Palzoloxodon, was finally referred in his article of 1934 to Loxodonta (Osborn, 1934.925, p. 2). Consequently the description, originally placed under Palxoloxodon above, has been removed to page 1286 of the present section, as in the case of Archidiskodon transvaalensis and A. sheppardi Dart, 1927, now regarded as Palzeoloxodonts rather than Archidiskodonts (see pp. 1284 and 1285 below).—Kditor]. TypicaL PaALMOLOXODON.—(Osborn, 1934.925, p. 14): “The type molars of the eight species referred to Palaeoloxodon . . . [below] are readily distinguished from Archidiskodon by the following five characters: (1) Dentine areas equal or exceed cement areas by relatively close compression of the ridge plates. (2) Absence of pre- and post-sinus central foldings, faint median expansion of the loxodont sinus. (3) Enamel relatively thin and more or less strongly and finely crimped. (4) Height of ridge plates increasing: P. kuhni=100 mm., P. wilmani=128 mm., P. archidiskodontoides = 145 mm., P. sheppardi=188 mm., P. transvaalensis=231 mm., P. hanekomi=259 mm. (5) Number of ridge plates. It seems probable that Dart’s type of A. sheppardi, dis- playing 1-13 ridge plates, is an LM2, in which case A. sheppardi becomes a synonym of A. transvaalensis Dart with 1-14 ridge plates.”’ See Dietrich, 1924, p. 24, where a length of 147 cm. is given as the greatest length.—Editor.] 1278 Palwoloxodon(?) andrewsi Dart, 1929! Figure 1139 Gong-Gong, Vaal River, South Africa. Middle terrace, 60-80 feet. Lower? Pleistocene.” (Osborn, 1934.925, p. 14): “A primitive or ancestral member of the Palaeoloxodon group may be this problematic A. andrewsi Dart (Fig. 5 [=Fig. 1139 of the present Memoir]), a type which on sectioning and very careful reéxamination by the present author, proves to be distinct both from A. planzfrons (Fig. 4 | = Fig. 876 of the present Memoir]) and A. subplanifrons (Fig. 1 [=Fig. 875 of the present Memoir]). The fragmentary type, f.LMs, displays the following characters: Ridge plate height =48 mm. est., estimated number of ridge-plates=6. Feeble pre-sinus fold; very prominent post-sinus fold. Valleys V-shaped. Estimated length=164 mm. Estimated breadth = 83 mm., estimated index=50. Enamel thick, crimped. It has been extremely difficult to restore this terribly shattered type specimen and deduce its outstanding characters . . . from the enamel folds which certainly belong at the front and back of the third inferior grinding tooth.” Archidiskodon andrewsi Dart, 1929. ‘‘Mammoths and Other Fossil Elephants of the Vaal and Limpopo Watersheds,” So. African Journ. Sci., Vol. XXVI, pp. 711-713. TyPr.— MeGregor Mus. 435, found by Mr. Luke Rademan and presented to the Museum by Mr. H. Rees. Cast Amer. Mus. 26968. [Re- garded by Professor Osborn as a left third inferior molar, 1.M3.] Horizon AND Locauiry.—‘‘Gong-Gong, Vaal River ‘Deep diggings at a depth of 80 feet,’ probably Middle Terrace. . . . ?Pliocene.’”’ Lower? Pleistocene.” Typr Ficgure.—Op. cit., 1929, fig. 14, p. 711 [= Fig. 1139 of the present Memoir]. Type DescripTion.—(Dart, op. cit., 1929, p. 712): “The fragments together embrace a length of 110 mm., the greatest width is approximately 80 mm. and the greatest height 60 mm. They include one fragment with two and a half plates and another with one and a half plates and a talon. The enamel is very thick (4-5 mm.). The plates are 17-22 mm. wide and display anterior and posterior buttresses as well developed as in A. [Archidiskodon| loxodontoides. Here the width of the plates is in the region of 25 mm. and by virtue of the abutment of adjoining buttresses upon one another the interlamellar cementum on either side of the tooth is separated into two parts as in A. loxodontoides.”’ “The enamel is markedly folded despite its thickness as in The distance between mid-points of adjacent ridge plates measured laterally is 1% in. The reduction (7-11 mm.) of interlamellar cementum on the medial side of the tooth and its almost complete absence on the A. loxodontoides, but in coarse fashion. lateral side in one fragment and its total absence in the other fragment together with the rapid contraction of the lamellar thickness (1 in. at base, in. at grinding surface) are witness to the Stegodontine characteristics of the molar—a feature to which OSBORN: THE PROBOSCIDEA attention has been frequently drawn in dealing with all these primitive mammoths. Further, despite the worn aspect of these fragments, I do not think that the true height was very greatly in excess of what the fragments portray.”’ “In form, measurements and general appearance, the tooth approximates more closely to A. loxodontoides than to any of the types hitherto discussed, but the coarser folding of its enamel, its One Centimeters 10 17 Yy nat. size McGregor Mus. 455 Type. Archidiskodon andrews. Dart. 1929 Type Or ?PALHOLOXODON ANDREWSI Vig. 1139. Restored type of “Archidiskodon” andrewsi Dart, 1929, MeGregor Museum 435, Kimberley, South Africa; cast Amer. Mus. 26968. Crown view restored with estimated 644 ridge-plates. Observe subequal cement and dentinal areas; pre- and post-sinus folds and two anterior ridges in contact; sharply V-shaped valleys between enamel ridges which penetrate about half the crown, thus differing widely from the enamel ridge-plates of A. subplanifrons. This is provisionally referred to Paleolorodon. One-half natural size. After Osborn, 1934.925, fig. 5, p. 13. reduced height, its lack and almost total absence of interlamellar cementum demonstrate the virtual certainty that we have here a still more primitive form not very far removed from the Stego- donts and apparently ancestrally related to A. vanalpheni and A. loxodontoides.”’ {It will be noted that the present species has been questionably referred by Professor Osborn to Paloloxodon (see Chap. XVI, pp. 984, 985 above, where it is provisionally listed under both the Metarchidiskodon griqua and Palzoloxodon transvaalensis groups).—Hditor.] °Cf. Table VIII, p. 984, of the present volume. THE LOXODONTINA: PALAXOLOXODON AND LOXODONTA OF SOUTH AFRICA 1279 Palwoloxodon hanekomi Dart, 1929 Figure 1140 Delpoort’s Hope, Vaal River, South Africa. Level unknown—Pleistocene. Archidiskodon hanekomi Dart, 1929. “‘Mammoths and Other Fossil Elephants of the Vaal and Limpopo Watersheds,” So. African Journ. Sci., Vol. XXVI, pp. 713-715. TyPE.— McGregor Mus. 2930. ‘‘?Third right upper molar.” Horizon AND Locatiry.—‘‘The old river bed of the Vaal River at a depth of 20 feet at Delpoort’s Hope [South Africa]. ?Upper Pleisto- cene.” Tyrr Ficurn.—Op. cit., figs. 15 and 16, p. 713 [= Fig. 1140 of the present Memoir]. Type DrscripTion.—(Dart. op. cit., 1929, pp. 714, 715): “This tooth was found by Major C. J. Hanekom on 23 October, 1927. It differs entirely in form from any hitherto discovered in South Africa being considerably shorter (200 mm.) than it is high (259 mm.). All specimens previously found have a length in excess of their height except in one case (A. [Palxoloxodon] trans- vaalensis) in which both measurements are virtually identical. The great height of this tooth (10% inches) equals that of the most gigantic form of Archidiskodont known, namely, A. ¢mperator Leidy of America. The greatest breadth (102 mm.) is virtually identical with that of A. [P.] sheppardi (100 mm.), but is not so great as that of A. [P.] transvaalensis (110 mm.). The number of plates also visible is 17 (Fig. 15 [=Fig. 1140 of the present Mem- oir]), that is, approximately identical with that of A. [P.] trans- vaalensis but in excess of A. [P.] sheppardi, where the number of lamellae is thirteen.” “Tn virtue of the hypsodonty of the specimen (Fig. 2 [= Fig. 1140 of the present Memoir]), there are only eight and a half plates in wear (Cf. figures of Mammonteus prigimius |primigenius| com- pressus, Osborn 1924). The irregularity of plate number in the worn area due to the curious nature of the enamel pattern—which forms an especially distinctive attribute of this particular species— is unparalleled in any South African mammoth type and recalls in some respects the irregularities of plate arrangements seen in Mammonteus (Elephas) primigenius (Cf. Zittel’s ‘Textbook of Palaeontology,’ Vol. 3, Fig. 349, 1925).” “The individual lamellae are as broad (11-15 mm.) as those of A. [P.] sheppardi, but do not reach so great a breadth as those of A. |P.] transvaalensis and the interlamellar cementum, unlike that in both those forms, is about as broad (10-12 mm.) as the lamellae. The lamellar walls are virtually parallel with one another, but only in the two anterior lamellae do they run entirely across the tooth and then only in a zig-zag fashion, demonstrating that each lamel- lar plate is virtually bifid to about 3 inches from its root base. These characters, coupled with the positive though not excessive fore and aft compression of the seventeen ridge plates and vertical elevation of the tooth or hypsodonty, show that we have in South Africa a further phase of mammoth evolution beyond what has heretofore been recorded for this country.” “The discovery of this exceedingly progressive form, whose hypsodont analogies are to be sought in America, renders it also probable that further discoveries of a convergent evolutionary nature between American and African forms will yet be made. .. . TYPE OF PALZOLOXODON HANEKOMI Tig. 1140. Type ?third right superior molar of Archidiskodon hanekomi Dart, 1929, from Delpoort’s Hope, South Africa. MeGregor Mus. 2930. After Dart, 1929, figs. 15 and 16, p. 718, side and crown views respectively. One-third natural size. Actually, being discovered at a depth of twenty feet in the river bed gravels at Delpoort’s Hope, it would seem that it belonged to an older geological horizon than A. [P.] transvaalensis and A. [P.] sheppardi, which were found at a depth of 4 to 5 feet in the river bed gravels higher up the river at Bloemhof. covery that these river bed gravels at Bloemhof present two dif- ferent gravel strata of very variable depth, separated from one another by two different phases of stone implement culture, Mr. Lowe’s dis- point to the necessity for similar investigations at Delpoort’s Hope and the possibility, despite the depth at which it was found, that A. [P.] hanekomi belongs to a still more recent phase of the Pleistocene than does A. [P.] transvaalensis.”’ 1280 Palwoloxodon yorki Dart, 1929 Figure 1141 Near Christiana, South Africa, Vaal River. Middle(?) Pleistocene. Pilgrimia yorki Dart, 1929. ‘‘“Mammoths and Other Fossil Elephants of the Vaal and Limpopo Watersheds,” So. African Journ. Sci., Vol. XX VI, pp. 719, 720. Typr.—MecGregor Mus. 4074, cast Amer. Mus. 22727. ‘Right lower (?third) molar.” Found by Mr. Alf. York during diamond digging operations in 1927. Horizon anp Locauiry.—‘‘Lowest stratum of the river bed gravels on the farm Vanasswegenshoek, O. F. 8., below Christiana, [South Africa], at a depth of within 6 feet... . ?Pleisto- cene.”’ Tyrer Ficgurr.—Op. cit., figs. 20 and 21, p. 719 [Fig. 1141 of the present Memoir]. Siete G LGD Typr OF PALAOLOXODON YORKI Pig. 1141. Type of Pilgrimia yorki Dart, 1929, from near Christiana, South Africa. “Right lower (?third) molar’? (McGregor Mus. 4074; cast. Amer. Mus. 22727). After Dart, 1929, figs. 20 and 21, p. 719, crown and side views, one-half natural size. Tyre Description.—(Dart, op. cit., 1929, pp. 719, 720): “This elephant tooth (Figs. 20, 21 [=Fig. 1141 of the present Memoir]), like the other remains found, is stony in nature and completely fossilised. It is a relatively diminutive tooth being 180 mm. long, 79 mm. broad, 115 mm. high, possessing 9 plates, all of which are in wear. There is no trace on the posterior plate or talon of a depression arising from the pressure of a tooth advancing behind it, as Falconer has pointed out (Palaeont. Memoirs, Vol. II, OSBORN: THE PROBOSCIDEA p. 294) is characteristic of a milk tooth. Hence the tooth must have been an adult one. . . . It approaches in number of ridge plates, enamel pattern, and tooth form to Z. priscus Faleoner and to H. trogontherti Pohl. There is exhibited a very slight tendency especially in the anterior crescents to throw out an anterior and posterior buttress, and hence to provide a vestigial loxodont sinus. The width of each lamella varies from the almost parallel regions where it is 12 mm., to 20 mm. in the region of the slight buttresses, but the lamellae are well separated by cementum across the whole grinding surface; the whole tooth is well eneased in the abundant cementum.” “The numerical seriation of this grinder of the lower jaw is a matter of question. It may be a second molar, but in any case it indicates by its reduced width, its symmetrical anteriorly-concave crescents, its slight tendency to single fore and aft buttresses, its more delicate enamel (2-3 mm. thick), and finer crimping a dis- tinctive type, so far as South Africa is concerned, whose relation- ships are with the most primitive Pilgrimia [= Palxoloxodon]. It is comparable in simplicity, but not in size, with EH. falconeri Busk. . . . Of known fossil forms outside Africa, it seems to ap- proximate most closely to Hlephas (Loxodon) priscus Falconer, not only in its dimensions and regularly crescentic plates with sinuous outline as viewed laterally, but also in the actual number of plates and the tendency to buttressing. Indeed, the appearances are such that A. [Palzoloxodon| yorki might well be ancestrally related to the #. (Loxodon) priscus of Falconer, as in A. [P.] yorki the processes leading to the production of such forms as F. priscus and E. trogontherii seem to be incipient.” “A. [P.] yorki is the simplest of the Pilgrimia [Palxoloxrodon| type yet recovered in Southern Africa.” Palwoloxodon wilmani Dart, 1929 Figure 1142 Below Christiana, Vaal River, South Africa. Pilgrimia wilmani Dart, 1929. ‘Mammoths and Other Fossil Elephants of the Vaal and Limpopo Watersheds,” So. African Journ. Sci., Vol. XXVI, pp. 720-722. TypE.— McGregor Mus. 4075, cast Amer. Mus. 22726. “Left lower (?third) molar.” Horizon AND Locauiry.—‘‘Lowest stratum of river bed gravels between the farms Vanasswegenshoek and Bloemheuvel, Transvaal, below Christiana at a depth of within 6 feet. . . ?Middle Pleistocene.” Type Ficgure.—Op. cit., fig. 22, p. 721 [Fig. 1142 of the present Memoir]. Tyrer Description.—(Dart, op. cit., 1929, pp. 721, 722): “This tooth fragment attains only the very narrow width of 70 mm. or 2%; in., but the height of the largest plate is 128 mm. It is, therefore, both narrower and higher than Pilgrimia |Palxoloxodon| yorki. Unfortunately the tooth is incomplete, the length of the fragment being 115 mm. and including seven plates only. There must have been in the original tooth at least one and probably two more plates posteriorly and another one, or two, more plates anteriorly. The ridge platenumber was ... +11 andapparently in excess of P. yorki (see Fig. 22 [Fig. 1142 of the present Memoir]). But the most striking dissimilarity between the two Middle(?) Pleistocene. THE LOXODONTINA: PALASOLOXODON AND LOXODONTA OF SOUTH AFRICA teeth lies in the character of the enamel plates which in this tooth are narrower (6 mm. in narrowest parts to 11 mm. in the slightly wider central parts), and are not so widely separated by the interlamellar cementum. Further, they run almost directly transversely across the grinding surface, displaying little if any tendency to crescentic outline on this aspect of the tooth. The lamallae are distinctly narrower laterally than medially (that is, the reverse of what is found in the modern African elephant and in Loxodonta antiqua zulu) where they are somewhat bulged, termi- nating in an anteriorly curved expanded portion almost as wide (9 mm.) as the widest central portion of the lamella.” “The same tendency towards a median buttress as was found in P. yorki is also encountered here, but to still less extent. The enamel also is more delicate (1-2 mm. thick) and is more finely crimped than in P. yorki. In sharp contrast also with P. yorkz, the tooth is not ensheathed in cementum but the ridge plates stand out to a depth of 4 to inch, except for the upper 1 to 1% inches of the lateral aspect, nor are the ridge plates much less obvious on the medial aspect of the tooth.” Typr OF PALAOLOXODON WILMANI Fig. 1142. Type of Pilgrimia wilmani Dart, 1929, from below Christiana, Transvaal, South Africa. “Left lower (?third) molar’ (McGregor Mus. 4075; cast Amer. Mus. 22726). After Dart, 1929, fig. 22, p. 721, one-half natural size. “There can be no question that this is another species of Pilgrimia [= Palxoloxodon|, which I shall denominate Pilgrimia wilmani, in honour of Miss Wilman, Director of the MeGregor Memorial Museum at Kimberley, who has been personally respon- sible for retrieving so many scientific treasures from the Vaal valley and permanently safeguarding them in that institution.” 1281 Palwoloxodon kuhni Dart, 1929 Vigure 1143 Pniel Estate, South Africa. ?River bed gravels. Middle(?) Pleistocene. Pilgrimia kuhni Dart, 1929. “Mammoths and Other Fossil Elephants of the Vaal and Limpopo Watersheds,” So. African Journ. Sci., Vol. XX VI, pp. 723, 724. Tyer.—McGregor Mus. 4144, cast Amer. Mus. 22725. ‘?Lower left molar.” | Hort- ZON AND Locauity.—[P. 723] ‘“Pniel Estate. ?River bed gravels. . . ?Pleistocene.”” [P. 724] “The site of discovery at Pniel must be fairly comparable with that at Vanasswegenshoek, and this indi- cates the necessity for stratigraphical study in the Pniel region in order to establish the correlation which is likely between these two widely separated sites along the Vaal valley. This is the more necessary in view of suggestions, which have been put forward from time to time, that the gravels of the river bed and of the terraces higher upstream are of a different age from the same gravel TyprE OF PALAOLOXODON KUHNI Fig. 1148. Type of Pilgrimia kuhni Dart, 1929, Pniel Estate, South Africa. ‘“?Lower left molar’ (McGregor Mus. 4144; cast Amer. Mus. 22725). After Dart, 1929, fig. 24, p. 725, one-third natural size. lower down stream. In the solution of this question the identifica- tion of elephant teeth types with particular gravels promises to be of premier importance.” Typr Ficurr.—Op. cit., fig. 24, p. 725 [Fig. 1143 of the present Memoir]. Type Descriprion.—(Dart, op. cit., 1929, pp. 723, 724): “The fragment consists of four plates and the greater portion of the anterior talon. Its dimensions (greatest length 96 mm., greatest width 75 mm. and greatest height 100 mm.) indicate to some extent its relationships. Its width and height are fairly closely comparable with those of certain specimens of H. antiquus Recki and in its general morphology it forms the closest approach to that form which I have hitherto seen in this country. In that form, however, the plates, as seen from the grinding aspect, are usually definitely crescentic, the horns of the crescents facing forwards. In this specimen they run transversely across the tooth. The anterior talon is vestigial, the most anterior true plate is continuous across the grinding surface, the second has a smaller left and a larger right island, while the third and the fourth have a central large island and internal and external smaller islets. These features indicate that the total length of the tooth could not have been great and the total number of plates in the tooth is unlikely to have exceeded seven or eight (see Fig. 24 [Fig. 1143 of the present Memoir]).”’ “The numerical seriation of the tooth is very doubtful. The enamel is very thin (2 mm.) and is, on the whole, very finely crimped.” 1282 OSBORN: THE Palwoloxodon archidiskodontoides Haughton, 1932 Figure 1144 Sydney-on-Vaal Breakwater, bed of the Vaal River, South Africa. Level unknown—Pleistocene. Pilgrimia archidiskodontoides Haughton, 1932. “On Some South African Fossil Proboscidea,”’ Trans. Roy. Soc. So. Africa, Vol. XXI, pp. 4-8. Coryprs.—“. . . a fragmentary skull with two worn molars that can be fitted into their sockets, a humerus which lacks the proximal end, the glenoid portion of a scapula, and the fairly complete left side of a pelvis.””, McGregor Museum, Kimberley, South Africa. Horizon AND LocaLtiry.— “Sydney-on-Vaal Breakwater in the bed of the Vaal River, 10% feet ‘below the maiden ground.’” Tyre Ficure.—Op. cit., Pls. ru [ = Fig. 1144 of the present Memoir]. Tyre Description.—(Haughton, op. cit., 1932, pp. 4-8): “In June 1930, Miss Wilman, the Curator of the McGregor Museum at Kimberley, forwarded to the writer for examination a collection of fragmentary mammalian remains which had been found by Mr. J.du Preez, jun., at the Sydney-on-Vaal Break- water in the bed of the Vaal River, 10% feet ‘below the maiden ground.’ In February 1931, thanks to the mediation of Major H. P. Tuckey, Miss Wilman received a further consignment, from the same locality, of material which was in the posses- sion of Mr. J. J. de Jong. Among the heterogeneous collec- tion of bones which made up these two consignments, the writer discovered some Elephantid remains—a fragmentary skull with two worn molars that can be fitted into their sockets, a humerus which lacks the proximal end, the glenoid portion of a scapula, and the fairly complete left side of a pel- vis. These are the bones that are briefly described and figured here.” “Teeth.— Both the left and right upper molars are preserved, PROBOSCIDEA both showing very worn grinding surfaces. From the fact that the socket into which the right tooth fits is bounded posteriorly by a wall of bone with a thickened and somewhat rugose ventral surface, and further that posterior to this wall is another, unfilled, socket, it is concluded that the teeth are the second permanent molars.” “The left molar has nine plates and a posterior talon preserved. The three anterior plates are worn down to the roots, so that the enamel of each has coalesced with that of the succeeding plate. The degree of wearing is greater on the inner than on the outer side of the tooth. As preserved, the length of the tooth is 148 mm.; the greatest breadth was probably 94 mm.; the maximum thick- ness of the plates (4th and 5th) is 16 mm.; the height of the posterior plate is about 145 mm. The length of the posterior six plates is 107 mm.; and six and a half plates occur in a length of 100 mm.” “There is no evidence of any tendency to the formation of a median loxodont buttress.” Tyrr or PALAOLOXODON ARCHIDISKODONTOIDES Fig. 1144. Museum, Kimberley, South Africa. Pl. 1, fig. 1. ?Second superior molar. About one-half natural size. Pl. u. Part of right humerus, anterior and posterior views. Type of Pilgrimia archidiskodontoides Haughton, 1932, from Sydney-on-Vaal Breakwater, bed of Vaal River, South Africa. After Haughton, 1932, Pl. 1, fig. 1, and Pl. 1, figs. 1 and 2. McGregor THE LOXODONTINA: PALASOLOXODON AND LOXODONTA OF SOUTH AFRICA “The enamel is of medium thickness (2.5-3 mm.), and is by no means strongly crimped.” “Tn the right molar the posterior talon is missing. Five plates occupy 80 mm., and there is very little interlamellar cement. The plates have their anterior and posterior faces roughly parallel, there being no marked antero-posterior thickening of the plate towards the root. Laterally there is a distance of 15 mm. from mid-point to mid-point of succeeding enamel ridge plates.” “Although they are wider than any of the three species de- scribed by Dart, these teeth seem—on account of their high lamel- lar frequency—to fall within the limits of the genus Pilgrimia [= Palzxoloxodon). .. . In Pilgrimia the frequency is 5-6 in P. yorki and 6.5 in P. wilmanz; and the length-lamellae ratio is 18.2 in the former and 16 in the latter. In the form under discussion the 1283 thicker ventrally than dorsally. In front of the molar, the palatal surface of the maxilla is strongly hollowed—a feature that is not seen in the skulls of Loxodonta africana that have been examined.” “Humerus.—Part of a right humerus is preserved, the bone lacking the proximal end. Its chief features can easily be discerned from the illustrations given. The bone is much more robust than that of Loxodonta africana. The shaft is thicker, and the deltoid crest much stronger and more prominent. The supinator crest is, proportionately, of about the same length; but, as in P. antiqua recki, its border is straight and has not the curved form with upper protuberance that is seen in L. africana.” “In actual size the bone is smaller than that of P. antiqua recki and that of P. antiqua andrewsi."! Its chief measurements are: Mice GIStANEMN Ml. «cys essere oe eyes vielsre wine eos ssa es Waduhetirochleararticularsurface: 5......02.-22.--.-02- 020.05 Height of top of supinator crest above distal end............. Minimum mtnickmessofishattie. ses deo. cc cle e sca et ces ce cose oe a (dae ae ITT | eeNe pee eS 265+ 350 350 227 ee 230 285 306 a Re to 267 410? 450 275 Ne ar: 125 145 167 80 figures are 6.5 and 15.4-15.6 respectively. The greater width of the two teeth described here tends to remove them from Pilgrimia as hitherto known in South Africa. Width, however, is a somewhat varying quantity within a species, as is evidenced by a study of a large number of molars of Loxodonta africana where the width varied from 60 mm. to 84 mm., and is dependent on the position and seriation of the tooth. Teeth from the lower jaw are always narrower than teeth from the upper jaw; and the upper molars of Pilgrimia york will certainly be found to be broader than 79 mm., which is the measurement given by the type lower molar. A breadth of 94 mm. is obviously not an impossibility for a Palgrimia upper tooth.” “That the form differs from any of the teeth of Pilgrimia hitherto described from South Africa is evident. In its breadth and in the tapering form of its plates it recalls certain forms of Archidiskodon such as A. yorki; and I propose, therefore, for the sake of convenience, to designate it by the new name Pilgrimia archidiskodontoides.” “Skull.—A portion of the maxillae, into which the two molars fit, is preserved. It shows that the palate was narrow and vaulted, and that the molars diverged posteriorly. In front the width between the molars is about 50 mm., and at the back it was about 90 mm. The vault of the palate is 60 mm. above the grinding surface of the teeth. The alveoli of the tooth in use and the suc- ceeding molar are separated by a wall of spongy bone, which is Comparative measurements of other forms are given in parallel columns. II—P. antiqua recki from Oldoway, East Africa. III—P. antiqua andrews |= Hesperoloxodon antiquus] from Upnor, England. 1V—Loxodonta africana from Addo.” “Although the head of the bone is not preserved, it can be seen that the inner border is more strongly bowed than in L. africana, and it is concluded that the caput humeri was larger or stood away further from the main axis of the bone than in the modern form.” “Pelvis.—The left side of the pelvis is almost entire, lacking most of the ischium, the symphyseal region, and the upper iliac crest.” “In shape, the ilium differs considerably from that of P. antiqua recki as figured by Dietrich. The pre-acetabular portion of the iliae plate is considerably longer; at the narrowest part of the shaft the outer face of the bone is practically flat, and above this the outer face is far less concave than in the East African form. From the ilium of L. africana this bone differs very considerably.” “The pubis has a much stouter shaft than in either P. antiqua recki or L. africana. In view of the broken nature of the borders of the specimen it is not possible to give many measurements; but the following show some of the differences between this specimen and that of a male L. africana from Addo. Shenexigenpthiof iliac blade. =: 0. coco. ieee eee eee es Distance from supra-acetabular ridge to spina anterior. ........ siiicknession lium at spina anterion...-.5.-..-.....-..28+eee- neighimoteacetabulumsa..22.0604.see sen. . o Widtheotacetabulimiasia ss cins.cabeat catod ee Meow asi tle Stan P. archidiskodontoides _\L. africana mm. mm. Sy SERRA ME Pe (As preserved) 680 646 | io ie eRe Lis 386 230 Re Ae ne late aU ta 145 85 PERU SAA re RRNA ESO, 214 | 150 Be eas On eee ea eee 200 (> eal4422 Equals Hesperoloxodon antiquus (sec footnote, p. 1222, above.)—Editor.] OSBORN: Palwoloxodontransvaalensis Dart, 1927 Figure 1145 From near Bloemhof, lowest terrace, Vaal River, South Africa. Pleistocene. Archidiskodon transvaalensis Dart, 1927. ‘“Mammoths and Man in the Transvaal.” Nature (Supplement), December 10, 1927, No. 3032, pp. 41-48. Typr.—Right third superior molar, r.M%, length 246 mm., breadth 110 mm., posterior height 47, fig. 6 (right), and 247 mm. Typr Ficure.—Op. cit., p. p. 48, fig. 7 (left). Dart’s Type oF PALAOLOXODON TRANSVAALENSIS! Fig. 1145. molar, r.M*, of the lowest Vaal River terrace gravels, near Bloemhof, South Modified after Dart’s photographs (cf. Dart, 1927, figs. Cast Amer. Mus. 27769. 14 ridge-plates, the maximum height of the Archidiskodon transvaalensis Dart. Type right third superior Africa. Pleistocene. 6, right, and 7, left), one-third natural size. This molar displays from 1 eleventh plate being 247 mm. Tyrer Description.—(Dart, op. cit., p. 47): “. . . the larger tooth was greater in every dimension. plates are missing, its present length equals that of the other tooth. It is 10 mm. broader and it is 50 mm. higher in its posterior portion. Although the anterior THE PROBOSCIDEA Apart from these points, the whole atmosphere of each tooth is different from that of the other. From the lateral aspect, despite its great height, the larger tooth has a massive squarish appearance, markedly different from the triangular form of this aspect in the smaller tooth. The individual plates are obscured in the larger tooth by a dense covering of cement over approximately the entire lower half (115 mm.) of this surface, and almost filling up the inter- lamellar clefts in the upper half, which are thus rendered broad and shallow. .. . From the grinding aspect, the larger tooth has a more bulging ovoid appearance than the narrower and more ellipsoidal appearance of the smaller tooth, as follows also from their re- Dart’s TYPE OF PALAOLOXODON SHEPPARD! Fig. 1146. Archidiskodon sheppardi Dart. Type left third superior molar, 1.M®, of the lowest Vaal River terrace gravels, near Bloemhof, South Africa. Modified after Dart’s photographs (ef. Dart, 1927, figs. 6, left, and 7, right), one-third natural size. This molar displays from 1-13 ridge-plates, as numbered, the height of the eleventh ridge-plate as preserved being 202 mm. Pleistocene. spective length and width measurements. Despite the fact that the total lengths of the two specimens are virtually identical, there are three (and perhaps more) additional lamelle in the larger than in the smaller tooth.” ‘Original specimens, formerly in the Ethnology Museum, University of the Witwatersrand, destroyed by fire.—Editor.] THE LOXODONTIN#A: PALAXOLOXODON AND LOXODONTA OF SOUTH AFRICA Palwoloxodon sheppardi Dart, 1927 Figure 1146 Near Bloemhof, lowest Vaal River terrace, South Africa. Pleistocene. Archidiskodon Sheppardi Dart, 1927. “Mammoths and Man in the Transvaal.’”’ Nature (Supplement), December 10, 1927, No. 3032, pp. 41-48. Typr.—Left third superior molar, 1.M$, length 246 mm., breadth 100 mm., posterior height 202 mm. Typr Ficurr.—Op. cit., p. 47, fig. 6 (left), and p. 48, fig. 7 (right). Type Description.—(Dart, op. cit., pp. 47 and 48): “In the smaller tooth the lamelle are covered with cement over ap- proximately the lower third (60 mm.) only of this surface, and above this point the interlamellar clefts are extremely deep (5-10 mm.) and narrow in appearance. The same features are repeated on the medial aspects of the teeth. . . . the individual lamelle are appreciably wider in the larger than in the smaller tooth, so that the interlamellar cement is more abundant in this tooth than in the former. There are also differences in form between the lamelle of both teeth, in that the narrower lamelle are more recurved poster- iorly at each end of the lamellz, and the lamine of each lamella possess a narrower or finer enamel and are more nearly parallel in the smaller than in the larger tooth. .. . In the smaller tooth the pattern presented by the digitations as they come into wear is considerably different, there being only three plates showing trans- ition stages from separate digitations to full plates. The most posterior of the three shows three small islets, the second shows four somewhat larger islets, and the third one very large medial islet and one small lateral islet. The remainder of the lamelle form complete single islands across the grinding surface of the tooth... . Sufficient differential characteristics between the teeth have been discussed to indicate that it is highly improbable that they belong- ed to the same species. Even if we looked upon the smaller tooth as being a second molar from a female, it is scarcely likely that there would be so great a gap between the two. In view, therefore, of the Sheppard brothers’ interest in securing the teeth and forwarding them for examination, I will denote the type indicated by this smaller upper and presumably third molar as Archidiskodon Sheppardi, sp. nov.” Dart concludes (p. 48): “It is evident, therefore, that the southern mammoths were represented in southern Africa by at least two distinct species of the genus [Archidiskodon], and that the line of their southerly migration is shown by the recovery of portion of a tooth of a nearly related species from the depth of 60-80 feet below the Nile at Khartum. .. . It has been shown that the lowest or mammoth |Archidiskodon] gravels of the Vaal bed are replete with evidences of the lower palolithic type of culture. They are therefore presumably pre-Bushman in orientation. The only pre- Bushman type known from extreme southern Africa so far is Bos- kop man. Containing, as they do, extinct forms of mammalian life, there is presumptive evidence furnished that these gravels will yet supply this, and perhaps other hitherto unidentified forms of human-kind, and show them to have been responsible for that culture. The recognition of extinct forms of mammalian life in the gravels of the river bed further enhances the age of the 60-80 ft. or mastodon terrace, and the evidences of paleolithic culture secured 1285 from this level and described by Hodkinson. The expectation of human remains there of great importance phylogenetically cannot be exaggerated, since this mastodon bed must reach back to a rather early phase of the Pleistocene. The 200-300 ft. terrace and any fossil mammalian remains or evidences of human culture at that level, which would appear to approximate if not actually to be situate entirely within the Pliocene, must be of premier anthropological importance.” NOTES ON ARCHIDISKODON [=PALAXOLOXODON] TRANSVAAL- ENSIS DART, 1927, AND ARCHIDISKODON [=PALHOLOXODON] SHEPPARDI DART, 1927 Referring to the site of discovery of two molars from the third or youngest terrace, or river bed gravel, Vaal River, near Bloemhof, South Africa (see above, p. 944, also Fig. 823), Dart continues (op. cit., 1927, p. 48): “The animal remains forwarded from Bloemhof consisted of two right [right and left] upper molars. . . . [p. 45] The present teeth have nothing in common with either Loxodonta griqua Haughton or with Elephas (Loxodonta) Zulu Scott. They represent an entirely different category altogether. . . [p. 46] They may, therefore, be included with those of H. meri- dionalis, E. planifrons, and EH. imperator amongst the southern mammoths in the generic phylum Archidiskodon of the subfamily Mammontine. .. . [p. 45] They are respectively right and left upper molars, and both are presumably third molars. Their measurements are as follows: Right. Left. [P. transvaalensis] [P. sheppardi| Third upper molar length 246 mm. 246 mm. i. “breadth Oe NO) K as “height of posterior portion 247 —** 202) “ The larger right superior molar (type of Archidiskodon [Palzoloxodon] transvaalensis) was found in a separate pit from the smaller left superior molar (type of A. [P.] sheppardi); they certainly belong to different individuals. There are thirteen ridge-plates in the left molar (sheppard?) and from sixteen to eighteen in the right molar (transvaalensis): “The lamelle [i.e., ridge-plates] are not compressed but are broad (17-19 mm. in the central portions, 13-15 mm. near the margin, and 15-17 mm. at the medial margin in the larger tooth, 7.e. right molar; 12-14 mm. in the central portion and 10-11 mm. near the inner and the outer margins of the smaller tooth, 7.e. left molar). The lamelle are considerably broader than the interlamellar dises of cementum; the cementum is nevertheless abundant in quantity— although not so abundant relatively as in H. planifrons and E. meridionalis types.”’ The enamel is definitely crimped. There is no tendency, as seen in #. [=Lozxodonta] zulu and Loxodonta [= Metarchidiskodon| griqua, to throw out a median posterior buttress or ‘loxodont sinus.’ The author rightly continues [p. 46] that these grinders belong to a genus not previously described from South Africa.! It remains to be determined whether the left (smaller tooth) represents a different species from the right (larger tooth); they present marked divergencies from one another in details, as shown in the accompanying type photographs (Figs. 1145 and 1146). Referred by Professor Osborn (Osborn, 1934.925, p. 2) to Palzolorodon.—Hditor.] 1286 Loxodonta zulu Scott, 1907 Tigures 1147, 1148 Zululand, South Africa, associated with the following extinct species: The Hippopotamus ponderosus, Bubalus andersoni, and Opsiceros simplicidens of Scott. Pleistocene. Speciric CHARACTERS.—This very large twelve or thirteen ridge-plated molar, identified by Scott as an 1.Ms, is relatively broad, length 265 mm., breadth 85 mm., height of posterior portion 81 mm.; it has therefore the relatively broad dimensions of re- ferred superior molars of Elephas antiquus, but the concave side of the ridge-plates (pointed forward) proves that this is a third left inferior molar, as identified by Scott. Consequently the ridge formula is apparently: INI armsact In deseribing this Upper Pliocene or Pleistocene species,! Scott observed that the type tooth does not resemble that of Loxodonta africana, because it lacks the characteristic s?nus loxodonte. On a large seale it resembles the types of Elephas antiquus Recki Diet- rich, 1916, of East Africa, and Elephas atlanticus Pomel, 1879, of Algeria, also the small Elephas jolensis Pomel, 1895, of Algeria, as well as the dwarfed species of Malta, Hlephas melitensis Falc., 1862, 1868, and Elephas mnaidrx Adams, 1870. Elephas (Loxodon) zulu Scott, 1907. ‘A Collection of Fossil Mammals from the Coast of Zululand.” Third Rept. Geol. Surv. Natal and Zululand, 1907, pp. 259-262. Typer.—Third right and left inferior molars, r.Ms, 1.M3. Horizon AND Locauiry.— Zululand, southeast coast of Africa. Tyre Ficgure.—Op. cit., Pl. xvi, fig. 6, Pl. xv, fig. 1. Tyrer Description.—(Op. cit., p. 261): “‘Indeed this tooth might almost be described as intermediate in character between Pohlig’s two groups, the Archidiscodonta and the Loxedonta. The third lower molar is very large, almost equalling in size that of the most gigantic modern African elephants, and in shape is elongate and rather narrow, though broader relatively than in FZ. africanus, so much so as to verge upon the laticoronate type. It will be ob- Types Lerr Tutrp INrERIOR Mouar oF LOXODONTA ZULU Fig. 1147. Same tooth as Pl. xvu., fig. 6." Less than one-half natural size. 85 mm. One of the two third inferior molars (left and right) constituting the type of Elephas (Loxodon) zulu Scott, 1907, Pl. xvi, fig. 1: “Left 3rd lower molar, crown-view; .. - Length 265 mm., breadth OSBORN: THE PROBOSCIDEA ; served from the table of measurements that while the breadth of crown equals the maximum recorded, even of the upper molars, for any individual of the existing African or Indian species, the length is considerably less than in these exceptionally gigantic individuals. ... It is plain, however, that the number of ridges, not including the talons, cannot have been less than 12, and may have been 13... . More significant than the number of the ridges is their shape, and one is immediately struck by the fact that in the fossil the pattern is decidedly less loxodont than in the recent African species; the median expansion of each lamina is less, and hence the successive ridges are more widely separated, while in the less abraded ridges the loxodont pattern is not displayed at all, the two enamel walls of each lamina being quite parallel. The enamel is very thick (pachyganal) and very strongly crimped. The lateral terminations of the lamin are either rounded or trifoliate, and each ridge has a feebly curved or crescentic shape, with the horns directed forward. The three posterior ridges have only the points of the digitations exposed, and of these there are four to each ridge, and those of successive plates are arranged in longitudinal rows in a manner suggestive of 2. meridionalis.” In several respects these teeth of #. zulu resemble the curious molars of E. antiquus, which were originally described by Falconer as HB. priscus. ... Measurements. Thirdllowenmolarslensth sere eee eee . 265 “s fS SWIthS ae co Gh ce sacle one 085 (a3 “ “ce height of posterior portion... .081”’ Scott regards this species as closely related to Hesperoloxodon antiquus and as possibly ancestral to Loxodonta africana. He concludes (op. cit., p. 262): “Should #. zulu eventually prove to be the actual ancestor of H. africanus, it would tend to give the latter a less isolated position, connecting it with Asiatic and European species. At all events, it is extremely interesting and important to find in South Africa an elephant with so many points of resem- blance to species characteristic of the northern hemisphere.” Reck (Sitzungsb. Ges. Naturf. Freunde zu Berlin, 1914, p. 307) remarks: “{1] Die Unter- schiede zum Elefas Zulu Scott endlich liegen hauptsichlich in den Punkten 1, 3 und 4 der Charakteristik des Oldowayelefanten. [2] Hinige der Zahne, niimlich die vorletzten Molaren erinnern stark an FE. antiquus Fale. aus dem europaischen Diluvium; doch entfernt die Gesamtheit aller Merk- male, besonders der M; den Elefanten von Oldoway vom Urelefanten, wiihrend die Stossziihne beide Elefanten wiederum einander nihern (s. unten). [3] E. Zulu ist der nur aus zwei zusammengehorigen M; Molaren bekannte ganz vereinzelt dastehende Fund eines dem neuen Elefanten noch am nichsten stehenden Tieres der Hlefas hysudricus-Reihe. Trotz- dem er primitiver und auch alter (Altdiluvial?) ist ‘(Professor Osborn first thought this species to be referable to Pal:coloxodon, but in his article of 1934 on the “Primitive Archidiskodon and Palacoloxodon of South Africa” (Osborn, 1934.925, p. 2) he definitely referred Elephas (Lorodon) zulu to Lorodonta.—Hditor. | THE LOXODONTINA: PALAOLOXODON als der Oldowayelefant, ist er doch schon zu viellamellig, um als direkte Ahnenform des Oldowayelefanten gelten zu k6nnen. lich ist das Material zu sicheren Schliissen zu diirftig. Frei- ” A thirteen Fig. 1148. ridge-plated third left inferior molar, 1.M3, from the Kaiso Bone-beds, near Lake Albert, Africa. After photograph kindly furnished the present author by Referred Loxodonta zulu (Brit. Mus. 12639). Dr. A. Tindell Hopwood (cf. Hopwood, 1926, Pl. 11, fig. 1). natural size. One-fourth Length 272 mm., breadth 65 mm. Loxodonta prima Dart, 1929 Figure 1149 Pilandsberg, Transvaal, bank of Rhenoster spruit tributary of the Limpo- po River, South Africa. ?Recent. Loxodonta prima Dart, 1929. ‘“Mammoths and Other Fossil Elephants of the Vaal and Limpopo Watersheds,” So. African Journ. Sci., Vol. XXVI, pp. 724-726. Typr.—McGregor Mus. 4077, cast Amer. Mus. 26987. ‘Left lower third molar {l.M,], fragment of right third molar, separated plates of upper molars.” Discovered by John Mostert. HorIzoN AND Locauiry.—Bank of Rhenoster spruit tributary of the Limpopo River, at a depth of 4 feet on the farm Nooitgedacht, Pilandsberg, Transvaal. ?Recent. Type Ficure.—Op. cit., figs. 25 and 26, p. 725 [Fig. 1149 of the present Memoir]. Tyre DescripTion.—(Dart, op. cit., 1929, pp. 725, 726): “The outstanding character of this tooth (Figs. 25, 26 [Fig. 1149 of the present Memoir]) is its comparability with the lving African species. It provides the first indubitable fossil evidence of a very close approximation to the distinctive lozenge-shaped lamellae of the living Loxodont grinding tooth. The absence of such a fossil type up to the present time according to Osborn a striking circumstance.’ The length of the reconstructed tooth is 4s AND LOXODONTA OF SOUTH AFRICA 1287 254 mm., breadth 74 mm., and height 142 mm. opposite the 7th plate, there being only 9 plates (6 plates in 6 inches), correspond- ing closely with L. africana and all in wear.” “There can be no doubt that in this narrow-crowned, loxodont- simulating, few-plated, mountain-inhabiting specimen here depict- ed, we have a long-sought ancestral type from which the modern African elephant tooth might reasonably be derived by a progres- sive widening-out of the loxodont sinus, for which reason I have named it Lorodonta prima. This fossil is of importance not only in demonstrating an advancing, though still simple Loxodont tooth pattern in the Transvaal, but also in revealing Africa, and possibly South Africa as the evolutionary home of the true Loxodonta. Its Fig. 1149. Type left third inferior molar, 1.M3, of Loxodonta prima Dart, 1929, figs. 25 and 26, p. 725, crown and side views respectively. McGregor Mus. 4077; cast Amer. Mus. 26987. One-third natural size. Found near Pilandsberg, Transvaal, South Africa. habitat in the elevated and relatively waterless Pilandsberg sug- gests the possible reasons for Loxodont persistence, namely, their becoming inured to more arid conditions and more active move- ment, as compared with their more ponderous Archidiskodont relatives.” Loxodonta africana var. obliqua Dart, 1929 Figure 1150 Valley of Steelpoort River tributary of Oliphants River, Northeast Transvaal, South Africa. ?Recent. Loxodonta africana var. obliqua Dart, 1929. ‘‘Mammoths and Other Fossil Elephants of the Vaal and Limpopo Watersheds,” So. African Journ. Sci., Vol. XXVI, pp. 726-728. yer McGregor Mus. 4078, cast Amer. Mus. 26988. ‘Right lower third molar [r.Ms].” Horizon anp Locauiry.—‘‘Valley of Steel- poort River tributary of Oliphants River, N.-E. Transvaal, on the farm Kranzkloof, at a depth of 17 feet... . ?Recent.” TYPE Ficurre.—Op. cit., figs. 27 and 28, p. 726 [Fig. 1150 of the present Memoir]. Tyre Description.—(Dart, op. cit., 1929, pp. 727, “This molar, forwarded to me in February, 1928, by Mr. C. Howard, was brittle superficially but nevertheless in an excellent 728): (ac 1288 state of preservation. It is a well worn tooth 256 mm. long, 76 mm. broad, and 137 mm. in height, possessing eleven complete and one rudimentary posterior ridge plates, 12 in all. Its measurements correspond very closely with those of Loxodonta prima just de- scribed, and also with those of the living form. But its corre- spondence with the living African elephant is demonstrated not only by its measurements but also by the number of enamel plates and the definite Loxodont character of the enamel sur- facesin wear. Indeed, its resemblances are so close as to render its separation from the existing species a matter of question.” Fig. 1150. third right inferior molar, r.M3, from the valley of the Steelpoort River, northeast Transvaal, South Africa. McGregor Mus. 4078; cast Amer. Mus. 26988. After Dart, 1929, figs. 27 and 28, p. 726, One-third natural size. Loxodonta africana var. obliqua Dart, 1929. Type crown and side views. “Certain features which perhaps are characteristics of variety rank, may, however, be referred to. It may be noted that in no case do the median buttresses impinge directly on one another but rather overlap uniformly to such an extent that the anterior buttresses impinge anteriorly on the posterior aspect of the medial portion of the ridge plate immediately in front of it, while the posterior buttresses impinge posteriorly on the anterior aspect of the lateral portion of the ridge plate immediately behind it. In this way the ridge plate rhombs are not regular but markedly skew or oblique in outline and appearance (igs. 27, 28 [Fig. 1150 of the present Memoir]). as being of special significance, seeing that a similar degree of obliquity due to overlapping of the rhombs is depicted in standard illustrations of the inferior molars of African elephants (e.g. Owen’s ‘Odontography,’ 1840-1845, Pl. 148, Zittel’s ‘Textbook of Palaeontology,’ 1925, Fig. 348), if it were not for the fact that in other illustrations (e.g. Faleoner’s ‘Palaeontological Memoirs,’ Vol. I should not have regarded this feature II, Plate 6, Fig. 1) of the same species, there is no evidence of such overlapping of the rhombs. In addition, there is present in this variety the full number of plates for the living species, together with a vestigial plate remnant posteriorly.” side views. OSBORN: THE PROBOSCIDEA “The salient enamel (3-4 mm. thick) is crimped, but so slightly that the general appearance is one of lack of crimping. The lateral terminations of the rhombs are not flattened but are on the contrary pointed while the medial terminations tend to be more rounded and flattened in contrast with the classical de- TypkE oF LOXODONTA SUBANTIQUA Fig. 1151. Type of Pilgrimia subantiqua Haughton, 1932, from Delport’s Hope, South Africa, “possibly a right lower molar, probably the second.” McGregor Mus. 4286. After Haughton, 1932, Pl. rv, figs. 1 and 2, crown and About two-fifths natural size. scription (Faleoner, 1868, ‘Palaeontological Memoirs’) of the living species.” “Whether these characteristics will prove in the long run adequate for the purpose of species differentiation it is difficult to ... In any ease, I have regarded the features presented as sufficiently distinctive to look upon it as a variety and to draw attention to these details in recording the presence of a fossil form extremely closely related to the living African elephant at great depth in the valley of the Steelpoort river in the Transvaal.” assess. Loxodonta subantiqua Haughton, 1932 Tigure 1151 Level unknown—Pleistocene. Pilgrimia subantiqua Haughton, 1932. “On Some South African Fossil Proboscidea,” Trans. Roy. Soc. So. Africa, Vol. XXI, pp. 8-10. Typn.—MeGregor Mus. 4286, presented by Mr. G. Barrell. “The tooth is possibly a right lower molar, probably the second.” Horizon AND Locauiry.—Delport’s Hope, half a mile from the Vaal River, South Africa. Found at Delport’s Hope, near Vaal River, Africa. THE LOXODONTIN: PALASOLOXODON OF JAPAN 1289 a depth of 40 feet in the ‘higher terrace.’ Pleistocene. Type giving a length-lamellae quotient of 19.5. The lamellar frequency Ficure.—Op. cit., Pl. rv, figs. 1 and 2 [Fig. 1151 of the present varies from 4.75 in the front part of the tooth to 5.75 in the back Memoir]. part, so that the individual lamellae increase in thickness towards - De uation, ee He Bee a Bee the root. They also taper in width upwards. The maximum Boo ts POsst PTY 2 night) ower molar, probably whe sec ond, thickness of a plate (4th) is 19 mm., and the minimum thickness of and its grinding surface is strongly concave in an antero-posterior direction. Eleven plates are preserved, of which the first has been worn down to the roots, and the last is just coming into wear. There was probably a posterior talon. The plates are fully covered with cementum both on their medial and lateral sides. The greatest length of the tooth as preserved, including the posterior “The tooth is low. The greatest height of any plate preserved, cementum, is 245 mm. Measured in a straight line, the ten and above the roots, is about 75 mm., and it is doubtful if the maximum a half plates visible on the grinding surface occupy 206 mm., height of any of the posterior plates exeeeded 85-90 mm.” interlamellar cementum is 4mm. The greatest width (including cementum) is 92.5 mm., and the maximum width of a plate at the grinding surface 79 mm. The enamel is not thick and is coarsely crimped.” VIII. LOXODONTINES OF JAPAN AND JAVA (Continued from Chap. XIV, pp. 901 to 909, and from p. 1185 of the present chapter) In this concluding historical and systematic section of the Loxodontinse we may review the original as well as one of the most recent treatments of the far eastern loxodonts of Japan and of Java, at a time when these countries constituted the eastern portion of the Asiatic continent and successively attracted the mastodonts, the stegodonts, and the true elephants chiefly of the genus Palzxoloxodon. History.— During the years since Dubois described (1908) his ‘Elephas hysudrindicus’ from the Kendeng for- mation of Java (now believed to be of Middle Pleistocene age), great changes in nomenclature have been made which are thoroughly set forth in the following systematic revision of eight species and subspecies! originally named and more or less fully described by Dubois, Makiyama, and Matsumoto, the total list to our present knowledge being as follows: PROBABLE RipGE ORIGINAL REFERENCE REFERENCE IN PRESENT Memoir Gro Loaic AGE FORMULA JAVA Elephas hysudrindicus Dubois, 1908 = Palxoloxodon hysudrindicus Middle(?) Pleistocene M 3 y'su JAPAN Elephas namadicus naumanni Maki- yama, 1924 = Palzxoloxodon namadicus naumanni Middle Pleistocene (?) M 3 73 Elephas namadicus namadi Maki- yama, 1924 = Palxoloxodon namadicus namadi Middle Pleistocene (?) M 3 7*** Euelephas protomammonteus Matsu- moto, 1924 = Palxoloxodon protomammonteus Upper Pliocene (?) WLS) area Loxodonta (Palxoloxodon) tokunagai Matsumoto, 1924 = Palxoloxodon tokunagai Upper Pliocene (?) WES eae Parelephas protomammonteus proxi- mus Matsumoto, 1926 = Palxoloxodon protomammonteus proximus Upper Pliocene (?) IMLS} a Loxodonta (Palxoloxodon) namadica (Yabez) Matsumoto, 1929 =Palxoloxodon namadicus yaber Middle Pleistocene M 3 eazemm Lox. (Pal.) Tokunagai junior, mut. Matsumoto, 1929 = Palxoloxodon tokunagai mut. junior Upper Pliocene(?) or Lower Pleistocene Mo 2\saan 'To these should possibly be added the following species described since this text was written and not examined by Professor Osborn: Parelephas |?Palxo- lorodon| protomammonteus matsumotoi Saheki, 1931, from Mishima, Province of Kazusa, Palxoloxodon yokohamanus Tokunaga, 1934, from Yokohama, and Palxoloxodon aomoriensis Tokunaga, 1936, from Tenjinbayashi, Aomori Prefecture.—Kditor. | 1290 OSBORN: THE PROBOSCIDEA Whereas Dubois and Makiyama gave merely a preliminary description of the three species named by them (B. hysudrindicus, E. naumanni, BE. namadi), Matsumoto named five species and contributed a succession of short articles and memoirs, fully listed in the Bibliography of the present Memoir; also in aid of the publication of the present Memoir, he sent the author in 1924 a letter giving his views at that time (June 24, 1924). His more mature views are expressed in his two memoirs entitled, ‘On Loxodonta (Palxoloxodon) namadica (Falconer and Cautley) in Japan,” with six plates (Matsumoto, 1929.1,) and “On Loxodonta (Paleoloxodon) tokunagai Matsumoto, with Remarks on the Descent of Loxodontine Elephants,”’ with one plate (Matsumoto, 1929.2). SUMMARY OF MATSUMOTO’S FINAL OBSERVATIONS AND THEORIES OF 1924 AND 1929 The present author differs widely from Matsumoto’s opinions and theories as to phylogeny, geographic distribution, and nomenclature; paleontologists are nevertheless deeply indebted to him for his great labors in setting forth the characteristics, geographic distribution, and geologic succession of the Japanese loxodonts. The following gives a summary of Matsumoto’s observations and theories. Marsumorto, 1924.—We owe to Doctor Matsumoto (letter, June 24, 1924) his geologic section (Fig. 790) of the fossil bearing formations of Japan and a series of valuable notes on the Japanese forms referable to Loxodonta | = Palxoloxodon| namadica. It appears that all varieties of L. namadica from Japan were distinguished by smaller and narrower grinding teeth than those of the typical L. namadica of the Nerbudda Valley. This relatively small and relatively narrow molar proportion is characteristic of all the insular species of the genus Loxodonta | = Palxolox- odon|; it is also a primitive character of all loxodontines. According to the 1924 observations of Matsumoto: (1) The parent forms of Loxodonta |= Palxoloxodon|} migrated into Japan early in Pleistocene time, namely, in the Calabrian-Villafranchian age, in which occurs an older type almost similar to Loxodonta [= Hesperoloxodon| ausonia in its evolutionary stage; this is perhaps of Upper Pliocene Calabrian or Villafranchian age; (2) in the succeeding Cromerian-Sicilian stage there occurs an ascending mutation, the Loxodonta namadica var. nawmanni of Makiyama [since made the “‘subgenotype”’ of Palzx- oloxodon by Matsumoto], almost similar to the type of L. |= Hesperoloxodon] antiqua which occurs in the Cromer Forest Bed; (3) a still more modern type, approaching more nearly the typical L. [= P.] namadica of India, occurs in the terrace gravels corresponding perhaps with the Monastirian or Tyrrhenian stage of Depéret. In descending geologic order these species appear to Matsumoto (letter, June 24, 1924) as follows: Ill. Loxodonta namadica ?mut.=a final mutation of the younger terrace gravels of ?Monastirian-Tyrrhenian age. Narrow toothed, like the species nawmanni, but thin ridged and thin enameled; loxodonty absent or very slight in the upper grinders, and slight loxodonty in the lower grinders; laminar frequency 7-8. Il. Loxodonta antiqua var. naumanni Makiyama = an older type, mutation of the Tokyo beds of ?Cromerian-Sicilian age. Al- most similar to the species of the Cromer Forest Bed Loxodonta antiqua, as well as to the L. hysudrin- dica of Java, in its evolutionary molar stage; laminar frequency 6-6.5. I. Older type, almost similar to Loxodonta ausonia of the Villafranchian=mutation of Minato, its geographical locality. Almost similar to L. ausonia in its evolutionary stage; crown low, Ms; about 120 mm.; laminar frequency 6-6.5; perhaps of Calabrian-Villafranchian age. According to the observations of Matsumoto, the narrow-toothed Japanese elephants differ from the typical broad-toothed Loxodonta |= Palzxoloxodon| namadica types of India and resemble the typical narrow-toothed L. |Hesperoloxodon| ausonia types of southern Europe, the Mediterranean Islands, and North Africa. THE LOXODONTINA: PALAAOLOXODON OF JAPAN 1291 Matsumoto, 1929.—The following passages are taken from Matsumoto’s Memoirs of 1929 (1929.1, 1929.2), entitled: On Loxodonta (Palzoloxodon) namadica (Falconer and Cautley) in Japan. Sei. Rept. Tohoku Imp. Uniy., Second Series (Geology), Vol. XIII, No. 1, (1929.1). On Loxodonta (Palxoloxodon) tokunagai Matsumoto, with Remarks on the Descent of Loxodontine Elephants. Sci. Rept. Tohoku Imp. Univ., Second Series (Geology), Vol. XIII, No. 1, (1929.2). (Matsumoto, 1929.1, p. 1): ‘“‘Assistant Professor Makiyama of the Kyéto Imperial University has really laid the cornerstone of further progress in the study of Hlephas namadicus Falconer & Cautley in Japan, of which two subspecies have been distinguished by him. Subsequently, his #. namadicus Naumanni was selected by the present writer to be the subgenotype of Palzoloxodon, which is referred to Loxodonta. The writer has now come to distinguish three races of the species in question in Japan. One of them, corresponding to the subspecies Nau- manni Makiyama, represents the Lower Pleistocene mutation of the species, while the other two appear to be characteristic of the Middle Pleistocene. Thus, the racial subdivision of the present species may play a part, the writer hopes, in the geological correlation of the Japanese Pleistocene.”’ (Matsumoto, 1929.2, pp. 7, 10): ‘‘An interesting archetypal Loxodontine elephant of Japan, named Loxo- donta (Palzoloxodon) Tokunagai by the writer, as well as the subgenus Palzxoloxodon created by him, has not yet been fully described in a European tongue. The writer here wishes to furnish a description of the elephant with a few remarks concerning the descent of Loxodonts. On this occasion, the writer has the pleasure to express his hearty thanks to the authorities of both the Imperial Museum of Uéno and the Geological Institute of the Kyéto Imperial University, by whom he was permitted to study the specimens described in the present report.” “Risk OF PALAOLOXODON”’ ‘As specially noticed by Lydekker [Footnote: ‘Lydekker, Brit. Mus. Cat. Foss. Mamm., Pt. IV., 1886, pp, 102, 103 & 106, text-fig. 25.’], a small form of very archetypal elephant is recorded from India under the name of Elephas planifrons. This form appears to be deviant from Archidiscodon in being narrow-toothed, though it appears to be closely related to the same in having the loxodont sinus of an obtuse type in the grinders. Again, by the first mentioned characteristic, it appears to be close to and by the second to be deviant from Palzxoloxodon. Phylogenically, it is, in all probability, ancestral to Palxoloxodon, and hence to the Loxodontine elephants as a whole. Thus, the affinity of this form appears to be stronger to the side of Palxoloxodon than to the side of Archidiscodon. A generic and a specific name have already been proposed to receive it [Footnote: ‘Matsumoto, on Leith-Adamsia Siwalikiensis, Jap. Journ. Geol. & Geogr., Vol. V., No. 4, 1926-1927.’]. Palzoloxodon, of which the most archetypal known form appears to be represented by Loz. (Pal.) Tokunagai, might have arisen from such an earlier form as represented by the aforementioned species through the partial perfection of the loxodonty of the grinders. No doubt, the Loxodonts as a whole were originally a narrow-toothed type, with a low ridge-formula. The acquisition of loxodonty might mechanically be correlated with the combined facts of their being narrow- toothed and long retaining a low ridge-formula. Palxoloxrodon appears to have originated in the region extending from India to Japan, and then to have been distributed over practically the entire southern Palearctic.” [Leith-Adamsia siwalikiensis Mats. is a synonym of Archidiskodon planifrons Fale. and Caut. (see Chap. XVI, p. 959, of the present Memoir). | ‘“PaL#OLOXODON AND LOXODONTA, S.S.”’ “The writer has never seen a molar of Palxoloxodon, which is more like those of Loxodonta, s.s., than the type- specimen of Loz. (Pal.) Tokwnagai, in being very narrow-crowned and in the very markedly lozenge-shaped disks 1292 OSBORN: THE PROBOSCIDEA of the well-worn ridges. If we suppose that the cheek-teeth of just this type had acquired hypsodonty of a very high degree, without increasing their width and the number of ridges, then we may obtain cheek-teeth of a type just characteristic of Loxodonta, s.s._ The modern Loxodonts might have originated in such a way, probably in Africa.”’ “SPECIFIC GROUPS IN PALHOLOXODON”’ “The most archetypal group in Palzoloxodon corresponds evidently to that represented by the phyla of Loz. (Pal.) prisca in Europe and of Loz. (Pal.) Tokunagai in Japan, showing the distinctive characteristics already stated. The question arises whether the group of pygmy Loxodonts of the Mediterranean islands and coasts, typified by Lox. (Pal.) melitensis (Falconer), is a close ally of the group just mentioned. The answer must be negative. Though the cheek-teeth of the group in question have a low ridge-formula, the disks of their ridges, as well as their general shape, appear to display an unmistakable similarity with those of the later phases of the phylum of Loz. (Pal.) antiqua. The low ridge-formula in this group may [be] due to degeneration as a result of having been dwarfed. Phylogenically, this group by itself is far from being a natural one. It can be considered natural only when it is taken as a group subordinate to the phylum of Loz. (Pal.) antiqua. By far the greatest part of Palzoloxodon is occupied by the group represented by the phyla of Loz. (Pal.) namadica in Southern to Eastern Asia and of Lox. (Pal.) antiqua in Europe. As indicated by the evolutionary tendencies observed in the mutations of the phylum of Loz. (Pal.) namadica in Japan, Lox. (Pal.) Tokunagai is probably ancestral to this phylum. Again, it appears probable, that the phylum of Loz. (Pal). antiqua, might also have had an almost similar form for its ancestor. It does not appear probable, however, that Lox. (Pal.) prisca was actually ancestral to that phylum, as a younger aspect appears to be present in the molars of this species in having the opposite loxodont sinus of two neighbouring ridges well-spaced.”’ OSBORN’S SUMMARY (1930) OF THE OBSERVATIONS OF MAKTYAMA (1924)! AND OF MATSUMOTO (1924-1929) From these descriptions and plates we conclude: (1) That Palzoloxodon is the only genus of true elephants that penetrated Japan; (2) that the primitive species P. tokunagai may have entered as early as Upper Pliocene time, and (3) that in ascending geologic levels there occur more progressive species, either indigenous or migrants from the P. namadicus stock of India. (4) In the present historic, geologic, and systematic revision, therefore, we shall present in each case Matsumoto’s opinions of 1924, and, in conclusion, give his opinions of 1929 under each species. Osborn, 1930: Osborn does not accept any of the phylogenetic or geographic theories expressed by Matsu- moto above; he regards Palzoloxodon as an entirely independent phylum originally derived from Africa and subsequently migrating through India to the Far East. He condenses from the invaluable observations of Maki- yama and of Matsumoto the following synopsis of their observations upon the ascending geologic successions and specific characteristics of the Japanese loxodonts: Palxoloxodon tokunagai Matsumoto, 1924, possibly of Upper Pliocene age, is regarded as the most archetypal group of Palzoloxodon in Japan, probably as ancestral to Palzoloxodon namadicus of southern and eastern Asia and to P. [Hesperoloxodon| antiquus of Europe (1929.2, pp. 10, 11). Estimated ridge formula of M 3 ;;x. Ridge frequency 5in 100mm. Length 295e mm., breadth 80 mm.; relatively low crowned. (No figure.) For the results of Makiyama’s recent studies, see his article “Japonic Proboscidea,”” Mem. Coll. Sci. Kyoto Imp. Univ., Ser. B, Vol. XIV, No. 1, Art. 1, May, 1938.—Editor.} THE LOXODONTINA: PALAXSOLOXODON OF JAPAN 1293 Palzxoloxodon tokunagai mut. junior Matsumoto, 1929, a mutation of Loxodonta (Palxoloxodon) tokunagat. Very primitive; either of Upper Pliocene or Lower Pleistocene age. Similar to Loxodonta (Palzoloxodon) prisca of Falconer. Differs from P. tokuwnagai which has molars of larger size, with slightly higher ridge formula and slightly more perfect lozenge-shaped dises of the ridges. Low crowned. Enamel thick. (Matsumoto, 1929.2, p. 10, also Pl. vu, fig. 1, type r.M2, our Fig. 1157.) Palzoloxodon protomammonteus Matsumoto, 1924, 1926, regarded as of Upper Pliocene (?), Basal Calabrian age (Fig. 1154). Lower molars small and narrow crowned. Estimated ridge-plate formula (M 3 ;.4;:) similar to that of the Lower Pleistocene Hesperoloxodon antiquus. Palzxoloxodon protomammonteus proximus Matsumoto, 1926. Upper Pliocene(?). Lower Calabrian age. Molars rather large and moderately wide. Type fragment, 1.M;, with 9+ ridge-plates. Ridge frequency 5 in 100mm. (Fig. 1155.) Palzxoloxodon namadicus naumanni Makiyama, 1924, characteristic of the Lower Pleistocene, very widely distributed. A narrow-toothed variety. Ridge frequency 5-6 in 100 mm. in M;, 6in 100 mm. in M*. Relatively broad crowned. Ridge formula of M 3 “47* [7%, Fig. 1152]. Length M* 303 mm., breadth 76 mm., max. height 216mm.; length M; 270 mm., breadth 72 mm. Palzxoloxodon namadicus yabei Matsumoto, 1929, characteristic of the Middle Pleistocene, very abundant. Estimated ridge formula of M 3 =;3; length 255 mm., width 74 mm.; ridge frequency 6.5-7 in 100 mm. (Fig. 1156.) Palxoloxodon namadicus namadi Makiyama, 1924, probably of Middle Pleistocene age. The largest and most numerous group of the Japanese loxodonts, representing a broad-toothed variety. Ridge frequency 5 in 100 mm. in M;. The type upper molar, r.M? (Fig. 1153), exhibits + 12+ ridge-plates indicating a larger total, as observed by Makiyama (1924, p. 263): ‘‘There is a loss of several posterior ridges, leaving fourteen in a space of 211 mm. The crown is very broad being 77 mm. in front, 90 mm. at the middle and 65 mm. posteriorly, measured on the second, fifth and fourteenth ridges respectively.” According to the above summary, the genus Palxoloxodon from Japan probably appeared in Upper Pliocene time as a narrow-toothed, coarse-enameled species, and ascended from Lower into Middle Pleistocene time, terminating in the species Palxoloxodon namadicus yabei, characteristic of the Middle Pleistocene and so closely resembling P. namadicus that it was first described by Matsumoto as Palxoloxodon namadicus typicus. This ascending order of phylogenetic succession, with broadening molar crowns and ridge-plates multiplying from M 3 was estimated (tokunagaz) to M 3 32 estimated (nawmannz) differs widely from that shown in figure 790, which represents Matsumoto’s earlier observations of the year 1924. The Japanese species of Palxoloxodon may all be derived from the same Upper Pliocene African stock, with narrow and low ridge formula, progressive in breadth and in ridge formula as follows: (Upper Pliocene) M 3 ;,; to (Lower Pleistocene) M 3 ;; to (Middle Pleistocene) M 3 73. TWO JAPANESE SUBSPECIES DESCRIBED BY MAKTYAMA (1924) Makiyama in a paper entitled ‘“Notes on a Fossil Elephant from Sahamma, Totomi” (1924.2, pp. 261, 262, 264) expresses the opinion that there are at least two varieties of Japanese fossil elephants, namely, the narrow toothed (e.g., Palzolorodon naumanni) and the broad toothed (e.g., P. namadz), which hitherto have been called Tusk Hie Nat size 44 Natural size ELEPHAS NAMADICUS NAUMANNI 7ype Kyoto /mp. Mus. M3 D L.Mjz sp ses Jee 10 11 12 13 8 53g Me a (e Genotypic Species (A-C) or PaLaotoxopon Matsumoto; (D) HESPEROLOXODON ANTIQUUS GERMANICUS OF TAUBACH All figures one-fourth natural size, same scale as figure 1073 excepting tusk (A) one-twelfth natural size Fig. 1152. Diagrammatic outline sketch of the type of Elephas namadicus nawmanni Makiyama, 1924 [=Palzxoloxodon namadicus naumanni], see Pls. xu, xt, xtv, Xv, and xv1, fig. 1, in comparison with Elephas antiquus |[Hesperolorodon antiquus germanicus], after Pohlig, 1888, Pl. 111, fig. 7, Pl. 1v, fig. 3. All figures one-fourth natural size, with the exception of the incisive tusk which is one-twelfth natural size. A, Incisive tusk, length 1930 mm. (PI. x1). Bl, Third right superior molar, exhibiting 14 wern and 5 unworn ridge-plates, total 19. Side view. (Pl. xm, fig. 1. Length 303 mm., breadth 76 mm., max. height 216 mm.) B2, Crown view cf same superior molar (PI. xu, fig. 2). Cl, Type inferior mandible, with M 3 in situ exhibiting 14 worn and 3 unworn ridge-plates, total 17. Superior view. Length of right ramus 405 mm., of left ramus 413 mm.; length of 1.M;,270mm.; max. breadth of same 72 mm. at fifth ridge (Pl. xv). C2, Same mandible, left lateral view (Pl. xv1, fig. 1). D, Elephas | =Hesperoloxedon| antiquus {germanicus] from Taubach, after Pohlig, 1888, Pl. 11, fig. 7, left Mi, crown view, and PI. tv, fig. 3, left Me, crown view. Compare figure 1075, type second left inferior molar, l.Me, exhibiting 12-13 ridge-plates, of ‘Hlephas antiquus’ Fale. Observe that the type of ‘Blephas namadicus naumanni’ Makiyama is decidedly inferior in size to the Palezolorodon namadicus of the Nerbudda, figure 1073, but that the grinders exceed in relative breadth those of Hesperoloxodon antiquus germanicus of Taubach (D, 1.M;, 1.My), as figured by Pohlig. 1294 THE LOXODONTINA: PALASOLOXODON OF JAPAN 1295 Elephas namadicus by many authors. Osborn, on the contrary, interprets the ‘narrow toothed’ as more primitive and geologically ancient, the ‘broad toothed’ as more progressive and geologically younger; progressive broadening of the grinding teeth and multiplying of the ridge-plates distinguish each line of ascent. NARROW TOOTHED.— (Op. cit., pp. 263, 264: “The second variety, to which the elephant of Sahamma belongs li.e., Palzoloxodon namadicus naumanni), is diagnosed by its narrower [76-79 mm.] bandy crown and lozenge- shaped dises. .. . I give here, a new subspecifie name Elephas namadicus Naumanni to the second variety, for which type the lower teeth from Sahamma should be chosen. . . . The second variety is apparently referable to E. antiquus in many respects. Therefore, it is not unreasonable that Brauns [Footnote: ‘Ueber Japanische Dilu- viale Siugetierer, Zeitsch. Deut. Geol. Ges., 1883, pp. 35-42.’] had supposed the possibility of direct migration of E. antiquus from Europe to Japan through Central Asia in the early Pleistocene age. He also pointed out some differences between his H#. antiquus in Japan and the types of HL’. namadicus in India.”’ BROAD TOOTHED.— (Op. cit., pp. 263, 264): ‘The first variety [Palxoloxodon namadicus namadi] is diagnosed by the broad [90 mm.] elliptical crown and less-crowded bandy discs. .. . The first variety referred to as typical E. namadicus should be called more strictly Hlephas namadicus namadi.’’ While Matsumoto (1929.1, p. 1) gives the original reference of these two subspecies as ‘‘Chikyti—The Globe, Vol. I, 1924, p. 381, Pl. vi,” this publication is not available to the present author and he is therefore citing from Makiyama’s supplementary descriptions in English (Makiyama, 1924.2) mentioned above. Palwoloxodon namadicus naumanni Makiyama, 1924! Figures 1152, 1189 From Sahamma, Tét6mi Province, Japan. Recorded from the Lower Pleistocene by both Makiyama and Matsumoto. Probably Middle Pleisto- cene. Genotypic species of Palzoloxodon Matsumoto, September 20, 1924. This subspecific name was applied by Professor Makiyama of the Kyéto Imperial University to an excellent medium-toothed type (Fig. 1152) rivaling in the dimensions of the teeth the ‘Elephas antiquus’ of Weimar (Fig. 1088B, M®*), in which the referred formula is M 3 45", the enamel thick and strongly crimped, the dentinal dises expanding mesially and coming into contact with extreme wear; the superior tusks strongly curved, the frontocranial structure unknown—on the whole, resembling the large Upper Pleistocene ‘#. antiquus [germanicus|’ of Weimar—distinguished both from the broad-toothed variety (maximum breadth 90 mm.) and from the extremely narrow-crowned variety. Elephas namadicus Naumanni Makiyama, 1924. ‘Notes on a Fossil Elephant from Sahamma, Tétémi.’’ Mem. College Sci. Kyéto Imp. Univ., Ser. B, Vol. I, No. 2, June 30, 1924, pp. 255— 264. Typr.—Cranium broken to pieces, leaving complete mandible with third inferior molars, also third superior molars, and inferior incisor. (Op. cit., p. 264): “I give here, a new subspecific name Hlephas namadicus Nawmanni to the second variety, for which type the lower teeth from Sahamma should be chosen.” Horizon anp Locauiry.—Excavated at Sahamma, about 12 kilometers northeast of Hamamatsu, T6t6mi Province, Japan, in 'Original description in Japanese: ‘“Chikyi—The Globe,”’ Vol. I, 1924, p. 381, Pl. vir (fide Matsumoto, 1929.1, p. 1). 1921. Typr Figurn.—Makiyama, op. cit., Pls. xi (incisor), xin (third right superior molar), x1v (third left superior molar), xv (mandible), xv1, fig. 1 (left lateral aspect of same mandible). SPECIFIC CHARACTERS. (cF. MAkIYAMA, PP. 260-264).— Superior incisor 1930 mm. in length, strongly upeurved (Pl. xm). Third left superior molar (Pl. xiv), length 286 mm., breadth 79 mm. at fourth ridge-plate, maximum ridge-plate height 217 mm., 16-4 ridge-plates in 286 mm.; third right superior molar (PI. xii), length 303 mm., breadth 76 mm., maximum ridge-plate height 216 mm., total ridge-plates 19, laminar frequency 17 in 303 mm., worn anterior plates exhibiting six lens-shaped dises with pro- nounced plications. Mandible (Pls. xv, xv1, fig. 1) with both third inferior molars in situ; symphyseal rostrum abrupt, subvertical; 1.M3, length 270 mm., breadth 72 mm., laminar frequency 17 ridge- plates in a space of 270 mm.; 14 anterior ridge-plates well worn “Showing a lozenge-shaped complete figure of enamel with minute but well-defined central angulations in touch with each other as obtained in #. africanus, and regularly crimped, comparatively thick enamel layers.” Comparisons.—In dimensions the upper right last molar, r.M’, is very like that of Hlephas antiquus |germanicus| from Weimar cited by Soergel (1913, Taf. vim). It also resembles the thick-plated variety of F. antiquus (ef. Leith Adams, 1877, Pt. I, p. 31). An M? deseribed but not figured, length 204 mm., breadth 72 mm., height 140 mm., differs in dimensions from those of LZ. antiquus, corresponding more nearly to those of H. meridionalis See also Chapter XXI of the present Memoir, p. 1408, under ‘1924 Elephas namadicus naumanni,” and p. 1413, under “1929 Elephas (Palzoloxodon) namadicus setoensis,’’ for Doctor Makiyama’s recent (1938) conclusions.—Editor. | 1296 (Soergel, 1913, Taf, v1); the dises are feebly rhombic and never touch at the central part. The mandible (Pl. xv1, fig. 1) in many respects resembles that of H. antiquus. Fryat Dracnosis (Marsumoto, 1929.1, ep. 2).—‘‘The race now under consideration can be diagnosed as follows: Cheek-teeth moderate in size, narrow-crowned. Well worn surface of the last upper molar shaped like an elongated ellipsoid, and that of the last lower molar band-like, with the outer border almost linear or concave. Frequency of ridges low, number- ing about 6 or less in a length of 100 mm. in the last upper molar and 5-6 or less in the lower. Mammille of the sum- mits of ridges stout and rather few, remaining proximally widely spaced to some extent. Disks of much or moder- ately worn ridges lozenge-shaped, without any marked differentiation of mesial portion and lateral arms. Even the lateral portions of disks are rather thick antero-posteriorly, the opposite loxodont sinus of the two neighbouring disks in the much or moderately worn portion of crown being in con- tact with or very closely set to each other. Layer of enamel rather thick; its plication being rather coarse, irregular and rather strong. .. . 4 Age: The present race appears, in all like- lihood, to be characteristic of the Lower Pleistocene, ranging in occurrence very probably from its base, as at Okine, up to 2. its very close, as at Tabata.” Palwoloxodon namadicus namadi Makiyama, 1924! Figures 1153, 1189 Dredged off the island of Sh6do, Sanuki Province, Japan. Probably Middle Pleistocene (fide Matsumoto, 1929.1, p. 4). Elephas namadicus namadi Makiyama, 1924. ‘Notes on a Fossil Elephant from Sahamma, T6t6mi.” Mem. College Sei., Kyéto Imp. Univ., Ser. B, Vol. I, No. 2, June 30, 1924, p. 264. Typr.—Last upper molar, right side, r.M’. Horizon AND Locautiry.—Dredged off the island of Shédo, Sanuki Province, Japan. Probably of Middle Pleistocene age (fide Matsumoto, 1929.1, p. 4). Typr Fiagurn.—Op. cit., Pl. xv1, fig. 2. Type Descriprion.—(Op. cit., 1924.2, pp. 263, 264): “An excellent example of the first variety is furnished by the last upper true molar... . There is a loss of several posterior ridges, leaving fourteen in a space of 211 mm. This crown is very broad being 77 ‘(Original description in Japanese: ‘“Chikyi—The Globe,” Vol. I, 1924, p. 381, Pl. vit (fide Matsumoto, 1929.1, p. 1). OSBORN: THE PROBOSCIDEA mm. in front, 90 mm. at the middle and 65 mm. posteriorly, meas- ured on the second, fifth and fourteenth ridges respectively. The grinding surface displays a characteristic oval outline and well- spaced bandy dises with minute median angulations. ‘The enamel layer is less well-crimped and thinner than that of the Sahamma TypE Mouar oF PALAOLOXODON NAMADICUS NAMADI Fig. 1153. Type r.M® of Elephas namadicus namadi Makiyama, 1924, exhib- iting 12 worn ridge-plates, one anterior ridge-plate missing, posterior ridge-plates not shown in photograph. One-half natural size. Breadth 90 mm. After Makiyama, 1924.2, Pl. xv1, fig. form... . The first variety referred to as typical #. namadicus, should be called more strictly Elephas namadicus namadi.” Frnaut Diacnosis (Matsumoto, 1929.1, p. 3).—‘“‘The present race is the largest group of the Japanese Loxodonts and represents the broad-toothed variety of the present species in Japan... . This race can be diagnosed as follows. Cheek-teeth large and rather broad. Well worn surface of the last lower molar shaped like an elongated ellipsoid. Frequency of ridges low, numbering about 5 or less in a length of 100 mm. in the last lower molar. Mammille of the summits of ridges stout and few. Disks of only the very strongly worn ridges lozenge-shaped as a whole; those of the moderately worn ridges consisting of a lozenge-shaped mesial portion and of nearly parallel-sided lateral arms. Interspaces between lateral arms of successive disks broad antero-posteriorly. Opposite loxodont sinus of the two neighbouring disks of mod- erately worn ridges widely separated from each other. Layer of enamel rather thick; its plication being coarse, irregular and feebly displayed.” See also Chapter XXI of the present Memoir, p. 1408, under ‘1924 Elephas namadicus naumanni,” and p. 1413 under “1929 Elephas (Palxoloxodon) namadicus setoensis,” for Doctor Maki- yama’s recent (1938) conclusions.—Editor.| THE LOXODONTINA: PALASOLOXODON OF JAPAN 1297 FIVE JAPANESE LOXODONTINES DESCRIBED BY MATSUMOTO (1924-1929) (Continued from Chap. XIV, pp. 906-908 of the present Memoir) During the period 1924-1929, Matsumoto described two species and three subspecies of the true elephants of Japan under the following names: Huelephas protomammonteus, 1924, Loxodonta (Palzxoloxodon) tokunagai, 1924, Parelephas protomammonteus proximus, 1926, Loxodonta (Palxoloxodon) namadicus yabei, 1929, and Loxo- donta (Palzoloxodon) tokunagai junior, mut., 1929. Matsumoto’s Memoir of 1924, ‘Preliminary Note on Fossil Elephants in Japan” (September 20), which contains the original descriptions of his subgenus Palxoloxodon, and of his species Huelephas protomammonteus and Loxodonta (Palxoloxodon) tokunagai, was published in the Japanese language; consequently the present author in the systematic description of the species and subspecies has cited freely from Matsumoto’s later Memoirs of 1926 and 1929 (which appeared in the English language) as given below. Osborn, 1929: Since the names Huelephas (Parelephas) protomammonteus (1924, 1926) and Parelephas protomammonteus proximus (1926) were assigned by Matsumoto under the impression that the genus Parelephas Osborn was represented in Japan, and since they were not included in the author’s subsequent memoirs of 1929, our conclusion is that both these specimens belong to Palzoloxodon. 1927 | = Palxoloxodon buski| below, p. 1333. Palwoloxodon protomammonteus Matsumoto, 1924, 1926! Figure 1154 Nagahama, Town of Minato, Kimitsu District, Province of Kazusa, Japan. Upper Pliocene (?), Lower Pleistocene. Euelephas protomammonteus Matsumoto, 1924. ‘Preliminary Note on Fossil Elephants in Japan.” Journ. Geol. Soc., Tokyo, 1924, Vol. XXXI, p. 262 (in Japanese language). Parelephas protomammonteus (Matsumoto) typicus Matsumoto, 1926. “On the Archetypal Mammoths from the Province of Kazusa.’”’ Sci. Rept. Tohoku Imp. Univ., (2), Geology, Vol. X, No. 2, pp. 43- 50. Typre.—A third inferior molar of the left side, 1.Ms, belonging to Mr. Natsume of the Town of Minato. Horizon AND Locauity.—Probably from the very base of the Narita series. Nagahama, Town of Minato, Kimitsu District, Province of Kazusa, Japan. Tyre Ficgure.—Matsumoto, 1926.2, Pl. XvIn, figs. 1, 2. Paratypr.—A third inferior molar of the right side, r.M;. From the same locality as the type; figured on Pl. xrx, figs. 1-3. In view of the fact that the original type description was published in the Japanese language, we may cite from Matsumoto’s Memoir of 1926, p. 50 (published in the English language) : “Stage I. Parel. protomammonteus (typicus). Basal Calabrian. Molars small and especially narrow-crowned. Inner and outer sides of ridges not very convex. Bases of ridges prominent, and valleys acutely pointed proximally. Basal cingula, and some- times also accessory columns, well developed.” Osborn, 1928: Matsumoto’s type figure and description appear to establish beyond question a strong resemblance to Palxoloxodon [Hesperoloxodon] antiquus typical form of the Lower Pleistocene of England; the ridge-plate formula is practically the same, i.e., M 3 yeu17- Matsumoto observes (op. cit., p. 44): “This molar is decidedly of a narrow-crowned type, much resembling in general See also Hlephas indicus Buski Mats., TyPr oF PALAOLOXODON PROTOMAMMONTEUS Vig. 1154. Type figure of Huelephas (Parelephas) protomammonteus (typicus), a third inferior molar of the left side, 1.M3, crown and internal aspects, one-third natural size. From Nagahama, Town of Minato, Kimitsu District, Province of Kazusa, Japan. After Matsumoto, 1926.2, Pl. xvmu, figs. 1 and 2, one-half natural size. Observe +16) ridge-plates. (Matsumoto, op. cit., p. 44): “The holotype, Ms, lacking its most anterior part, consists of sixteen ridges and a posterior talon. Its original ridge formula might probably be about 17. Its length, as preserved, is 275 mm.; and its original length might probably be some 300 mm. The maximal width is 75 mm. at the seventh ridge, as estimated above, and the height of the crown is 1384 mm. at the eleventh ridge, as estimated above, which had just com- menced to wear. The frequency of ridges in a length of 100 mm. is 5'-6 at the sides and about 7 at the grinding surface.” See Chapter XXI of the present Memoir, p. 1408, note under ‘1924 Hlephas namadicus naumanni Makiyama.”’—Editor.] 1298 shape the corresponding ones of the phyla of Loxodonta (Palxo- lorodon) namadica and antiqua, in contrast to Parel. trogontheric and higher mammoths. Consequently, the sides of the ridges in fore and aft views are only weakly convex, instead of being so strongly so as in the two last-mentioned species, and are not so strongly convergent toward the base as in the same. The ridges, except the very posterior ones, are curved forward in lateral views, quite as those of the lower molars of Parel. trogontherit.”’ Osborn agrees in the close resemblance of Palxoloxodon proto- mammonteus to Hesperoloxodon antiquus, but does not agree that it is related to the Parelephas trogontherii phylum. Palewoloxodon tokunagai Matsumoto, 1929 Soyama, Gokayama, Hira-mura, Higashi-Tonami District, Province of Etch, Japan. Recorded as of Upper Pliocene or Lower Pleistocene age. Loxodonta (Palzoloxodon) tokunagai Matsumoto, 1924. ‘Pre- liminary Note on Fossil Elephants in Japan.” Journ. Geol. Soc., Tokyo, Vol. XXXI, September 20, 1924, p. 267 (Japanese lan- guage). Supplementary Description (1929.2): “On Loxodonta (Palzoloxodon) tokunagai Matsumoto, with Remarks on the De- scent of Loxodontine Elephants.”’ Sci. Rept. Téhoku Imp. Univ., (2), Geology, Vol. XIII, No. 1, pp. 7-10 (English language). Typr.—A third inferior molar of the right side, r.M;. Originally belonging to the Imperial Museum of Uéno, numbered 2208—now in the Museum of Education at Ochanomizu, Tokyo. Hort- ZON AND LocaLity.—Soyama, Gokayama, Hira-mura, Higashi- Tonami District, Province of Etcht, Japan. Of Upper Pliocene or Lower Pleistocene age. Tyrer Ficurre.—The present author has been unable to locate a type figure. Tyrer Description.—(Cited from Matsumoto’s Memoir of 1929.2, p. 8): ‘Description of the type-specimen: This last molar consists, as preserved, of eleven ridges and the single-columned posterior talon. Judging from the general shape of the molar, two more ridges in all likelihood, might have originally been present anterior to the most anterior ridge as preserved. Thus, the ridge- formula appears to be nearly X13. Its length as preserved measures 258 mm.; its original length can be estimated as about 295 mm. Its full maximal width is 80 mm. at the ninth ridge as preserved, which is quite abnormal in having a superfluous column standing out on its inner side. Not including this superfluous column, however, the true maximal width of the molar measures 74 mm. at the sixth to eighth ridges as preserved.”’ “The frequency of ridges in a length of 100 mm. is about 5 on both the inner and outer sides, as well as at the grinding surface. l’rom an inner or outer view, the ridges are seen gradually narrowed distally, though the proximal ends of the valleys, with the exception of the most posterior ones, are of a rather obtuse type. The curvature of the ridges varies from very to rather slight for a lower molar.” “The smallness of the transverse width of the more anterior ridges is quite remarkable; and the disks of the well worn ridges are very markedly lozenge-shaped,—to such a degree that they remind us of those in Lorodonta, s.s. The tips of the anterior loxodont sinus of one ridge and the posterior loxodont sinus of the one immediately preceding in the well worn portion of the molar are in close contact with each other. ... The layer of enamel is very OSBORN: THE PROBOSCIDEA thick, measuring 2.5-3 mm. across; and its plication is very coarse and indistinct. There is a distinct, linear, transverse streak, or even fissure, in each area of cement between two disks at the grinding surface.” Palwoloxodon protomammonteus proximus Matsumoto, 1926 Figure 1155 Lower part of the Narita Series. Isone, Kokubo, Onuki-mura, Kimitsu District, Province of Kazusa, Japan. Upper Pliocene (?). Referring to Matsumoto’s definition of Huelephas (Parelephas) protomammonteus (typicus), that species is regarded as basal Calabrian, whereas the following species belongs on a higher level, namely, lower Calabrian (Matsumoto, 1926.2, p. 50): “Stage II. Parel. protomammonteus Calabrian.” “Molars rather large and moderately wide. Inner and outer sides of ridges not very convex. Bases of ridges more or less promi- nent, and valleys acutely pointed proximally. Basal cingula appar- ently not well developed.” Parelephas protomammonteus proximus, mut. nov. Matsumoto, 1926. “On the Archetypal Mammoths from the Province of Kazusa.” Sci. Rept. Tohoku Imp. Univ., (2), Geology, Vol. X, No. 2, pp. 48-50. Typr.—Fragment of a third inferior molar of the left side, 1.M;, belonging to Mr. Kat6 of Kokubo. Horizon AND Locauity.—‘This specimen appears to have been derived, very probably, from a certain lower part of the Narita Series.’’ Isone, Kokubo, Onuki-mura, Kimitsu District, Province of Kazusa, Japan. Tyrer Ficure.—Matsumoto, op. cit., 1926, Pl. xxiv, figs. 1, 2. Typrk Descrretron.—In addition to Matsumoto’s type figure and type description quoted below in the legend, Matsumoto compares this stage with his Huelephas (Parelephas) protomam- monteus as follows: The type molar of Parelephas protomam- proximus. Lower TYPE OF PALAOLOXODON PROTOMAMMONTEUS PROXIMUS Fig. 1155. Type figure of Parelephas protomammonteus proximus, mut. Matsumoto, 1926.2, Pl. xxrv, fig. 1, one-half natural size, from Isone, Frag- nov. Kokubo, Onuki-mura, Kimitsu District, province of Kazusa, Japan. ment of a left inferior molar, 1.M3, crown view. (Matsumoto, 1926.2, p. 48): “This specimen, representing the anterior part of Mg, consists of nine ridges and measures 168 mm. in length as preserved. Its maximal width is 88 mm. at the seventh ridge as preserved, and its height is 128 mm. at the last ridge as preserved, which is slightly worn. The frequency of ridges in 100 mm. is 5.” THE LOXODONTIN#: PALAMOLOXODON OF JAPAN monteus proximus is distinguished from the type molar of Parele- phas protomammonteus by its greater width and by the slightly less prominent bases of its ridges; as a whole P. proximus is a mutation intermediate between P. protomammonteus and P. trogontherii. This phylogenetic opinion is expressed by Matsumoto on page 50 (op. cit., 1926.2) as follows: Stage III. Parel. trogontherii. Calabrian to Tyrrhenian. Stage Il. Parel. protomammonteus proximus. Lower Calabrian. Stage I. Parel. protomammonteus (typicus). Basal Calabrian. Ossporn, 1930.-In the above description Matsumoto ex- presses the opinion, not shared by Osborn, that these specimens are referable to the Parelephas trogontherii phylum. Osborn re- gards them rather as loxodonts referable to the Palzxoloxodon phylum and quite distinct from any of the phyla of the Mam- montine. Both in the ridge formule, and in the proportions and the structure of the ridge-plates these types appear to agree quite closely with Hesperolorodon antiquus and Palxoloxodon namadicus and to be very distinct from the Parelephas trogontherti phylum. Palwoloxodon namadicus yabei Matsumoto 1929! Figure 1156 Inland Sea, Japan. Recorded as of Middle Pleistocene age. Loxodonta (Palzxolorodon) namadica (Yabe7) Matsumoto, 1929. “On Loxodonta (Palxolorodon) namadica (Falconer and Cautley) in Japan.”” Sci. Rept. Tohoku Imp. Univ., (2), Geology, Vol. XIII, No. 1, pp. 4 and 5. Typr.—‘Right ramus of mandible, bearing last molar in situ; belonging to the Second High School.” Horizon AnD Locauiry.——-Inland Sea. ‘The present race appears to be characteristic of the Middle Pleisto- cene.”” Typr Figure.—Op. cit., 1929.1, Pl. mt, fig. 2, and Pei. Tyrer Descriprion.—(Op. cit., 1929.1, p. 4): “The molar of the type-specimen is nearly complete, save that the most anterior portion of the crown, corresponding to the most anterior root, is broken away. The lost portion mentioned very probably consisted of the anterior talon and first two ridges. Since this molar, as preserved, contains fifteen complete ridges and the posterior talon, it is almost probable, that its original ridge-formula corresponds to x17. Its total length, including the broken portion, above the margin of the jaw, is about 255 mm. Its maximal width is 74 mm. at the eighth ridge. Its frequency of ridges in 100 mm. counts Goa 7.7 REFERRED SPECIMEN.—‘‘Penultimate upper molar of right side; belonging to our Institute of Geology and Paleontology.” The locality of the referred specimen is Sorachi, Uryti District, Province of Ishikari, Hokkaid6. Matsumoto (op. cir., 1929, p.5): “This race can be diagnos- ed as follows. Cheek-teeth moderate in size, narrow-crowned. Well worn surface of the last lower molar shaped like an elongated 1299 ellipsoid. Frequency of ridges rather high, that of the last lower molar of the type-specimen counting 6.5-7 in a length of 100 mm. Mammille of the summits of ridges slender, tending to be numer- ous and closely set; the clefts between the mammille extending proximally to a considerable height. Disks of the much worn ridges lozenge-shaped; those of the moderately worn ridges con- sisting of an expanded mesial portion and of nearly parallel-sided lateral arms, which are characteristically thin antero-posteriorly. Opposite loxodont sinus of the two neigh- bouring disks in the moderately worn portion of the crown moderately or very widely sep- arated from each other. Layer of enamel thin; its plication being fine, irregular and strong. . . . The present race is by far the most common Proboscidea in Japan.” Type Or PALAOLOXODON NAMADICUS YABHI Tig. 1156. Type right ramus of mandible, containing r.Mg am situ, of Loxodonta (Palzoloxodon) namadica (Yabei) Matsumoto, 1929. Ridge-plates of r.M3 34-7728, length 255 mm., breadth 74 mm., ridge frequency 6.5-7 in 100 mm. After Matsumoto, 1929.1, Pl. 1m, fig. 2, and Pl. tv. Jaw one- fourth natural size; molar two-fifths natural size. Palewoloxodon (Archidiskodon?) tokunagai mut. junior Matsumoto, 1929 Figure 1157 Japan, precise locality unknown. Pliocene or Lower Pleistocene. Probably upper part of the Upper The thick enamel, the widely spread ridge-plates, and the alleged ‘Upper Pliocene’ or ‘Lower Pleistocene’(?) age suggest a possible reference to Archidiskodon (ef. pp. 957-959, A. plani- See Chapter X XI of the present Memoir, p. 1408, note under ‘1924 Elephas namadicus nawmanni Makiyama.”—KHditor.] 1300 frons of the Siwaliks). The side view of the type (Matsumoto, 1929.2, Pl. vu, fig. 2) displays a relatively low-crowned, widely ridge-plated molar. Lor. (Pal.) Tokunagai junior, mut. nov. Matsumoto, 1929. “Qn Loxodonta (Palzxoloxrodon) tokunagai Matsumoto, with Re- marks on the Descent of Loxodontine Elephants.”’ Sci. Rept. T0- hoku Imp. Univ., (2), Geology, Vol. XIII, No. 1, p. 10. TYPE.— Second inferior molar of the right side, r.Me, belonging to the Geological Institute of the Kyéto Imperial University. | Hor1zon AND Locauity.—Japan, precise locality unknown. ‘‘. . . judging from the less but still archetypal feature, and from a comparison of Primitive Type Mouar oF PALaoLOXxoDON (ARCHIDISKODON?) TOKUNAGAI MUT. JUNIOR Fig. 1157. cene or Lower Pleistocene(?). After Matsumoto, 1929.2, Pl. vir, fig. 1. fifth ridge as preserved. Second inferior molar of the right side, r.Mo, of Lorodonta (Palxolorodon) Tokunagai junior mut. Matsumoto, 1929. Original in the Geological Institute of the Kyéto Imperial University. Upper Plio- Two-thirds natural size. exhibits +1114 ridge-plates. Length as preserved 223 mm., estimated length about 240 mm., breadth 73 mm. at OSBORN: THE PROBOSCIDEA the specimen to the Lower Pleistocene mutation—Nauwmanni Makiyama—of Loz. (Pal.) namadica in Japan, as well as from the degree of fossilisation, the. . . specimen can probably be referred to either an upper part of the Upper Pliocene or to the Lower Pleisto- cene.” Tyre Ficurr.—Op. cit., 1929.2, Pl. vit, figs. 1, 2. Tyrr Description.—(Op. cit., 1929.2, p. 10): In a comparison of the type of the present mutation, namely, Loxrodonta (Palzoloxo- don) tokunagai junior, with the type of Loxodonta (Palzoloxodon) tokunagai, Matsumoto notes the following differences which can be observed between them: (1) The more anterior portion of the crown of the mutation junior is not so narrow as that of tokunagai, on the whole, it appears to be less nar- row crowned; (2) the disks of ridges are less lozenge shaped; (3) the oppo- site loxodont sinus of the two neigh- bouring disks of the well worn ridges are closely set in the junior mutation, whereas they are in contact with each other in tokunagaz; (4) the plication of enamel is stronger, and (5) the layer of enamel appears to be thinner in the junior mutation than in tokunagaz, though this difference in the absolute thickness should be underestimated in the present case. “The writer pro- poses, in passing, to refer the muta- tion represented by the . . . specimen to Lox. (Pal.) Tokunagat junior, mut. noy.” This molar JAPANESE SPECIES DESCRIBED BY SAHEKI AND TOKUNAGA (1931, 1934) Parelephas protomammonteus matsumotoi Saheki, 1931! Figure 1158 From Mishima, Kimitsu District, Chiba Prefecture, Province of Kazusa, Japan. [The deseription and figure of this subspecies were not studied by Professor Osborn. referred Inasmuch, however, as he Parelephas protomammonteus and P. protomam- monteus proximus of Matsumoto to the genus Palxoloxodon, the present subspecies has been inserted in this section under the generic designation of the original author, namely, Par- elephas.—Editor. | Parelephas protomammonteus (Matsumoto) matsumotoi n. var. Saheki, 1931. ‘On Parelephas protomammonteus (Matsumoto) Recently Found in the Province of Kazusa.”’ Japanese Journ. Geol. and Geog., Vol. VIII, No. 3, pp. 125-129, Pl. xv, 1 text fig. Typr.—'. fragment of a left mandibular ramus, including symphysis, and bearing Ms, in situ.” Horizon AND Locauity.—From Mishima, Kimitsu. District, Chiba Prefecture, Province of Kazusa, Japan. (Op. cit., pp. 128, 129): “Lower Calabrian in age. The conglomerate of the region undoubtedly belongs to the Fig. 1158. protomammonteus matsumotoi Saheki, 1931, Pl. xv, fig. 1, one-fourth natural size. From Mishima, Province of Kazusa, Japan. Tyre oF PARELEPHAS PROTOMAMMONTEUS MATSUMOTOL Portion of left mandibular ramus, with Mg in situ, of Parelephas base of the Sanuki bed of Dr. H. Matsumoto, [Footnote: ‘The name “Sanuki bed” here used is different from what Mr. I’. Ueda calls by the same name, but may correspond to his “Umegase bed.” ’] which is considered to be the basal horizon of the Narita series; to the Upper Miura series of Dr. H. Yabe and See Chapter XXI of the present Memoir, p. 1416, note under “1931 Parelephas protomammonteus (Matsumoto) matsumotot Saheki.”—Editor.] THE LOXODONTINA: PALAXOLOXODON OF JAPAN Mr. 8. Nomura; to the Sasage bed of Mr. J. Makiyama; to the Higashi-Higasa bed of Mr. Y. Otsuka, and to the Umegase bed of Mr. F. Ueda. As to the age of the bed, the first proposes that it is Calabrian and correlates it to the Naganuma bed of the Miura Peninsula, while the last says it is Lower Pliocene and compares it to the Koshiba bed of Lower Pliocene in Miura. But accord- ing to Dr. O. Abel, the Elephas group first appeared in Middle Pliocene, while nearly all the other paleontologists state that it first appeared in later Pliocene. According to Dr. H. Matsumoto, ‘any well-established true Elephant is as yet unknown throughout the world before the earlier boundary of Calabrian.’ Moreover, Stegodon orientalis Owen (typicus) which is found associated with Parelephas protomammonteus (Matsumoto) typicus Matsumoto in this case, ... has never been found to be older than Upper Pliocene. [Footnote: ‘Mr. J. Makiyama stated that Stegodon orientalis in our country, is Pleistocene, (Proceedings of the Third Pan-Pacific Science Congress).’]” “In view of the foregoing, I am convinced that the bed is Calabrian in age, and is to be correlated to the Naganuma bed of the Miura Peninsula, ... as Dr. H. Matsumoto does. As to its stratigraphical position, however, I am inclined to favour Mr. F. Ueda’s views, separating it from the so-called Sanuki bed of Narita series as another unit of geologic formation [Footnote: ‘This concerns the problem of the boundary between Pliocene and Pleistocene. The conspicuous unconformity between Shimosuye- yoshi bed, i.e. the lower part of Narita series in the Miura Peninsula site and the Naganuma bed has been observed recently by Mr. Y. Otsuka, as well as by Mr. J. Makiyama.’]—the Umegase bed as it may be called.” Tyre Figurr.—QOp. cit., Pl. xv, figs. 1 and 3 [Fig. 1158 of the present Memoir]. Typr Derscription.—(Saheki, op. cit., 1931, pp. 125-129): “Since 1928, the conglomerate bed exposed at the river cliff of the Koito, has been worked for grit in the construction of prefectural roads newly opened in the village of Mishima, Kimitsu district, Chiba Prefecture. In the course of the work in December, 1928, the left half of a mandibular ramus, bearing a cheek tooth and some pieces of bone of a certain mammal, was found in the conglomerate bed at Higashi-Higasa of the village.” “The mandibular ramus of the specimen measures as follows (in mm.): “Mandibular angle seems to be conspicuously smaller than P. protomammonteus (Matsumoto) typicus Matsumoto.” “Tength from anterior end of symphysis to posterior end of M3 Length of symphysis 455 7 -a Width of ramus just anterior to base of ascending bar 160 Depth of symphysis 75” “The symphysis is very short-spouted. The ridges of diaste- mata are rather concavely arched, instead of being linear as in those of P. protomammonteus (Matsumoto) typicus Matsumoto, and sloping down obliquely from the anterior ends of the alveolar margins to that of the symphysis more gently than those of P. protomammonteus (Matsumoto) typicus Matsumoto. The anterior end of the alveolar margin lies at a distance anterior to the posterior end of the symphysis.” 1301 “Three anterior mental foramina are present, though Matsu- moto’s specimen is reported to have four—all along the ridge of the diastemata—the middle one being the largest.” “The Mz; of this mandibular ramus measures 287 mm. in length as preserved, lacking only some lower part of the posterior talon, consisting of eighteen ridges and a posterior talon. Its maximal width is 76 mm. at the seventh ridge, and its height is ca. 130 mm. at the twelfth ridge, which just shows signs of wearing. The frequency of ridges in 100 mm. is 6 on the inner side, 7 on the outer side, and less than 6 at the grinding surface.” “The worn surface of the molar is oval and more highly concaved than that of P. protomammonteus (Matsumoto) typicus Matsumoto, as also the mid-ridges which are concaved considera- bly to the anterior. The layer of enamel of the rather slightly worn ridges on the grinding surface is irregularly wavy; that of the more worn ridges is irregularly and coarsely plicated; while that of the still more worn ridges is gently waving and their enamel layer shows regular, fine, and weak plication.” “The layer of enamel is 1.8—2 mm. thick.” Palwoloxodon yokohamanus Tokunaga, 1934! Figure 1159 From Yokohama, Japan. Lower(?) Pleistocene. [The following text has been prepared from a translation kindly furnished by Mr. Ushinosuke Narahara of the American Museum. The original description by Doctor Tokunaga was not seen by Professor Osborn.—Editor. | Palzoloxodon yokohamanus Tokunaga, 1934, pp. 363-371, “Fossil Elephant Teeth found at Yokohama and Kakio, Kanagawa Fig. 1159. Type second right superior molar, r.M?, of Palxoloxodon yokohamanus Tokunaga, 1934, Pl. vu, fig. 1. Length of molar as preserved 178mm. From Yokohama, Japan. Prefecture,” Journ. Geog., Vol. XLVI, No. 546, July, 1934 (in Japanese). Typr.—A second right superior molar, r.M’. Horizon anv Locaurry.—Found at the mouth of the Tsurumi- gawa in 1931, Yokohama, Japan. Lower Pleistocene. TYPE Ficure.—Op. cit., Pl. vut, figs. 1 and 2. The author states that the molar is of a dark brown color and very shiny. It has ten preserved ridge-plates, two probably lacking, making a total of 12. Length of molar as preserved 178 mm., great- est breadth 76 mm., maximum height 177 mm., 5, 6 ridge-plates in100mm. Regarded by the author as of Lower Pleistocene age. See Chapter XXII of the present Memoir, p. 1408, note under ‘1924 Elephas namadicus naumanni Makiyama.”—Editor.] 1302 OSBORN: THE PROBOSCIDEA JAVANESE SPECIES DESCRIBED BY DUBOIS Palwzoloxodon hysudrindicus Dubois, 1908 The long and narrow right and left third inferior molars (B 122 83 and B 122 E 2C, Dubois Coll., Leiden) are beautifully preserved and, coming from the same individual, firmly establish ate the dimensions, the number of the ridge-plates, and the low crown Kendeng-Schichten, Pithecanthropus zone, Lower (or Middle?) Pleisto- of this progressive mid-Pleistocene species; rM3; measures 324 cene of Java. This zone, originally determined as Upper Pliocene, has recently mm., 1.M3;, 297 mm., as reproduced one-third natural size. The (Dietrich) been regarded as Middle Pleistocene, because it also contains the ridge-plate formula (M 3 ea) surpasses that of Palxoloxodon vary progressive mpecies /Stegedot arr awana: : Ae namadicus naumanni (M 3 +2), the genotypic species of Palzxo- The cotypes below (Fig. 1160), reproduced from original : photographs kindly sent to the present author by Doctor Dubois, are of great interest as positively demonstrating that ‘Hlephas hysudrindicus’ is a very progressive species of Palxoloxodon with Figure 1160 Contemporary of the Trinil Ape Man, Pithecanthropus, Middle(?) Pleisto- loxodon Matsumoto. a high ridge-plate formula: pee P. hysudrindicus M 3 yo9x:- I9F 9 10,11 12:13:14 15.16, 1% 18y!9 ee 5:6 7 8 9101213 14154617 18 * Coryrrs or PALZOLOXODON HYSUDRINDICUS. CONTEMPORARY WITH PITHECANTHROPUS Compare type of Hesperoloxodon antiquus ausonius (lig. 1087). Vig. 1160. Blephas hysudrindicus Dubois, 1908, after photographs of the original cotypes kindly furnished by Dr. Eugen Dubois for this Memeir. Un- iformly reduced to one-third natural size. L.M®, a third left. superior molar (B 85 Dubois Coll., Leiden) from the Kendeng deposits, Java, of an aged individual with ridge-plates 5-16 more or less worn, ridge-plates 1-4 completely worn away, ridge-plates 17-19-+ still unworn. Length 209 mm., reduced to 70 mm. or one-third natural size. External and crown views. R.Msg, a third right inferior molar (B 122 E3 Dubois Coll., Leiden) from the Kendeng deposits, Java, of a middle-aged individual with ridge-plates 1-8 more or less fully wern, ridge-plates 9-19!4 completely unworn. This grinder measures 324 mm. in length; it is reduced to 108 mm. or one-third natural size. Crown views. L..Mg, a third inferior grinder (B 122 E 2C Dubois Coll., Leiden) from the Kendeng deposits, Java, of a more aged individual with ridge-plates 3-11 fully or partly worn and ridge-plates 1-3 worn away or imperfect, ridge-plates 12-19 unworn. This grinder measures 297 mm. in length; it is reduced to 99 mm. or one-third natural size. Crown and lateral aspects. THE LOXODONTINE: The cotype superior molar (B85 Dubois Coll.) of Palzxolorodon hysudrindicus (Fig. 1160) exhibits, especially on ridge-plates 5 to 12, enamel foldings or plications more numerous and deeper than those of the type of P. namadicus namadi (Fig. 1153); these plications resemble those of the ‘Hlephas namadicus’ (Fig. 1189) de- scribed by Lydekker, 1886.2, as “from the Pleistocene between Kanagawa and Tokio (Yedo), Japan”; it was this crimping or plication and the absence of the ‘loxodont sinus’ which led Dubois to relate these grinders to Elephas indicus. Dubois (1908) named this species as among his ‘“‘Kendeng- fossilien”; Stremme (1911) remarked that Dubois had no Elephas remains from Trinil in his large collection. Dubois (1908) distin- guished Elephas hysudrindicus as standing near Hlephas hysudricus, but still more close to the living Hlephas indicus; consequently he named it Hlephas hysudrindicus. Stremme (1911, p. 144), however, remarked on its closer relationship to Hlephas [= Hesperoloxodon| antiquus: “In bezug auf die Zahl der Joche steht dieser dem ZL. antiquus niiher als dem rezenten FE. indicus.” Elephas hysudrindicus Dubois, 1908. ‘Das Geologische Alter der Kendeng-Oder Trinil-Fauna.” Tijdschr. Nederl. Aardr. Genoots. Amsterdam, Tweedie Serie, Deel XXVB, No. 6, p. 1257. Cotyrrs.—Molar teeth with lamellae not exceeding 194; skull subsequently found (see description below), but type specimen not clearly designated or figured. HorR1Izon AND Locauiry.—Kendeng-Schichten, Java, Middle? Pleistocene. Coryer Ficures.—Not published by the author (see Fig. 1160 of the present Memoir). OrIGINAL DescripTion.—(Dubois, 1908, p. 1257): “Auch von Elephas liegt unter meinen Kendengfossilien nur eine ernzige Art vor, wortiber mich genaue Durchsicht der vielen gesammelten Molaren und besonders auch Bekanntwerden mit der Schadelform belehrt hat. Es ist diese eine dem Hlephas hysudricus sehr nahe stehende, jedoch noch mehr als letzterer sich dem jetzt lebenden Elephas indicus anniherende Art, die ich deshalb als Hlephas hysudrindicus n. sp. bezeichnen will. Hatte man die Art aus den Siwalik-Schichten bereits als die vermutliche Stammform des lebenden asiatischen Elephanten erkannt, durch die neue Art von Java wird diese verwandtschaftliche Beziehung noch klarer. Die grosse Ahnlichkeit der Molaren hatte mich anfanglich sogar dazu PALHOLOXODON OF JAVA 1303 verfiihrt die Anwesenheit des #. indicus unter der Kendeng- Fauna anzunehmen, obgleich die Lamellen-Zahl der fossilen javanischen Molaren nicht tiber 19 geht, also viel geringer ist als bei #. indicus; der (spiiter bekannt gewordene) Schidel nihert sich aber bedeutend mehr der Siwalik- als der lebenden Art, namentlich durch sein Profil, durch die gréssere laterale Ent- wickelung der parietofrontalen Hécker, und durch die geraden Alveolen fiir die Stosszihne, doch ist die Breite zwischen den Schlafengruben bereits grésser geworden; hierin nahert sich die javanische ausgestorbene betriichtlich der lebenden indischen Art, als deren unmittelbaren Stammvater wir die erstere wohl un- zweifelhaft anzusehen haben.” (Stremme, 1911, p. 144): “Ein wichtiges Leitfossil wire eventuell Hlephas, dessen Zahnbruchstiick Janensch dem EHlephas antiquus am niichsten stellt. Das Stiick stammt nicht von Trinil; auch Dubois hat keine Hlephas-Reste von Trinil in seiner grossen Sammlung. Dubois selbst ist nicht geneigt, an Altersunterschiede der Fundstellen bei diesem eigenartigen Fehlen zu denken, sondern er halt die Lebensweise der Elefanten in bezug auf die Verteilung ihrer Reste fiir massgebend. Dubois hat nach seinem gr jsseren Material des Elefanten diesen als Elephas hysudrindicus n. sp. bezeichnet und stellt ihn namentlich nach dem Vergleiche der Schidel nahe an Elephas hysudricus aus den Siwalik-Schichten vom Pendschab und den subhimalajischen Bergen und aus dem Alt- pleistociin des Narbadatales.” Stremme also ‘observes (op. cit., p. 143): “Der Elephanten- zahn zeigt nach Janensch mehr loxodonten Charakter als Hlephas indicus und erinnert mehr an Hlephas antiquus. (Der Vertreter des FE. antiquus in Indien, nach Leith Adams Elephas namadicus Fale. aus dem Altpleistociin des Narbadatales, wird aber ausdriicklich von Janensch als verschieden bezeichnet.)’”? Janensch (‘Die Proboscidier-Schiidel der Trinil-Expeditions-Sammlung,” 1911, p. 194) does not comment directly on the characters of the species E. hysudrindicus Dubois beyond the observation quoted from Stremme above. Osborn, 1930: This progressive stage of Palxolorodon, like that of Stegodon airdwana, is of great anthropological interest as establishing the Middle Pleistocene age of Prthecanthropus erectus, first pointed out by Dietrich. IX. SUMMARY OF GEOGRAPHIC DISTRIBUTION ALONG THE EASTERN COAST OF ASIA Sees — —— —— = y \so" Wo : Wy Branget Altogether, the monographic re- searches of Matsumoto and Makiyama supplement those of Falconer, of Owen, and of Osborn and prove that primi- tive species of the genus Palxoloxodon spread eastward across all of southern Asia during Upper Pliocene time until their march was arrested by the an- da cient Pliocene shore lines of Japan and of the East Indies, including Burma, Java, and Sumatra. It is in the Tolan & ni uric Kendeng stratum of Java, now regard- ed as of Middle Pleistocene age (as fully treated above, Chap. XIV, the Stego- dontine, especially p. 887) that we find another advanced stage of Palzxoloxo- don, to which Dubois applied the name ‘Elephas hysudrindicus.’ The bathymetric maps of Chisholm and Leete (Fig. 1161) and of Yabe (Fig. 1162) are extremely interesting and im- portant in displaying three features, namely: (1) The ancient coast of northern and southern Asia and of the East In- dies during late Pliocene and early to mid-Pleistocene times; (2) the sueces- sive northerly range over the Bering Strait region of Archidiskodon, (?)Par- elephas, and Mammonteus to North America; (3) the southerly and south- sasterly range of Stegolophodon, Stego- don, and Palzoloxodon, and finally of Elephas. Fig. 1161. Japan as part of the Asiatic continent in Plio-Pleistocene time, showing in white the broad true continental border on the 100-fathom or 600-foot line. Reproduced by permission after Longmans’ New School Atlas. This shows clearly the migration lines through Sumatra, Borneo, the Philippine Islands, Japan, and Kamchatka. Compare with the more bbe ae Wl eS | recent figure after Yabe (Fig. 1162). THE LOXODONTINA:: GEOGRAPHIC DISTRIBUTION ALONG ASIATIC COAST 1305 The three outstanding facts are: The geologically oldest or Lower Pliocene! range of the Stegodonts” covers an area from the type locality of Stegolophodon latidens on the Irrawaddy River, Burma, 1300 miles southeast, to the northern Bruni district of Borneo, namely, Brit. Mus. M.2498, described by Lydekker (1886.2, pp. 75, 76, Fig. 19) as ‘Mastodon latidens’ but which the present author made the type of Stegolophodon lydekkeri in Vol. I, p. 700, of the present Memoir. Of Upper Pliocene and Lower Pleistocene age is the transcontinental range of descendants of North African species of Palzoloxodon, such as Palzxoloxodon jolensis and P. atlanticus of Pomel, and P. recki of Dietrich, to Japan; this must have occurred in late Pliocene or early Pleistocene time, because the more primitive Japanese stage described by Matsumoto, namely, P. tokwnagai (M 3 ;;43-,), corresponds closely in ridge formula with the more primitive North African species and those of the Mediterranean Islands. Finally there is the Palzoloxodon namadicus stage (M 3 +2) of the Middle to Upper Pleistocene of India, 6 which corresponds closely with the P. namadicus naumanni of Makiyama, although the latter has a higher ridge formula, namely, M 3 +%. Of the true Stegodonts, the Stegodon elephantoides of the Irrawaddy River (M 3 ,;4,), including Stegodon cliftii of Falconer (1.M 1 with ,,, ridge-crests), may be ancestral to the Stegodon insignis birmanicus of Burma (M3 5,,) and to the S. orientalis grangeri (M 3 ,;;) of the Yangtze River, China. Fig. 1162. Japan as part of the Asiatic continent in Plio-Pleistocene time and two landlocked internal basins. After Yabe, 1929, map, p. 169, who describes the latest land connection of the Japanese Islands with the Asiatic continent as follows (p. 168): “The sea floor now encircling the Japanese Islands to the depth of some 720 m., in the average, was once a land surface and the land submerged below the sea level in a time geologically not much remote from the present. Prior to this great submergence of land, the Japanese Islands were some 720 m. [2375 ft.] more elevated than they are at present and directly connected with the Asiatic continent, as the annexed map shows. The 720 m. line, which marks the true continental border of eastern Asia more properly than the 160 m. line, follows closely the present Pacific coasts of Taiwan, the Riukiu Group, Kytsht, Shikoku, Honshi Hokkaidé, the Chishima Group and Kamchatka. The entire region lying inside this line is regarded once to have been a dry land, with the exception of two extensive basins—one occupying the greater part of the Japan Sea and the other [the] southernmost part of the Okhotsk Sea—, and a narrow, linear one lying along the inner border of the island are of the Riukiu Group. All these basins, now more than 700 m. deep, are thought by the writer as having existed there as entirely or almost land-locked basins during the continental stage of the Japanese Islands; .. . It was at the time of the maximal extension of land in the continental stage of the Japanese Islands that the land took the outline stated above. . . . In other words, the continental stage of the Japanese Islands continued from the time of land emergence to the 90 m. line to the time of land submergence to the same line.” \ VJ YY, LA ‘(Probably Lower Pleistocene—see note on page 824 above.—Editor. | *{See footnote on page 837 above.—Hditor.] 9081 “AGNVYY JO “LT ALVIg Ad NOTA) NI daHdVuUNOLOHd SLNVHdAIY AIVWAA GNV AIVJY “SQOIGNI SVHda 1h aqduudday “E9ll “Ply CHAPTER XX THE SUBFAMILY ELEPHANTINAE (SUPERFAMILY ELEPHANTOIDEA), OF EASTERN ASIA, INCLUDING THE RECENT ELEPHAS AND RELATED LOWERS PEBISTOCENE SPECIES CLASSICAL REFERENCES TO ELEPHAS. ESTABLISHMENT BY LINNUS OF THE GENUS AND SPECIES ELEPHAS tNpIcus (1735-1758). CHARACTERS OF THE SUBFAMILY ELEPHANTINA AND INCLUDED SPECIES AND SUBSPECIES OR GEOGRAPHIC VARIETIES, BY CORSE, DE BLAINVILLE, FALCONER, TEMMINCK, MATSUMOTO, AND OTHERS (1799-1927). FosstL LOWER PLEISTOCENE AND UPPER PLIOCENE SPECIES DESCRIBED BY FALCONER AND BY OSBORN (1845-1930). I. Hisrortcan Inrropuction Aanp NoMENCLATURE (850? Elephas indicus sumatranus Temminck. B.C.-1936). Elephas indicus hirsutus Lydekker. 1. Faleoner (1868) on the specific unity and vertebral Elephas indicus Buski Matsumoto [=?Palxoloxodon formule of the Asiatic elephants. busk7]. 2. Corse, de Blainville, and Faleoner on characters of the 2. Distinctions and measurements of the Indian elephant. eographic varieties. . , Be ine. Se or subspecific forms, continental and JTL, CaMenensms OP mai UEpan Puuoemia uy Ibowar Fai 330: CENE Species HypseLEPHAS HYSUDRICUS AND PLATELE- PHAS PLATYCEPHALUS. 1. Falconer’s original descriptions of Elephas [Hypselephas| hysudricus. 2. Observations of Osborn on the fourteen specimens col- lected by Barnum Brown (1922) referred to Hypsele- insular. 4. Fossil forms more or less closely related to Hlephas indicus. 5. Names of species and subspecies of the subfamily Elephantine in order of description. Il. CHARACTERS OF THE SUBFAMILY HLEPHANTIN 2 AND INCLUD- phas hysudricus. ED GENERA AND SPECIES. 3. Cranial characters and affinities of Hypselephas hysudri- Genus Elephas. CUS. 1. Systematic description of species of Elephas. Juvenile crania with dentition. EKlephas indicus Linneus. 4. Genus Platelephas. Elephas indicus ceylanicus de Blainville. Platelephas platycephalus. Elephas indicus bengalensis de Blainville. APPENDIX. It is a striking circumstance that we have not as yet discovered the direct ancestry of either of the typical living elephants, namely, the Hlephas indicus of India or the Loxodonta africana of Africa. Both of these typical living species, around which center historical as well as zoological references from the time of Homer (850? B.C.) to the present time, are still lacking the phylogenetic background of their history. All attempts both by zoologists and paleontologists to trace back either Hlephas indicus or Loxodonta africana to previously living or fossil forms! are shown in the present Memoir to be problematical, because neither in the cranium, the teeth, nor the skeleton do any of the known fossil elephants meet all the phylogenetic conditions. Even the Platelephas platycephalus of the Upper Pliocene and the Hypselephas hysudricus of the Lower Pleistocene of India, very fully described below in the present chapter, fail to meet all the conditions ancestral to the true H. indicus. The conclusion is inevitable, that in some as yet unexplored region of Asia the direct ancestors of H. indicus were slowly evolving, while in some unexplored area, probably west of central Africa, the very conservative ancestors of L. africana were also slowly evolving. In brief, we now know the phylogeny of most of the other subfamilies of the Elephantide better than we know the phylogeny of the Elephantine, typified by the genus Elephas. ‘According to Pomel (1895) the true Loxodonta africana occurs in comparatively recent deposits of North Africa, and Dart (1929) refers to the discovery in the Pilandsberg (Transvaal) of a primitive form of L. africana (cf. Loxodonta prima Dart, 1929, Chap. XTX above, p. 1287). —Editor.] 1307 1308 OSBORN: THE PROBOSCIDEA The early historical references or allusions to this most majestic of all living quadrupeds of the Asiatie con- tinent are full of fascination and interest.! The uses of these animals in industry, in transportation, in art, and in war would fill volumes. But here, as a fitting conclusion to the previous nineteen chapters of the history of the Proboscidea, we must confine ourselves rigidly to the slow emergence and clarification of zoological nomenclature and anatomical analysis from the time of the Greeks onward to the present time. As in the Loxodontine (Chap. XIX) and in the Mammontinz (Chap. XVIII), the strict application of the Rules of the International Commission on Zoological Nomenclature (Congress of Berlin, 1901, and of Monaco, 1913) to types and descriptions of the early part of the nineteenth century is beset with insuperable difficulties, as exemplified in the classic case of whether the Indian elephant should be called ‘“Klephas indicus’ or ‘Elephas maximus.’ The grounds for the adoption of ‘Elephas maximus’ are herewith shown to be either precarious or absolutely untenable. I. HISTORICAL INTRODUCTION AND NOMENCLATURE (850? B.C.-1936) The very ancient name ‘elephant’ has risen in rank, becoming the type of the subfamily Elephantine, family Elephantide, superfamily Elephantoidea. History or NoMENCLATURE.—Hlephas is a Greek name of uncertain origin, subsequently Latinized. It is treated by Liddell and Scott (1883) and by James A. H. Murray (1891) as follows: (Liddell and Scott, 1883, p. 454): €Médas, avtos, 6 the elephant, first mentioned by Hat. as a native of Africa, 3.114., 4.191; whereas Arist. H. A. 2.1, 45 treats only of Elephas Indicus, cf. 9.1, 30, etc., though the African is mentioned by him in Cael. 2.14,19:—not generally known in Greece till the time of Alexander, Paus. 1.12,4. II. known to Hom. only as the name for the elephant’s tusk, ivory, U1. 5. 582, and so Hes. and Pind.; for ivory was brought by Phoenician traffic to Greece long before the animal was known to Greek travellers; Hat. calls the tusks more accurately €\é@avros bdovtes, 3. 971:—Hom. brings false dreams through an ivory gate, v. sub €Mépatpouat, .. . (Pott and others refer to the Hebr. Eleph (ox), and compare bos Lucas, the old Lat. name of the elephant, Lucret. 5. 1301; as Paus. (9.21, 2) calls a rhinoceros raupos At@tomtxds. On the other hand the Hebr. name for the animal, ibdh, recalls the Skt. zbhas, which is identical with the latter part of €\-épas, and the first part of the Lat. eb-wr, whence 7v-oire, etc.) (Murray, 1891, III, Pt. 2, p.84): Elephant... Gr. edépas (gen. €d€pavros). The refashioning of the word after Lat. seems to have taken place earlier in Eng. than in Fr., the Fr. forms with el- being cited only from 15th ec. . . . Of the ultimate etymology nothing is really known. As the Gr. word is found (though only in sense ‘ivory’) in Homer and Hesiod, it seems unlikely that it can be, as some have supposed, of Indian origin. | The resemblance in sound to Heb. . . . eleph ‘ox’ has given rise to a suggestion of derivation from some Phcenician or Punic compound of that word; others have conjectured that the word may be African. Elephas was named as a genus by Ray (1693, p. 131 [=p. 123 of authors]), and was included in the “Systema Nature” of Linnzus in 1735 in the same division of quadrupeds with the rhinoceros (see below facsimile of portion of page 10 of the First Edition of the ‘Systema Nature’”’ of Linnzus, 1735). The genus Hlephas Linn. dates from 1735, the species indicus dates from 1754, the species maximus dates from 1758, as shown in the three facsimile Linnean definitions from (1) the ‘Systema Nature,’ First Edition, 1735, p. 10; (2) Memoir of the Museum Adolphi Friderici Regis, Holmiz !Stockholm], 1754, p. 11, and finally in the Tenth Edition of the “Systema Nature,” 1758, p. 33. Dr. C. Davies Sherborn of the British Museum, the greatest living authority on generic and specific names, believes that it is impossible to determine which elephant, African or Asiatic, Linnzeus had in mind in proposing the term maximus, a name without standing in Sherborn’s opinion: ‘Kunz, George F’., 1916. Ivory and the Elephant in Art, in Archeology and in Science. THE ELEPHANTINA: HISTORICAL INTRODUCTION AND NOMENCLATURE 1309 (Sherborn, 1929, letter of July 14) ‘‘In reply to your enquiry about Elephas maximus, Linnzeus. Linneus’ type seems to have been Ray, Quadr. 131. Blainville, Eléphants (p. 47) in his ‘Ostéographie,’ says Ray saw a young elephant in Florence. Now Ray gave no locality for his specimen but described it in much detail. In the Mus. Adolf. Frid. in 1754 Linnzeus says ‘Indien’ both in his Latin and his Swedish version and why or on what authority he puts the word Zeylansk in opposition to ‘indicus’ passes my understanding. So far as I know there is no authority for Zeylansk and in the S. N., ed. XIT, 48, Linnzeus clearly spatchcocks the whole lot into one species both for Asia and Africa and considers that there is only one Elephant. And as that beast has been more or less of a domesticated animal since before Alexander the Great, I don’t think Linnzeus was far wrong in such a guess. In such a case it appears to me that you have no option but to say ‘INDETERMINATE’ and proceed to the next person who divided the subject, e.g. Blumenbach in 1797 for ‘africanus’ and Cuvier in 1798 for ‘indicus.’ Unless one can definitely say what beast Linnzeus saw when he gave the name ‘maximus,’ which apparently is now a thing impossible to decide, it appears to me that ‘maximus’ has no standing and must be swept aside.’”’ [See Loénnberg (letter of 1929) quoted below, also footnote on page 1323]. In 1754 the specific name zndicus appears and in our present language this was the original genotypic species of Hlephas; it was universally used as such by all writers on the elephant family until 1901. Unfortunately, and for some reason unknown to us, in the Tenth Edition of the “Systema Naturz’’ Linnzeus substituted the name Elephas maximus Linn. for that of H. indicus Linn., which, with all other names in the Tenth Edition, was officially —_—— — — —~—-—_—_—_——_——— | dS | Equus. Mamme 2 inguinales, Equus. yy SC | q Pedes integri. | Afinus. | 8s ei | Onager. = = 2 | Zebra. : ts) & ts | Eipeoper mts Mamma 2 inguinales ( Arif}.) | Equus marinus. | Gs Z| | Pedes quadrifidi. aa eS Pa | FS | Elephas. Mammz 2. pettorales. Elephas. <5 > | P | Pedes 5. callis inttructi. ? Rhinoceros. | | = s Sus, Mammze to. abdominales. | Sus. Eon: | Pedes biungulati: raro fimplices. Aper. 2 | Porcus, | at | | Barbyrouffa. ey || | | Tajacu. | Fig. 1164. Facsimile of portion of page 10, in same size, of the First Edition of Linnzeus’ “Systema Nature,” 1735, in which appears his first definition of the genus EHlephas, bracketed with ?Rhinoceros. QUADR. JUMENTA. FYRFOT. BOSKAP. JUMENTA. BOSKAP. ELEPHAS. ELEPHANT. indicus. ELEPHAS. Raj. qvadr. £23. Sy/ft. ELEPHANT. = Raj. qvadr. 123. Zeylansk.. Nat, it. Syft, Nat. it. Habisat in INDIA. Bor uti INDIEN. Denres varii, magni. TAWDER af detta djur af attkillig ftorlek. Fig. 1165. Facsimile of portion of page 11, reduced in size, of Linneus’ Memoir of the Museum Adolphi Friderici Regis, Stockholm, 1754, in which first appears the species Hlephas indicus, habitat India. adopted by the Fifth International Congress of Zoology held at Berlin in 1901 (see Verhandlungen des V. Inter- nationalen Zoologen-Congresses zu Berlin, 12.16. August, 1901, published 1902, p. 967). In the present Memoir, to avoid confusion with the entire literature prior to 1901, the earlier specific name Elephas indicus Linn. is used throughout as a ‘Linnean collective species’ embracing several geographic subspecies. We have seen repeatedly in the present Memoir that the rules of the Congress of 1901 are impracticable when one attempts to apply them to early stages of paleontological nomenclature. We must remember that Linnzus 1310 OSBORN: THE PROBOSCIDEA believed that the number of created species was limited; in his mind the Indian and African elephants were of the same species. This is indicated in the following letter from Professor Einar Lénnberg (July 31, 1929): Linna&us’ Type or ELepHas rnpicus (LONNBERG, LETTER, JULY 31, 1929).—“‘In reply to your letter concerning the type of Linneus’ Hlephas indicus in Museum Adolphi Friderici Regis, 1754, I may say that it is not a skeleton nor a stuffed animal but a foetus, which still is kept in this Museum. The King had bought it together with several other natural history specimens from Holland, where it once was kept in ‘Museum Sebae.’ It isof an African Elephant, probably from West Africa. The reason why Linnzus speaks of India and Ceylon is probably that he quotes Rajus in the first rank. I have not looked up what Rajus says, but you can easily find that yourself after the quotation: ‘Raj. quadr. 123. In Systema Nature, ed. X., Rajus is also quoted in the first rank and Seba only in the third rank. But Seba’s specimen is the same as that of Museum Adolphi Friderici Regis, 1754. Seba has figured his specimen in his ‘Thesaurus,’ Vol. I, tab. CXI, fig. 1, and he says that it is ‘Een ongeboren Olifant uit Africa.’ From the above is evident, that the name Hlephas maximus L. 1758 refers as well to the Indian (Rajus quoted first) as the African (Seba quoted as third) Elephant.” Genus (Linnaus, 1735, 1754, 1758).—The genus Hlephas Linneus dates from 1735 (“Systema Nature,” First Edition, p. 10) without mention of a genotypic species. In 1754 (Memoir, Mus. Adolphi Friderici Reg., Stockholm, p. 11) reappears the generic name Klephas with the species indicus, habitat India (Zeylansk), indicat- ing that Linnzeus based the species on the Ceylon variety of elephant. In 1758 (‘Systema Nature,’ Tenth Edition, p. 33) again appears the generic name EHlephas, habitat India (Zeylonz), with the new specific name Elephas maximus. This edition (known as the Editio Decima, Reformata) was reprinted in 1901 and adopted as the standard by the Fifth International Congress of Zoology of 1901, held at Berlin. MAMMALIA BRUTA. Elephas. 33 In 1795 the name EHlephantus was used by Geoffroy I. BRRwCLA and Cuvier, and in 1801 was cited by Cuvier and Lacé- péde as Hlephantus indicus, as Cuvier continued to ig- Dentes Primores nulls utringue. nore the terminology of the “Systema Nature’ of 3 F Linnzeus. 5. ELEPHAS, Dentes Primores nulli. 4 Laniarié fuperiores elon- Grnorypic Specitus.—The species indicus of Lin- ati, i sy tage Probofcis \ongiffima, prehenfilis nzeus, which we select as genotypic, is important, be- ? ig 5 . Corpus nudivicuium, cause of the recent substitution by the Congress of 1901 maximus. 1, ELEPHAS. Raj. quadr.123. Syft nat. y1. Seb. muf.1. of Hlephas maximus Linnzeus (1758) for the original Seo hed name Hlephas indicus Linneeus (1754). Why did Lin- Elephantus. Gejn. guadr. 377. Aldr. quadr. |. 1. ¢. 9. : oe Fonft. quadr. 30 ¢. 78. f. 9. neus abandon the appropriate name EHlephas indicus Habitat in Zeylonz paludofis ad Amnes, edit Ramos, and substitute the inappropriate name Hlephas mazi- Cocos, Guilaudine femina, Frumentum. 9 : BA eye : : ain mus? Is not the explanation found in his belief that the Maximum quadrupes. Oculi parvt. Dentes Laniarii f~ ; 3 ‘ periores exferti (Ebur). Aures amplifiime, pendule, Indian and the African elephant were of the same species? dentate; att. angl. 277. p. 1051. Cutis crafifima, callofa. Mamme 2 juata pedtas. Ungues in apicibus First.—There is little doubt! that the genotypic loLorum pedum. Genua flexilia. Collum breve, , Sc ahs : : Probofcis longifinta, extenfilis, acute odorams, loco ma- species Hlephas indicus Linn., 1754 (=EHlephas maxi- nis ipfi inferviens; ea cibum potumque baurit, boftem- mys Linn., 1758) was a domesticated elephant from the gue pellit; ea preecifa occiditur ; murem metuit {ub i . i ‘ 5 sc forno ob trachew infertionem. Retro co & mingit. island of Ceylonin which occurs the native or indigenous Portat domos, reGore collo infidente; in bello arma- : -< invi PPPs tonlnnlchle ietty ain tS atlautere Fariee small, typical Ceylonese variety named by de Blainville Sus occidster , caterum prudens, docilis, (1845) Elephas indicus ceylanicus. This subspecies of 1 si alae tds TCs ita «a € . Fig. 1166. Facsimile of page 33, in same size, of Linnzus’ original Tenth ( eylon is cl arly distinguished by its small, slender Edition of the “Systema Natur,” 1758, in which Elephas maximus is substi- tusks pointing obliquely downwards, by its narrow, tuted for Elephas indicus, and the genus Elephas and species maximus are p 5 c : defined. Habitat in Zeylone. elevated head, with narrow narial openings, by its gentle "In 1758, as shown in the accompanying facsimile of page 33 of the Tenth Edition of the “Systema Nature,’ Linnxus follows Ray, as of first rank: “maximus. 1. Evepnas. Raj. quadr...... Habitat in Zeylonx.”’ THE ELEPHANTINA: HISTORICAL INTRODUCTION AND NOMENCLATURE 1311 characters which would now entitle it to distinct subspecific rank from the large, broad-headed, and otherwise different elephant of Bengal (see com- parative figures 1171 and 1170). The subspecies ceylanicus, moreover (Osborn, 1930), resembles the small, slender-tusked EHlephas indicus var. Mukna of Falconer and Cautley (1847). disposition facilitating domestication, and by its typical 20 dorsal vertebree Second.—The characters of the Ceylonese genotypic species Elephas indicus (=H. maximus Linn.) are, how- ever, confused by the fact that the larger Bengal elephant was imported into Ceylon, and it may have been one of these larger Bengalese types of India imported into Ceylon which Linnzeus had in mind in applying his second specific name Hlephas maximus. On this point Lydekker observed (1916, p. 82): The name EHlephas maximus is typified by a Ceylon elephant, and at first sight it would seem that Ceylon should be taken as the typical locality of the species. But there are two races of elephants in that island; one—probably indigenous—in which the tusks of the males are insignificant, and the other—almost certainly introdueed—in which they are large. Now it seems almost certain that the big-tusked race formed the type of Linne’s species; and if so, Ceylon will not be the typical locality. Unfortunately, it is uncertain whether the tusker-race was imported from the Indian mainland . . . or from Lower Burma. . . . Under these circumstances the writer [Lydekker] has considered it advisable to take southern India (say Mysore) as the type locality, and to regard the small tusked Ceylon form as a second race. Third.—From the above historic résumé it appears technically that Linnzeus named as the genotypic species the Ceylonese variety Elephas indicus ceylanicus; that Linneus’ type specimen, however, was according to Loénnberg (see letter, July 31, 1929, on p. 1310) the foetus of an African elephant (Loxodonta africana); that in the substitution of Elephas maximus for Elephas indicus he had both the African and the Indian elephant in mind, apparently in the belief that they constituted a single created species. For these reasons it is preferable to retain the name Elephas indicus. InDIAN ELEPHANT GROUP Fig. 1167. The specimens in this group are from the hills in the Province of Mysore, India, about 35 miles south of the city of Mysore. They were shot in the spring of 1923, by Mr. Arthur S. Vernay, who presented them to the American Museum of Natural History. While they are not record specimens (see Fig. 1194, where one of the tallest elephants is given an estimated height of 10 ft. 6 in.), they are very fine examples and form an imposing central group in the Vernay-Faunthorpe Hall. The male (Amer. Mus. 54453) is at the left; the female (Amer. Mus. 54452) at the right. Estimated Measurements (See caption to Fig. 1194) Male Female Skeletal height at shoulder 2558 mm. or 8 ft. 4% in. Height in the flesh 2720 Sie 2490 mm. or 8 ft. 2 in. Seapula 676 2 254 Humerus 879 2 10% Ulna 673 2 2 1312 OSBORN: THE PROBOSCIDEA 1. FALCONER (1868) ON THE SPECIFIC UNITY AND VERTEBRAL FORMULA OF THE ASIATIC ELEPHANTS (Continued from pp. 930, 931 of the present Memoir) As to specific unity, Falconer (‘‘Palzeontological Memoirs,” Vol. IT, pp. 267-270), after prolonged residence in Ceylon and India and very careful study, erroneously concluded that there were no constant specific dif- ferences either in the vertebral formule or in the structure of the cranium between the Bengal, or continental Indian elephant, and the insular elephant of Ceylon, as appears in the following summary of his observations and his conclusion that there is but a single species of Asiatic elephant: Sumatran Elephant: 20 dorsals and 20 pairs of ribs (Schlegel)—constant. Ceylon Elephant: 20 dorsals and 20 pairs of ribs (Peter Camper, Cuvier, de Blainville). Brought to Paris in 1795 4 lumbars; total dorso-lumbars 24. Indian Elephant [Bengal?]: 19 dorsals and 19 pairs of ribs (Schlegel); by no means certain that the Schlegel states that ‘‘all the Indian [Bengalese] number is constantly limited to 19. elephants which he had examined had, [See 20 dorsals cited below, ‘Duvaucel’ and ‘Choonee.’] without exception, only 19 dorsal verte- bre and 19 pairs of ribs.” Bengal Elephant: 20 dorsals. Schlegel meets this exceptional case by the hypothesis that the Duvaucel male skeleton in Paris Museum live animal may have been imported from Ceylon into Bengal. {see de Blainville, 1839-1864, Pl. 11 —Fig. 1170 of the present Memoir]. Bengal Elephant: 20 dorsals and 20 pairs of ribs. “Choonee.”’ Imported from Bengal in 1810, on board E. I. C. ship “Astell”’ by Capt. Hay. College of Surgeons. “This case, coupled with the Duvaucel skeleton in the ‘Jardin des Plantes,’ seems to establish, without searching for others, that the Continental Elephant of Northern India varies in the number of its dorsal vertebra from 19 to 20, as the African varies from 20 to 21.” Footnote: ‘The ingenious view advanced by Prof. Schlegel regarding the inverse relation between the number of laminz in the molars and the number of dorsal vertebre in the different species (supra, p. 263), does not appear to be tenable against the evidence adduced above, of the numerical variability in the living species.” Brit. Mus. and Indian elephants the crania are ‘‘so closely similar, that, in a museum, Killed in jungles on banks of Ganges, at no | without a record, the mere form will not instruct the observer whence the Bengal Elephant: | These two skulls agree in general form and proportions. In the Ceylon great distance from Meerut, in May, 1833 “lial came—whether continental or insular.” Ceylon Elephant: {Falconer’s statement is not supported by de Blainville’s plate showing College Surgeons 2656. the Ceylon and Bengal crania in profile, as reproduced in our Fig. 1170.] “The hypothesis entertained by Professor Schlegel, upon the statement of Diard, that Ceylon Elephants are frequently imported into Bengal is, I am satisfied, untenable. . . . On a review, therefore, of the whole case, the evidence in every aspect appears to fail in showing that the Elephant of Ceylon and Sumatra is of a species distinct from the Continental Indian form. ... The result of this range of observation, combined with long osteological study, has been to establish the conviction in my mind that there is but a single species of Asiatic Elephant at present known, modified, doubtless, according to his more north- ern or southern habitat, but not to an extent exceeding that of a slight geographical variety.” In Falconer’s opinion the cases above cited establish the fact that Hlephas indicus, including the continental and insular varieties, varies in the number of dorsal vertebrae from 19 to 20, in contrast to the African elephant which varies from 20 to 21. Osborn, 1929: From the above detailed observations of Falconer, together with those cited from Falconer on pp. 930, 931 of Chapter XV of the present Memoir, it appears probable that: (1) In the insular Ceylon ele- THE ELEPHANTIN#: HISTORICAL INTRODUCTION AND NOMENCLATURE 1313 phant, Elephas indicus ceylanicus de Blain., there is a constant number of 20 dorsal vertebra and 20 pairs of ribs together with certain constant differences in the cranium, tusks, etc., clearly establishing the subspecies; (2) in the continental form the vertebral formula is not constant, 19 dorsals and 19 pairs of ribs are recorded in certain skeletons of Bengal, while 20 dorsals and 20 pairs of ribs are recorded in other cases. (3) This would indicate that on the continent there are two hereditary blood strains, one characterized by 19 dorsals, the other by 20, possibly arising from the race indigenous to Bengal mingled with the race indigenous to Ceylon. (4) It would be interesting to ascertain whether there is a correlation between these two vertebral strains and the two cranial strains respec- tively known as ‘Mukna’ and ‘Dauntela.’ (5) The case may be parallel with that seen in the Equide, namely: (a) Pure Arab strain of horses (Equus caballus africanus), in which there are 23 dorso-lumbar vertebree, (b) the pure northern strain (Hquus caballus nordicus), in which there are 24 dorso-lumbar vertebre; there are 5 lumbar vertebre in the Arab strain and 6 in the Nordic strain, the rib-bearing dorsals being 18 in each subspecies. 2. CORSE, DE BLAINVILLE, AND FALCONER ON CHARACTERS OF THE GEOGRAPHIC VARIETIES (Continued more in detail on pages 1325-1333 of the present chapter) It appears that the subspecific forms of Elephas indicus are less numerous and somewhat less diverse than those of Loxodonta africana described above in Chapter XIX. The obvious explanation is that there is far less physiographic variation in the range of the Indian as compared with the African elephant. The geographic varieties or subspecies of the Indian elephant which have been successively named are: In 1841: Hlephas indicus Isodactylus Hodgson, of Nepal, Tarai, N. India. Elephas indicus Heterodactylus Hodgson, of Nepal, Tarai, N. India. While Hodgson did not name the above subspecies until 1841, he observed as early as 1832 the differences in the Indian elephant inhabiting the Ceylon and Bengal regions: (Hodgson, 1832, pp. 344, 345): “The elephant and rhinoceros abound in the forest and hills of the lower region of Nepal, where they breed, and have their fixed abode; and whence, in the season of the rains, they constantly issue into the cultivated parts of the Tarai to feed upon the rice crops. Both these genera are entirely unknown to the central and northern regions. The elephant is that so well known as the Indian variety, and as such is contra-distinguished from the African variety. But it may be questioned, if there be not two distinct varieties or species in India alone, viz. the Ceylonese, and that of the saul forest [Bengal]. The former differs materially from the latter by having a smaller lighter head, which is carried more elevated, and by higher forequarters. It is also said to be larger, and of a more generous and bold temper. The difference of size, how- ever, is certainly a mistake. I cannot speak to the point of temper.” In 1845: Elephas indicus ceylanicus de Blainville, the elephant of the island of Ceylon. Elephas indicus bengalensis de Blainville, the elephant of Bengal, continental. De Blainville figures in Pl. m1 [Fig. 1170 of the present Memoir] of his ‘“Ostéographie”’ of 1839-1864 the two subspecies from Ceylon and Bengal respectively, describing them briefly in the legend as follows: E. Indicus Ceylanicus, 4 grandes défenses. De profil, avee la mandibule en place; sexe inconnu. E. Indicus Bengalensis, i grandes défenses. De profil, avec la mandibule en place. D’un individu male dont le squelette, préparé par M. Duvaucel, a été envoyé par lui au Muséum. Geographic distinctions between the two continental races or varieties in India, known as the ‘Dauntela’ and ‘Mukna’ (as fully cited below) had been observed and very clearly stated by Corse as early as 1799 and 1314 OSBORN: accepted and amplified by Geoffroy St. Hilaire and Frédéric Cuvier in 1825 (also cited below). (1868, Vol. II, p. 257): THE PROBOSCIDEA ‘aleoner states 3ut even in the Sal forests of North Western India, at the extreme northern limit of the species at the present day, the difference of slender-built and squat-built Elephants is well known, being expressed by Corse, for the Bengal variety, under the designation of ‘mirghi,’ or Cervine [merghee, slender built, of Corse] for the former, and ‘Koomarea’ [koomareah, deep bodied, squat built, of Corse] for the latter, or when the characters are combined ‘Sunkareah.’ Falconer and Cautley accordingly gave geographic varietal names to these two very distinct continental forms of the Indian elephant, Known among the natives as ‘Dauntela’ and ‘Mukna,’ and published in his plates the following designations of these varieties: In 1847: Elephas indicus (Dauntela var.), the Dauntela variety [=deep bodied, koomareah of Corse]. Elephas indicus (Mukna var.), the Mukna variety [=slender built, merghee of Corse]. Temminck designated the Sumatran elephant under the following specific name and held that by its verte- bral and other characters it was thoroughly entitled to specific distinction from the Indian elephant, a point disputed by Falconer in 1868: In 1847: EHlephas sumatranus, the elephant of Sumatra. ELEPHAS INDICUS SUMATRANUS, THE SUMATRAN SUBSPECIES Fig. 1168. Pair of young elephants from Sumatra (Llephas swmatranus) captive (1921) in the Zoological Park of Washington. presented by Secretary Charles D. Walcott. (Broili, letter, August 12, 1929): In the State Zoological Museum of Munich there is an adult mounted specimen of Llephas sumatranus Temminck, the skeleton of which is not mounted but the skull is on exhibition. Head Preparator Kusthardt of the zoological collection writes (August 10, 1929) Photograph kindly that the adult mounted specimen came from Sumatra in 1907; that the young mounted specimen (six months of age) exhibits dark brown hair and red woolly hair. Fig. 1169. The Sumatran elephant, apparently a female, living in the Amsterdam Zoological Gardens, August, 1913. After a photograph kindly presented by Mr. Graham Renshaw, inscribed ‘““lephas indicus sumatrensis, Amsterdam Zoological Gardens, August, 1913.” (Letter of transmittal, November 21, 1923): “The Sumatran elephant was remarkable for the tessellation of the hide; as you see it was not a large animal . .. Buttikofer in the Guide to the Rotterdam Zoo figured another Sumatran elephant.” THE ELEPHANTINA: HISTORICAL INTRODUCTION AND NOMENCLATURE 1315 3. LIVING SPECIFIC OR SUBSPECIFIC FORMS, CONTINENTAL AND INSULAR (Continued more in detail on pages 1323 to 1333 of the present chapter). Falconer (1863, p. 81), in addition to his analysis of the vertebral characters, discusses very fully the unity or plurality of species among the existing Asiatic elephants, namely, types from Ceylon, Nepal, Bengal, and Sumatra, and decides in favor of the unity of the species Elephas indicus. He points out (1868, Vol. II, p. 258) the geo- eraphic variations which distinguish not only the Ceylonese and the Bengalese elephants, but the effects of climate and breeding in different regions, namely, Assam, Silhet, Chittagong, Tipperah, or Cuttack. He concludes that the specific distinctions of Temminck and Schlegel founded upon external characters, as in the case of Hlephas sumatranus, completely fail. He also believes that H. sumatranus is barely distinguished by the characters of the ridge lamine. He points out (p. 260) that while the typical ridge formula of H. indicus is: Dp 24 Dp 3s Dp 42 M1 M22 M3 the last true molar, M;, never shows less than 20 ridge-plates, commonly about 22 ridge-plates, but sometimes in the lower jaw attaining as many as 27 ridge-plates. He remarks (p. 269) ‘‘the evidence in every aspect appears to fail in showing that the Elephant of Ceylon [E. indicus ceylanicus] and of Sumatra |Z. swmatranus] is of a species distinct from the Continental Indian form [H. indicus]. Again (p. 270) he observes: ‘The result of this range of observation, combined with long osteological study, has been to establish the conviction in my mind that there is but a single species of Asiatic Elephant at present known, modified, doubtless, according to his more northern or southern habitat, but not to an extent exceeding that of a slight geographical variety.” He admits that the Ceylon elephants are occasionally imported into Bengal but does not admit, as claimed by Schlegel, that this is an explanation of the variation in the vertebral and rib formule of the Indian elephants. Falconer (1863), while inclined to accept Darwin’s theory of the evolution of species, does not recognize the very great importance of geographic isolation and of insulation in causing real geographic variation and subspecifie and varietal evolution both in external and internal characters. He is inclined to maintain the Linnean idea of species and to oppose the more modern idea of geographic subspecies and geologic ascending mutations. Osborn believes that the wide variations in cranial and vertebral characters as well as in dental and dermal characters and in the shape of the external ear support the subdivision of Elephas indicus into at least four out of the large number (12) of geographic varieties or subspecifie forms successively named, as follows: COLLECTIVE SPECIES ELEPHAS INDICUS LINNASUS, 1754 Synonyms: Elephas maximus Linneus, 1758; Elephas asiaticus Blumenbach, 1797; Elephantus indicus (Cuvier), 1801. Varieties and subspecies: Elephas indicus Isodactylus Hodgson, 1841 (named, without definition). Nepal, Tarai, N. India. Elephas indicus Heterodactylus Hodgson, 1841 (named, without definition). Nepal, Tarai, N. India. Elephas indicus ceylanicus de Blainville, 1845 (figure and plate description), the Ceylonese variety. Elephas indicus bengalensis de Blainville, 1845 (figure and plate description), the Bengalese variety. Elephas indicus var. Dauntela Fale. and Caut., 1847 (figure and plate description, without subspecific name). Compare Elephas indicus bengalensis de Blainville. Elephas indicus var. Mukna Fale. and Caut., 1847 (figure and plate description, without subspecific name). Compare Elephas indicus ceylanicus de Blainville. Elephas sumatranus Temminck, 1847. The Sumatran variety. Elephas maximus zeylanicus (Lydekker), 1907, 1916. Identical with Elephas indicus ceylanicus de Blainville, 1845. Elephas maximus maximus (Lydekker), 1916. Compare Elephas indicus bengalensis de Blainville, 1845. Elephas maximus sumatranus (Lydekker), 1900, 1916. Identical with Hlephas indicus sumatranus Temminck. Elephas maximus hirsutus Lydekker, 1914, 1916 (= Elephas indicus hirsutus). Malay variety. Elephas indicus Buski Matsumoto, 1927 (= ?Palzolorodon buskt). E.INDICUS CEYLANICUS. EsINDICUS BENGALE Ds BuaInvitLe’s Types or Mate EvepHas INDICUS CEYLANICUS AND E. INDICUS BENGALENSIS Fig. 1170. The narrower-skulled, slender-tusked Ceylon [ef. Mukna var.] and the broader-skulled, large-tusked Bengal [ef. Dauntela var.] races of the Indian elephant. After de Blainville, 1839-1864 [1845], Pl. 11, one-eighth natural size. Subspecies Elephas indicus ceylanicus. Summit of cranium more pointed (hypsicephalic); orbits relatively closer to condyles (eyrtocephalic). Premaxil- lary sockets longer; mandible shallower and more slender. The original of this cranium is not in the Muséum National d’Histoire Naturelle, Paris; Doctor Anthony reports (letter, July 16, 1930) that he does not know where this fine cranium is preserved; it is not the same as the female cranium in the Paris Museum shown in figure 1172 of the present Memoir. Subspecies Elephas indicus bengalensis. Occipitofrontal crest more rounded, less elevated; condyles to orbits relatively less cyrtocephalic. Premaxil- laries shorter, tusks broader; inframaxillaries more powerful. Mus. d’Hist. Nat. A.8016. 1316 sz THE ELEPHANTIN#: HISTORICAL INTRODUCTION AND NOMENCLATURE 1317 The discovery and naming of these geographic varieties and subspecies are recited above and continued more in detail on pages 1327 to 1334 of the present Memoir. Since all the above varietal and subspecific names were given prior to the adoption of the modern system of nomenclature, we await monographic research and comparison of the living varieties and subspecies of Hlephas; meanwhile those that seem best entitled to consideration at present are the following: 1845 Hlephas indicus ceylanicus de Blainville Syn.: Hlephas maximus zeylanicus Lydekker, 1916 Compare Elephas indicus var. Mukna! Fale. and Caut., 1847 1845 Hlephas indicus bengalensis de Blainville Syn.: Hlephas maximus maximus Lydekker, 1916 Compare Elephas indicus var. Dauntela! Fale. and Caut., 1847 1847 Elephas [indicus] sumatranus Temminck Syn.: Hlephas maximus sumatranus Lydekker, 1916 1916 Hlephas maximus hirsutus Lydekker [= EF. indicus hirsutus|? \ \ i o S a aNSTS r = eee \ ; . ELEPHAS INDICUS BENGALENSIS TYPE AND E. INDICUS CEYLANICUS XXII. A. E. INDICUS.(DAUNTELA mr) / iN Fig. 1172. Evep U ? XUILB E.INDICUS (MUKNA war) REFERRED. Paris MusEUM : After special photographs kindly taken under the direction of Dr. R. Anthony Fig. 1171. The broader, large-tusked DauNTELA and narrower, small- tusked MuKNA varieties of Hlephas indicus. After Falconer and Cautley, (eon) ed facial view of the cranium of Hlephas indicus bengalensis 1846 [1847], Pl. xum, figs. xxrm.a (Dauntela) and xxr.p (Mukna), one- (Mus. d’Hist. Nat. A.8016) described by Dr. R. Anthony (Paris, letter, July sixteenth natural size. Falconer’s distinctions between the Dauntela and 16, 1930) as follows: Cranium of a male Bengal eleubem same as that Mukna varieties are still more clearly shown in figure 1204 below. figured in profile by de Blainville in his “Ostéographie, 1839- 1864, Pl. m1 DaunTeLa var. Cranium relatively broader (brachycephalic), and less [Fig. 1170 of the present Memoir]. Found along the Ganges River by Duvaucel elevated. Premaxillariesshorter, with deep median excavation, highly charac- 1 1824. In this cranium the left BESTE is broken away and missing. teristic of the Elephantine. Compare Hypselephas hysudricus. The measurement h to h’ is 925 mm. or 3 ft. in. Reproduced one-sixteenth Muxna var. Cranium relatively narrower in proportion to its height; natural size. 3 ; seit i Q acl lew she crani Elephas indicus lant occiput more elevated, anterior nares narrower. Premaxillaries more elongate; (Right) BU eer air Oi wae Snag ® ee repens ees) fealeless divergent (Mus. d’Hist. Nat. A.8014). Of this specimen Doctor Anthony writes (Paris, July 16, 1930): A female born in Ceylon, brought from Holland, lived in the Jardin des Plantes 1797-1816; dissected by Cuvier and Rousseau; figured by The resemblance of Elephas indicus ceylanicus de de Blainville in his ‘“Ostéographie,”’ 1839-1864, Pls. 1 and u. Total facial : : 2 2 height of cranium h to h! 975 mm. or 3 ft. 2% in. Reproduced one-sixteenth Blain. to the EH. indicus var. Mukna Fale. and Caut. is _ natural size. afforded by the drawings of the cranium (side view) by de Blainville and Falconer and Cautley figured fully below. A fine front view of the cranium of bengalensis is compared with the front aspect of the cranium of ceylanicus (Fig. 1172), as described by Doctor Anthony in the legend above. Referring to the use of the terms “mukna” and “dauntelah,’’ Mr. Dunbar Brander commented (November, 1930) that he did not like a classification which used these two native terms, since a tuskless elephant (mukna) was apt to appear in any region and the term had no diagnostic value for any character other than tusklessness.—Editor.] *[See footnote under description of hirsutus on page 1332 below.—Editor.] 1318 OSBORN: THE PROBOSCIDEA The resemblance of the Hlephas indicus bengalensis de Blain. to the EL. indicus var. Dauntela Fale. and Caut. is also supported by a comparison of the crania and figures (lateral view). Further comparison should be made of the front views of the crania in these two subspecies and varieties. - HYPSELEPHAS~ PLAT ELEPHAS Fig. 1173. Geographic distribution of the principal species and subspecies (living and extinct) of Elephas, also of Hypselephas and Platelephas, accord- ing to the numbers given in the list on the opposite page. The white dots within the black areas represent the approximate localities where the types were discovered; these dots each carry a number in a circle, representing the chronoiogic sequence of type description. The + locates some of the principal referred specimens. 4. FOSSIL FORMS MORE OR LESS CLOSELY RELATED TO ELEPHAS INDICUS It is remarkable that no fossil Pliocene ancestors of the recent Indian elephant have as yet been discovered. While the cotypes of Hlephas hysudricus, attributed by Pilgrim to the Boulder Conglomerate zone, are in part recorded as found near Moginand, Simla Hills (Fig. 1196), none of the specimens personally collected by Barnum Brown in 1922 was actually found in the Boulder Conglomerate zone; they all appear as if washed or eroded out of this zone with more or less adherent gravel or concretionary material; they were found in hollows or ravines cut into the underlying Pinjor horizon. 1846 [1845]. Hlephas hysudricus Falconer and Cautley, of the Lower Pleistocene, found ‘below the con- glomerates’ of India, shows few resemblances in the cranium to the /. indicus (Dauntela var.) of Falconer and Cautley, and no very marked resemblances in the grinding teeth; this species appears to be unique. 1908. Hlephas hysudrindicus. Dubois described this species as theoretically intermediate between Hlephas hysudricus and Ef. indicus, but Stremme rightly regards this animal as more nearly related to ‘H. namadicus.’ Osborn treats this animal (Chap XIX) as belonging to Palewoloxodon, namely, Palxoloxodon hysudrindicus. 1927. Elephas indicus Buski. Matsumoto, having become convinced of the occurrence in Japan of the true Asiatie elephant in the fossil state (first suggested by Busk in 1868), described a first superior molar from the Ninohe District as the type of this subspecies. Osborn regards this tooth as referable to Palxoloxodon. 1929. LHlephas platycephalus. Osborn in the present Memoir redescribes this species from the Upper Pliocene or Lower Pleistocene of India, recorded by Barnum Brown as from ‘below the conglomerates,’ a level THE ELEPHANTINA: HISTORICAL INTRODUCTION AND NOMENCLATURE 1319 higher than that in which Archidiskodon planifrons occurs and lower than that in which EF. (Hypselephas) occurs. As shown in figure 1174, this is a very ancient and primitive animal. It is now made the genotype of Platelephas. 1929. Elephas platycephalus angustidens. Based upon a single second superior molar, r.M?, originally mistaken by Osborn for a third inferior molar, which now proves to belong to Hlephas | = Hypselephas| hysudricus. oO. NAMES OF SPECIES AND SUBSPECIES OF THE SUBFAMILY ELEPHANTINA IN ORDER OF DESCRIPTION See Figure 1173 A summary of the names applied to the above living and fossil forms actually or apparently related to Elephas is as follows: COLLECTIVE SPECIES Species Subspecies e2) 9) 9. Species 10. Subspecies 11. as 12: 13. Species 14. 15. 16. 17. 18. WwWnrR Gn Gr ps 1754 1758 1797 1801 1811 1841 1841 1845 1845 1847 1847 1847 1900, 1916 1907 1914, 1916 1916 1845, 1846 1915 1915 [1927 1929 1929 LIVING SPECIES ORIGINAL NAME Elephas indicus Linneus Elephas maximus Linnzeus Elephas asiaticus Blumenbach Elephantus indicus (Cuvier) Elephas gigas Perry Elephas indicus Isodactylus Hodgson Elephas indicus Heterodactylus Hodgson Elephas indicus ceylanicus de Blainville Elephas indicus bengalensis de Blainville Elephas indicus (Dauntela var.) Falconer and Cautley Elephas indicus (Mukna var.) Elephas sumatranus Temminck Elephas maximus sumatranus (Lydekker) Elephas maximus zeylanicus (Lydekker) Elephas maximus hirsutus Lydekker Elephas maximus maximus (Lydekker) SPECIFIC REFERENCE IN PRESENT MEMOIR Elephas indicus Elephas indicus Elephas indicus Elephas indicus Elephas indicus Indeterminate Indeterminate Elephas indicus ceylanicus Elephas indicus bengalensis Elephas indicus var. Dauntela Elephas indicus var. Mukna Elephas indicus sumatranus Elephas indicus sumatranus Elephas indicus ceylanicus Elephas indicus hirsutus' Hlephas indicus bengalensis EXTINCT SPECIES Elephas hysudricus Falconer and Cautley Elephas hysudricus hysudricus Soergel Elephas hysudricus primitivus Soergel Elephas indicus Buski Matsumoto Elephas platycephalus Osborn Elephas platycephalus angustidens Osborn Hypselephas hysudricus [Not determined by the present author] [Not determined by the present author] ?Palzoloxodon buski this chapter, p. 1333] Platelephas platycephalus Hypselephas hysudricus TYPE HABITAT Ceylon, India Ceylon, India (2) Nepal, India Nepal, India Ceylon, India Bengal, India Bengal, India Bengal, India Sumatra, Palembang District Sumatra, Deli [Labuan Deli?| Ceylon, India Malay Peninsula, Negri Sembilan Mysore,” India Upper Siwaliks, India Siswan, India Chandigarh, India See footnote on page 1332 below, where reasons are given for regarding this subspecies as of doubtful validity.—Editor. | *{Lydekker states on page 82 of his ‘Catalogue of the Ungulate Mammals in the British Museum (Natural History), 1916, that he has chosen Mysore as the typical locality of Linneus’ “Hlephas maximus.” Professor Osborn has chosen Ceylon, for the reasons stated above.—Kditor. | 1320 OSBORN: THE PROBOSCIDEA d uJ RNS Vim ‘ 7 Fic. 1174. Resrorations or PLATELEPHAS PLATYCEPHALUS, HyPSELEPHAS HYSUDRICUS, AND ELupeHAs INpIcUS. ApriL, 1930. All three heads reduced to a uniform one twenty-fourth scale (Left) Platelephas platycephalus drawn directly (Center) Hypselephas hysudricus _ profile (Right) Hlephas indicus, a middle-aged female. on the cranium, slightly crushed downwards; ears drawn directly on Falconer’s outline of the cranium Observe the relatively high position of the eye of supposed primitive size, occipitofrontal line (Fig. 1204). Observe the extremely low position of midway between the lip and the summit of the drawn directly on top of cranium. Observe the the eye, corresponding with the occipital concavity, cranium; also the lack of the occipitofrontal crest eye in normal position. namely, quite close to the premaxillary socket. and the presence of the prominent occipitofrontal The ear is given a slender elephantine outline, as in convexity and muscular ridge for the supratempo- E. indicus. The small tusks correspond with the rals. relatively small alveoli in this specimen. II. CHARACTERS OF THE SUBFAMILY ELEPHANTINA AND INCLUDED GENERA AND SPECIES SurpeERFAMILY: ELEPHANTOIDEA Osborn, 1921 FaMILy: ELEPHANTID& Gray, 1821 SuBraAMILy: ELEPHANTINZ Osborn, 1910 Original reference: ‘“The Age of Mammals in Europe, Asia and North America,’’ Osborn (1910.346, p. 558); also Osborn, 1918.468, p. 135. Compare: Elephantina Bonaparte, 1838; Elephantini Winge, 1906. SuBFAMILY CHaracters.—(1) Skull brachycephalic, progressive from platycephalic (Hlephas |Platele- phas| platycephalus) to hypsicephalie (HZ. indicus). (2) Jaws, including rostrum, progressively abbrevi- ating and deepening, completely brevirostral in recent time (H#. indicus). (3) Premaxillaries and tusk alveoli relatively narrow, in contrast to the broad rostrum of the Loxodontine. (4) Upper tusks upturned, out-turned, finally inturned, attaining moderate length, as compared with Loxodonta. (5) Grinding teeth progressively hypsodont. Ridge-plates increasingly compressed and enamel finely folded (EZ. indicus). (6) Ridge-plate formula of third molar multiplying from M 3 1° (platycephalus), to »32Ss-15 (hysudricus), to 24s; (indicus). (7) Inferior ridge-plates exceeding superior ridge-plates in number (E. indicus), in contrast to the Mammontine (Parelephas, M 3 2°-2°). (8) Including the progressive E. indicus, readily distinguished by its cranial axes and contours from all the known members of the Loxodontine (Chap. XIX), and Mammontine (Chaps. XVI-XVIII). (9) Provisionally including three extinct specific if not generic phyla, represented by EH. [Platelephas| platycephalus, E. |Hypselephas| hysudricus, and E. indicus, not constituting a single ascending phylum but probably polyphyletic. DovustruL RELATIONSHIPS.—The above definition of the subfamily Elephantine is continued from Chapter II, pp. 11-13 and 16, also from Chapter XV, pp. 913, 915, and 918. It is provisional and heterogeneous, because Elephas hysudricus Fale. and Caut. and £. platycephalus Osborn are at present known by cranial characters only and appear to represent generic or subgeneric phyla distinct from the true Elephas. There are three separate lines of cranial adaptation in the fossil and living species embraced within the subfamily Elephantine. This is in THE ELEPHANTINA: HISTORICAL INTRODUCTION AND NOMENCLATURE 1321 contrast to the cranial uniformity which prevails in the Mammontine and the clearly defined phylogenetic suc- cession observable in the three included genera (Archidiskodon, Parelephas, Mammonteus). These three generic or subgeneric lines may be compared and distinguished phyletically as follows: Phylum I PLATELEPHAS Osborn Typified by Elephas platycephalus. Upper Pliocene or Lower Pleistocene. Cranium relatively elongate, dolicho- cephalic, and platycephalic, occipital condyles somewhat above level of grind- ing surface of molars. Premaxillaries greatly elongated in front of molars, somewhat divergent; tusks unknown. Orbits large, elevated, near frontal profile. Grinding teeth relatively low, ridge- plates directly transverse, as in Elephas, no rudiment of ‘loxodont sinus.’ Ridge- plate formula: M 3 164% Habits unknown, probably like those of Elephas indicus. Very primitive in cranial structure and in the limited number of ridge-plates. Phylum II HyYPsELEPHAS Osborn Typified by Elephas hysudricus and E. platycephalus angustidens. Lower Pleis- tocene. Cranium elevated or hypsicephalic, condyles well raised above grinding sur- face of molars, occiput elevated with broadly transverse frontal crest, front- als deeply coneave. Premaxillaries relatively narrow or laterally compressed, not deeply extended below front of molars; tusks relatively straight, incurved, somewhat divergent at base; rostrum of lower jaw elongate, prominent, ramus shallow. Orbits large, depressed, near maxillary rostrum. Grinding teeth with ridge-plates con- vexo-concave, reversed above and below, trace of a median ‘loxodont sinus,’ of less height but otherwise as in Hlephas. Ridge-plate formula: M 3 17 Sree 19 Habits unknown, but from the tusks and grinding teeth probably similar to those of Elephas indicus. Progressive in cranial structure and in the somewhat larger number of ridge- plates, although the molar crowns are still low. Phylum III EvLepHas Linneus Typified by Elephas indicus. Extreme Upper Pleistocene and Recent. Cranium bathycephalic, eyrtocepahlic, hypsicephalic, oecipitofrontal dome more or less rounded, not acute, with expanding diploé, frontals gently coneave. Premaxillaries relatively narrow, sub- vertical, borders relatively close to front of molars; tusks relatively straight, in- curved, used in feeding and uprooting habits, unlike the adult Mammonteus. Jaw with extremely abbreviated rostrum, Orbits large, elevated. Grinding teeth with finely plicated enamel, not rising to the extreme hyp- sodonty of Parelephas, Archidiskodon, or Mammonteus. Ridge-plates multiplying: Known habits chiefly browsing, crush- ing of coarse leafage and herbage, secondarily grazing. Highly progressive and distinctive in cranial structure, with maximum number of plicated ridge-plates. I. As the Pleistocene is now recognized as extending over approximately a million years of geologic time, Platelephas platycephalus was nearly contemporary with Archidiskodon planifrons and with Hesperoloxodon antiquus of far western Europe and was separated by an enormous interval of geologic time from the recent Elephas indicus, but despite its very primitive cranial structure it would be rash to disbar it entirely from the ancestry of the modern FH. indicus. Il. Similarly Hypselephas hysudricus of the Lower Pleistocene is perhaps a million years older than Hlephas indicus but its cranium does not appear to be evolving in the direction of that of H. indicus. III. Therefore the recent species Hlephas indicus of southeastern Asia is left without a known ancestral form, just as its living contemporary Loxodonta africana is left without a known ancestral form.! In both eases this ancestral time gap will probably be filled by discoveries in northerly unexplored regions of Eurasia and of Africa respectively. In view, therefore, of the uncertainty regarding the ancestral relationships of Platelephas platycephalus and of Hypselephas hysudricus to Elephas indicus, we may at present define the genus Elephas from the characters preserved in the genotypic species /. indicus and in its geographic varieties and subspecies, as follows. See Loxodonta prima Dart, 1929, also L. africana var. obliqua Dart, 1929, Chapter XIX, pp. 1287, 1288 of the present. Memoir.—Kditor.] 1322 OSBORN: THE PROBOSCIDEA A Herp or Witp BenGat ELEPHANTS (ELEPHAS INDICUS BENGALENSIS) Fig. 1175. A herd of wild elephants in a bamboo jungle of Mysore. Photograph by Mr. Wiehle of Bangalore, originally published in The [llustrated London News of January 8, 1910, and reproduced by courtesy of Mr. Bruce 8. Ingram, Editor. The title of this remarkable photograph is “Quietly dreaming in the jungle.’ The giant bull to the left of the center illustrates the extreme prominence of the occipitofrontal convexities. SuPERFAMILY: ELEPHANTOIDEA Osborn, 1921 FAMILY: ELEPHANTID Gray, 1821 SUBFAMILY: HLEPHANTINA Osborn, 1910 Genus: ELEPHAS Linneus, 1735-1758 Pleistocene and Recent of Siam and India: Ceylon, Bengal, Burma, and Sumatra. Syn.: Huelephas Falconer (in part), 1857; Hlephantus Cuvier and E. Geoffroy, 1795; Polydiskodon Pohlig (in part), 1888. Genotypic species: Hlephas indicus of Ceylon. Geographic varieties and subspecies: Hlephas indicus ceylanicus, Elephas indicus bengalensis, Elephas indicus sumatranus, Elephas indicus hirsutus. GENERIC CHARACTERS.—(Linneus, 1758, p. 33): “Maximum quadrupes. Oculi parvi. Dentes Laniaril superiores exserti (Kbur). Aures amplissime, pendulex, dentate; act. angl. 277. p. 1051. Cutis crassissima, callosa. Mamme 2 juxta pectus. Ungues in apicibus loborum pedum. Genua flexilia. Col- lum breve. Proboscis longissima, eaxtensilis, acute odorans, loco manus vipsi inserviens; ea cubum potumque haurit, hostemque pellit; ea precisa occiditur; murem metuit sub somno ob trachex insertionem. Retro coit & mingit. Portat domos, rectore collo insidente; in bello armatur falcibus; vulnusculo inter axin & atlantem Juriosus occiditur; cxeterum prudens, docilis. (Osborn, 1924): Genotypic species Hlephas indicus (= maximus). Cranium widely distinct in form from that of Loxodonta africana, of Archidiskodon, of Parelephas, of Mammonteus, namely, relatively acrocephalic, hypsicephalic, moderately bathycephalic. Frontals gently concave, occiput decidedly convex (male), fronto-occipital crest uniformly convex, moderately elevated (female), i.e., acrocephalic. Molars of intermediate breadth, absence of ‘loxodont sinus’ in Elephas indicus, faintly indicated in Hyps- elephas hysudricus; moderately compressed enamel ridges of intermediate thickness, extremely crimped or sinuous in #7, indicus; more ridge-plates in inferior than in superior molars; ridge-plate formula of E. indicus, M 3 524;, of H. hysudricus (?) ref., M 3 +%*,,. Dorsal or rib-bearing vertebre 19-20. Digits of manus with five horny sheaths or nails; digits of pes with either four or five horny sheaths or nails. Digital formula: Manus 5, pes 4-5. As shown in the cranial and dental sections of the present Memoir, the genus Hlephas may be clearly defined as readily distinguishable both in cranial and dental characters not only from Loxodonta but from Archidiskodon, Parelephas, and Mammonteus. THE ELEPHANTINA:: ELEPHAS 1323 RipGe-PLATe COMPARISON WITH PARELEPHAS.—It is a striking fact that Faleoner’s ridge formula (Hlephas indicus), namely, M 3,74, assigns a higher number of ridge-plates to the last ¢nfertor molar than to the last superior molar; this reverses the condition observed in Parelephas progressus in which the ridge formula is M33 7—3. If this difference proves to be constant, it affords additional means of distinguishing species of Parelephas from species of Hlephas, as follows: In Parelephas there are more ridge-plates in the swpervor molars than in the inferior, whereas in Hlephas indicus there are more ridge-plates in the inferior molars than in the supervor. 1. SYSTEMATIC DESCRIPTION OF SPECIES OF ELEPHAS ELEPHAS INDICUS Linneus, 1735-1754, Collective Species! Figures 794, 796-800, 802-814, 816, 893, 912, 992, 995, 1042-1044, 1081, 1082, 1084, 1109, 1112, 1129, 1163, 1174, 1176-1178, 1190-1194, 1234, 1243, 1244 Syn: Hlephas marimus Linn., 1758; Hlephas asiaticus Blumenbach, 1797; Hlephantus indicus Cuvier, 1801 (in Cuvier and Lacépéde); Hlephas gigas Perry, 1811; EF. (Buelephas) indicus Faleoner, 1857. The history, nomenclature, and general characters of this collective species are fully set forth in the present chapter, pages 1308 to 1321 above. It is shown that Linnzeus (1754, p. 11—see facsimile, p. 1309 above) in naming Elephas indicus, while depend- ing upon Ray, Seba, and others, was very indefinite as to the type locality, mentioning both the continent of Asia and the island of Ceylon. When he changed the specific name to ‘maximus’ (1758, p. 33) he may have had in mind the African elephant, of which he had a foetal specimen, but he subsequently states ‘‘Habitat in Zeylonae.”’ Consequently we are inclined to the technical opinion that Linnzus’ type, both of his 1754 description of Elephas indicus and of his 1758 description of Hlephas maximus, was the Ceylon animal subsequently named by de Blain- ville (1845) Elephas indicus ceylanicus. If this historic interpretation is correct, H. indicus ceylanicus is not only the type of the collective species Hlephas indicus, but it is genotypic of the genus Hlephas itself. The main descriptions, figures, measurements, and characterizations by all other authors have been of the better known continental variety, to which de Blainville (1845) applied the subspecifie name Hlephas indicus bengalensis. It becomes necessary, therefore, to review the observations and history of opinion on the specific forms of the collective species ‘Hlephas indicus.’ [ELEPHAS GIGAS Perry, 1811 (=syNonym oF E. INDICUS, FIDE CoLBERT).—This species is described (p. Ii) ”) and figured (plate opposite that page) in George Perry’s article of 1811 in the ‘‘Areana,” a portion of which description is cited herewith: ‘Natural Order—Mammalia. Species—Elephas gigas. Generic Character—No fore-teeth in either jaw; the tusks of the upper are elongated and projecting, none in the lower; the proboscis or trunk very long and prehensile; the body armed with a very thick skin, covered with a few scattered hairs.” “The Elephant may justly be considered as the largest and strongest animal at present known, and is plenti- fully found in a wild state in the extensive regions of Africa and Asia.” “There is also found a second and different species, which is said to reside in the kingdom of Thibet, and being much smaller and of an opposite form, is to be considered as a separate animal from the above, under the title or Name of the Elephas socotrus [indeterminable from the description].”” Neither species determined by the present author.—Editor. | ‘Ray, in his description of Elephas (1693, pp. 131, 132), speaks of the elephant of Sumatra in reference to its weight, and to the skeleton in Florence in ref- erence to the number of ribs, characters of the tusks, etc. Seba (1734, p. 175, Pl. CXI) gives a very full description of the Foetus Elephantis Africani ineditus, which Lonnberg refers to as the type. In this early stage of zoology no one dreamed of selecting any particular specimen and designating it as the type. 1324 OSBORN: THE PROBOSCIDEA FALCONER DESCRIBES THE CONTINENTAL BREEDS OF BENGAL AND ASSAM The characters of these continental breeds of Indian elephant were more profoundly and thoroughly studied by Falconer than by any previous or subsequent writer. Falconer’s discussion of the vertebral formulz as well as of the varietal and subspecific characters is fully given above in the present chapter; he was not inclined to admit the constancy of the subspecific characters claimed by Schlegel, Temminck, and other observers. Conse- quently the following citations from Falconer refer to Hlephas indicus as a ‘Linnzan collective species.’ Fatconer (1868), DenraL CHARAcTERS.—In comparison with the grinding teeth of Hlephas |Hypselephas| hysudricus (Figs. 1197, 1198, of this Memoir), Falconer’s description of the superior grinding teeth of Hlephas indicus is as follows (Falconer, 1868, I, p. 78): “The existing Asiatic elephant, #. Indicus, furnishes the next modification represented in this plate. Fig. 2, Pl. v. (or fig. 2a, Pl. 1., F.A.S.), shows a section of the penultimate upper molar of this species. The gradual attenuation of the plates, successively exhibited from H. insignis to EB. Hysudricus, is here carried to excess, eighteen of these divisions being comprised within the space oceupied by about nine in the equivalent tooth of the African species. They are produced vertically in the same proportion, the height of the middle plate being about three-fourths of the entire length of the tooth; they, in fact, represent parallel perpendicular lamelle, of nearly uniform thickness from the base to the apex, interstratified with layers of cement of nearly the same thickness. The layer of enamel is attenuated into a thin transversely un- dulated brittle plate, the surface of which is deeply wrinkled with strix, for the firm cohesion of the cement. The general character of the section is a pectinated arrangement of the lobes like the teeth of a comb, which contrasts strongly with the chevron-formed ridges of E. insignis, and the cuneiform plates of E. planifrons. The mass of ivory at the base of the tooth is much thinner than in the cor- responding molar of #. Hysudricus, bearing but a very slender proportion to the height of the tooth; and numerous small and distinct fangs are given off from its inferior edge. This tooth had been some time in use, the anterior part of the crown being worn off as far as the ninth plate. The plane of the truncated portion is very oblique, being inclined nearly at a right angle to the coronal surface of the unworn portion. This specimen is 8.2 inches in length.” RipGe Formuta or EvLepHas inpiIcus (FALCONER, 1868).— Falconer appears to have finally based the ridge formula of M3; on an Indian elephant from Assam (Falconer, 1846, p. 43 [1845, fig. 20], with 27 ridge-plates); it is probable, therefore, that Falconer’s observations on the grinding teeth refer chiefly to the northern race, Elephas indicus bengalensis (cf. Falconer, ‘Paleontological Memoirs,” Vol. I, p. 422, Pl. 1): Hlephas indicus. Plate 1, fig. 2b, Indian elephant from Assam. Mz, with 27 ridges, length of crown about 15 in. The final formula of Falconer (1868) is consistent with that of Faleoner (1863), namely: H#. indicus of Assam, M3,,. Thus Falconer corrected his earliest ridge formula (1857, p. 315) of Blephas indicus and substituted (1863, p. 65) the follow- ing collective ridge formula of 2. (Huelephas) indicus: Dp 24 Dp 3$ Dp 422 M 122 M 222 M3524, Falconer considers and repeatedly states that the ridge formula of Hlephas [Mammonteus| primigenius is closely similar to that of his subgeneric group #. (Hueleph.) indicus (ef. the ridge- plate formule of the ascending mutations of M. primigenius above (p. 1138), namely, M 336 to M3 we )27/° Fig. 1176. (Falconer, 1868, Vol. I, p. 422): ‘“Hlephas indicus. section of unusually large specimen of last lower molar of an Indian Elephant The entire length of the crown is about fifteen inches, and it includes as many as twenty-seven ridges, Vertical from Assam, in India House [London] collection. of which the anterior thirteen are more or less abraded. The first five or six ridges incline a little forwards, while the posterior ridges incline so much in an opposite direction, that the hindermost are nearly horizontal, producing the flabelliform character that so readily distinguishes in most instances the last The same disposition and proportions of the dental substances are observed as in the upper grinder.’”’ After Falconer and Cautley, 1846 [1845], Pl. 1, fig. 2b. One-fourth natural size. from the penultimate lower molar. Inrertor Monar, M3, AssAM Bresp.—(Falconer, 1846, p. 43): “Fig. 2b represents the section of a very fine specimen of the last inferior molar of the existing Indian Elephant of Assam, from the collection at the India House. It is an unusually large speci- men, showing as many as twenty-seven plates, the anterior twelve of which have been in use. Precisely the same disposition of the dental substances is observed in this case as in the upper grinder, and they are developed in the same relative proportions. The vertical height of the plates is still greater than in the correspond- ing lower molar of 2. hysudricus. The upper surface is concave, and the under very convex. The anterior plates are nearly vertical, while the posterior gradually slope backwards till they become almost horizontal in the hindmost portion, with a corresponding gradual diminution in their height. This is a mechanical arrange- ment arising from the contracted diameter of the posterior part of the dental canal, in which the back part of the tooth is developed, close under the condyle, the plates being disposed so as to occupy the least vertical space. The basal mass of ivory between the plates and the fangs is reduced to a small quantity. This tooth measures 15% inches long in a straight line.” THE ELEPHANTINA: ELEPHAS RIDGE FORMUL OF ASSAM, CEYLON, AND SUMATRAN FORMS Falconer ‘Paleontological Memoirs,” 1868, Vol. IT, pp. 256, 260. Since the posterior ridge-plates of M%, M; develop very late in life, the ridge-plate formula is difficult to determine. Schlegel, Temminck, and other authorities contend that the very high ridge- plate formula of the Assam breed, M 3 5,745, is not obtained in the insular breeds of Ceylon and Sumatra, to which they assign not only a lesser ridge-plate formula but a greater thickness of the enamel ridge-plates, as quoted by Falconer (1868, IT, pp. 256, 260) : “(B.) Molar teeth—Ribbons (discs of wear) in form like those of the Indian species, 7.e. the enamel-plates highly crimped, parallel, and free from the rhomb-shaped expansion of the African Elephant; but the ribbons wider (in the direction of the long axis), and con- sequently less numerous than in the Indian species; the difference being in the ratio of 3 or 4:1 in the Sumatran, and 4 or 6:1 in the Continental Indian form (Schlegel in Temminck). Ribbons of enamel nearly [or] quite as wide as in the African Elephant. (C. L. Buonaparte.)”’ (Schlegel in Falconer, op. cit., p. 260): ‘‘ ‘The lamine of the teeth afford another distinction which however is less apparent to the eye than that taken from the number of vertebrae. These lamine, or bands, in #. Swmatranus, are wider (or if one may so say, broader in the direction of the long axis of the teeth) than in E. Indicus.’”’ Vig. 1177. Type r.M! of Elephas asiaticus Blumenbach, 1797 (Blumenbach 1797.2, No. 19): “B. vom Asiatischen Elephanten, ... Nur bilden jene beym Elephas asiaticus geschlingelte an beiden Enden paarweis zusammenlaufende Linien; hingegen beym africanus rautenformige Leisten. Diese Zihne der beiderlei Elephanten sind nach Originalen im hiesigen academischen Museum gezeichnet.”” Inverted to show natural position of molar. Blumenbach’s type figure is apparently a first superior molar of the right side, r.M!, outer side convex, inner side plane or slightly concave, exhibiting twelve ridge-plates, as in Falconer’s typical ridge-plate formula of Elephas indicus. Falconer, after the examination of a very large quantity of materials in India and Europe, concludes that the ridge formula runs thus (op. cit., p. 260): Milk molars. True molars. 4, 8, 12 ; Csi, 20-84 4, 8, 12 \ 12, 16, 20-24 [-27) This he regards as typical, stating (cf. p. 261) that neither Schlegel nor any of the other advocates of distinct specific ridge formule have proved that either the Ceylon or the Sumatran species shows a lesser number than M 2 75. He concludes: ‘“These 1325 instances prove, so far as they go, that the ridge-formula is the same in the Ceylon and Sumatran form as in the Indian.” He also contests the alleged differences in the width of the enamel bands and shape of the dises, stating as regards Elephas sumatranus that the supposed width of the enamel bands is due to the obliquity of the section (ef. p. 262). Falconer’s entire discussion (op. cit., pp. 256-270) of the unity or plurality of species confirms his conviction that there is but a single species of Asiatic elephant, modified only to the extent of a slight geographical variety. Fic. 238. — Coupe d’une derniére molaire inférieure d’Elephas indicus, au 1/4 de grandeur. Fig. 1178. Section of a partly worn third inferior molar of Elephas indicus. After Gaudry, 1878, p. 179, fig. 238, one-fourth natural size. Observe that six or more anterior ridge-plates have been worn off, the total number being twenty-four or more. DISTINCTIONS BETWEEN THE VARIETIES OF THE INDIAN ELEPHANT, CORSE (1799), GEOFFROY SAINT-HILAIRE AND FREDERIC CUVIER (1825), TEMMINCK (1847), DE BLAINVILLE (1839-1864), AND FALCONER (1867, 1868) (Continued from p. 1313 of the present chapter) It is not clear from the following descriptions by Corse (1799) and by Geoffroy and Cuvier (1825) in what part of India these two varieties occur; probably they refer to the continental varieties of Bengal. The derivation of the words mooknah, dauntelah, and pullung daunt, as given by Corse, is as follows: “Probably from mookh, the mouth or face . . . Dauntelah signifies toothy; having large or fine teeth. . . Pullung signifies a bed or cot, and daunt, teeth; and, from the tusks projecting so regularly, and being a little curved and elevated at the extremities, the natives suppose a man might lie on them at his ease, as on a bed [i.e., Pullung].”’ (Corse, 1799, p. 208): ‘After premising these general observa- tions, I may here observe, that elephants have two tusks, in the upper jaw only; but those in some of the females are so small as not to appear beyond the lip, whilst in others they are almost as large as in one variety of the male, named mooknah. . . . The largest tusks, from which the best ivory is supplied, are taken from that species of male named dauntelah, . . . in consequence of his large tusks, and whose countenance, from this circumstance, is the most opposite, in appearance, to that of the mooknah; which, as I have just observed, is hardly to be distinguished, by his head, 1326 OSBORN: THE PROBOSCIDEA ELEPHAS INDICUS BENGALENSIS Fig. 1179. A famous individual known as the ‘giant tusker of Udiapur.’ His tusks grew to such a length that he was unable to lie down and they were shortened at the extremities and encased in metal. After photograph by F. D. Fayrer, originally published in Asia Magazine, Vol. 29, June, 1929. Reproduced by courtesy of “Mondiale” through Asia Magazine. Scale approximately one-fiftieth natural size. from a female elephant. Though there is a material difference in the appearance of a mooknah and a dauntelah, as well as in the value of the tusks, yet, if they are of the same cast, (zat), size, and disposition, and perfect, that is, free from any defect or blemish, there is scarcely any difference in their price... . There must be five nails on each of his fore feet, and four on each of the hind ones, making eighteen in all; his head well set on, and carried rather high. The arch or curve of his back rising gradually from the shoulder to the middle, and thence descending to the insertion of the tail; and all his joints firm and strong. There are several other points, of less consequence, which are taken notice of by the natives as well as Europeans. The dauntelah is generally more daring, and less manageable, than the mooknah; for this reason, until the temper and disposition of the two species are ascertained, Europeans will prefer the mooknah; but the natives, who are fond of show, generally take their chance, and prefer the dauntelah; which, when known to be of a mild and gentle disposition, will always be preferred, both by Europeans and natives. The varieties between the mooknah and dauntelah are considerable, and for these there are appropriate names, according as the form of the tusks varies from the projecting horizontal, but rather elevated, curve of the pullung daunt .. . of the perfect dawntelah, to the nearly straight tusks of the mooknah, which point directly down- wards.” (Geoffroy Saint-Hilaire and F. Cuvier, 1825, p. 7, Eléphant d’Asie): ‘Nous avons déja vu que les femelles des Indes n’ont jamais que de trés-courtes défenses: il y a des males qui n’en ont pas de plus longues, sans qu’on en sache la raison. On les appelle Mookna. Ceux qui les ont longues se nomment Dauntelah, du mot daunt qui est le méme que notre mot dent. Cette différence n’en apporte pas dans le prix. Lorsqu’on ne connait pas le caractére d’un Eléphant, les Européens aiment mieux l’acheter sans grandes défenses, parce qu’il aura moins de moyens de nuire s’il se trouve méchant: mais les Indiens préférent assez les individus 4 longues défenses, pour s’exposer a tous les risques. Lorsque le bon naturel de animal est connu, les deux nations |’aiment mieux avec de grandes défenses. Il y a une infinité de variétés parmi les Daunte- lahs, par rapport 4 la direction et 4 la courbure de leurs défenses. Les plus estimés sont ceux ou elles approchent le plus de la direc- tion horizontale. Les princes indiens ont aussi un respect super- stitieux pour les Dauntelahs qui n’ont qu’une défense, comme cela arrive quelquefois.”’ Osborn, 1930: From the facts and comparisons cited in the early part of the present chapter and given more in detail above, it appears probable that the continental Indian varieties, as ob- served by Corse, Falconer, Geoffroy, and others, represent the descendants of two wild indigenous races or subspecies, including the larger, more vigorous northern form (var. Dauntela) and the smaller, more slender southern form (var. Mukna), which, freely interbreeding, according to Corse, have given rise to a large number of intermediate forms. It also appears probable, as sug- gested by Schlegel, that the smaller, more slender continental form is related to the Hlephas indicus ceylanicus of de Blainville, while the larger, more vigorous form is related to the HL. indicus bengalensis of de Blainville. On further research it is possible that these two subspecies may be clearly defined not only by the cranial characters described below, but by the vertebral formule enumerated above (p. 1312), as well as by the number of ridge- plates in the grinding teeth. It seems premature, however, to attempt subspecifie definition at present. THE ELEPHANTINA: ELEPHAS 1327 DE BLAINVILLE (1839-1864) SEPARATES THE BENGAL AND CEYLON BREEDS From the measurements and characters assigned by Corse, Geoffroy and F. Cuvier, de Blainville, and Faleoner, may be deduced the following: Elephas indicus ceylanicus de Blainville, 1845 Figures 1170-1172, 1180, 1204, 1226 Island of Ceylon and mainland of India. Compare Elephas indicus var. Mukna Fale. and Caut., 1847 Var. Muxna Fate. (cr. E. Inpicus CEYLANICUS DE BLAIN.) See Falconer, ‘Paleontological Memoirs,” 1868, I, p. 477, and 1845, Pls. -m1 of de Blainville’s ‘“Ostéographie,”’ 1839-1864. Small slender tusks, nearly straight and pointing obliquely downwards. Head smaller, hard to be distinguished from female elephant; premaxillary rostrum narrow, more elongate. Cranial dome more pointed, less brachycephalic, more hypsi- cephalic (see Figs. 1180, 1204). Narial openings narrower and smaller (see Fig. 1171). Five nails on each of the fore feet and four on each of the hind feet. Of gentle disposition; more manageable. Elephasindicus bengalensis de Blainville, 1845 Figures 801, 936, 963, 1013, 1170-1172, 1175, 1179, 1180, 1204 Chiefly Bengal, Assam. Compare Elephas indicus var. Dauntela Fale. and Caut., 1847. Var. DAUNTELA Fatc. (cr. E. INDICUS BENGALENSIS DE BLAIN.) See Falconer, “‘Palzeontological Memoirs,” 1868, I, p. 477, and 1845, Pls. 1-111 of de Blainville’s ““Ostéographie,”’ 1839-1864. Large and fine tusks, projecting horizontal, but rather elevat- ed, curve of the perfect Dauntela, known as the ‘‘Pullung daunt.”’ Head larger, premaxillary rostrum broader, less elongate. Cranial dome more rounded, more brachycephalic; frontals more convex (see Fig. 1204). Narial openings broader (see Fig. 1171). Five nails on each of the fore feet and four on each of the hind feet. Of more daring disposition; less manageable. CONTINENTAL AND INSULAR SUBSPECIES (DE BLAINVILLE, 1839—1864).—Preceding Falconer’s distinction between the Elephas indicus (Mukna var.) and the #. indicus (Dauntela var.) were de Blainville’s observations and his superb plates of 1845. When we compare these plates and the subspecifie descriptions of de Blain- ville with the descriptions of Corse, Geoffroy Saint-Hilaire and F. Cuvier, and Falconer, we are struck by an obvious cranial re- semblance, either due to affinity or to analogy, namely: Elephas indicus ceylanicus de Blainville, 1845, resembles Elephas indicus (= Mukna var.) Fale. and Caut., 1847. Elephas indicus bengalensis de Blainyille, 1845, resembles Elephas indicus (=Dauntela var.) Fale. and Caut., 1847. Elephas indicus ceylanicus male adult All 1.16 nat. size Elephas indicus Elephas indicus female juv. male adult Elephas indicus bengalensis male adult ComPaRISON OF ELEPHAS INDICUS CEYLANICUS (A, FEMALE, JUVENILE; C, MALE) WITH ADULT Matess (B, D) or E. INDICUS BENGALENSIS Fig. 1180. Male, female, and geographic characters in the skull of Zlephas indicus. All figures copied, with identifications, from de Blainville’s ‘“Ostéo- graphie,”’ 1839-1864, reduced to one-sixteenth natural size. Compare frontal aspect of crania (Fig. 1172). A, Female of Elephas indicus (?) ceylanicus, with tusks in place. The frontal curvature and extreme hypsicephaly remind us strongly of the profile of the juvenile Hypselephas hysudricus (Fig. 1213). B, Male of Elephas indicus (?)bengalensis, with adult tusks in place. C, Adult male skull of Elephas indicus ceylanicus, the geographic type of Ceylon. D, Adult male skull of Hlephas indicus bengalensis, the geographic type of the Bengal district. Observe the marked resemblance of the extremely hypsicephalic and eyrtocephalic cranium (A) to that of the juvenile Hypselephas hysudricus cranium (Figs. 1213, 1214), as well as to that of the Elephas indicus displayed in figure 797. 1328 OSBORN: THE PROBOSCIDEA Dr BLAINVILLE’s Priortry.—According to de Blainville’s figures of the Ceylon and Bengal crania, which represent individu- als of approximately the same age, de Blainville (1845) separated these two crania subspecifically as types of two subspecies, namely: Elephas indicus ceylanicus and Elephas indicus bengalensis; this subspecific separation is fully justified by the more or less hypsicephalic characters of the skull, and differences in the jaws and tusks; all these characters are widely different in these two geographic subspecies. As indicated above, these subspecifie char- acters appear to be the same as those which separate Elephas indicus (Dauntela var.) from Elephas indicus (Mukna var.) of Falconer. From this comparison, it would appear that Faleoner’s two varietal names (‘Mukna’ and ‘Dauntela’) proposed in 1847 may prove to be synonymous with the two subspecific names of de Blainville proposed in 1845. ONTOGENETIC AND SexuAL CuHaracters.—De Blainville’s observations (1839-1864) on the sexual characters of Hlephas indicus are clearly displayed in figure 1180, copied from his plates. We observe in figure 1180A a female apparently of Hlephas indicus ceylanicus, with extremely hypsicephalic head; in figure 1180C we observe an adult male of the same subspecies, #. indicus ceylanicus. In figure 1180B we observe an adult male, apparently of Elephas indicus bengalensis, with its rounded superior occipito- frontal dome; in figure 1180D we observe another adult male of E. indicus bengalensis, with the same rounded dame but with more massive cranial proportions. The crania of these subspecies, both male and female, are distinguished: (1) By numerous differences in the profile both of the cranium and of the jaws; (2) by the abbreviation of the jaw (C), the oldest individual figured; (8) by the uniformly convex fronto-occipital profile (B and D), growth stages of HE. indicus bengalensis; (4) by the more pointed and hypsicephalic fronto- occipital profile (A and C), growth stages of #. indicus ceylanicus. Fatconer’s Two Varteties.—It remained for Falconer also to name and distinguish these two continental varieties, signi- fying that he considered them of permanent varietal or subspecific value, as follows (footnote, Falconer, 1867, p. 57, and 1868, I, p. 477): “Comparison between Mukna and Dauntela varieties of Elephas Indicus. Mukna Dauntela. (big head). Inches Inches Extreme length of cranium............... 40.5 41.5 Width between zygomatics............... 29.5 29.75 Ditto post-orbitary processes.......... 22.5 24.0 Length from niche of occiput to tips of nasals. 22.0 21.0 Greatestiwidthvol OCcipiutins) seen eee 31.0 30.5 Widthrofinasalfopening? +). ..8 as aaene ae 14.0 15.0 Dept nom Gibto rs: att och cee Ao Beto BY. 73) Widthiof tisk-sheaths'2) 4.2.0. sheen 14.25 17.5 INaTLOWaWIGUnOM DOW, A era eee Oo SO 13.25 DepthvoMorbittrers 2s aches Ae ene 6.5 6.5 Height from condyles to occiput. .......... 22.25 22.0 ACTOSS CONG YIES Fp. ml os aio ete eee 8.75 8.5 [See footnote 1 on page 1317 above.—Kditor.] From condyles to tip of tusk-sheath........ 33.0 35.25 From ditto to anterior margin of molar aly eoluses yak scone ee Rene 22.5 22).25 From anterior margin occipital hole to pos- terion border palates. s-esmasessees see 12.0 11.5 ikength oftpalatesaa ee Cee ee 8.0 9.5 Depth of head from condyles to frontal sur- face at middle, opposite nasal opening... 23.5 23.5 Height from diastemal surface to bulge of OCCipUtS A. eer ae ee 30.75 31.25 Length of condyloid surface............... 6.5 6.25 From ear-hole to top of occiput........... 19.0 19.0 Length of anterior tooth, upper jaw........ 8.37 Width ditto ditto Number of plates, about ten Length of anterior tooth, lower jaw........ 9. Width ditto ditto pre tcne tact: Sion tirs 3.5 ¢dropped out 37 \dropped out Falconer also stated in this same footnote that ‘“The plates of teeth in the Mukna variety slope greatly backwards and are exces- sively and finely crimped; those of Dauntela are much less crimped.” Osborn, 1930: We observe that Falconer omits the most distinctive bathycephalic measurement, namely, (a) from the sum- mit of the occiput to the occlusal surface of the superior grinders, as compared with (b) occipital condyles to the orbital level of the frontals. These two measurements are shown in figures 805 and 806 and give us what may be called the bathycephalic index of the skull. SuMMARY OF OsBorRN, 1930: (1) Prior to Falconer’s de- scription (1847, 1867), de Blainville in 1845 separated the insular Ceylon animal as a subspecies, Hlephas indicus ceylanicus, from the mainland Bengal animal, subspecies Hlephas indicus bengalen- sis. (2) Falconer (1847) distinguished the two varieties ‘Dauntela’ and ‘Mukna.’ (3) Lydekker observes (Ency. Brit. 11th Ed., p. 259) that the insular ‘Ceylon animal, which is generally tuskless, may be the typical #. maximus [of Linneus], in which case the Indian race will be LE. maximus indicus.”’ (4) It appears probable that the Ceylon animal (2. indicus ceylanicus) with slender tusks was introduced into India by breeders, giving rise by crossing to the two varieties or hybrids designated by Falconer as Dauntela var. and as Mukna var.'! This theory was suggested by Schlegel, but Falconer (1868), as quoted above, did not admit that the insular Ceylon breed ever exerted any considerable influence on the north- ern Bengalese breeds, nor did he admit the specifie distinction of the Ceylonese or Sumatran elephants. (5) As shown by Osborn in a close comparison of the crania, Faleconer’s Mukna var. has a cranium closely similar to that of de Blainville’s H. indicus ceylanicus, while Faleoner’s Dauntela var. has a cranium more similar to de Blainville’s 2. indicus bengalensis. (6) Another ex- planation is that the island of Ceylon did not separate from the mainland until recent geologic time and that the H. indicus ceylanicus ranged through continental southern India. This problem of the continental and insular races and sub- species, however, requires further investigation from the original materials. THE ELEPHANTINA:: Elephas indicus sumatranus Temminck, 1847 Figures 1168, 1169, 1181-1185 District of Palembang, Island of Sumatra. The profound cranial differences which divide the Ceylon and Bengal elephants from each other as well as from the Sumatran elephant in all probability will be found to differentiate the Suma- tran and other still undiscovered extinct types. By comparison with the evolution of other Pleistocene ungulates it appears prob- able that a very long period of time separated these continental and insular subspecies and species from each other, a period of time equivalent perhaps to nearly half of Pleistocene time or 500,000 years, during which through isolation and segregation the sub- specific and specific characters were thoroughly founded. Here again monographic research is essential before we can reach a final conclusion. It is interesting to note that this Sumatran species, which we may now regard as a subspecies, was partly distinguished by the Dutch naturalist Temminck as early as 1847. He pointed out that Hlephas sumatranus has one more dorsal vertebra, i.e., 20, than FE. indicus, i.e., 19; that the free portion of the intermaxil- laries is shorter, the nasal cavities are shorter, the space between the orbits is narrower, the posterior portion of the cranium is broader than in EZ. indicus, the grinders have the narrower propor- tions of Loxodonta africana, the ridge-plate formula is intermediate between that of L. africana and that of F. indicus, and that por- tions of the #. sumatranus grinders and ridge-plate formula are intermediate to those of L. africana, while the shape of the plates is like that of the plates of #. indicus. Elephas sumatranus Temminck 1847. ‘‘Coup-d’Oeil Général sur les Possessions Néerlandaises dans L’Inde Archipélagique,”’ II, 1847, p. 91. Derscription.—Temminck distinguishes Elephas swmatranus from the African and Indian species as follows: ELEPHAS 1329 namely, the number of dorsal vertebre and the ridge-plate formula, may not be constant or valid. Consequently we are inclined to the opinion that Hlephas sumatranus is a case of geographic isolation or insulation with characters of not more than sub- specific value. Fig. 1181. Elephas indicus sumatranus from Deli {Labuan Deli?], Sumatra. Oblique front view of head, showing small and peculiarly formed ears and small tusks of a young male. Photographed in the Rotterdam Zoological Gardens. After Lydekker, 1916, Vol. V, fig. 24, p. 83, through the courtesy of the British Museum (Natural History), August 10, 1929. Characters distinguishing Ii. swma- tranus from FE. indicus and Loxodonta africana (cf. Tem- minck, 1847, pp. 91, 92): Vertebral formula: Cenvicalsyer ore een Morsalsreeriet ase oes ae oe as HEMI DATS MS oie eines Baste SAGrAIS Mee ies ak inate C@audalsmey vee tee es cies sires Sree ese sok cars a Mioatingeribstepsrrscrece) 4-2 oh Elephas sumatranus Enamel ridges thick crenu- lated, lamelle broad, less numerous; ratio 3 or 4 lamella to 1 of width of grinder; 6 lamelle in 12 centimeters Subsequent research and comparison, especially by Falconer, both as regards the vertebral and the ridge-plate formule, appear to demonstrate that the two characters of supposed specific value, Elephas indicus Enamel ridges thin, crenu- lated, Jlamelle narrow; ratio 4 or 6 to 1 of width of grinder; 6 to 8 lamella in 12 centimeters Loxodonta africana Enamel bandslozengeshaped, broad as in #. sumatranus; ratio 3 or 4 lamelle to 1 of width; 6 lamella in 12 cen- timeters Director E. D. van Oort of the Leiden Museum (letter from Dr. Max Weber, August 15, 1929) informs us that the “Catalogue ostéologique des Mammiféres,”’ y published by Jentink, in 1887, 1See Sclater’s translation of Schlegel’s paper on ‘““The Sumatran Elephant,’ Nat. Hist. Rev., 1862, p. 75, where it is stated that the number of caudal vertebrx in both H. sumatranus and ZL. indicus is usually thirty-three, but in very young examples sometimes only thirty.—Editor.] 1182. Hlephas sumatranus Temminck, cotype male and female crania from Palembang, Sumatra, in the Leiden Museum. Compare Jentink, Fig. Regu en 1845 de M. J. C. Baud (ef. Temminck, Coup d’ceil poss. Néerl. Inde arch., Cat. Ostéol. des Mamm., Mus. d’Hist. Nat., Tome IX, 1887, p. 169. Tome II, 1847, p. 91). After photographs kindly furnished by Director E. D. van Oort. A, Male cranium. Cat. a. Height of occipital crest to tip of premaxillaries 973 mm. B, Female cranium. Cat. b. Height of occipital crest to tip of premaxillaries 832 mm. A ig. 1188. Hlephas sumatranus crania from Sumatra in the Munich Museum. After photographs kindly furnished by Dr. Hermann Diirck and Herr Gustav Kiisthardt, November 13, 1930. A, Adult cranium from Batang Serangan, Unterlangkat, east coast of Sumatra, belonging to the mounted specimen represented in figure 1184. Observe the depressed and prominent position of the lower border of the orbits, as compared with Hlephas indicus bengalensis (ig. 1170), approaching rather E. indicus ceylanicus (Fig. 1170), and widely contrasting with the Loxodontine (Fig. 1108A-C); also the slender tusks, although a male, and extreme bathy- cephaly (750 mm. x 550 mm.). B, Infantile cranium from Sumatra (exact locality unknown to the present author), belonging to the mounted specimen represented in figure 1185. Compare the position of the external ear with that in the cranial profile of the adult 2. sumatranus skull (lig. 1184). 1330 THE SUMATRAN ELEPHANT IN THE Municu Museum After photographs and measurements by Dr. Hermann Diirck and Herr Gustav Kiisthardt (November 13, 1930) Fig. 1184. This specimen was measured immediately after it was shot at Batang Serangan, Cranium, right lateral view of same individual as that Unterlangkat, east coast of Sumatra. shown in figure 1183A. Total length, tip of tusk to tip of tail Height of shoulder above forelimb 294 Height above hind quarters 290 Length of ear to point 70 Breadth of ear 89 Total length of tusk 130 Observe slender tusks, although a male, and extreme bathycephaly (750 mm. x 550 mm.). INFANTILE SumaTRAN EverHant 1x THE Munich Museum Vig. 1185. Observe the triangular form of the ear, the hairy covering at the top of the head and of the back, and the infantile profile of the cranium (Fig. 1183B), in contrast to the type of Elephas maximus hirsulus (Pig. 1187), in which the hair covers the entire body. After photograph and measurements by Dr. Hermann Dirck and Herr Gustav Kiisthardt. ELEPHANTINA: 790 cm. =23 ft. 3% in. ELEPHAS 1331 less than three skeletons of JZ. sumatranus and several skulls, also one from Bor- enumerates not neo. The elephants of Borneo are restricted to the northeasterly part of the island; they are not indigenous but are feral, having been sent origin- ally as a present to one of the Rajahs. The ele- phant of Sumatra (van Heurn, 1929), long an economic factor in the life of the island, has re- cently been trained for military purposes. Coryprs.—Four skeletons of the wild Suma- tran elephant procured in August, 1845, from the district of Palembang, Sumatra, and forwarded to the Royal Museum at Leiden (see Schlegel, Verslagen en Mededeelingen der [Koninklijke Akademie van Wetenschappen, Afd. Natuur., XII, 1861, p. 101; translation by Sclater, Nat. Hist. Rev., London, 1862, p. 72); characters described by Schlegel and communicated to Temminck, who published his species in 1847 (“Coup deel,’ Vol. II, p. 91) under the name Elephas sumatranus, as cited below. ORIGINAL DescRIPTION (TEMMINCK, 1847, Vou. II, p. 91).—The cotype skeletons of Hlephas sumatranus are in the Leiden Museum. We may cite from the original description, as follows: (Op. = 2 ue cit., footnote 1, Chap. III) “Elephas Sumatranus Seay nob. ressemble par la forme générale du crane A = 9 lil Veléphant du continent de |’Asie; mais la partie 4 3ys libre des intermaxillaires est beaucoup plus courte et plus étroite; les cavités nasales sont beaucoup moins larges; l’espace entre les orbites des yeux est plus étroit; la partie postérieure du crane au contraire est plus large que dans Vespéce du continent.” “Les macheliére se rapprochent, par la forme de leur couronne, plutét de Vespéce asiatique que de celle qui est propre a l’afrique; c’est-a-dire que leur couronne offre la forme de rubans ondoyés et non pas en losange: mais ces rubans sont de la largeur de ceux qu’on voit 4 la couronne des dents de ?Eléphant d’Afrique; ils sont conséquemment moins nombreux que dans celui du continent de |’Asie. en arriére, comparées a celles prises dans la direction transversale et laterale, sont en raison de 3 ou 441; tandis que dans |’ Eléphant Les dimensions de ces rubans, dans la direction d’avant du continent elles sont comme 4 ou 641. La longueur totale de six de ces rubans, dans l’espéce nouvelle de Sumatra, ainsi que dans celle d’Afrique, est d’environ 12 centimétres, tandis que cette longueur n’est que de 8 4 10 cent. dans l’espéce du continent de |’Asie.”’ “Tes autres formes ostéologiques sont & peu pres les mémes dans les trois especes; mais il y a différence dans le nombre des os dont le squelette se compose, ainsi que le tableau comparatif ci-joint l’éprouve.”’ Marertats.—Lydekker (1916, p. 83) treats the animal as Elephas maximus sumatranus and figures a male elephant of this subspecies from Deli, Sumatra, formerly living in the Rotterdam Zoological Gardens (see Fig. 1181 of the present Memoir). Graham Renshaw (letter, July 27, 1925) writes: “In the Munich Museum there is said to be a male of Hlephas indicus sumatranus stuffed, from Lower Landak, Hast Sumatra. The skull 1332 of the extraordinary elephant with tusks sweeping round in a huge curve, mammoth fashion, was in the Indian Pavilion at the Wembley Exhibition, where I saw it. You will remember it was shot by Theobald and figured some years ago in the Bulletin of the New York Zoological Society.” OSBORN: THE PROBOSCIDEA triangular, form of the ear, the early date at which its upper margin is bent over, and the presence in the young condition—at least, In some cases—of a thick coat of black and in part bristly hair.” (Lydekker, 1914.2, pp. 285-288): ‘Thanks to the Trustees of Fig. 1186. Burmese elephant mounted by the taxidermist Mr. H. Brazener of Manchester, England. After photograph by Mr. G. Cameron, presented for the purposes of the present Memoir by Mr. Graham Renshaw (see letter of March 30, 1924). While comparatively a small animal, superficially it does not present any clear differences from the northern Elephas indicus bengalensis. One-thirtieth natural size. Elephas indicus hirsutus Lydekker, 1914! Figure 1187 From the Kuala Pila District, Negri Sembilan Province, Malay Peninsula. EBlephas maximus hirsutus Lydekker, 1914. Abstract Proc. Zool. Soc. London, Vol. I, No. 180, March 17, 1914, p. 20. Paper read March 17, 1914, under the title, ““The Malay Race of the Indian Elephant, Hlephas maximus hirsutus,”’ Proc. Zool. Soc. London, Vol. I, Art. 18, pp. 285-288. Typrn.— Young female Negri Sembilan Elephant, formerly living in the Gardens of the Zoological Society of London. British Museum (14.2.16.1). Locauity.—Kuala Pila district of the Negri Sembilan province, Malay Peninsula. Typr Figure.—Lydekker, 1914.2, p. 285, text fig. 1 (taken when alive in the Society’s Gardens); 1916, p. 84, fig. 25, as mounted in the British Museum. Tyre Description.—(Lydekker, 1914.1, p. 20): “Hlephas max- imus hirsutus, subsp. n., characterized by the square, instead of the estate of the late Mr. Rowland Ward, the Natural History Branch of the British Museum has received the mounted skin of the young Malay Elephant which died in the Society’s Gardens during the latter part of last year. While yet alive, the extraordi- nary hairiness of this animal attracted the attention of naturalists; and this feature, coupled with a peculiarity in the form of the ears, seems so well marked and so distinctive as to justify the recognition of the Malay Elephant as a distinet local race of the Asiatic species .. . this Elephant came from the Kuala Pila district of the Negri Sembilan province of the Malay Peninsula, and is believed to have been about three years old at the time of its death. As shown in the accompanying photograph from life (text fig. 1), it has a somewhat stunted appearance—suggestive, at first sight, of its belonging to a small race,—the height of the specimen, as mounted, being about 3 feet 8 inches. This stunted appearance may, however, be merely due to the effects of early captivity, for Mr. T. R. Hubback, in his book on Elephant and Seladang Hunting (The differentiating characters ascribed to this subspecies are the excessive quantity of hair and the peculiarity of the ears; but as these characters are not known to persist in any adult specimens, it is considered a subspecies of doubtful validity. 1938).—Editor.] (Note prepared by Dr. Barnum Brown, September 19, {In 1932, Professor Osborn stated in a letter to the editor that the Curator of the Raffles Museum regarded Lydekker’s hairy type of the Sumatran Elephant (2. hirsutus) as pathological. Professor Osborn added, however, that this subspecies was very small and its ears peculiar.—Editor.] THE ELEPHANTINA: ELEPHAS AND PALAXOLOXODON in the Federated Malay States, .. . records that the Elephants of the Negri Sembilan are of ordinary size, although of late years most of the big bulls have been killed off. He also mentions that practically all the bulls are tuskers, and very generally have one tusk much smaller than the other. . . . The tail is considerably longer than in the Indian calf, but since, according to Sanderson, there is considerable variation in this respect among Indian Ele- phants—which may or may not eventually prove to be of racial value,—I do not for the present propose to take any account of this feature. The great difference in the contour of the ear, coupled with the excessive development (at least in some instances) of black and in part bristly hair in the juvenile condition, seems sufficient to justify the separation of the Elephant of the Malay Peninsula as a distinct race, under the name of Hlephas maximus hirsutus.”’ Type OF ELEPHAS INDICUS HIRSUTUS Fig. 1187. Type of Elephas maximus hirsutus, from Negri Sembilan, Malay Peninsula, formerly in the Gardens of the Zoological Society, Regent's Park, London. After Lydekker, 1916, Vol. V, p. 84, fig. 25, reproduced through the courtesy of the British Museum (Natural History), August 10, 1929. Elephas indicus Buski Matsumoto, 1927! [=?Palewoloxodon buski| Figure 1188 Type: Ninohe District, Province of Mutsu, Japan. Post-Pleistocene to Recent. Referred localities (Matsumoto, 1927, p. 58): “Over forty miles from the sea-shore between Kanagawa and Toky6 (Leith Adams); Yedobashi, Tokyo (Naumann); Province of Mino (Tokunaga); Sapporo, Province of Ishikari, Hokkaid6é (Tokunaga); . . . Prefecture of Wakayama.” Matsumoto (1927, p. 57) remarks that the ‘‘occurrence of the Asiatic elephant in a fossil state in Japan was stated for the first time by Leith Adams and communicated by Busk [Footnote: ‘Leith Adams: Has the Asiatic Elephant Been Found in a Fossil State? Quart. Journ. Geol. Soc., London, Vol. XXIV, 1868, pp. 1333 496-498, text-fig.;—Busk, Additional Remarks, 7d., pp. 498, 499.’]. According to Busk, a certain fossil elephant from Japan is referred to Elephas indicus, representing, however, a form with teeth some- what larger than the average of the existing one. Quite recently the writer has come to be fully convinced about this first record, which appears to have been overlooked so long, and proposes here to call the Japanese form in question to the credit of Busk, as follows. Elephas indicus Linné Buski, subsp. nov.” This interesting specimen, as described by Matsumoto, is as follows: Elephas indicus Linné Buski Matsumoto, 1927. ‘On a New Fossil Race of the Asiatic Elephant in Japan.” Sci. Rept. Téhoku Imp. Univ., Second Series (Geology), Vol. X, No. 3, pp. 57, 58. Typr.—‘Antepenultimate upper molar of left side (I.M'].”” Inst. Geol. and Paleont. 7266. HorIzoN AND Locauiry.—(Op. cit., p. 58): “Judging from the exceedingly feeble fossilization of the remains, as far as examined by the writer, the present form may probably belong to a very late geological age, such as the Post-Monastirian. The occurrence of the present form may indicate a warmer climate, that being well in accord with certain geological evidence already known.’ From Ninohe District, Province of Mutsu, Japan. Type Figure.—Op. cit., 1927, Pl. xxvut, figs. 2 and 3. Typr Description.—(Matsumoto, 1927, p. 57): ‘The type- specimen consists of eleven ridges besides the anterior and posterior talons, and measures 155 mm. in length, 60 mm. in the greatest width at the third ridge, and 154 mm. in the greatest height of crown at the seventh ridge, which was just ready to commence to wear. The middle part of the crown is peculiarly narrowed, being narrower than both anterior and posterior parts. Ina palatal view, the crown is more or less bent inwards. Its frequency of ridges in a length of 100 mm. is about 8 on both the sides and about 7 on the grinding surface. The ridges are only very weakly flexuous; their inner and outer sides in a fore-and-aft view are not very markedly convex, being almost parallel at the greater middle part of the height of the ridges. The anterior and the posterior side of the moderately to strongly worn ridges at the grinding surface are nearly parallel, without any loxodont sinus. The plication of enamel of those ridges is almost uniform from end to end, and is very fine, regular, and strong, as a very distinctive characteristic of the present species. The figure of enamel seen in the rather Evepuas rNpicus Buskt Matsumoto, 1927 [=?PALa#OLOXODON BUSK1] Fig. 1188. Type figure of Elephas indicus buski Matsumoto, 1927, PI. XXvul, fig. 3, one-half natural size, from Ninohe District, Province of Mutsu, Japan. Original in Institute of Geology and Palzontology, Tohoku Imperial University (No. 7266). A first superior molar of the left side, 1.M!, crown view. 1Osborn, 1930: At first disposed to place this type near Elephas indicus, comparison with figure 1177, an M! of E. indicus (=asiaticus) proves that such reference is doubtful. It appears rather to belong near Palaoloxodon namadicus naumanni, P. namadicus namadi, or P. hysudrindicus. 1334 slightly worn ridges is mesially laminar and laterally annular. The worn surface of the sixth ridge, which is very slightly worn, con- sists of five mammille arranged in a transverse row. The layer of enamel is considerably thin, measuring about 1-1.5 mm. in thick- ness on the anterior and the posterior side of the ridges.” LeitH ADAMS AND Busk (1868).—(1) In 1868, as mentioned above, page 1062, Leith Adams (1868, p. 497) described and figured a ‘Fossil Tooth of Elephas Indicus from Japan,” discovered forty miles inland between Kanagawa and Jeddo. He rightly interpreted this tooth (Fig. 1189 of present Memoir) as an r.M?, describing it as follows: “This penultimate true molar, right side, upper jaw, although partaking of the characters of H. Armeniacus, Fale., differs from the latter in its thick plaits, their less approxi- mation, the festooning being carried round the loops of the disks of wear, and the total absence of any mesial expansion.”’ (2) Busk (op. cit., p. 498), after comparing the tooth with #. [Parelephas] Armeniacus Fale. and EF. [P.] columbi Fale., and finally with a corresponding fossil tooth [in his opinion an 1.M! of F. indicus], concludes as follows: ‘‘But these differences [referring to his comparison with #. indicus] do not appear to be of much im- portance, and there seems to be every reason to believe that the Japanese fossil tooth belonged to a form of E. indicus, with teeth somewhat larger than the average of the existing one.’”’ (3) The teeth described and figured by Adams and Busk (1868) and by Matsumoto (1927) are not dissimilar in respect to thickness and the interspacing of the enamel ridge-plates, but in the type of Elephas indicus buski the enamel foldings or plications are much closer and more compact and consequently more numerous. The ridge-plates in the Adams-Busk grinder (Fig. 1189) also in the type of EB. indicus buski (Fig. 1188) are much more widely interspaced OSBORN: THE PROBOSCIDEA than in Dubois’ type superior molar (Fig. 1160) of Palxoloxodon hysudrindicus. Osborn, 1930: From the above comparisons Osborn is in- clined to agree with Lydekker that the above grinding teeth, namely, the type of Hlephas indicus buski (Fig. 1188) and #, namadicus ref, Lydekker (Fig. 1189), belong either to Z. [Palzxoloxo- don| namadicus naumanni, E. |P.| namadicus namadi, or to some even more progressive species of Palxoloxodon, e.g., EH. [P.| hysudrindicus. Seconp Ricur Superrtor MOLAR REFERRED TO PAL.ROLOXODON Fig. 1189. A second right superior molar, r.M?, probably from the Middle Pleistocene of Japan, between Kanagawa and Tokio (Yedo). After Lydekker, 1886.2, p. 168, fig. 29, about one-half natural size. Original in the Museum at St. John, New Brunswick, Canada. This molar, first figured by Leith Adams and Busk (1868, p. 497), and interpreted as a ‘Fossil Tooth of Elephas Indicus from Japan,” was compared by the same authors (op. cit., p. 498) with the H. [Parelephas| armeniacus Fale. of Armenia and of China. Length 187 mm., width 76 mm., ‘height 111 mm. The present author regards it as referable either to Pal«oloxodon namadicus naumanni, P. namadicus namadi, or P. hysudrindicus. Observe crimping or plication of the enamel and absence of ‘loxodont sinus,’ resem- blance to Hlephas indicus grinders. Observe similar plication or crimping in the type (Fig. 1160) of P. hysudrindicus. 2. DISTINCTIONS AND MEASUREMENTS OF THE INDIAN ELEPHANT Comparison has been made in Chapter XV between the Indian and the African elephants as regards dental, cranial, and vertebral structure and prevailing feeding habits. As to dental structure, the very earliest observers remarked the profound differences between the crown view of the molars of the African and of the Asiatic ele- phants, finally figured and described (1846) by Richard Owen (Fig. 1191). Itis not possible that the African ele- phant, with its coarse-plated, relatively low-crowned grinders, can attain the same age as the Indian elephant, with its very high and long-lived grinding teeth. Detailed observations by Falconer (fully discussed in Chap. XV) on the food of the Indian and African ele- phants indicate that the straight tusks of the African elephant, as in all other Loxodontines, are used as crowbars in uprooting trees as well as in combat. It does not appear that the shorter and more slender tusks of the Indian elephant are used to so great a degree for such purposes; they are invariably more slender and more strongly upcurved, and those of the old Indian bulls may attain great length (Fig. 1179). Doubtless for a very long period of time the Asiatic elephant was hunted for its ivory, even before it was domesticated, both in India and Ceylon. The ivory tusks, both of the Indian and Ceylonese varieties, are capable of carrying very heavy logs, held in place across the tusks by the trunk. In longitudinal section (Fig. 1192) it appears that the low-browed African cranium, in perfect harmony with its low-crowned grinders, is in very wide contrast to the high-browed Indian cranium, with its very high grinding teeth. On the principle explained in Chapter XV, the Indian cranium (Fig. 800) is far more bathycephalic than poe ES aah hed i OCA hau ho hb fiipad. eG, shirts Fig. 1190. Superior view of the head of a young African elephant (left) and of an adult Indian elephant (right). After Geoffroy Saint-Hilaire and I'rédéric Cuvier, 1825, Livr. LI, LI, one twenty-fourth natural size. The superior view of the head of the adult Hlephas indicus should be compared with the superior view of the heads of the adult male (Loxodonta africana albertensis) and female (Loxodonta africana peeli), as mounted in the American Museum collection and photographed from above in figure 1063 of this Memoir. The very marked differences between the male or female Loxodonta and the male or female Elephas are indicated in this comparison. Loxodonta is distinguished by the rounder occiput, lacking the pronounced air sinuses of E. indicus, also by the greater elongation of the antorbital or facial region of the cranium. The differences between the cranium of Loxodonta and of Elephas are quite as pronounced as the differences between the grinding teeth displayed in figure 1191. § qrind@ surf. Bs ~ ELEPHAS INDICUS LOXODONTA AFRICANA Asiatic Elephant. African Elephant. 3, nat. size. Fig. 1191. Crown view of the third inferior molar of the right side of: Fig. 88, Loxodonta africana; Fig. 89, Elephas indicus, one-third natural size. After Owen, “A History of British Fossil Mammals and Birds,” 1846, pp. 280-232: “Thus in the African Elephant, (fig. 88), in which the lozenge-shaped plates are always much fewer and thicker than the flattened ones in the Asiatic species, the variation which can be detected in any number of the grinders of the same size is very slight. ... In the molars of the Asiatic Elephant, (fig. 89), which, besides the difference in the shape of the plates, have always thinner and more numerous plates than those of the African species, a greater amount of variation in both these characters obtains; . . . and the like caution is still more requisite in the comparison of the molars of the Mammoth (Hlephas primigenius), which, having normally more numerous and thinner plates than in the existing Asiatic Elephant, present a much greater range of variety.” AIS OF M3 SUrFacp Fig. 1192. Contrasts of (right) the low-browed African cranium (Amer. Mus. Dept. Mam. 51939) with (left) the high-browed Indian cranium (Amer. Mus. Dept. Mam. 54261). Compare figures 805, 806, 800, 1061, and 1053. The brain, deeply embedded below the cranial air cells, is seen to be slightly larger in the Indian elephant (6686 ccm.) then in the African elephant (6651 ccm.). Bath., the vertical bathycephalic diameter from summit of occiput to crown of grinding teeth. S.oc.—S. oc., the supraoccipital axis (=occipital plane). B. oc.—B. oc., the basioccipital axis (=basioccipital plane). Compare figure 1112. Resp.—Resp., the respiratory axis from the anterior to the posterior nares (=respiratory plane). Pal.—Pal., palatal axis of the roof of the mouth. M°—M5, grinding surface of the superior molar teeth. 1335 1336 OSBORN: THE PROBOSCIDEA the African cranium (Fig. 1061), yet the African elephant, as shown in the measurements (legend to Fig. 1053) of the growing “‘KKhartum”’ (Loxodonta africana oxyotis) attains its greater height very rapidly, namely, from a height of 4 ft. 14 in. in October, 1906, to a height of 10 ft. 8% in. in January, 1930, or an average annual growth of 3% in. in twenty-four years. The extreme shoulder height of the African elephant, in the flesh, is estimated at 11 ft. 64in. (see footnote 2 on page 1022, caption to Fig. 912), whereas, as shown in the observations below, there are only two records (neither of which is absolutely reliable) of the attainment by the Indian elephant of a greater height than 10 ft. 6 in., the average being about 9 feet. SHOULDER HEIGHTS OF THE INDIAN ELEPHANT Throughout this Memoir the estimates of shoulder heights are based on the fully extended forelimb, with certain allowance for the cartilages between the bones, for the foot pads, and for the great muscles of the neck. As compared either with the African or with the great extinct elephants of Pleistocene time, the Indian elephant is generally inferior in stature; it exceeds Mammonteus primi- genius in height by a foot to eighteen inches; the somewhat doubtful ‘record’ of the Indian shoulder height, as shown in figure 1194, namely, 10 feet 6 inches, agrees with the highest figure given by Corse below, while the average large Indian elephant rarely exceeds 9 feet. The observations of Corse (1799, p. 35) chiefly relate to the heights of domesticated elephants and are very interesting to cite in this connection: (Cf. Corse, 1799, p. 35): “elephants attain their full size be- tween eighteen and twenty-four years of age. . . . In India, the height of females is, in general, from seven to eight feet; and that of males, from eight to ten feet, measured at the shoulder.” One elephant, on good authority, exceeding ten feet, was a male, meas- uring as follows: Irom the top of the shoulder, perpendicular height 10 ft. 6 in. Irom the top of the head, when set up, PES 2. From the front of the face to the insertion of the tail 15 “ 11“ Of 150 Bengal elephants only a few males attained 9 feet 6 inches, not one of them attained 10 feet. The Ceylon elephants are neither higher nor superior in any respect to those of Bengal. Contrary to prevailing opinion, the elephant is in no sense Uneunicrapism. Raprocrarn or Riaut Poor or YounG plantigrade; as shown in the radiograph reproduced herewith a aus, ae nD. : (Fig. 1193) the foot is wnguligrade; the phalanges are subver- Fig. 1193. This beautiful radiograph of the pes of a young 3 ‘ ; elephant taken from the inner side of the foot exhibits clearly the tical in position, the terminal phalanges are greatly reduced in complete inner or first digit, including Ph. Tin Phed. Mts. I, Cu., Nav., Astrag., Cal., Ep., and Tibia. The remaining four digits are faintly seen in outline. The pes is shown to be unguligrade, part of the foot from injury, while a posterior elastic pad is i.e., to rest upon the ungues surrounding the tips of the terminal ; : i . phalanges, in this case Ph. I7; it proves that Elephas is in no sense pressed down in a walking or running gait. plantigrade. It also exhibits the cartilaginous interspaces, which add considerably to the height of the skeleton. The scale is In the accompanying figure (Fig. 1194) are given the esti- approximately two-fifths natural size. Reproduced through the . , ; courtesy of Dr. G. M. Vevers of the London Zoological Society. mated shoulder heights of three elephants in the American size and are encased in horny sheaths which protect the front THE ELEPHANTINA: ELEPHAS 1337 Museum collection, ranging from 8 ft. 4% inches to 8 feet 8% inches, as compared with that of a very large male measuring 10 feet 6 inches in the flesh,’ according to Rowland Ward’s “Records of Big Game”’ and a measure- ment taken from Dollman-Bather, 1927. This 10 feet 6 inches maximum agrees with the single exceptionally large specimen reported by Corse. ERERIARAS INDICUS 2566 1 sant S in ip ual , “" ‘ ta “o“ 85 8 4%’ 10° 6 «” " ia “ i SAMSON GUNDA VERNAY 5 RECORD A./7.39082 A. /7.3908/ A. N71, 54453 Fig. 1194. EstimatTep SHOULDER HEIGHTS Or THE INDIAN ELEPHANT, SKELETAL AND FLESH [Professor Osborn’s method, used in the present Memoir, of estimating the height in the flesh is to add six and a third per cent. to the skeletal height.— Editor. | Small male, “Samson” (Amer. Mus. 39082), skeletal height to summit of scapula SKELETON Lesa 2566 mm. 8 ft. 5 in. Larger male, ““Gunda” (Amer. Mus. 39081), about twenty-one years, total extended limb segments, skeletal height Male wild elephant, Vernay Coll. (Amer. Mus. 54453), skeletal height 2660 mm. 8 ft. 8°4 in. 2558 mm. 8 ft. 434 in. height in flesh 2720 mm. 8 ft. 11 in. Very large male, alleged record (Ward, also Dollman-Bather, in Bather, 1927, p. 104) in the flesh (skeleton not measured) 3200 mm. 10 ft. 6in.! According to Rowland Ward (‘‘Records of Big Game,” 1922, p. 468), the record shoulder height of the large male wild Indian elephant is 10 feet 6 inches {A subsequent record (Ward, edition of 1928, p. 451) gives a height of 10 ft. 8 in.—Kditor.] 1This “record” is not thoroughly authentic, as shown below in citations from Corse and others; it is very difficult to measure a prostrate elephant accurately. The record of the large Haslemere specimen exceeds this, the living height at the withers being recorded as 11 ft. 1 in. The following is cited from The Haslemere Museum Gazette, Vol. I, No. 1, May, 1906, p. 13: ‘It is that of a very large Ceylon elephant, shot by Mr. W. H. Varian, at Chalampi Madua, in the North Coast Province of Ceylon, in 1882... . The following measurements of this animal, taken immediately after death, 11 feet 9 inches; height at withers, 11 feet 1 inch; ... Height at arch of back, length from the tip of the tail to the tip of the trunk, 26 feet; girth of the body at thickest part, 22 feet 4 inches; weight about 8 tons.” The measurements of the large Haslemere cranium agree closely with those of the two erania given by Falconer (1868, Vol. I, p. 477). 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ERLIMOLIOND suapy -u0d pue { ‘(auq sf (ereweq pue -snBup Uuopo}aq suapyy souvpspurs -1B18M) STE neg) stim “vijay ‘suyeuler = -snbup = wopoy suaprysndup orey ayy jo oreyy 24} JO gurid YatM sem0%s 5 e5v}S (QoIINW 5 -9Qv4jAT UIA vopojaqn4}a TL, soUO}spues souozspues pues ompjsnovl = JeddQ) — ynesey FA ‘(speq_—-eseq) *(speq yeseq) a ey jsouttodd ysounredd a ‘ones [[nesey a O38}S [VITUE = NYBAMIG IIMOT says yuloM | reyyouryg “TL UvIzIATSH = — = — —— !~ aa eel FI (A) Fre] — poztusoo 4 2 -a1 Ueeq jou Ed, S aan —- poled asuarluryd x z WEYSH LON ]RT styg 03 9qu (4) wenvayolh A 2 ILlapengy -dojol SOQ etay ‘sou0j3 s)dojaque [jvus puv pug Jo -op spoq ouvyy -paes Aysour ‘oeualuryd Unisay) see, soa | (spoq oppprte) ° 0 =, “sullas ucytn | J -ohy ‘abeqe Wury) iO -1vd]td © 94D qvyyourlyy 'T uviuoyoy, —_--- --— SSS el S$ |—_—_—_]} 9) aurpuod = |-———_———— afaynvs uopob ~faT10) Sd}OKJ -aS ‘xhtayoffosy MOSATT 5» UMajinns uopobay UEnILpUL wnt ‘.quojspues ‘souoj}spues pue -2yyohyy = ‘ahsous pues soyeys (spaq toddn) a 8 ve os stvpo ‘oDe}s Wen oe “ula ‘9BV4S BEN ~ se ‘sayeys Hefoy | Ieyyourey “TL uepyjvaes asuys = = ous 943 03 3 ol pivvaop an nD pusjxo Avul Zz = guq ‘[us3qnop 3S rd 9004s qyuny, = Jamo] suopijin). uopollag aa sodojajue = -puvs pus sdep> “Up2]4bd DIAC, pie snjobne n OBIE] ‘untayjopn) ‘(ureqiaomn UIE sau0}s -409 Wopojaqns}a |, -eH ‘uounddiy St SIU} JO 4y0NIT SouOPSpUUS Zunosducusyx ye |-pues ‘euuing wi step pure “upqi) sng suapryv} JMO] 249) STITH puv - s~ cee CONGVIES eee oe 1085 925 : 4 Height of cranium, grinders to mid- Fig. 1218. Right cranial profile of Platelephas platycephalus (...... ) and ; | nee, oy » of Hypselephas hysudricus (_____), one-sixteenth natural size. fronta Region: Fee eee ee 605e 645 This figure illustrates the primitive platycephaly of the Platelephas Width across midtemporal region. . 332 334 platycephalus type as compared with the relative hypsicephaly of the Hypsele- Length of tusks (outer curve)...... 2245 phas hysudricus type. Both crania may be regarded as adult and fully de- Mranaverse cianieter. of) husk. > at veloped. Observe that P. platycephalus does not in the least resemble the ; 7 x Me ‘ 0 ui At a juvenile profile of H. hysudricus (Wig. 1213); it is much more primitive (ef. emergence, i.e., width of alveoli 120—-135e 117 Figs. 1207, 1209). THE ELEPHANTINA:: PLATELEPHAS 1361 grinders, the distance being 507 mm., as compared with 325 mm. the anterior nares of the space between the premaxillary sockets in #. indicus. and the tusks; (4) and finally the great prominence of the orbital Special features are: (1) Very deep excavation of the superior rims, indicating that the eyes were set very wide apart. border of the occiput, totally unlike that of either Archidiskodon Platelephas platycephalus, therefore, is totally different in or Elephas; (2) the relative breadth of the anterior nares, measur- profile as well as in superior and palatal aspects from either Archi- ing transversely 435 mm.; (3) the very deep excavation just below diskodon planifrons or Elephas indicus. Amer Mus. /98/8 D ' ' ' ' ' \ \ Typr CRANIUM OF PLATELEPHAS PLATYCEPHALUS Vig. 1219. Four aspects of the type cranium of Platelephas platycephalus (Amer. Mus. 19818), discovered by Barnum Brown in 1922 above the Pinjor horizon of the Upper Siwaliks, ‘below conglomerates.’ One-twelfth natural size. Compare Osborn, 1929.797, p. 22, fig. 21. A, Left lateral aspect, exhibiting orbits, auditory openings, and occipital condyles on practically the same level. Al, Palatal view, exhibiting slightly broadened premaxillary rostrum, maxillary prominence containing fractured M® with 16} ridge-plates, deeply in- dented posterior nares, and relatively low plane of occipital condyles. A2, Superior aspect, exhibiting moderately broadened premaxillaries, flattened fronto-occipital plane with deeply indented supra-occipital border without rugosity, and relatively long and narrow cranial proportions. A3, Posterior aspect, exhibiting moderately small, elevated occipital condyles when cranium is placed in exactly the same plane as (A). Compare with widely different cranial profile of Hypselephas hysudricus (Fig. 1205) and of E. indicus ceylanicus or E. indicus bengalensis (Fig. 1170), in which the occipital condyles are elevated above the level of the orbits. APPENDIX TO CHAPTER XX [In a recent Memoir (1936) by Pére Teilhard de Chardin and Dr. C. C. Young, ‘‘On the Mammalian Remains from the Archaeological Site of Anyang,’ Pal. Sinica, (C), XII, Fase. 1, pp. 1, 52, and 53) they have given their views regarding the existence in historical times of the wild Indian elephant in China, from which the following has been extracted. The reader is also referred to the map on page 1594 which gives the range of Elephas indicus, its varieties and subspecies, as far as known.—KEditor. | Close to the modern city of Changteho (N. Honan) are buried under the mud of the Chinese maritime plain the remains of Anyang, the latter city being the old capital of the Shang dynasty (Cirea 1400 B.C.—1100 B.C.). Remains of Elephant are not uncommon in Anyang (fragmentary young skulls and limb-bones). We only have at our disposal an incomplete lower molar of a young individual, probably a tooth number 3. Six lamellae are preserved, occupying alength of 68mm. Thickness of one lamella5mm.; breadth48mm. The lamellae are widely separated and set rather oblique- ly. As much as we can judge from this unsatisfactory specimen, we most probably are dealing here with an Elephas indicus, and not at all with #. primigenius, as supposed by Matsumoto (1916), basing on a centrum of vertebra and Hopwood (1934), using fragmentary molars. This reference of the Anyang Elephant to the Mammoth group was only possible when the deposits were still held as being of Pleistocene age. The presence of #. indicus in Honan once more raises the question as to how far north this form was living in China, and as to how long it lasted there, in historical times. Of course, paleontologically, we have no evidence to suggest as yet that the species ever existed in China proper. Its closest ally, HZ. namadicus, has never been found in the post-Loessic deposits. And no Elephas (but only Stegodon) occur in the Szechuan Pleistocene fissures. But historically it has been stated in some Chinese texts which were recovered in Anyang that ‘the king hunted and killed an Elephas(?)’. A critical discussion of those texts was given by Dr. Chang (1926). Dr. Chang concluded that there are no historical evidence proving the presence of wild Elephant and Rhinoceros in N. China since historical times. This view seems to be so far the most conservative and the best supported by facts. We shall therefore admit here that the Anyang Elephant, just like the Tapir of the same locality, represents a southern type imported to the city as a tribute, possibly for hunting purposes. . . It has to be noted in this connection that these exotic animals were brought and kept alive. Both in the case of the Elephant and the Tapir, our fossils belong to young individuals represented not only by some ivory but also by perfectly useless bones. 1362 CHAPTER XX] NOMENCLATURE OF THE PROBOSCIDEA HIsToRY OF NOMENCLATURE OF THE MASTODON AND THE MAMMOTH. CHRONOLOGIC LISTS OF SUPERFAMILIES (1921-1935), FAMILIES (1821-1937), SUBFAMILIES (1838-1937), GENERA AND SPECIES (1735-1939). 1. History of nomenclature of the Mastodon and the 4. List of subfamilies. Mammoth. 5. Genera and species of the Proboscidea in the 2. List of superfamilies. order named by the authors in the original 3. List of families. descriptions. Professor Osborn regarded nomenclature as “‘the tool rather than the master of paleontologic thought” and believed that ‘‘no technical principles should override the work of the early discoverers and naturalists.’”’ As repeatedly observed in the present Memoir (see especially Vol. I, Chap. I, pp. 5-13, also Vol. II, p. 1173), he thought it “impracticable in palzontology to apply all the principles of nomenclature established in zoology and botany, because the classification of the imperfectly known fossil forms is ever changing with our increasing knowledge of origins, adaptive radiations, and phyletic successions. Such mutability does not disturb the nomen- clature of living animals and plants, in which priority of adequate description, figure, and definition is the chief concern of systematists.”’ Doctor Hopwood in his Memoir of 1935 on the ‘Fossil Proboscidea from China” (p. 11) states that he has “attempted to make of the Rules [International Rules of Zoological Nomenclature] a useful servant, rather than to allow them to become a blind, unreasoning, master.’’ The extent of the difficulties encountered in Proboscidean paleontology may be judged from the accompanying nomenclature of the Mastodon and the Mammoth. This chapter was not completed by the author, but has been compiled by the editor from materials left by Professor Osborn, under the conditions mentioned in the “‘Publication Note” on page viii of Volume I. 1. HISTORY OF NOMENCLATURE OF THE MASTODON AND THE MAMMOTH 1788 The genus Mammonteus Osborn (1924.633, p. 2; Mammonteum Camper, 1788, p. 251) is of doubtful validity; in fact, it is possible that Professor Osborn would have abandoned it in his final revision of the present Volume and adopted Mammuthus Burnett, 1830, as the reader may conclude from the following account of the nomenclature of the northern or woolly mammoth and of the true mastodon (Mastodon americanus), the descriptive literature of which is so involved that the history of one is inseparable from that of the other. Consequently the subject is treated chronologically, dating from 1788 to the present time. Tue AMERICAN MaAstopon (MASTODON AMERICANUS) 1792 Kerr.—In 1792, p. 116, Robert Kerr proposed the name Hlephas americanus for tusks and grinders found at Big Bone Lick near the banks of the Ohio River, Boone County, Kentucky. 1797 BiumMENnBACH.—(1) In 1797 Blumenbach in his “Abbildungen Naturhistorische Gegenstiinde,’’ em- ployed the term Ohio-Incognitum, which first occurs on the back of the title-page following other specific names, 7.e., ‘11. Sima troglodytes ...19. Backenziahne von fossilen Ohio-Incognitum, und von den beiderlei Gattungen des Elephantengeschlechts.”” The name again appears opposite figure 194. 1363 1364 1799 1803 1806 1808 1814 1816 1817 1868 1869 1902 1904— 1905 1920 OSBORN: THE PROBOSCIDEA In a letter from C. W. Andrews to Professor Osborn (June 6, 1922) he quotes C. Davies Sherborn as follows: ‘‘The words ‘Ohio incognitum’ are not used in a generic or specific sense; they mean simply ‘the Ohio incognitum’ as is easily seen by the second page of the description, and are quite invalid.” (See Article 25 of the International Rules of Zoological Nomenclature.) (2) Subsequently (1799.1, p. 698) Blumenbach in his sixth edition of the ‘““‘Handbuch der Natur- geschichte” assigned the name Mammut ohioticum to the same animal. In the French edition of the “Handbuch” (translated in 1803 by Soulange Artaud under the supervision of Blumenbach) mammout Ohioticum appears on page 408, also Ohio-Incognitum occurs under the figure 19A, first used in the 1797 “Abbildungen”’ of Blumenbach (see Vol. I of the present Memoir, caption to fig. 113, where it is stated by Professor Osborn that it is “technically a type figure’’). Dr. Edwin H. Colbert in a note to the editor (1937) makes the point that as Kerr, the author of Elephas {= Mastodon] americanus, did not include a figure in his description, we cannot be sure that figure 113 just referred to is a figure of the type. Cuvirr.—It was 1806 before Cuvier applied ‘““Le Grand Mastodonte”’ (1806.2) and “Mastodonte de Ohio (1806.3) to the American Mastodon, which in 1817 (see below) he redeseribed under the specific designation of Mastodon giganteum. FiscHER DE WALDHEIM.—The following generic names were applied to the Mastodon by Fischer de Waldheim: Harpagmothervum. Mastotherium. OxEeN.—The final form Mastodon was published by Oken in 1816, p. 789, citing Cuvier’s five classic species (‘‘Lehrbuch der Naturgeschichte,” Theil III, p. 789). Cuvier.—Although anticipated by Oken by one year (see above, 1816) in the use of the term Mastodon (see 1817, ‘“‘Le Régne Animal,” I, p. 233, for description of Mastodon giganteum), Cuvier has been regarded as the author of this genus, since he first used the French form Mastodonte in his description of 1806. Lreipy.—-Leidy (1868, p. 175) was the first to use the name Mastodon americanus, embracing the Mastodon ohioticus or M. gigantewm of authors. Leidy in 1869 (p. 392) lists Mamontewm Camper among the synonyms of Mastodon americanus. Hay.—Mammut Blumenbach was revived by Hay (1902, p. 707) and adopted by many subsequent authors. In his last work (1930, p. 623) Hay names the new subfamily Mammutine to include the Masto- donts, although he places it under the Elephantide without recognition of the family Mastodontide. TrovEssarT.—The term Mammut Blumenbach was regarded by Trouessart (p. 600) as a misno- mer: “Le nom barbare de ‘Mammut,’ basé par Blumenbach sur une erreur grossiére (l’identité du Masto- donte de Vv Ohio avec |’ Elephas primigenius), n’a aucun droit, malgré sa priorité, a étre substitué 4 celui de ‘Mastodon,’ genre bien caractérisé par Cuvier.” ALLEN (excerpts from letter of Dr. Joel A. Allen to Professor Osborn, dated November 16, 1920): In your letter (October 22, 1920) you ask whether I consider Blumenbach’s name Mammut entitled to recognition. 1 certainly do, although with regret that such name was . . .ever proposed as a generic designation. Neither its barba- rous form nor the fact of his misconception that it was the mammoth renders it invalid . . . . All modern codes or rules of nomenclature declare that such names are not to be rejected on the ground that they are nonclassical, and that incor- rectly formed classical names are not subject to emendation. 1935 1936 1799 1799 1830 1837 1845 1848 1850 1888 NOMENCLATURE OF THE PROBOSCIDEA 1365 Horwoop.— Doctor Hopwood substitutes Mastodon for Mammut, as stated in his ‘‘Fossil Proboscidea from China,” 1935, p. 42: “To retain Cuvier’s generic name instead of Blumenbach’s, is to follow con- venience rather than rule. The generic term Mammut and the vernacular Mammoth are so much alike that one or other should be suppressed in the interests of clarity. It is very much easier to discourage the use of an unfamiliar name than a well-known popular one, so that, whilst acknowledging Blumenbach’s prior claim, I have given the preference to Cuvier.” OsBORN’S CONCLUSIONS IN VOLUME I, pp. 6 AND 7, OF THE PRESENT Memorr.—For the reason that Mammut (signifying ‘earth-burrower’ and suggesting the mammoth) was employed to describe the Masto- don; that if a vernacular name were to be accepted Ohio-Incognitum had priority, and lastly that inasmuch as Cuvier’s ‘““Mastodonte” (Mastodon) had been used throughout the literature of the past century, Professor Osborn committed himself to the genus Mastodon in the following citation from his Memoir (Vol. I, p. 6): ‘Consequently to rob Cuvier of his clear conception of grinding tooth structure, which he termed Mastodonte, and to substitute the barbaric term Mammut, signifying ‘earth-burrower,’ would be gross injustice to the founder of vertebrate paleontology.” Tue NorTHERN OR WOOLLY Mammotu (MAMMONTEUS PRIMIGENIUS) BLUMENBACH.—In the same article in which Blumenbach described Mammut ohioticum, namely, in the ““Handbuch der Naturgeschichte,” sixth edition (1799.1, p. 698), he also described Elephas primigenius (p. 697), stating that there are abundant remains in Germany, but he mistakenly gives as an example a skeletal specimen from Burg-Tonna (now in the Gotha Museum), which proves to belong to Hlephas | = Hesperoloxodon| antiquus Falconer and not to the northern Mammoth. Cuvier.—In the autumn of the same year that Blumenbach described Elephas primigenius (1799), Cuvier assigned the name Elephas mammonteus to the mammoth; subsequently, however, he adopted Blumenbach’s name EF. primigenius. The following is a list of the names assigned to the Mammoth (Hlephas primigenius) : Mammuthus Burnett, Quart. Journ. Sci. Lit. & Art, XXVIII, p. 352. Dicyclotherium K. Geoffroy St. Hilaire, Compt. Rend. Acad. Sci., Paris, pp. 119, 120, fig. 1. Mammontheum de Blainville, 1839-1864, ‘‘Ostéographie,” p. 237. Cheirolites von Meyer, (in Bronn’s “Handbuch einer Gesch. d. Natur.,”’ II, Index Pal., p. 286). Synodontherium Costa, Palaeont. del Regno di Napoli, Pt. I, pp. 271-275, Tav. m1, figs. 1-4. Polydiskodon Pohlig, Nova Acta Leop. Carol. Deutsch Akad., LITT, pp. 138, 252. With the removal by Professor Osborn of Mammut Blumenbach from the valid genera of the Probo- scidea, and the adoption of Mastodon Cuvier, there arose the question of the correct genus to which Hlephas primigenius, the northern Mammoth, should be referred. Professor Osborn chose Camper’s name Mam- monteum, 1788, which, like Mammut, had reference to a specimen of the American Mastodon and not to 1366 OSBORN: THE PROBOSCIDEA the Mammoth. The revival of Mammonteum (Mammonteus) by Professor Osborn occurs in his article “Parelephas in Relation to Phyla and Genera of the Family Elephantide’”’ (Osborn, 1924.633, p. 2) as follows: 1924 By many authors all the generic phyla of the mammoths are still referred to the genus Hlephas. Such reference, from our present knowledge, is inconsistent with the fact that none of the mammoths contains the ancestral characters of Elephas. (1) We thus revive the ill-defined name Mammonteus Camper for the Elephas primigenius phylum, which Depéret and Mayet have traced back to the Upper Pliocene Elephas primigenius astensis of northern Italy and into the Lower Pleistocene 2. meridionalis cromerensis of the Forest Bed of Cromer. Here this line terminates in the typical FE. primigenius of western Europe and onward into the progressive new subspecies Mammonteus primigenius com- pressus of North America. 1934 Preparatory to the final revision of the present Volume of the Memoir, Professor Osborn, in 1934, called the attention of Dr. A. Tindell Hopwood to the Mammuthus of Burnett (1830) as having priority over Dicyclotherium Geoffroy (1837). Doctor Hopwood adopted the name Mammuthus in his Memoir on 1935 ‘The Fossil Proboscidea from China,” 1935, p. 98, from which we cite: Remarks.—With the progress of research on the Proboscidea, it has become ever clearer that the Mammoth is distinct from the Indian Elephant, which typifies the genus Elephas. Professor Osborn has sought to give expression to this result by reviving an alleged genus of Peter Camper’s and emending the name. If I do not adopt the same name, it is because Camper was not referring to the Mammoth, and because he was using the word ‘Mamonteum’ as a vernacular. In the section ‘De ossibus Mamonteis’ Camper (1788, p. 259) uses ‘Mamonteum’ in an adjectival sense throughout. Not only so, but he also makes it quite clear that he is referring to an animal from America, and Pallas contributes a foot-note on p. 261 in which he explains that whereas in Russia the term ‘Mammontean bones’ is common- ly applied to the bones of Elephants found in superficial deposits, it has suited Camper to apply the name to bones found in America. With so much evidence as to Camper’s meaning, it is not possible to argue that the phrases on p. 251 of the same work refer to the Mammoth as distinct from the Elephant. Several names have been applied to the Mammoth, but the first that is valid, in so far as it possesses a genotype, appears to be Mammuthus Burnett, 1830. Under ‘Elephantide, Elephant-kind’ he groups the following,— Genera. Species. lephas. Indicus. Indian Elephant. Africanus. African. Mammuthus. Borealis. Vossil Mammoth. Meridionalis. Mastodon. Giganteum. Gigantic. Angustidens. Lesser. From this table it is clear that he used the word Mammoth in the sense generally accepted today, and did not apply it to the American Mastodon after the manner of the majority of English and American authors at the beginning of the nineteenth century, (cf. Cuvier, Ossemens foss., Ed. 1, Vol. II, Art. Sur le grand mastodonte), and that his Mammuthus borealis is the equivalent of Hlephas primigenius Blum. Doctor Hopwood then described a molar from Mongolia (regd. M10941) and fragment of another from Honan (regd. M14102) which he refers to Mammuthus primigenius (Blumenbach). The following excerpts from the correspondence between Doctor Hopwood and Professor Osborn are self explanatory: September 10, 1935. Professor Osborn wrote as follows: ‘Among the Elephantide I regret that you substituted Mammuthus Burnett for Mammonteus Camper... If we stood on technicalities, the specific name Hlephas primigenius is based on a species indubitably belonging to Hesperoloxodon antiquus ... Technical methods would compel us to alter the entire nomenclature of the fossil Proboscidea. As in religion the spirit rather than the letter is important.” NOMENCLATURE OF THE PROBOSCIDEA 1367 October 19, 1935. Doctor Hopwood replied to Professor Osborn: “With regard to the substitution of Mammuthus for Mammonteus, I should not have done this had it been possible to show that Camper was referring to a European fossil, but the evidence that the reference is to the American Mastodon is so strong that I had no other alternative. The case of EL. primigenius and H. antiquus is hardly parallel; Blumenbach was naming a new species, and later authors have misinterpreted him. The error is theirs, not Blumenbach’s.”’ October 25, 1935. Professor Osborn replied at once as follows: ‘‘Your point that Camper was referring to the American Mastodon when he used the name Mammonteus is a very strong one and I shallimmediately refresh my memory on this point, although I fear it may be difficult to correct it now.” October 29, 1935. Five days later he added the following postscript to the foregoing letter. “I am looking up Camper’s use of the name Mammonteus. At the time, it was believed that there was only one extinct elephant, the true Mammoth in the Old World and the Mastodon in the New. First allusions to the American Mastodon speak of it as the Mammoth. It is interesting to note that Kerr gave the name Elephas americanus in 1792, several years before Blumenbach gave the name Hlephas primigenius in 1799. I already have the complete bibliography of names applied to the European Mammoth and shall give them in my Monograph. Perhaps the first properly defined generic name was Dicyclotherium. Palmer, ‘Index Generum Animalium,’ 1904, p. 397, gives Mammont as applied to Mastodon ameri- canus, Mammuthus Burnett. Burnett simply gives the name and the species Mammuthus borealis, a species not listed in Sherborn’s Index nor anywhere defined. I doubt if Mammont, Mammuthus, or Mammut can be adopted on any rules of nomenclature, since a genus must rest on a type species. At present I prefer the Latin name Mammonteum Camper, which rests, according to Camper’s description, both on the specimens found in Siberia and in North America, but I shall keep an open mind and settle this question in Volume IT.” Professor Osborn’s death occurred November 6, 1935. 1937 Concuiusions.—A recent note from Dr. Edwin H. Colbert to the editor reads as follows: 1830 Mammuthus Burnett Type designated as Mammuthus borealis, which is a synonym of Elephas primigenius Blumenbach. Therefore the designation becomes Mammuthus primigenius (Blumenbach). Hopwood’s arguments (1935) are valid. 1937 Dr. William Berryman Scott in his recent (1937) revision of ‘‘A History of Land Mammals in the Western Hemisphere,” adopts Mammuthus Burnett. Owing to the fact that Mammonteus has appeared so frequently in the literature of the past decade and has been used throughout the entire text of the present Memoir, it has been decided to retain the name herein, with the explanatory notes above, despite the evidence in favor of its abolishment.—Kditor. | 2. LIST OF SUPERFAMILIES YEAR NAME AUTHOR BIBLIOGRAPHIC REFERENCE REFERENCE IN PRESENT MeEmMorIrR 1921 M(@RITHERIOIDEA Osborn, Amer. Mus. Novitates, No. 1, p. 2. See M@(:RITHERIOIDEA also Volume I, p. 24, of the present Memoir. Includes the family Meeritheriide of Andrews, 1906. 1921 DINOTHERIOIDEA Osborn, loc. cit. DEINOTHERIOIDEA Includes the family Curtognati [=Curtognathide, this Memoir] of Kaup, 1833. 1921 MASTODONTOIDEA Osborn, loc. cit. MASTODONTOIDEA Includes the families Mastodontide of Girard, 1852, and the Bunomastodontide of Osborn, 1921. 1921 KLEPHANTOIDEA Osborn, loc. cit. ELEPHANTOIDEA Includes the family Elephantide Gray, 1821. 1935 STEGODONTOIDEA Osborn, Proc. Nat. Acad. Sci., X XI, No. 6, p. 408, STEGODONTOIDEA also fig. 2; Vol. I, pp. 22, 25, and Vol. II, p. 807, of the present Memoir. Includes the family Stegodontide2 Young-Hopwood, 1935. 1368 YEAR NAME AUTHOR 1821 ELEPHANTIDAS Gray 1821 1833 1842 1845 1850 1906 1906 1921 1927 1929 1929 MASTODONAD® Gray CURTOGNATI Kaup ELEPHASIDE A Lesson DINOTHERID A! Bonaparte DINOTHERIID Bonaparte MASTODONTIDAE Girard M@ERITHERIIDAS Andrews PALMOMASTODONTIDAS Andrews BUNOMASTODON TID AE Osborn AMEBELODONTIDA® Barbour MAMMUTID® Cabrera GOMPHOTHERIIDAS Cabrera OSBORN: THE PROBOSCIDEA 3. LIST OF FAMILIES BIBLIOGRAPHIC REFERENCE London Medical Repository, XV, No. 88, p. 305. 1838, Bonaparte, Nuoy. Ann. Sci. Nat. Bologna, Anno I, Tom. II, p. 112 1850, Bonaparte, ‘“‘Conspectus Systematis Mastozoologia. Mam- malia.” 1852, Girard, Proc. Amer. Assoc. Ady. Sei., for the year 1851, pp. 326, 328. 1891, Zittel, “Handbuch der Palacontologie,” p. 458. 1910, Osborn, “The Age of Mammals,” p. 558. London Medical Repository, XV, No. 88, p. 306. Neues Jahrb. Min., p. 516. Dinotheriide Bonaparte, 1850, “Nouveau Tableau du Régne Animal,” p. 156. Embraces one Mastodon, ‘‘Elephas arvernensis” Croizet and Jobert. “Catalogo metodico dei Mammiferi Kuropei,” p. 4 (fide Palmer, 1904, p. 738). “Conspectus Systematis Mastozoologiw. Mammalia.” 1918, Osborn, Bull. Geol. Soc. Amer., X XIX, p. 134. In Vol. I, pp. 26, 82, 83, this Memoir, changed to Cur- tognathide, based on family Curtognati Kaup, 1833. Proce. Amer. Assoc. Ady. Sci., for the year 1851, pp. 326, 328. 1918, Osborn, Bull. Geol. Soc. Amer., X XIX, p. 134 (in part). “A Descriptive Catalogue of the Tertiary Vertebrata of the Fayim, Egypt,” p. 99. Op. cit., p. 180. Regarded by the present author as invalid, because founded on the generic characters of Phiomia rather than Palzomastodon. Amer. Mus. Novitates, No. 1, pp. 2, 4. Replaced the subfamily Bunomastodontine Osborn, 1918, pp. 134-136 (see Vol. I, p. 27, this Memoir). Bunomastodonti- dx inadmissible, however, under the rules of zoological nomen- clature (see Scott, 1937, p. 287). See also Trilophodontide Simpson, 1931, p. 1869, and Gomphotheriidie Cabrera, 1929, below, this list. Neb. State Mus., Bull. 18, I, p. 131. Subsequently changed by Barbour (1929.2, p. 139) to Amebelodontine. Rev. Mus. La Plata, XXXII, p. 74. Tbid., p. 75. REFERENCE IN PRESENT MeEmMoIR ELEPHANTIDA BLEPHANTIDA® ELEPHANTIDA ELEPHANTID/ ELEPHANTIDA® (in part), also Mastodontide, Buno- mastodontide, Serridenti- de, Humboldtide, Stego- dontide ELEPHANTID (in part), also Curtognathide, Masto- dontide, Bunomastodonti- de, Stegodontide MASTODONTIDA CURTOGNATHIDAs ELEPHANTIDA CURTOGNATHIDAS CURTOGNATHIDA# CURTOGNATHIDAs MASTODONTIDAD MASTODONTIDAS M@RITHERITD. BUNOMASTODON TIDE BUNOMASTODONTIDAL BUNOMASTODONTIDA® (subfam. AMEBELO- DONTIN@) MASTODONTIDA® BUNOMASTODONTIDAL (in part), also Serridentide, Humboldtide NOMENCLATURE OF THE PROBOSCIDEA 1369 1931 TRILOPHODONTIDAi Simpson Bull. Amer. Mus. Nat. Hist., LIX, Art. V, p. 281. BUNOMASTODONTID A Substituted for inadmissible name Bunomastodontide. Adopted by Scott, 1937, pp. 267, 280, 287. [Doctor Simpson now prefers Gomphotheriidze Cabrera, 1929 (see p. 1525 below).— Editor. | 1935 HUMBOLDTID<%® Osborn Proc. Nat. Acad. Sci., X XI, No. 6, fig. 2 (name only). HUMBOLDTID 2 1936, Osborn, this Memoir, Vol. I, pp. 575, 722. Without a type genus. Stegomastodontide substituted by Scott, 1937 (see below, this list). 1935 SERRIDENTID<® Osborn Proc. Nat. Acad. Sci., X XI, No. 6, fig. 2 (name only). SERRIDENTIDAi 1936, Osborn, this Memoir, Vol. I, p. 729. 1935 STEGODONTIDA Young Pal. Sinica, Ser. C, IX, Fasc. 2, p. 5. STEGODONTID 2 1935, Hopwood, Pal. Sinica, Ser. C, LX, Fase. 3, p. 71. Included both Stegolophodon and Stegodon. Restricted in present Memoir to Stegodon. 1935 DIBUNODONTIDA® Hopwood Pal. Sinica, Ser. C, IX, Fase. 3, pp. 11, 55. BUNOMASTODONTIDA8 See especially Hay, 1925, Journ. Wash. Acad. Sci., XV, p. (Gin part, 7. e. only the 382. Brevirostrine of Osborn) 1937 STEGOMASTODONTID< Scott “A History of Land Mammals in the Western Hemi- HUMBOLDTIDAi sphere,” pp. 267, 281, 294. Substituted for Humboldtide, which is without a type genus. 4. LIST OF SUBFAMILIES EFERENCE IN ESE YEAR NAME AUTHOR BIBLIOGRAPHIC REFERENCE REFERENC PRESENT MEMOIR 1838 ELEPHANTINA Bonaparte Nuoy. Ann. Sci. Nat. Bologna, Anno I, Tom. ELEPHANTIN® II, p. 112; 1850, “Conspectus Systematis Mastozoologia. Mammalia.” 1841 DrnorHEeRINA Bonaparte Trans. Linn. Soc. London, XVIII, p. 253. DEINOTHERIIN © 1850 DrnorHERIINA Bonaparte “Conspectus Systematis Mastozoologie. Mam- DEINOTHERIIN® malia.”’ 1869 MasropontiIna Brandt Mém. Acad. Imp. Sci. St. Pétersb., (VII), MASTODONTIN XIV, No. 1, p. 35. 1906 MarirHeriNni Winge “Jordfundne og nulevende Hovdyr (Ungulata) MarITHERIIN.2® fra Lagoa Santa, Minas Geraes, Brasilien,” p. 172. 1906 DrnorHEritni Winge Loc. cit. DEINOTHERIIN® 1906 ELmPHANTINI Winge Loe. cit. KLEPHANTIN 2 1910 DinorHERIINa Osborn “The Age of Mammals,” p. 558. DEINOTHERIIN © 1910 MasropontTin® Osborn Loc. cit. MASTODONTIN 2 1910 ELEPHANtTiIna Osborn Loc. cit.; 1918, Bull. Geol. Soc. Amer., XXTX, ELEPHANTINAD p. 135. 1918 BuNoMASTODONTIN® Osborn Bull. Geol. Soe. Amer., XXIX, pp. 134-136. Replaced by BUNOMASTODONTID® (in part) 1918 SreGopontinz Osborn Ibid., pp. 135, 136. STEGODONTIN © 1918 LoxopontiIna Osborn Ibid., pp. 135, 136. LOXODONTIN © 1370 YEAR 1918 1918 1918 1918 1921 1921 1921 1923 1923 1927 1927 1928 1929 1929 1929 1929 1929 NAME AUTHOR EUELEPHANTIN2® Osborn LONGIROSTRIN® Osborn RHYNCHOROSTRIN2& Osborn BREVIROSTRIN® Osborn MAMMONTIN Osborn SERRIDENTIN® Osborn NovrorostRInz Osborn Ma@rITHERIINAE Winge-Osborn ZYGOLOPHODONTIN Osborn ARCHIDISKODONTIN® Dietrich PARELEPHANTIN Dietrich PLATYBELODONTIN Borissiak AMEBELODONTIN2 Barbour RHYNCHOTHERIIN® Cabrera GOMPHOTHERIIN Cabrera CUVIERONIIN2 Cabrera ANANCIN-® Cabrera OSBORN: THE PROBOSCIDEA BIBLIOGRAPHIC REFERENCE Ibid., p. 136. Invalid because the genus Huelephas is invalid (see Chap. XIX, p. 1177, of present Memoir). Tbid., p. 136. Without type genus. See Trilophodon- tine Scott, 1937, p. 1371 below, this list. Ibid., p. 136. See Rhynchotheriine Cabrera, 1929, on this page below; Scott, 1987, pp. 267, 280, 292, also adopts the form Rhynchotheriine. Ibid., p. 136. See Pentalophodontine Scott, 1937, p. 1371 below, this list. Amer. Mus. Novitates, No. 1, pp. 1, 14. Should Mammonteus prove to be invalid, this would leave the subfamily Mammontine without a type genus. Bull. Geol. Soe. Amer., XX XII, p. 3380. Tbid., p. 330. Without type genus. See Cordillerionine Scott, 1937, p. 1371 below, this list. Amer. Mus. Novitates, No. 99, p. 1. Loc. cit. Neues Jahrb. Min., I, Abth. B (Referate), p. 313. Loe. cit. Ann. Soe. Paléont. Russie, VII, p. 119. Neb. State Mus., Bull. 16, I, p. 139. Rev. Mus. La Plata, XXXII, p. 75. Loc. cit. Ibid., p. 76. Ibid., p. 76. REFERENCE IN PRESENT MeEMOIR INVALID LONGIROSTRIN © RHYNCHOROSTRIN BREVIROSTRIN MAMMONTIN® SERRIDENTIN NOTOROSTRIN #® MaRrITHERIIN ZYGOLCPHODONTIN | MAMMONTIN® (in part)| [MAMMONTIN® (in part)| PLATYBELODONTIN 22 AMEBELODONTIN® RHYNCHOROSTRIN 28 LONGIROSTRIN (in part) TETRALOPHODONTIN® SERRIDENTIN NoTOROSTRIN#® (in part) NOTIOMASTODONTIN® Nororostrin&, Hum- BOLDTIN®, and BREVIROSTRIN © (in part) NOMENCLATURE OF THE PROBOSCIDEA 1371 YEAR NAME AUTHOR BIBLIOGRAPHIC REFERENCE REFERENCE IN PRESENT Memoir 1930 MammutTin® Hay “Fossil Vertebrata of North America,” II, p. [MaAsropoNnTo1pEA of 623. Osborn| Name without definition but indicates the inclusion of the subfamilies of the Mastodon- toidea of Osborn. 1932 TrTRALOPHODONTIN#® van der Maarel ‘Contribution to the Knowledge of the Fossil TETRALOPHODONTIN 2 Mammalian Fauna of Java,” p. 108. 1934 HumBoLpTIN® Osborn Proc. Amer. Phil. Soc., LX XIV, No. 4, p. 277, | HUMBOLDTIN2® fig. 3, and p. 283, also Vol. I, p. 575, of present Memoir. 1935 GNATHALODONTIN# Barbour and Neb. State Mus., Bull. 42, I, p. 395. GNATHABELODONTIN 28 Sternberg [emend. Osborn Gnatha- belodontinz| 1936 PALMOMASTODONTIN® Osborn This Memoir, Vol. I, p. 691. PALMHOMASTODONTIN 1936 STEGOLOPHODONTIN® Osborn Op. cit., p. 700. STEGOLOPHODONTIN 1936 NorTIoMASTODONTIN® Osborn Op. cit., p. 730. NOTIOMASTODONTIN 4 1937 TRILOPHODONTIN® Scott “A History of Land Mammals in the Western LONGIROSTRIN Hemisphere,’ pp. 267, 280. Substituted for Longirostrine which is without a type genus. 1937 CoRDILLERIONIN Scott Loc. cit. NOTOROSTRIN 22 Substituted for Notorostrine which is without a type genus. 1937 PENTALOPHODONTIN® Scott Op. cit., pp. 267, 292. BREVIROSTRIN © Founded on Pentalophodon Falconer, 1857, 1865. Compare Anancine Cabrera, 1929, above, this list. 1937 STEGOMASTODONTIN® Scott Op. cit., pp. 267, 281. HUMBOLDTIN © Replaces Humboldtinz which is without a type genus. GENERA AND SPECIES OF THE PROBOSCIDEA IN THE ORDER NAMED BY THE AUTHORS IN THE ORIGINAL DESCRIPTIONS? The following Tables present a revised chronologic list (1735-1939) of generic and specific names as originally and subsequently spelled by their authors, which rest upon the authority of the authors mentioned, or cited (e.g., fide) in cases where the original references were not available to the present author. cr LIST OF GENERA Generic Reference Year Genus Author Bibliographic Reference Genotypic Species in Present Memoir 1735-1758 Hlephas Linneus, ‘Systema Nature,” 1st to 9th editions. Elephas indicus Elephas Linnzus [In the 10th edition of the “Systema Nature’’ (known as the Decima Reformata), 1758, p. 33, Linnzeus substituted the name Elephas maximus Linn. for that of H. indicus Linn., which, with all other names in that edition, was officially adopted by the Fifth In- ternational Congress of Zoology, held in Berlin in 1901.—Kditor.} For first use of Hlephas, see John Ray, 1693, p. 131 [p. 123 of authors]. The author of the present Memoir refused to recognize privately printed publications (unless on sale, bearing the name of the publisher) as a source of systematic names; consequently such references have been omitted from the present list.—Editor. | 1372 OSBORN: THE PROBOSCIDEA Generic Reference Year Genus Author Bibliographic Reference Genotypic Species in Present Memoir 1788 Mammonteum Camper, Nova Acta Acad. Sci. Imp. Petropol., II, for [Refers to an animal Mammonteus! the year 1784, p. 251; Mammonteus Osborn, 1924, from America (ef. Hop- Amer. Mus. Novitates, No. 152, p. 2. wood, 1935, p. 98).] Camper’s designation of Mammonteum is cited above. Hay (1902, p. 708) and J. A. Allen (letter, Nov. 23, 1920) do not regard it as valid. Allen writes: “the names of the fossil mammals he [Camper] discusses are vernacular names rendered into Latin. They have no nomenclatural significance.” Osborn nevertheless adopts Mammonteum Camper, 1788, in preference to Dicyclotherium Geoffroy, 1837, as a distinct generic name assigned to E. primigenius. While Allen in the above letter objects to Mammonteum as a vernacular name rendered into Latin, in a letter (Nov. 15, 1920) on the subject of barbarous names as generic names, he nevertheless agrees with Palmer (‘‘Index Generum Mammalium,”’ 1904, pp. 45, 46) and says: ‘‘All modern codes of [or] rules of nomenclature declare that such names are not to be rejected on the ground that they are nonclassical, and that incorrectly formed classical names are not subject to emendation.”’ See also Allen’s note on the word “Mammut.” 1795 Elephantus Cuvier and Geoffroy, Revue Enecyclopédique (or Mag. En- Genotype not given in Hlephas cyclopédique), II, (6), p. 189. 1795, but in 1801 (Cuvier and Lacépéde) indicus is mentioned 1797 Ohio Blumenbach, Abbild. naturhist. Gegens., Heft 2, No. 19, fig. A, Ohdo-incognitum Mastodon and back of title-page. Blumenbach Not a valid generic description. See introduction to this chapter. 1799 Mammut Blumenbach, Handb. d. Natur., 6th edition, p. 698. Mammut ohioticum Mastodon Blumenbach A vernacular name preoccupied by Ohio for E. primigenius. Allen, however, writes (letter, Nov. 15, 1920): ‘In your letter you ask whether I consider Blumenbach’s name Mammut entitled to recognition. I certainly do, although with regret that such name was not [sic] ever proposed as a generic designation. Neither its barbarous form nor the fact of his misconception that it was the mammoth renders it invalid.” Murray, 1908, VI, Pt. 2, p.98: “MAMMOTH. ... / Also 8 mammuth, mamant, maman, mamont, Mammon, mammot (mammoht), 8-2 mammouth. [a. Russian, ... Mammor, mammot, whence mammotovot kost, mammoth’s bones (Ludolf, Gram. Russ. 1696, p. 92); now MamMauTb, mamant. Hence also F. mammouth,...mamant,...mammont. The word is of obscure origin; the alleged Tartar word mama ‘earth’ (usually cited as the ety- mon) is not known to exist.]”’ 1805 [1806] Mastodonte G. Cuvier Ann. Mus. d’Hist. Nat., VIII, pp. 270, 272, 298. Le Grand Mastodonte Mastodon Cuvier By courtesy to Cuvier restricted Falconer and Cautley, 1846, letterpress, p. 18: “But in his second extended and to M. americanus elaborate memoir, published in 1805 [1806], which formed the groundwork of what he has written on the subject in the ‘Ossemens Fossiles,’ Cuvier separated the Elephants with mammillated molars from the ordinary forms with lamelliform molars, and united the former into a genus which he designated Mastodon, taking the North American species, under the name of M. giganteum, as the type [Footnote: ‘Annales du Muséum d’ Histoire Naturelle, tom. viil. “Sur le grand Mastodonte.”’ ’].”’ See also de Blainville (1834-69, p. 245); and Leidy (1869, p. 393): ‘‘The earliest date at which I have been able to find the name of Mastodon systematically expressed, is in the work here quoted (Cuvier, 1817, p. 233]. Previously, Cuvier appears only to have used the gallicized term of Mastodonte. Bronn, in the 3d edition of the Lethaea Geognostica, page 820, credits Mastodon to Cuvier as early as 1805, but does not give the reference.” [See Oken, 1916, this chapter, p. 1364 above.] 1808 Harpagmotherium Fischer de Waldheim Prog. d’Invit. Séance, Pub. Harpagmotherium cana- Mastodon Soc. Imp. Nat. Moscou, September, pp. 19, 20 (fide dense Fischer de Palmer, 1904, p. 311). Waldheim Compare Sherborn, ‘‘Index Animalium,” who gives on page 1022 (Sect. 2, Pt. 5, 1924) Harpagonotherium canadense, and on page 2915 (Sect. 2, Pt. 12, 1927) Harpagmotherium. '|Mammuthus Burnett, 1830 (see below, this list) selected by Dr. A. Tindell Hopwood, 1935, p. 27. For complete historical account of the names Mam- monteus, Mammut, Mastodonte, Mastodon, up to the year 1935, see pp. 1363-1367 above.—Editor.] Year 1814 1816 1817 1817 1825-1827 1829 1830 1830 1837 1837 NOMENCLATURE OF THE PROBOSCIDEA Bibliographic Reference Genotypic Species “Zoognosia,” III, p. 337. Genus Author Mastotheriwm Fischer de Waldheim Species cited: M. megalodon (Cuv.) leptodon (Cuv.) microdon (Cuv.) hyodon (Cuv.) humboldtii (Cuv.) Mastodon Oken Lehrbuch Natur., Theil III, Abth. 2, p. 789. First use of the term Mastodon for Cuvier’s Mastodonte, citing Cuvier’s five classic species. “Le Régne Animal,” I, pp. 232, 233. Mastodon gigantewm Cuvier, Mastodon angustidens Cuvier Mastodon G. Cuvier Mastodontum de Blainville Nouv. Dict. Hist. Nat., IX, p. 276. Loxodonte F. Cuvier “Hist. Nat. Mamm.,” III, Livr. LI, LII, with [=Loxodonta, 2 pp. text, 1827, 1828, Zool. Journ., London, III, p. 1827, 1828] 140 (unsigned review). Geoffroy Saint-Hilaire, Etienne, et Cuvier, Frédéric, 1824-1829 [1825], (Loxodonte), p. 2: “Je proposerai pour nom générique de cette espéce, le mot de Loxodonte, qui peut rappeler le caractére de ses dents, les losanges qu’on apergoit sur leur coupe.”’ A review (unsigned) of this work appeared in the Zool. Journ., London, 1827, 1828, III, p. 140, noticing the ‘dismemberment of the genus Elephas, for the purpose of establish- ing a new one under the name of Loxodonta. . . . For the Elephant of Asia he [Cuvier] retains the original generic name Hlephas. The surfaces of its molar teeth present fascie of enamel irregularly festooned; while in those of the African Elephant, the type of the new genus Lozodonta, the enamel is disposed in lozenges. In addition to this striking distinction derived from the dentary system, M. F. Cuvier also enumerates the other characters which have hitherto been regarded as specific. The smaller, more elongated, and less irregular head of the African animal when compared with the Asiatic; the rounded forehead of the former, strongly contrasted with the deep depression in the middle of that of the latter; the ear of the former also twice the extent, while the tail is only half the length, &c.”’ Isis, [XXII], Heft IV, p. 401, Taf. 1. Elephas africanus Blumenbach Deinotherium gi- Deinotherium Kaup ganteum Kaup Mammuthus borealis Burnett Mammuthus Burnett Quart. Journ. Sci., London, July-December, 1829, p. 352. {See introduction to the present chapter, pp. 1366 and 1367.] Tetracaulodon Godman ‘Trans. Amer. Phil. Soc., N.8., II, p. 484. Tetracaulodon masto- dontoidewm Godman Gomphother‘um Burmeister Handb. d. Natur., p. 795. Not cited. Burmeister, 1837, p. 795: ‘Mastodon. Wie Elephas, aber die Backzihne mit 2 Reihen kegelf6rmiger Hécker.—Von mehreren untergegangenen Arten findet man Knochen in Nordamerika, besonders am Ohio, daher Ohiothier. Stossziihne in beiden Kiefern besass die gleichfalls untergegangene Gatt. GOMPHOTHERIUM.” Osborn, 1922: The name Gomphotherium is invalid: (1) Because no genotypic species except the Ohio mastodon is mentioned; (2) because several different genera of masto- donts display the same character, namely, ‘“‘Stosszihne in beiden Kiefern,” e. g., T'etracaulo- don Godman, a four-tusked true Mastodon americanus. Matthew (unpublished manu- script of 1918) states: “Apparently the author [Burmeister] had primarily in mind certain specimens of the American mastodon which retain the lower tusks; but for these the name Tetracaulodon Godman had been proposed in 1830.” (3) Confusion as to mastodons with four tusks is demonstrated further by the fact that Kaup first used Godman’s term in describing his Eppelsheim species, viz., T'etracaulodon longirostre. (4) It is consequently clear that we cannot be certain what animal Burmeister had in mind. (See note under Gamphotherium Gloger below.) Hay adopted the name in 1917 (see Gomphotherium gratum and G. elegans, p. 1405 below). Dicyclotheriuwm E. Geoffroy Saint-Hilaire Compt. Rend. Acad. Sci., Hlephas primigenius Paris, IV, No. 4, pp. 119, 120, fig. 1. Blumenbach This name assigned in reference to the fact that the genus was supposed to pass through two cycles of time. Assigned by Geoffroy in an important communication on the influence of climate on evolution. 1373 Generic Reference in Present Memoir Mastodon americanus Trilophodon angusti- dens Turicius tapiroides Cordillerion andium Cuvieronius humboldtii Mastodon Cuv. Mastodon [Mastodon Cuv.| Loxodonta Deinotherium Mammonteus Mastodon Mastodon Mammonteus 1374 Year 1840? 1841 1841 1841 L846 OSBORN: THE PROBOSCIDEA Genus Author Bibliographic Reference Genotypic Species Missourium Koch ‘‘Fossil Remains,’”’ pp. 1, 2. Missourium Kochi Koch First described as Koch’s Missourian in 1839 (American Journal of Science, XX XVII, pp. 191, 192). In 1841 Koch renamed it the Missouri Leviathan, or Leviathan missouri, and in 1843 he described it as the Missourium Theristocaulodon or Leviathan missouriensis. Cymatotherium Kaup “Akten der Urwelt,” pp. 11-14, Tab. tv. Cymatotherium antiquum Kaup {Professor Osborn did not regard this as a proboscidean molar. It was originally listed by Kaup as a Sirenian, but neither Dr. G. G. Simpson nor Dr. E. H. Colbert considers it as such; they think it possible that it may be an embryonic tooth of a proboscidean.— Editor.) Leviathan Koch ‘Description of the Missourium, or Missouri Levia- Leviathan Missouri than,” ete., London, p. 17. (= Leviathan missouri- ensis = Missourium theristocaulodon) Gamphotherium Gloger ‘“‘Gemeinniitziges Hand- u. Hilfsbuch Natur- Mastodon angustidens gesch.,”’ I, pp. xxxii, 119. See also Oldfield Thomas, Cuvier 1895, Ann. Mag. Nat. Hist., (6), XV, pp. 191, 192. Gloger, 1841, p. 119: ‘‘Letztere theilte mit ihm noch ein anderes, welches man fiiglich Schnabel-Mammuth nennen kann, (Gamphotherium angustidens,) obwohl bei ihm die unteren Stosszihne selbst nicht bloss klein blieben, sondern auch nur in der Jugend vorhand- en waren und dann bald fiir immer ausfielen.”’ Gamphotherium is regarded as a misspelling of Gomphotherium Burmeister, 1837, by most authors (Matthew, Hay, Allen); Oldfield Thomas, however, writes (letter, Oct. 20, 1920): “Gamphotherium Gloger has as its genotype, by monotypy, G. angustidens—no other species being referred to. There appears to be no reason to suppose the word has any special connection with Gomphotheriwm, either as misspelling or correction. . . it may have been a misprint for Gomphotherium of Burmeister, but there is no evidence for this, & the a occurs equally in Gloger’s Systematic Index, in the body of the work, and in the alphabetical index at the end. . . The genotype of Trilophodon Falc. & Caut., 1846, is quite clearly ohioticus, no other species being mentioned in the paragraphs referring to it.’”” Hopwood, 1935, pp. 138, 14: “Burmeister’s genus Gomphotherium was originally diagnosed thus, ‘Stossziihne in beiden Kiefern besass die gleichfalls untergegangene Gatt. Gomphotherium.’ (Burmeister, loc. cit.), but he mentioned no species as belonging to this genus. The late Dr. O. P. Hay (Hay, 1923, [1923.2] p. 109), regarded this as coming under Opinion 46 of the International Commission on Zoological Nomenclature and adopted it as a valid genus with genolectotype M. angustidens Cuvier (cf. Cope & Matthew, 1915, expl. to pl. exx). Since the American Mastodon occasionally has mandibular tusks, it is clear that Burmeister’s diagnosis does not distinguish Gomphotherium from Mastodon under which he expressly mentions ‘Ohiothier’, 7. e., Mastodon americanus. On this we may regard Gomphotherium as invalid, because the diagnosis is inadequate. Not only is it in- valid, but it also antedates Gamphotherium Gloger. Oldfield Thomas (1895, p. 189) regards this latter as a new generic name, and (op. cit., p. 191) italicises it as a name which is not a simple synonym of an earlier name. The spelling, however, is quite clearly either an error of transcription, or a lapsus calami, or even a misprint, and the word should be written Gomphotherium, thus becoming a synonym of Gomphotherium Burmeister. The next name, Trilophodon, was originally proposed for the species M. ohioticus and M. angustidens. Of these the former is the genotype of Mastodon, hence the latter only remains in T’rilophodon and may be regarded as the genotype of that genus. For these reasons T’rilophodon is here adopted as a valid genus to include all the bunolophodont proboscideans grouped round Mastodon angustidens.”’ Trilophodon (Section name) Falconer and Cautley ‘Fauna Antiqua Sivalensis,” letterpress, p. 54: “Mastodon. Sect. Trilophodon.—M. Ohioticus.—The next degree of deviation from the ordinary dental rule is presented by Mastodon Ohioticus.” In 1847 Falconer and Cautley (Pls. xtm—xtv) use the same term, e. g., Generic Reference in Present Memoir Mastodon SIRENIAN? Mastodon Trilophodon Trilophodon Collective genus. By courtesy to Falconer restricted to M. an- gustidens (see Trilopho- don Fale., 1857). Year 1847 1847-1857 1847-1857 1847 1848 Genus Author NOMENCLATURE OF THE PROBOSCIDEA Bibliographic Reference Mastodon Sect. Trilophodon, to include the species: III. Mastodon tapiroides, IV. M. ohioticus, V. M. angustidens, VI. M. andium; the same species are listed by Falconer (1867, p. 56) as “I. Trilopho- dontes,”’ with the addition of Deinotheriwm gigan- teum and D. indicum. Genotypic Species This does not appear as a generic definition; see also T'etralophodon Falconer and Cautley below. Compare also Warren (1852, p. 139): “The Mastodons are separated into two groups; one called Trilophodon, and the other, not particularly named, which might be called Tetralophodon.’’ Defined by Falconer as a subgenus of Mastodon (Falconer, 1857, pp. 318, 316, and Synop. Tab. opp. p. 319 with addition of four species and transfer of andium to Tetralophodon). Tetralophodon (Section name) Falconer and Cautley Op. cit., Pls. XLII-XLV, (as distinguished from Sect. Trilopho- don) applied as a section of Mastodon to include the species: VII. M. perimensis, VIII. M. sivalensis, IX. M. arvernensis, X. M. longirostris, XI. M. latidens; these five species in the order named are again listed by Falconer (1867, p. 56) as: “II. Tetralophodontes,” as distinguished from “I. Trilo- phodontes” and “III. Stegodontes.”’ This designation of the Section Tetralophodon is not a generic definition; see also Trilophodon above. Defined by Falconer as a subgenus of Mastodon (Falconer, 1857, pp. 313, 316, and Synop. Tab. opp. p. 319 with addition of andiwm). See Vol. I, Chap. IX, p. 348, of the present Memoir. Stegodon Falconer and Cautley Op. cit., Pl. xuu, figs. xm—xv. Also Falconer, 1857, Quart. Journ. Geol. Soc. London, XIII, pp. 314, 318, and Synop. Tab. opp. p. 319. Falconer, 1857, p. 314: “To this group we have assigned the subgeneric name of Stegodon (Footnote: ‘From oTéyn tectum, and OO0Us dens, having reference to the gable-end form of the section of the ridges.’]. It is limited to extinct forms confined at present to the Indian Tertiaries. The Elephas cliftic Fale. & Caut., E. bombifrons Fale. & Caut., E. ganesa (?) Fale. & Caut., E. insignis Fale. & Caut. Stegodons constitute the intermediate group of the Proboscidea from which the other species diverge through their dental characters, on the one side into the Mastodons, and on the other into the typical Elephants.” Stegodon is admirably characterized by Falconer by four species belonging in the same phylum. It preoccupies Emmenodon Cope, 1889. Loxodon Falconer and Cautley Op. cit., [1847, Pls. xu, xuiv] (name). Falconer, 1857, Quart. Journ. Geol. Soc. London, XIII, pp. 315, 318, Synop. Tab. opp. p. 319; 1865, ibid., X XI, p. 263. Elephas planifrons Falconer & Cautley, E. africanus Blumen- bach, EL. priscus(?) Goldfuss Loxodon Falconer, 1847, 1857, preoccupied by Loxodon Miiller and Henle, 1841, for a genus of sharks. Compare Lozodonte F. Cuvier, 1825, and Loxodonta (unsigned review of F. Cuvier, 1827, 1828—see above, this list). Elasmodon Falconer and Cautley Op. cit., Pl. xuu. A subgeneric name based upon species now known to belong to five different genera (see p. 1177 of the present Memoir), namely, Elephas [Hesperoloxodon|] antiquus, E. [Hypsele- phas] hysudricus, E. |Archidiskodon] meridionalis, E. [Palzoloxodon| namadicus, E. indicus, and E. [Mammonteus] primigenius. Name, moreover, preoccupied by Elasmodus, replaced by Euelephas Falconer, 1857, p. 315 (see Zuelephas, p. 1376, below). Cheirolites von Meyer (In Bronn’s Handb. einer Gesch. d. Natur, III, Index Pal., p. 286). Elephas primigenius Blumenbach Compare Owen, 1846, p. 228: “A separate plate [of molar tooth], with its digital processes, offers a rude resemblance to a hand, and such specimens have been figured by the older collectors of petrifactions, under the name of ‘Cheirolites,’ as the fossilized hand of a monkey ora child.” 1375 Generic Reference in Present Memoir Tetralophodon Collective genus. By courtesy to Falconer restricted to M. longi- rostris (see Tetralopho- don Fale., 1857). Stegodon Loxodonta INVALID, PREOCCUPIED Mammonteus 1376 Year 1850 1857 1857 1857-1865 OSBORN: THE PROBOSCIDEA Author Bibliographic Reference Genotypic Species Synodontherium Costa Atti Accad. Pontaniana, V, Pt. I, pp. 271-275, Tay. 1, figs. 1-4. Genus Tooth belonging to EHlephas primigenius (fide Leidy, Marschall). cory Mastodon latidens Clift, M. arver- nensis Croizet & Jobert, M. siva- lensis Cautley Tetralophodon Warren he Mastodon Giganteus of North America,” p. 139. Collective genus invalid because based upon specimens now known to belong to three different genera. Moreover, preoccupied by T'etralophodon Falconer and Cautley, 1846 {1847]. (In Dorlhae, Ann. Soc. Agric. Puy, XTX, for 1854, p. 507). Anancus macroplus Aymard (=M. arvernensis) Anancus Aymard Anancus is the first valid generic name applied to a member of this phylum. A preced- ing invalid collective name is T'etralophodon Warren. Succeeding synonymous terms are Tetralophodon Falconer and Cautley (in part), Pentalophodon Falconer, Bunolophodon Vacek (in part), Slegomastodon Pohlig, Rhabdobunus Hay, Dibunodon Schlesinger. The name Anancus first appeared in a list in Dorlhac (1855, p. 507) which he says was borrowed from a verbal communication to the above Society by Aymard in 1855; later in 1859, p. 493, Lartet established its validity by making Mastodon arvernensis the type. Mastodon angustidens and other trilopho- donts, as listed above under Trilophodon, 1846 (Section mame) Trilophodon Faleoner Quart. Journ. Geol. Soe. London, XIII, p. 313; defined as a subgenus, p. 316 and Synop. Tab. opp. p. 319. [See above, this list, under Gamphotheriwm, 1841, for conclusions of Dr. Hopwood regarding validity of T’rilopho- don.] Elephas Hysudricus Falconer & Cautley, antiquus Falconer & Cautley, merzdionalis Nesti, Namadicus Falconer & Cautley, indicus Linnzeus, primigenius Blumen- bach Euelephas Faleoner Quart. Journ. Geol. Soe. London, XIII, pp. 315- 318, Synop. Tab. opp. p. 319, subgenus of Hlephas. Falconer, 1857, p. 315, footnote: Substitution of Huele- phas for Elasmodon. Sclater in 1900, p. 317, erroneously specified Elephas planifrons as the genotype of Huelephas, because this species was not included in the original defi- nition of Hlasmodon (see Chap. XIX, p. 1177 above). Mastodon longirostris and other species, as listed above under Tetralophodon, 1846 Tetralophodon Falconer Quart. Journ. Geol. Soc. London, XIII, pp. 313, 317, and Synop. Tab. opp. p. 319, defined as a subgenus of Mastodon, with the doubtful inclusion of Andium at this time. [1847] (Section name). Quart. Journ. Geol. Soc. London, XIII, p.314; No species cited in 1857; Mastodon sivalensis referred to Tetralophodon, 1857; As a subgenus of Mastodon, name without genotype in made the genotype, 1857, name with genotype (WV. sivalensis) in 1865. Falconer, 1865. 1857, p. 314: “Tor reasons which will be explained in the sequel, it would seem that there has existed in nature another subgeneric group of Mastodon, of which only a single form is at present known, in which the crowns of the ‘intermediate molars’ are divided upon a quinary ridge-formula. This group in our arrangement would be characterized, in harmony with the others, as Pentalophodon.” In Falconer’s paper of 1857, M. sivalensis is repeatedly placed in the subgenus Tetralo- phodon; Valconer states on page 317 that: ‘Mastodon Sivalensis, although with five- ridged ‘intermediate molars,’ is provisionally included under T'etralophodon.” (See also Synop. Tab. opp. p. 319.) Tinally, in 1865, p. 262, Falconer adopts the name Pentalopho- don with the genotypic species M. (Tetralophodon) sivalensis. Pentalophodon Valconer 1865, ibid., X XI, p. 262. Generic Reference in Present Memoir Mammonteus INVALID, PREOCCUPIED Anancus Trilophodon By courtesy to Fal- coner restricted to M. angustidens INVALID SUBGENERIC NAME Tetralophodon By courtesy to Fal- coner restricted to M. longirostris Pentalophodon Collective genus. By courtesy to Falconer and Cautley restrict- ed to M. sivalensis Year 1867-1883 Leptodon Gunn 1868 1868 1877 1877 Genus Author NOMENCLATURE OF THE PROBOSCIDEA Bibliographic Reference Genotypic Species Geol. Mag., IV, p. 422 (name only); 1883. Geol. Mag., Leptodon minor, L. Dec. II, N.S., X, p. 458. giganteus Gunn Gunn, 1867, p. 422: ‘(b) the necessity for establishing a new species, called by Mr. Gunn Leptodon, from the fineness of the enamel.’’ Gunn, 1883, p. 458: ‘‘submitted to a meeting of the Geological Society at Somerset House in 1867, under the names of Lepto- don minor and Leptodon giganteus.’’ Type: E. (Leptodon) antiquus; Forest bed stage, England. Generic name Leptodon preoccupied by Gaudry, 1860, for a genus of chalico- theres; by Sundevall, 1835, for a genus of birds. Antoletherium Falconer ‘‘Paleeont. Mem.” [editor: Murchison], I, p. 416, Pl. xxxrv, figs. 1, 2. Footnote on p. 416: “It does not appear from Dr. Falconer’s notes that he had seen the specimen, which is here described from a drawing made by Col. Baker.—[{Ed.]” Rhynchotherium Falconer (MS. 1856); ‘‘Paleont. Mem.,” II, pp. 74, Cast of lower jaw from Cenobasileus Cope 75. See also Falconer, 1863, pp. 44, 56, 60, and Mexico, in Geneva Osborn, 1918.468, p. 136. Museum Falconer, 1868, II, pp. 74, 75: “Extract of Letter from Dr. Falconer to M. Lartet, September 12, 1856. ‘At Genoa [Geneva] I saw a cast of a large lower jaw of a Mastodon from Mexico, with an enormous bec abruptly deflected downwards and containing one very large lower incisor. The beak is much thicker than in M. (Trilophodon) angustidens and larger than in M. (Tetralophodon) longirostris. You know that every one (Laurillard, Gervais, &c.) has insisted on the absence of the lower incisors from both of the South American species. The outline of the jaw resembles very much the figure in Alcide D’Orbigny’s Voyage, described by Laurillard as M. Andium. The specimen is unpublished material, and I was therefore only allowed to examine it very cursorily. The Genoese paleontologists had provisionally named it Rhynchotherium, from the enormous develop- ment of the beak, approaching Dinotherium.’” Search for this cast was unavailing for many years, but finally (1931) it was located at the Natural History Museum in Geneva, Switzerland (see Vol. I, p. 477). In the meantime Osborn (1921.515, p. 5) validated this genus: “The original genotype may be termed Rhynchotherium tlascalz, new species, from the locality Tlascala.’’ Compare figure 448. Proc. Amer. Phil. Soe., XVI, pp. 584, 585 (Pal. C. tremontigerus Cope Bull. 24). (=M. americanus, fide Hay, 1902, p. Based on an artifact; name withdrawn by Cope in 1889 708). (Amer. Nat., XXIII, p. 207). Zygolophodon Vacek Abhand. geol. Reichsanst., VII, Heft IV, p. 45. Mastodon borsoni Hays, Wien. M. turicensis Schinz, M. tapi- A clear and admirable designation of the four species roides Cuvier, M. which together appeared to constitute a single distinct genus. pyrenaicus Lartet Osborn, however, in 1926 (1926.706) removed the species M. turicensis, making it the type of a new genus T'uricius; in the present Memoir he assigns M. tapiroides Cuvier to the genus Turicius (see Vol. I, pp. 203, 217) leaving M. borsont and M. pyrenaicus as typical of Zygolophodon (see Vol. I, Chap. VII). Hopwood, 1935, p. 42: ‘Vacek (1877) used the genus Mastodon strictly in the Cuvieri- an sense, and sub-divided it into two groups, Zygolophodon and Bunolophodon, including in the former, M. tapiroides Cuv., M. pyrenaicus Lartet, ‘Form von Baltaviir’, “M. Turicen- sis von Pikermi’, M. Borsoni Hays, and M. ohioticus. Of these species Matthew (1918, p. 200) selected M. tapiroides as the genotype. Osborn (1926 [1926.706]) has taken M. borsont as genotype, but under the rules of priority this selection is inadmissible. . . . The first mention of M. tapiroides is in Desmarest (1822, p. 386) who latinised Cuvier’s vernacular, and took as his type the specimen figured by Cuvier. Hence, the genotype of Zygolophodon is M. tapiroides Desmarest, spec. indet., and thus Zygolophodon remains an indeterminate genus.” See also this Memoir, Vol. I, p. 203, footnote. Bunolophodon Vacek Loc. cit. Mastodon atticus Wagner, M. longi- rostrisplkaup, M. angustidens Cuvier, M. arvernensis Croizet and Jobert, M. pentelici Gaudry 1377 Generic Reference in Present Memoir Hesperoloxodon Deinotherium Rhynchotherium Nov A PROBOSCIDEAN Zygolophodon By courtesy to Vacek restricted to M. borsoni Collective genus. InVALID= Anancus, =Trilophodon, ete. 1378 OSBORN: THE PROBOSCIDEA Generic Reference Year Genus Author Bibliographic Reference Genotypic Species in Present Memoir 1882 Notelephas Owen Proc. Roy. Soe. London, XX XIII, p. 448; also Phil. Notelephas australis Drproropon, a Trans. Roy. Soc. London, 1883, CLX XIII, Pt. III, Owen marsupial 1s tlc See note under Notelephas australis, 1882 (p. 13897 below), from Jack and Etheridge, 1892, p. 683. 1884 Dibelodon Cope Proce. Amer. Phil. Soc., XXII, pp. 2-8. Mastodon shepardi INVALID Leidy Invalid because preoccupied by Rhynchotherium (see this Memoir, I, p. 525). 1884 Tetrabelodon Cope Ibid., pp. 4-5. Mastodon angustidens INVALID Cuvier Invalid because preoccupied by T'rilophodon (see this Memoir, I, pp. 249, 525). 1885 Archidiskodonten Pohlig Zeitschr. deutsch. geol. Ges., XXXVII, p. Hlephas planifrons Archidiskodon 1027. Falconer and Cautley 1888 Archidiskodon Pohlig Nova Acta Leop. Carol., LIII, pp. 138, 252. Elephas meridionalis Archidiskodon Nesti, #. planifrons Falconer and Cautley Pohlig, 1885, p. 1027: “8. Ich theile die Elephanten nach Kronenformen und Lamel- lenzahlen der Molaren ein in Archidiskodonten (E. planifrons, E. meridionalis), Loxodonten (BE. africanus, ?E. antiquus) und Polydiskodonten (E. primigenius, E. indicus, etc.), die Stegodonten mit Clift wieder zu Mastodon ziihlend.” An excellent generic distinction of two very closely related species (H#. planifrons, E. meridionalis), to which may now be added Elephas imperator. 1885 Loxodonten Pohlig Zeitschr. deutsch. geol. Ges., XX XVII, p. 1027. Elephas africanus Loxodonta Blumenbach, ?2L. antiquus Fale. & Caut. 1888 Loxo-(disko)-donten Pohlig Nova Acta Leop. Carol., LILI, pp. 138, 252. Hlephas africanus Loxo(-disko-)don Blumenbach, £. priscus Goldfuss, E. antiquus Fale. & Caut. Proposed as a group or sectional name, equivalent to the Loxodontins of Osborn. See note under Archidiskodon, 1885-1888, above, first paragraph. 1885 Polydiskodonten Pohlig Zeitschr. deutsch. geol. Ges., XXXVII, p. Hlephas primigenius INVALID 1027. Blumenbach, LH. indicus Linneus 1888 Polydiskodon Pohlig Nova Acta Leop. Carol., LIII, pp. 138, 252. Elephas primigenius Blumenbach, £. indicus Linneeus, LE. (2) namadicus Fale. & Caut. Proposed in 1885 as a group or sectional name; in 1888 as a genus. See note under Archidiskodon, 1885-1888, above, first paragraph. Collective genus invalid, because based on species now known to belong to three dif- ferent genera, Mammonteus, Elephas, and Palzxoloxodon. 1885 Stegodonten Pohlig Zeitschr. deutsch. geol. Ges., XX XVII, p. 1027. No species cited 1888 Stego(-lopho-)don Pohlig Nova Acta Leop. Carol., LIII, p. 252. Proposed in 1885 as a group or sectional name. See note under Archidiskodon, 1885- 1888, above, first paragraph. 1889 Emmenodon Cope Amer. Nat., XXIII, p. 194. Elephas cliftti Fale. & INvauip Caut., Mastodon ele- Invalid because based upon two species which are phantoides Clift typical members of the genus Stegodon. Year 1901 1901 1902 1912 1912 1912 1914 1914-1917 1914 1917 1917 1917 NOMENCLATURE OF THE PROBOSCIDEA Genus Author Bibliographic Reference Genotypic Species Palexomastodon Andrews Zoologist (4), V, Aug. 15, pp. 318, 319 (name Palxomastodon beadnelli only); Tageblatt V Internat. Zoologen-Cong., Ber- Andrews lin, No. 6, Aug. 16, p. 4 (published vol. 1902, p. 528); Geol. Mag., Dec. IV, N.S., VIII, pp. 400-409. Meritherium Andrews Tageblatt V Internat. Zoologen-Cong., Berlin, Mceritheriwm lyonsi No. 6, Aug. 16, p. 4 (name only); Geol. Mag., Dee. Andrews IV, N.S., VIII, pp. 400-409. Phiomia Andrews and Beadnell “A Preliminary Note on Some New Phiomia serridens Mammals from Upper Eocene of Egypt,’’ Surv. Andrews and Bead- Dept., Pub. Works Ministry, pp. 1-9. nell Promastodon Pohlig Bull. Soe. belge Géol., X XVI, Procés-verbaux, Puerco mammal p. 192. Pohlig, 1912, p. 192: ‘“.. .illustrée aussi par celui de Puerco (prototype de Promastodon Pohlig).’’ Founded on a supposed missing link in the American Paleocene Puerco; possi- bly Cope’s Polymastodon, a multituberculate. This invalid description of Promastodon precedes (p. 192) the valid generic definition of Stegomastodon cf. mirificum, page 193, as cited below. Stegomastodon Pohlig Jbid., p. 193. Mastodon mirificus Leidy Pohlig, 1912, p. 193: ‘. . .c’est prouvé, notamment, par une superbe dent de Stego- mastodon ef. mirificum Leidy (prototype de Stegomastodon Pohlig).”’ Hemimastodon Pilgrim Mem. Geol. Surv. India (Pal. Indica), N. 8., Tetrabelodon crepusculi IV, Mem. 2, pp. 17-21. Pilgrim Type Tetrabelodon crepusculi Pilgrim, 1908 (p. 157) subsequently named Hemimasto- don, 1912. Eubelodon Barbour Univ. Neb. Studies, XIV, p. 194. Hubelodon morrilli Barbour Megabelodon Barbour Neb. Geol. Surv., IV, Pt. 14, p. 217 (as a sub- Tetrabelodon lulli genus); 1917, zbrd., IV, Pt. 30, p. 512 (raised to Barbour higher rank). Barbour, 1914.2, p. 217: “For this new species, we wish to propose the name Tetra- belodon lulli, for Professor Lull. It is not unlikely that it is entitled to rank as a new subgenus at least, and it might not be amiss to propose the name Megabelodon.” Barbour, 1917, p. 512: “In the case of Tetrabelodon lulli, the sub-generic title Megabelodon . . . was proposed, but it seems entitled to higher rank [i. e., genus].”’ Rhabdobunus Hay lowa Geol. Surv. Ann. Rept. for 1912, XXIII, pp. Mastodon mirificus 59, 373. Leidy Preoccupied by Stegomastodon Pohlig, 1912, above (cf. also Hay, 1930, Publ. Carnegie Inst. Wash., No. 390, p. 633). Stegolophodon Schlesinger Denk. Naturhist. Hofmus., I, p. 115 (as a Mastodon latidens subgenus). Clift Schlesinger, 1917, p. 115, footnote: ‘Ich schlage fiir WM. latidens, das sich durch seine kurze Symphyse von dem Subgenus Bunolophodon, durch seinen Molarenbau von Dibuno- don entfernt, den Untergattungsnamen Stegolophodon vor. Der Name bringt einerseits die nahen Beziehungen zum Genus Stegodon, anderseits die Loslésung der Untergattung von Bunolophodon und ihre Sonderstellung gegeniiber Dibunodon zum Ausdruck.’’ See Stego(-lopho-)don Pohlig, 1888, above, this list. Dibunodon Schlesinger Jbid., p. 124 (as a subgenus, with definition). Mastodon (Dibunodon) arvernense Croiz. & Jobert Choerolophodon Schlesinger Jbid., p. 181 (as a subgenus). Mastodon pentelicus Gaudry and Lartet Type: Mastodon (Choerolophodon) pentelict Gaudry and Lartet. Schlesinger, 1917, p. 181: “Schédel niedrig, langgestreckt mit zwei aufwirts und auswarts geschwungenen schmelzbandlosen Stosszihnen. Unterkiefer mit missig langer Symphyse ohne untere Inzisoren. Molaren choerodont (hochgradig suid).’’ Sce text and figures, Vol. I, Chap. VIII, this Memoir. Generic Reference in Present Memoir Palxomastodon Meritherium Phiomia INVALID Stegomastodon SUINA Eubelodon Megabelodon Stegomastodon Stegolophodon Anancus Trilophodon (Choerolophodon) 1379 1380 OSBORN: THE PROBOSCIDEA Generic Reference Year Genus Author Bibliographic Reference Genotypic Species in Present Memoir 1917 Genomastodon Barbour Neb. Geol. Sury., IV, Pt. 30, p. 512. Tetrabelodon willistoni Trilophodon Barbour, 7. lulli (Genomastodon) Barbour, 1917, p. 512: “Longirostral mastodons seem Barbour, 7’. osborni to have reached their maximum in such forms as Tetrabelo- Barbour don willistoni, lulli, osborni, and the like. For the present at least, the above may be grouped under a new and distinct genus, Genomastodon. . . .«In the case of Tetrabelodon lulli, the sub-generic title Megabelodon . . . was proposed, but it seems entitled to higher rank [1. e., genus].’”’ Tetrabelodon lulli, therefore, is preoccupied by Megabelodon lulli; by the process of elimination, Genomastodon applies to T. willistoni and T. osborni only. 1917 Mastelephas Barbour Loe. cit. No definition and no Nomen NupUM genotypic species given. 1922 Miomastodon Osborn Amer. Mus. Novitates, No. 49, p. 4. Mastodon merriami Miomastodon Osborn 1923 Cuvieronius Osborn Amer. Mus. Novitates, No. 99, p. 1. Mastodon humboldtii Cuvieronius Cuvier (in Desmar- Cabrera (1929) erroneously refers Mastodon humboldtii est) to the genus Stegomastodon Pohlig, which, as shown above, is based on Leidy’s ‘Mastodon mirificus,’ wholly distinct from M. humboldtiiz. 1923 Serridentinus Osborn Ibid., p. 2. Mastodon productus Serridentinus .Cope, M. serridens Osborn, this Memoir, I, pp. 285, 286: ‘‘(December, Cope, M. floridanus 1932) On very close reéxamination of the cast (Amer. Mus. Leidy, M. obscurus 1909) of Leidy’s classic type (Figs. 232, 233) of ‘Mastodon’ Leidy, Serridentinus obscurus, also of the original type molar fragments of ‘Ser- simplicidens Osborn ridentinus’ simplicidens Osborn, these two species prove to belong in the genus Trilophodon because of the presence of ‘central conules’ uniting as central trefoils with the median conules. . . . Osborn, 1933: Final study of the veritable type of Mastodon obscurus Leidy proves that it does not belong in the genus Serridentinus Osborn, but in the related though distinct genus Trilophodon Falconer. The phylogenetic position of 7’. obscurus appears to be distinct from the species Mastodon (= Serridentinus) productus of Cope and M. [ =Ocalientinus (Ser.)] floridanus of Leidy.” 1923-1926 Prostegodon Matsumoto (In Osborn, Amer. Mus. Novitates, No. 99, p. Mastodon latidens Stegolophodon 2). Chift Subsequently proposed by Matsumoto (1924.3, pp. 324-327) as a subgenus of Stegodon; preoccupied by Stegolophodon Schlesinger, 1917, a genus also based upon the Mastodon latidens of Clift. This description appeared in the Japanese language. In 1926.1, p. 9, Matsumoto published his English text on this genus. 1924 Harpagonotherium Vischer de Waldheim, 1808 (Fide Sherborn, 1924, Pt. V, p. 1022). 1924 Parastegodon Matsumoto Journ. Geol. Soc. Tokyo, XXXI, pp. 256, Hlephas aurore Archidiskodon? 257, 262 (in Japanese); 1929, Sci. Rept. Tohoku Mats., 1915, or a progres- Imp. Univ., (2), Geology, XIII, No. 1, pp. 13-15 1918 sive Stegodon (in English). Elephas (Parastegodon) aurorx, well established on an excellent type, is, like Stegodon mindanensis, either a highly progressive member of the Stegodon phylum or a primitive member of the Archidiskodon phylum, a point which can only be determined positively by the discovery of the cranium of Parastegodon aurore. 1924 Palzxoloxodon Matsumoto Journ. Geol. Soc. Tokyo, XX XI, pp. 255— Elephas namadicus Palzxoloxodon 272 (in Japanese); 1929, Sci. Rept. Téhoku Imp. naumanni Makiyama Univ., (2), Geology, XIII, No. 1, pp. 7, 10 (in English). 1924 Sivalikia Osborn Amer. Mus. Novitates, No. 152, p. 2. Elephas namadicus Palzxoloxodon ’ Faleoner and Cautley Year 1924 1924 1924 1924 1924 1926 1926 1926 1927 1927 1928 1929 1929 1929 1930 1931 1931 Genus Author Pilgrimia Osborn NOMENCLATURE OF THE PROBOSCIDEA Bibliographic Reference Loe. cit. Mammonteus Osborn Loc. cit. Parelephas Osborn Compare Mammonteum Camper, 1788, above, this list. Tbid., p. 4. Morrillia Osborn Jbid., No. 154, p. 1. Lydekkeria Osborn Raised to rank of genus, this Memoir, I, pp. 690, 739. Ibid., p. 2. See Volume I, p. 353, where it is stated that possibly Lydekkeria may become a synonym of Tetralophodon. Pliomastodon Osborn Ibid., No. 238, p. 1. Turicius Osborn Ibid., p. 3. Cordillerion Osborn Leith-Adamsia Matsumoto Japanese Journ. Geol. and Geog., V, No. 4, Amebelodon Barbour Platybelodon Borissiak Ann. Soc. Paléont., Russie, VII, for the year Synconolophus Osborn Notiomastodon Cabrera Rev. Mus. La Plata, XXXII, pp. 90, 91. Torynobelodon Barbour Prodinotherium Ehik Geol. Hungarica (Palaeont. Ser.), Fase. 6, pp. Teleobunomastodon Revilliod Hesperoloxodon Osborn Separated from Zygolophodon by Osborn. Ibid., p. 15. The generic name Cordillerion replaces the preoccupied name Dibelodon, as used by Cope and Lull, and Mastodon humboldtii Cuy. In 1923 Osborn proposed Cuvieronius to embrace M. humboldtii. Cabrera (1929, p. 90) erroneously holds that the geno- Genotypic Species Elephas falconeri Busk; other species cited, E. melitensis Fal- coner, EL. mnaidre Adams, E. antiquus Recki Dietrich Elephas jeffersonii Osborn Tetralophodon barbourt Osborn Mastodon (Trilopho- don) falconert Lydekker Mastodon (Miomasto- don) matthewi Osborn Mastodon turicense Schinz Mastodonte des Cor- diliéres Cuvier = Mastodon cordil- lerarum Desmarest = Mastodon andium Cuvier type of Cordillerion (Mastodon andium Cuv.) belongs to the same genus as the genotype of Cuvieronius (M. humboldtii Cuv.), and hence are alike referable to Stegomastodon Poh- lig based on M. mirificus Leidy. 1929, p. 90, for Cordillerion Osborn. Art. XII. 1927, pp. 105-120. 3-21. 21. Neb. State Mus., Bull. 13, I, pp. 131-134. Amer. Mus. Novitates, No. 393, pp. 9, 10. Neb. State Mus., Bull. 16, I, pp. 147-153. Mém. Soe. Paléont. Suisse, LI, pp. 20, Amer. Mus. Novitates, No. 460, p. 21. He accordingly substitutes Notiomastodon gen. nov., Leith-Adamsia siwalik- zensis Matsumoto Amebelodon fricki Barbour Platybelodon Danovi Borissiak Synconolophus dhok- pathanensis Osborn Notiomastodon ornatus Cabrera Torynobelodon loomisi Barbour Prodinothervum hungaricum Ehik Mastodon bolivianus Philippi Palzoloxodon antiquus italicus Osborn Generic Reference in Present Memoir Palzxoloxodon Mammonteus Parelephas Morrillia Tetralophodon (Lydekkeria) Pliomastodon Turicius Cordillerion Archidiskodon Amebelodon Platybelodon Synconolophus Notiomastodon Torynobelodon Deinotherium Cordillerion(?) Hesperoloxodon 1381 1382 Year 1933 1933 1933 1933 1933 1933 1933 1933 1934 1934-1941 1934-1941 1935 Year 1754 1792 OSBORN: THE PROBOSCIDEA Bibliographic Reference Bull. Amer. Mus. Nat. Hist., LIX, Art. LX, pp. Genotypic Species Blickotherium. blicki Genus Author Blickotherium Frick 505, 509, 515, 527-531. Frick Aybelodon Frick Ibid., pp. 505, 527, 532. Aybelodon hondurensis Frick Ocalientinus Frick Ibid., pp. 505, 576, 579. Ocalientinus ojo- caliensis Frick Trobelodon taoensis Frick Tatabelodon rio- grandensis Frick Trobelodon Frick Tbid., pp. 505, 576, 580. Tatabelodon Frick Tbid., pp. 505, 576, 581. Paraplatybelodon Frick TIbid., p. 592. Serbelodon Frick Tbid., pp. 506, 592, 594. Serbelodon barbour- ensis Frick Cryptomastodon von Koenigswald Wetenschappelijke Mededeelingen, Dienst Mijnbouw Nederl.-Indié, I Teil, No. 23, pp. Natit iil. Amer. Mus. Novitates, No. 741, pp. 2, 12. Cryptomastodon mar- tini von Koenigswald Loxodonta griqua Haughton Metarchidiskodon Osborn Proc. Amer. Phil. Soc., LX XIV, p. 285 (name only); 1936, this Memoir, I, p. 12; 1941, op. cit., II, p. 1340. Elephas hysudricus Falconer and Cautley and Elephas platycephalus an- gustidens Osborn Hypselephas Osborn Proc. Amer. Phil. Soc., LX XIV, p. 285 (name only); 1936, this Memoir, I, p. 12, Pl. x1; 1941, op. cit., II, p. 1358. Gnathabelodon Barbour and Sternberg Neb. State Mus., Bull. 42, I, pp. 395, 396. Elephas platycephalus Osborn Platelephas Osborn Gnathabelodon thorper Barbour and Stern- berg LIST OF SPECIES, SUBSPECIES, AND VARIETIES Author Elephas indicus Linneus Name “Systema Nature,” p. 11. Hab.: India, Ceylon. This name was first used by Linneus in 1754, but in the 1758 edition of the “Systema Nature,”’ p. 33, he used Hlephas maximus for the Ceylon elephant. Elephas maximus Linneus “Systema Nature,” edition 10, p. 33. Hab.: India, Ceylon. Sherborn (letter, July 14, 1929) is of the opinion that maximus should be ‘“‘swept aside.”’ Elefante Indiano Giovanni Targioni Tozzetti Elephas americanus Kerr ‘‘Animal Kingdom of Linneus,” I, p. 116. Lick(?), Boone County, Kentucky, near Ohio River. Leidy, 1869, p. 392, footnote: ‘Cuvier, in the works quoted [Tab]. Elem. Hist. Nat. (an. 6) 1798, 149; Mem. Inst. Nat. Sci. (An VII [1799], 19, 21.], and De Blainville, in his Osteog. Gen. Elephants, 327, 245, attribute this name to Pennant. Falconer and Cautley, in the Fauna Antiq. Sival., 17, also observe, ‘that Pennant first ventured in 1793 to desig- nate the American fossil animal, in a systematic work, as a species of Elephant by the name of E. americanus.’ 1 have been unable to find the name thus expressed in any of the works of Pennant, nearer than the words ‘American Elephant,’ which occur in the Synopsis of Quadrupeds of 1771 and in both editions of the History of Quadrupeds, that of 1781 and 1793.” Type loc.: Big Bone Generic Reference in Present Memoir Blickotherium Aybelodon Ocalientinus Trobelodon Trilophodon (Tatabelodon) NOMEN NUDUM Serbelodon ?SIRENIAN Metarchidiskodon Hypselephas Platelephas Gnathabelodon Reference in Present Memoir Klephas indicus (pre-Linnean) Elephas indicus Elephas |Archidisko- don| meridionalis Nesti, 1825 (fide Weithofer, 1890, p. 133) Mastodon americanus Year 1796 1797 1797 1798 1798 1799 1799 1799 1801 1803 NOMENCLATURE OF THE PROBOSCIDEA Name Author Elephas Mammoth Cuvier [and Geoffroy] Falconer, 1868, II, p. 158: ‘Cuvier was undoubtedly the first to characterize the ex- tinet species with exactness, in his joint memoir with Geoffroy, under the name of Elephas Mammoth, in the year 1796 [Footnote: ‘Mém. de l'Institut, 1™° Classe, tom. 1i.’].’’ Ohio-Incognitum Blumenbach Abbild. naturhist. Gengens., No. 19. Hab.: Ohio River, North America. Elephas asiaticus Blumenbach ‘“‘Handb. d. Natur.,’’ 5th edition, p. 124. Hab.: India. Type fig.: Blumenbach, Abbild. naturhist. Gegens., No. 19, fig. B. Elephas africanus Blumenbach ‘Handb. d. Natur.,’”’ 5th edition, p. 125. Hab.: South Africa. Type fig.: Blumenbach, Abbild. naturhist. Gegens., No. 19, fig. C. Not in the edition of the “Handbuch der Naturgeschichte”’ of 1779 [First Edition] but in the Fifth Edition (1797); in ‘“Abbild. Naturhist. Gegenstande,’’ Heft 2, No. 19, fig. C, the name is already used. . . The arrangement of the lamelle would indicate its Cape origin. It can only be a question of the Cape Colony and the Congo, perhaps of the French Congo. The confluence of the anterior lamelle is peculiar. (Matschie, letter, 1921.) Sherborn (1902, Pt. I, p. 22) lists Z. africanus as in the Fifth Edition of the ““Hand- buch” (1797.1, p. 125), but reference is not made to a figure in this edition. In 1923, Pt. IT, p. 135, he cites EZ. africanus in Blumenbach, Man. Hist. Nat., I, 1803, p. 155. Elephas indicus Cuvier ‘Tabl. Elémen. Hist. Nat. Animaux,” p. 148. Hab.: India. See Cuvier’s figure, with description, 1799, Mém. Inst. Nat. Sci. et Arts. Scei., Mathém. et Phys., II, Fructidor, an VII [1799], p. 21, and Plates (11, figs. 1G 22 eiveie sl verge I wie cfios 2) Elephas capensis Cuvier ‘‘Tabl. Elémen. Hist. Nat. Animaux,” p. 149. Hab.: South Africa. Type fig.: Cuvier, 1799, Mém. Inst. Nat. Sci. et Arts. Sci., Mathém. et Phys., II, Fructidor, an VII [1799], Pl. m1 and PI. rv, fig. 2. Elephas capensis G. Cuvier, “Tableau Elémen.,” 1798. Cuvier had at his disposal one skeleton from the Senegal and one skull from the Cape [‘‘Cap de Bonne-Espérance,’’ Cuvier, Mém. Inst. (de France) National des Sciences et Arts, sometimes called the Acadé- mie des Sciences, Vol. II, année 7 (1799?)]. (Matschie, letter, 1921.) Elephas primigenius Blumenbach ‘‘Handb. d. Natur.,” 6th edition, p. 697. Hab.: Siberia and northern Germany. No original type figure. Mammut ohioticum Blumenbach Op. cit., p. 698. Hab.: Ohio River, North America. Elephas mammonteus Cuvier (1796 MS.), published August-September, 1799, Mém. Inst. Nat. Sci. et Arts. Sci., Mathém. et Phys., II, Fructidor, an VII [1799], p. 21, Pls. v, fig. 2, and v1, fig. 1. Falconer, 1868, II, p. 158: ‘In the same year [1796] he [Cuvier] read a memoir [MS.] at the first public meeting of the ‘Institute,’ but which was not published until 1806 [1799, p- 21], in which the diagnostic marks are very pointedly expressed under the designation of Elephas Mammonteus: ... [subsequently—1806, Ann. Mus. hist. nat., VIII, p. 264] he abandoned the name E. Mammonteus of his [MS.] memoir of 1796, and adopted the desig- nation of Elephas primigenius, proposed by Blumenbach [Footnote: ‘Voigt’s Mag. 1803, Band v, p. 16.”], in 1803 which is that now generally accepted among paleontologists.” See, however, Bibliography, Cuvier, 1799.1. Elephantus indicus Cuvier and Lacépéde “La Ménagerie du Muséum National d’Histoire naturelle ou Les Animaux Vivants,” An X. Hab.: India. Elephas macrocephalus A. Camper Original text of 1802 not available. Atlas of 1803 gives the name Hlephas macrocepha- lus under Fig. 18. Sherborn, 1922-1933, “Index Animalium” (1928, p. 3760) gives the follow- ing reference: ‘‘Elephas macrocephalus,’ Camper, (Luvres de P. Camper, IT, 1803, 18, f. n. [non usu spec.}.” 1383 Reference in Present Memoir Mammonteus primi- genius Mastodon americanus Elephas indicus Loxodonta africana Name preoccupied by Linneus, 1754 Loxodonta africana capensis Mammonteus primi- genius Mastodon americanus Mammonteus primi- genius Elephas indicus Mastodon americanus 1384 Year 1806 1806 1806 1806 1806 1806 1807 1808 1808 1808 1809 1811 1811 1814 1814 OSBORN: THE PROBOSCIDEA Author Le Grand Mastodonte Cuvier Ann. Mus. hist. nat., VIII, pp. 270, 293 (Cuvier, 1806.2). Hab.: Ohio River, North America. Type fig.: Cuvier, zbed., Pl. 49 [1], figs. 1-5. Cuvier, ibid., pp. 270, 272: “Animal trés-voisin de 1’éléphant, mais 4 mécheliéres hérissées de gros tubercules, dont on trouve les os en divers endroits des deux continens, et surtout prés des bords de l’Ohio, dans l’Amérique Septentrionale, improprement nommé Mammouth par les Anglais et par les habitans des Etats-Unis. . ..Nous empruntons le nom de mastodonte de deux mots grecs qui signifient dents mammelonnées, et qui expriment par conséquent son principal caractére.”’ Cf. Mastodonte de Vv Ohio Cuvier, below, this list. Name Mastodonte de VOhio Cuvier Tbid., p. 412 (Cuvier, 1806.3). Hab.: Ohio River, North America. Cf. Le Grand Mastodonte Cuvier, 1806, above, and Mastodon giganteum Cuvier, 1817, below, this list. Mastodonte a@ dents étroites Cuvier Loc. cit. Type loc.: Simorre, l'rance. Type fig.: Cuvier, 2bid., Pl. 66 [1], fig. 4. Cf. Mastodon angustidens Cuvier, 1817, below, this list. Petit mastodonte Cuvier Ibid., p. 4138. Type loc.: Montabusard, France. rbid., Pl. 68 [11], fig. 6. Loc. cit. Type loe.: Saxony, Germany. Type fig.: Cuvier, zbid., Pl. 67 [ur], fig. 11. Cuvier applied the term “Petit mastodonte” to two distinct types, from Montabusard and Saxony, to which he subsequently assigned the names Mastodonte tapiroide, 1821, and M. minutus, 1824, respectively (see below, this list, and also M. parvus, 1834). Type fig.: Cuvier, Mastodonte des Cordiliéres Cuvier Loc. cit. Type loe.: Near voleano of Imbaburra, Quito, Heua- dor. Type fig.: Cuvier, zbéd., Pl. 67 [1], fig. 1. This is the single type which Cuvier subsequently (1824) called Mastodon andium (see below) and which Desmarest called M. cordillerarum. The first distinctive generic name applied to these animals was Mastotheriwm Fischer (1814). Subsequent names: Dzibelodon Cope (1884.2), Cordillerion Osborn (1926). See List of Genera above. Mastodonte humboldien Cuvier Loc. cit. Type loc.: Near Concepcion, Chile. Type fig.: Cuvier, ibid., Pl. 67 [1], fig. 5. Cf. Mastodon humboldtit Cuvier, in Desmarest, 1818-1824, below, this list. Blephas mammouth Link Beschr. Nat. Samml. Univ. Rostock (4) 1807, 3 (fide Sherborn, 1928, p. 3845). Blephas primevus Blamenbach (In Adams, translation from the French by Sir Joseph Banks, Phil. Mag. (Tilloch), X XIX, London, p. 152); cited by Tilesius, 1815, p. 452. Hab.: Lena River, Siberia. Cf. Elephas brachyramphus Brandt, 1832, below, this list. EKlephas minimus Nesti Attributed by Falconer and Cautley, 1846, “Fauna Antiqua Sivalensis,” letterpress, p. 13, to Nesti, but not found by the present author in the original reference. Name abandon- ed by Nesti. Harpagmotherium canadense Fischer de Waldheim Prog. d’Invit. Séance, Pub. Soc. Imp. Nat. Moscou, September, p. 19. Hab.: Ohio River, North America. Mém. Soe. Imp. Nat. Moscou, II, p. Mastotherium ohioticum (Blum.) Fischer de Waldheim 252. Elephas gigas G. Perry “Arcana,” p. li and plate. Elephas socotrus G. Perry Op. cit., p. Ii. Or Mastotherium megalodon Fischer de Waldheim Hab.: Ohio River, North America. Cf. Le Grand Mastodonte and Mastodonte de V Ohio Cuvier, 1806, above, this list. Loognosia,” p. 340. Mastotherium leptodon Vischer de Waldheim Loe. cit. Type loe.: Simorre, France. Cf. Mastodonte a dents étroites Cuvier, 1806, above, this list. Reference in Present Memoir Mastodon americanus Mastodon americanus Trilophodon anqusti- dens Turicius tapiroides Trilophodon angusti- dens minutus Cordillerion andium Cuvieronius humboldtii Mammonteus primi- genius Mammonteus primi- genius Mastodon americanus [Mastodon americanus} [Elephas indicus] [Indeterminate] Mastodon americanus Trilophodon angusti- dens Year 1814 1814 1814 1814 1814 1814 1814 1814 1815 1817 1817 1818 1818 1820 1821 1821 1824 NOMENCLATURE OF THE PROBOSCIDEA Author Mastotherium microdon Fischer de Waldheim Loc. cit. Type loc.: Montabusard, France. Cf. Petit mastodonte, i petites dents, Cuvier, 1806, above, also Mastodon tapiroides Cuvier, in Desmarest, 1822, below, this list. Mastotherium hyodon Fischer de Waldheim Op. cit., p. 341. Cf. Le Mastodonte des Cordiligres Cuvier, 1806, above, this list. Name Hab.: Cordilleras. Mastotherium humboldtii Fischer de Waldheim Loe. cit. Cf. Mastodonte humboldien Cuvier, 1806, above, and Mastodon humboldtii Cuvier, in Desmarest, 1818-1824, below, this list. Mastodon Macrodon Rafinesque Specchio Sci., II, No. 12, p. 182. Hab.: North America. Mastodon rhomboides Rafinesque Loc. cit. Hab.: Cordilleras. Mastodon humboldianus Rafinesque Loc. cit. Hab.: South America. Mastodon Senodon Rafinesque Loc. cit. Hab.: France and Europe. Loc. cit. Hab.: Europe. Elephas primordialis Blumenbach (In Tilesius, Mém. Acad. Imp. Sci. St. Pétersb., (V), V, p. 470). Hab.: Germany and Siberia. See also EL. primordialis Brayley, 1831, Phil. Mag., IX, pp. 411-418; Bronn’s Neues Jahrb., 1833, p. 372, and the “Index paleontologicus” of von Meyer and Géppert, in Bronn’s “Handbuch einer Geschichte der Natur,’ 1848, III, p. 455. Hab.: Eschscholtz Bay, Alaska. Mastodon giganteum Cuvier ‘“‘Le Régne Animal,” p. 233. County, Kentucky, North America. Cf. Le Grand Mastodonte Cuvier, 1806, Mastodonte de VOhio Cuvier, 1806, Masto- therium megalodon Fischer, 1814, above, also Mastodon maximus Cuvier, 1824, Mastodon ohioticum, 1832, and Mastodon americanus Leidy, 1868, below, this list. Mastodon angustidens Cuvier Loc. cit. Type loc.: Simorre, France. Type fig.: Cuvier, 1806, Ann. Mus. hist. nat., VIII, Pl. 66 [1], fig. 4. Not until the year 1817 did Cuvier substitute the name Mastodon angustidens for his “Mastodonte A dents étroites” of 1806. Cf. Mastotherium leptodon Fischer, 1814, above, this list. (In Desmarest, Nouv. Dict. d’Hist. Nat., XIX, p. 446). Mastodon microdon Rafinesque Type loe.: Big Bone Lick, Boone Mastodon minor Cuvier Cf. Petit Mastodonte Cuvier, 1806, above, and Mastodon minutus Cuvier, 1824, below, this list. Mastodon humboldtii Cuvier (In Desmarest, ibid., p. 447); Cuvier, 1821-1824, ‘“‘Ossemens Fos- siles,”’ V, Pt. 2, p. 527. Type loc.: Near Concepcion, Chile. Type de- scription: Cuvier, 1806, Ann. Mus. hist. nat., VIII, p. 413. Type fig.: Cuvier, zbid., Pl. 67 [1], fig. 5. Cf. Mastodonte humboldien Cuvier, 1806, also Mastotherium humboldtii Fischer, 1814, above, this list. Elephas jubatus Schlotheim ‘Die Petrefactenkunde,” p. 4. Hab.: Germany. Mastodonte tapiroide Cuvier ‘“Ossemens Fossiles,”’ I, p. 268. Type loc.: Caleaire de Monta- busard, France. Cf. Petit mastodonte Cuvier, 1806, above, this list. Not until the year 1821 did Cuvier replace “Petit: mastodonte”’ with the name tapiroide, subsequently written tapiroides (Des- marest, 1822, p. 386; Cuvier, 1821-1824, V, Pt. 2, p. 527). Elephas priscus Goldfuss Nova Acta Acad. Leop. Carol., X, Pt. II, Pl. xutv. (?)Cologne, Germany. Falconer (1868, II, p. 94) retained the name Elephas (Loxod.) priscus [ = Hesperoloxo- don antiquus of present Memoir] for undoubted Pleistocene fossil teeth from Gray’s Thur- rock and elsewhere, although he states (p. 95) that the actual type of Goldfuss, 1821, “conveyed to my mind a corresponding impression that the molar was probably of modern origin.” Subsequently Falconer (cf. Leith Adams, 1877-1881, pp. 1, 2) abandoned the name Elephas priscus Goldfuss, and it is now considered (fide Pohlig) as a synonym of Loxodonta africana. Type loe.: Near 1385 Reference in Present Memoir Turicius tapiroides Cordillerion andium Cuvieronius humboldtii Mastodon americanus Cordillerion andium Cuvieronius humboldtii Trilophodon angusti- dens (?)Turicius tapiroides Mammonteus primi- genius Mastodon americanus Trilophodon anqusti- dens Trilophodon angusti- dens minutus Cuvieronius humboldtii Mammonteus primi- genius Turicius tapiroides Loxodonta africana 1823 1824 1824 1824 1824 1824 1825 1828 1828 1828 1829 1829 OSBORN: THE PROBOSCIDEA Name Author Mastodon cordillerarum Desmarest ‘‘Mammalogie,’’ Seconde partie, p. 385. Cf. Mastodonte des Cordilitres Cuvier, 1806, and Mastotherium hyodon Vischer, 1814, above, also M. andium Cuvier, 1824, below, this list. Mastodon minus Desmarest Op. cit., p. 386. Cf. Petit mastodonte Cuvier, 1806, above, and Mastodon minutus Cuvier, 1824, below, this list. Mastodon tapiroides Cuvier (In Desmarest, loc. cit.). Also Cuvier, 1821-1824, ‘“Ossemens Fos- siles,” V, Pt. 2, p. 527. Type loe.: Montabusard, France. Type descrip- tion: Cuvier, 1806, Ann. Mus. hist. nat., VIII, p. 411. Type fig.: Cuvier, ibid., Pl. 68 [111], fig. 6. Cf. Petit mastodonte Cuvier, 1806, Mastotheriwm microdon Fischer, 1814, Mastodonte tapiroide Cuvier, 1821, above, this list. Elephas antiquitatis Kriiger ‘‘Geschichte der Urwelt,” p. 832. Type loc.: Thiede, Germany. Type fig.: Breislak, 1820, ‘‘Lehrbuch der Geologie,” II, p. 428. Mastodon maximus Cuvier ‘Ossemens Fossiles,’’ V, Pt. 2, p. 527. Type loc.: Big Bone Lick, Boone County, Kentucky. Cf. Mastodon giganteum Cuvier, 1817, above, this list. Mastodon Andium Cuvier Loc. cit. Type loec.: Near Voleano of Imbaburra, Quito, Ecuador. Type description: Cuvier, 1806, Ann. Mus. hist. nat., VIII, pp. 411, 4138. Type fig.: Cuvier, 1806, zbzd., Pl. 67 [1], fig. 1. Cf. Mastodonte des Cordilitres Cuvier, 1806, above, this list. The name Mastodon andium is erroneously dated by Trouessart and others as 1806. We cannot find the name earlier than 1824. It is thus technically preoccupied by Masto- therium hyodon Fischer, 1814, and Mastodon cordillerarum Desmarest, 1822, but the name Mastodon andium Cuvier is adopted, following de Blainville and Falconer (see footnote on p. 122 of Vol. I of the present Memoir). Mastodon humboldii Cuvier ‘“Ossemens Fossiles,” V, Pt. 2, p. 527. Cf. Mastodon humboldtii Cuvier, in Desmarest, 1818-1824, above, this list. Mastodon minutus Cuvier Loc. cit. Type loc.: Saxony. Typedescription: Cuvier, 1806, Ann. Mus. hist. nat., VIII, p. 411. Type fig.: Cuvier, zbzd., Pl. 67 [nm], fig. 11. Cf. Petit mastodonte Cuvier, 1806, M. minor Cuvier, in Desmarest, 1818, above, and M. parvus Cuvier, in Hays, 1834, below, this list. Mastodon turicense Schinz ‘Naturgesch. u. Abbild. d. Saugethiere,” p. 278. Type loc.: Elgg, Canton Zurich, Switzerland. Type fig.: Schinz, 1833, Denk. schweiz. Ges. Naturw., I, Abth. 2, Taf. 1, fig. 1, and p. 59. Elephas meridionalis Nesti Nuoy. Giorn. Lett., XI, No. 24, p. 211. Type loc.: Val d’Arno supérieure, northern Italy. Type figs.: Nesti, zbzd., Tav. 1, figs. 1, 2 (lectotype cranium ©); Tav. 1, fig. 3 (cotype cranium A). Mastodon arvernensis Croizet and Jobert ‘‘Ossemens Fossiles .. . Puy-de-Dome,” p. 188. Type loe.: Perrier, Auvergne, France. Cotype figs.: Croizet and Jobert, op. cvt., Pl. 1, figs. 1-4, Pl. 1} fig. 7. Mastodon latidens Clift Trans. Geol. Soc. London, (2), II, Pt. III, pp. 369-375. Type loc.: Near Yenangyaung, Irrawaddy River, Burma. Lectotype and cotype figs. : Clift, zbid., Pl. xxxvu, fig. 1, and Pl. xxxvm, fig. 1. Mastodon elephantoides Clift Ibid., pp. 372, 373. Type loc.: Near Yenangyaung, Irrawaddy River, Burma. Lectotype fig.: Clift, zbid., Pl. xxxvuitt, fig. 2. Deinotherium giganteum Kaup Isis, |X XII], Heft IV, p. 401. Type loe.: Eppelsheim, Germany. Type fig.: Kaup, zbid., Taf. 1. Elephas mammonteus Fischer de Waldheim Nouv. Mém. Soc. Imp. Nat. Moscou, I, pp. 285, 286. Hab.: Russia. Cf. Elephas mammonteus Cuvier (MS. 1796, published 1799) above, this list. Reference in Present Memoir Cordillerion andium Trilophodon angusti- dens minutus Turicius tapiroides Hesperoloxodon antiquus germanicus Mastodon americanus Cordillerion andium Cuvieronius humboldtii Trilophodon angusti- dens minutus Turicius turicensis Archidiskodon meridion- alis Anancus arvernensis Stegolophodon latidens Stegodon elephantoides Deinotherium gigan- leum Mammonteus primi- genius Year 1829 1829 1829 1829 1829 1830 1830 1831 1831 1831 1831-1833 1832 1832 1832 1832 1832 1832 1832 NOMENCLATURE OF THE PROBOSCIDEA Name Author Hlephas paniscus Fischer de Waldheim bid., pp. 285, 289. Hab.: Volga, Russia. Written also panicus. Elephas periboletes Fischer de Waldheim — [bid., pp. 285, 290. Type loc.: Podolia, Russia. Type fig.: Fischer, zbid., Tab. xvi, fig. 1. Also written proboletes. Elephas pygmexus Fischer de Waldheim Jbid., pp. 285, 292. Type loc.: District of Calomna and of Zwenigorod, Russia. Type fig.: Fischer, /béd., Tab. xvn, fig. 2. Elephas campylotes Fischer de Waldheim Jbid., pp. 285, 291. Type loe.: Borders of the Bug, Russia. Hlephas Kamenskii Fischer de Waldheim [bid., p. 276. Hab.: Siberia. Mammuthus borealis Burnett Quart. Journ. Sci., London, p. 352. T. [Tetracaulodon| Mastodontoideum Godman Trans. Amer. Phil. Soc., N. 8., III, pp. 478-485. Type loc.: Near Newburgh, Orange County, New York. Type figs.: God- man, 7bd., Pls. xvi, Xvi. M. |Mastodon| intermedius Eichwald “Zoologia Spec.,” III, p. 361. Type loc.: Volhynia, Russia. Type: Maxilla (no figure found). Paratype: Jaw (supplementary de- scription, 1835, Nova Acta Acad. Leop. Carol., XVII, p. 737, Pls. uv, Lex). Dinothertum maximum Kaup ‘‘Fossil Saugeth. Rheinhessens.”’ (Fide de Blainville, 1839-1864, “Ostéographie,” p. 18, on Dinotherium; fide von Meyer, 1832, ‘‘Paleologica z. Geschichte der Erde,” p. 78). Type loc.: Eppelsheim, Germany. Deinothertum Bavaricum von Meyer Neues Jahrb. Min., p. 297. Type loe.: Gmiind, Bavaria. Supplementary description: von Meyer, 1832, Nova Acta Acad. Leop. Carol., XVI, Pt. Il, p. 487, Tab. xxxtv, figs. 12-15, Tab. xxxv1. Elephas primordialis Brayley Cf. Elephas primordialis Blumenbach, in Tilesius, 1815, above, this list. Elephas primordialis Brayley, 1831 (without name); in Bronn, 1833, who attributes the name to Brayley. Mammut Sibiricum von Meyer ‘‘Paleologica z. Geschichte der Erde,” p. 64. Von Meyer attributes this name to Schlotheim. Dinotherium maximum von Meyer (Ex MS. Kaup), ‘“Paleologica z. Geschichte der Erde,” p. 78. Type loe.: Eppelsheim, Germany. Dinotherium Cwvieri Kaup ‘Description d’Ossements Fossiles,’’ Cahier I, pp. 2, 14. Hab.: Comminge, Carlat-le-Comte, Chevilly, France. Tetracaulodon longirostre Kaup Isis, [XX V], Heft VI, p. 628. Type loc.: Eppelsheim, Germany. Type fig.: Kaup, zbid., Taf. x1, fig. A. At first thought to be referable to Mastodon angustidens. Kaup, however, substituted the name T'etracaulodon longirostre, which subsequently (1835, p. 65) he changed to Masto- don longirostris (see below, this list) by which name it has since been known. Its reference in the present Memoir is to T'etralophodon longirostris. Mastodon ohioticum (In Bronn, Neues Jahrb. Min., p. 355. In Gervais, 1848-1852, Zool. Pal. Frangaises, I, p. 187.) Elephas brachyramphus Brandt Mém. Acad. Imp. Sci. St. Pétersb., (6), II, Math. et Phys., Bull. Sci., No. 2, p. xi. Type loc.: Mer glaciale, near mouth of Lena River, Siberia. Type: Adams skeleton in the Zoological Museum of the Academy of Sciences, Leningrad, U.S. 8. R. Figured by Tilesius, 1815, Mém. Acad. Imp. Sci. St. Pétersb., (V), V, Tab. x. Described by Adams, 1808, Phil. Mag. (Tilloch), X XIX, and Tilesius, 1815. Cf. Elephas primxvus Blumenbach, in Adams, 1808, above, this list. Elephas homotaphrus Brandt Ibid., p. xii (name only). 1387 Reference in Present Memoir Mammonteus primi- genius Mammonteus primi- genius Mammonteus primi- genius Mammonteus primi- genius Mammonteus primi- genius [| Mammonteus(?) primigenius]| Mastodon americanus Anancus intermedius Deinotherium gigan- teum Deinotherium bavari- cum Mammonteus primi- genius Deinotherium gigan- teum Deinotherium cuvieri Tetralophodon longi- rostris Mastodon americanus Mammonteus primi- genus 1388 Year 1832 1832 1832 1832 1832 1833 1834 1834 1834-1843 1834 1834 1834 OSBORN: THE PROBOSCIDEA Name Author Elephas giganteus Brandt Loe. cit. Type loc.: Indigirka River, Siberia. Type: Messerschmidt cranium. Figured by Breyne, 1741, Phil. Trans. Roy.Soc. London, XL, PI. 1, figs. 1, 1; and by Cuvier, 1806, ““Ossemens Fossiles,” Pl. 39, fig. 1, and PI. 41, fig. 11.F. Elephas commutatus Brandt Loe. cit. Type loc.: ‘“Bords du Volga.” Type: “Crane du Comte Mussin Pusehkin.”’ Figured by Cuvier, 1825, ‘“‘Recherches Ossemens Fos- siles,”’ 3d edition, I, Pl. rx, fig. 7, and p. 179. Elephas stenotoechus Brandt TIbid., p. xiii. No locality given. ‘Type in Zoological Museum of the Academy of Sciences, Leningrad, U.S. 8. R. Elephas platytaphrus Brandt Ibid., p. xiv. No locality given. Type in Zoological Museum of the Academy of Sciences, Leningrad, U. 8. 8. R. Type fig.: Cuvier, 1825, ‘Recherches Ossemens Fossiles,’’ 3d edition, I, Pl. rx, figs. 5, 6. Elephas affinis Brandt Ibid., p. xiv. The history of this specimen (a skeleton in the Zoological Museum of the Academy of Sciences, Leningrad, U. 8. 8S. R.) is given in Oken’s Isis of 1832, XXV, Heft X, pp. 1111— 1114, by a writer signing himself “Ein Naturforscher in St. Petersburg.’’ He states that this skeleton together with other materials was originally discovered in a hole beneath an oven of a house in St. Petersburg which was built upon ground formerly a swamp and which was torn down in 1828. This anonymous reviewer refers to the author of the description as “Herr B” and expresses doubt as to the specimen being a fossil. See also Neues Jahrb. Min., 1838, p. 611. Dinotherium medium Kaup Neues Jahrb. Min. p. 419. Type loc.: Eppelsheim, Germany. Type fig.: Kaup, zbid., Taf. vu, fig. 1, reproduced in Kaup, 1835, ‘‘De- scription d’Ossements Fossiles,” Cahier IV, Add. Tab. 1. Supplementary description: Kaup, 1833, Neues Jahrb. Min., p. 509. M. |Mastodon\ parvus Cuvier (In Hays, Trans. Amer. Phil. Soc., N.8., IV, p. 333.) Cf. Petit mastodonte Cuvier, 1806, of Saxony, above, this list. M. |Mastodon| Borsoni Hays Ibid., p. 334. Type loc.: Near Villanova d’Astica, Piedmont, Italy. Type fig.: Borson, 1823, Mem. Accad. Sci. Torino, XX VII, Tav. 1 (as M. giganteum). M. |Mastodon| Cwvieri Hays Ibid., pp. 322, 323, 334. Fragment of lower jaw in cabinet of the American Philosophical Society. Type fig.: Hays, zbid., Pl. xxiv. M. |Mastodon|\ Jeffersoni Hays Ibid., pp. 323, 334. Fragment of right lower jaw and portion of left ramus in cabinet of the American Philosophical Society. Type fig.: Hays, ibid., Pl. xxv. T. [Tetracaulodon\ Collinsii Hays Ibid., pp. 326, 327, 334. Portion of right lower jaw in cabinet of the American Philosophical Society. Type fig.: Hays, ibid., Pl. xxvut. T. |Tetracaulodon| Godmani Hays Ibid., pp. 327, 334. Fragment of right lower jaw in cabinet of the American Philosophical Society. Type fig.: Hays, zbéd., Pl. xx1x. Mastodon Chapmani Hays Ibid., explanation of Pl. xxur, figs. 3 and 4, p. 338 (without name); name used by Hays, 1843, Proc. Amer. Phil. Soc., II, p. 270. Type loe.: Unrecorded locality in the United States. Type lost or misplaced. Type fig.: Hays, 1834, Trans. Amer. Phil. Soc., N.S., IV, Pl. xx, figs. 3 and 4. Elephas macrorynchus Morren “Mém. Ossemens Fossiles Eléphans Belg.,”’ p. 23. Type loc.: Tamise, Belgium. Type fig.: Morren, op. cit., Pl. 1, figs. 1-4. Mastodon dubsus Kaup and Scholl ‘Verzeichniss der Gypsabgiisse von den ausgezeichnetsten urweltlichen Thierresten des Grossherzoglichen Museum zu Darmstadt,” p. 22. Type loc.: Eppelsheim, Germany. Mastodon dubius is a synonym of M. longirostris (fide Kaup, 1835, p. 77). Mastodon grandis Kaup and Scholl Op. cit., p. 25. Type loc.: Eppelsheim, Germany. Type fig.: Kaup, 1835, ‘Description d’Ossements Fossiles,” Pl. xv, fig. 9. Mastodon grandis is a synonym of M. longirostris (fide Kaup, 1835, p. 77). Reference in Present Memoir Mammonteus primi- genius Mammonteus primi- genius Mammonteus primi- genius Mammonteus primi- genius Deinotherium medium Zygolophodon borsoni Mastodon americanus Mastodon americanus Mastodon americanus Mastodon americanus Stegomastodon chap- mant Mammonteus primi- genius Tetralophodon longi- rostris Tetralophodon longi- rostris Year 1835 1835 1835 1835 1835 1836 1836 1836 1836 1838 1840(?) 1841 1841 1841 1841 1841 NOMENCLATURE OF THE PROBOSCIDEA Author Elephas odontotyrannus Eichwald Nova Acta Acad. Leop. Carol., XVII, p. 723. Type loe.: Banks of Nieman River, Dist. of Novogrodek, gouv. Vilna, Russia. Type fig.: Eichwald, zbed., Pl. uxin, figs. 1, 2. Name Mastodon podolicum Eichwald Ibid., p. 736. Typeloc.: Near Tultschin, Podolia, Russia. Type figs.: Eichwald, zbzd., Pls. tvr, Lv1. See Vol. I, p. 85 (footnote) of the present Memoir. Dinotherium proavum Eichwald TIbid., p. 741. Hab.: Podolia, Russia. ibid., Pl. ux, figs. 1-5. First mentioned as 7.(Tapirus) proavus by Eichwald in 1827 (Naturhistorische Skizze, p. 239—not available to the present author). Described as Tapirus proavus in ‘‘Zoologia Specialis,” 1831, III, pp. 353, 360. See de Blainville, 1839-1864, ‘Du Dinotherium,”’ pp. 12; 19. Dinotheriwm uralense Eichwald Jbid., p. 742. Hab.: Ural Mts. See Pallas, 1777, p. 213, Tab. rx, fig. 4; also de Blainville, 1839-1864, p. 19; and Lar- tet, 1859, p. 482. Type fig.: Eichwald, Mastodon longirostris Kaup ‘Description d’Ossements Fossiles,”’ Cahier IV, p. 65. Cf. Tetracaulodon longirostre Kaup, 1832, Mastodon dubius Kaup, 1834, and Mastodon grandis Kaup, 1834, above, this list. Dinotherium secundarium Kaup (In Lartet, Bull. Soe. géol. France, (1), VII, p. 218.) loe.: Simorre, France. Cited also by de Blainville, 1839-1864, ‘“Du Dinotherium,” p. 19. Original description by Kaup not found by the present author. Type M. tapiroides-minus Lartet Loc. cit. Type loe.: Simorre, France. M. angustidens minus Lartet Loc. cit. Type loc.: Simorre, France. M. |Mastodon] Sivalensis Cautley Journ. Asiatic Soc. Bengal, V, p. 294. Type loe.: Doab Canal, vicinity of Nahun, India. Type fig.: Cautley, ibzd., Pl. x1, figs. 2, 3. E. [Elephas] jacksoni Mather “First Annual Rept. Geol. Survey Ohio,” pp. 96, 97 (notice of dis- covery, without name); Amer. Journ. Sci., (1), XX XIV, p. 358 (description, without name); zbdd., pp. 362-364 (final description, with name). Type loc.: Salt Creek, Jackson County, Ohio. Type fig.: Mather, Amer. Journ. Sci., (1), XXXIV, p. 363, fig. A. Present location of type specimen unknown. Missourium kochii Koch ‘‘Fossil Remains,” p. 2. Type loc.: 22 miles south of St. Louis, Jeffer- son County, vicinity of Sulphur Springs, Missouri. The type skull was found in May, 1839, and first described (Amer. Journ. Sci., (1), XXXVII, pp. 191, 192) as Koch’s Missourian; in a subsequent paper (‘Fossil Remains,” supposedly of date 1840, p. 2) Koch published a supplementary description, assigning the name Missourium kochii. See Horner, 1840, Proc. Amer. Phil. Soc., I, pp. 279-283, for remarks on the collection of Koch. Leviathan Missourti Koch ‘‘Description of the Missourium, or Missouri Leviathan,” p. 13. Type loc.: Near the shores of the river La Pomme de Terre, a tributary of the Osage River, Benton County, Missouri. Skeleton excavated in March, 1840, and regarded by Koch as belonging to the same genus, namely, Missouriwm; he named it, however, Missouri Leviathan (1841, p. 13). Changed in 1843 to Missourium Theristocaulodon (see below, this list). T. (Tetracaulodon| Osagii Koch ‘Description of the Missourium, or Missouri Leviathan,” another edition, p. 1. Hab.: Missouri. Hab.: Missouri. Tetracaulodon Tapyroides Koch Loc. cit. Elephas indicus Isodactylus Hodgson Journ. Asiatic Soc. Bengal, N. S., X, p. 907 (name without definition). Elephas indicus Heterodactylus Hodgson Loc. cit. (name without definition). 1389 Reference in Present Memoir Mammonteus primi- genius Deinotherium podoli- cum Deinotherium proavus Deinotherium uralense Tetralophodon longi- rostris Deinotherium secun- darium(?) Turicius tapiroides- minus Trilophodon angusti- dens minutus Pentalophodon siva- lensis Parelephas jacksoni Mastodon americanus Mastodon americanus Mastodon americanus Mastodon americanus Indeterminate Indeterminate 1390 Year 1841 1841 1842 1842 1842 1842 1842 1843 1843 1843 1844 1845 1845 1845 OSBORN: THE PROBOSCIDEA Name Author Dinotherium Kénigii Kaup “Akten der Urwelt,” pp. 49, 50. Type loc.: Eppelsheim, Germany. Dinotherium minutum von Meyer Neues Jahrb. Min., p. 459. Type loc.: Mésskirch, Germany. T. [Tetracaulodon\ kochii Koch Proc. Geol. Soc. London, III, p. 715. Hab.: Missouri. Koch, 1842, p. 715: “It does not require a close examination. . . to perceive that the animal to which these remains belonged was neither male, female, nor young Mastodon, or Missourium, the whole inner and outer conformation of the upper tusks showing that they were calculated to be used in harmony with the lower tusk in grubbing and rooting.” Tetracaulodon Haysii Grant Tbid., ILI, Pt. Il, p. 771. Cf. Koch, 1845, p. 29. Tetracaulodon Bucklandi Grant Loe. cit. Cf. Koch, 1845, p. 29. Mastodon Brasiliensis Lund (In Lesson, ‘“‘Nouv. Tabl. Régne Animal,” p. 157.) Type loc.: Valley of the Velhas River, Province of Minas Geraes, Brazil. Mastodon Brasiliensis, the specific name of which is attributed to Lund by Lesson, appears as ‘‘Mastodon sp.’”’ in Lund (1889, p. 133), compared with M. andium and M. humboldtii but net figured by Lund. See Lund, 1839, pp. 117, 129, 130, 133 (‘12. Mastodon sp.’”’). E. [Elephas| americanus De Kay ‘Natural History of New York,” p. 101. Type loc.: Irondi- quoit River, Monroe County, near Rochester, New York. Was in cabinet of the Lyceum of Natural History, New York, but was destroyed by fire. Type fig.: De Kay, op. cit., Pl. xxxm, fig. 2. Missourium Theristocaulodon Koch 15, frontispiece. First named by Koch, 1841, Leviathan Missouri. “Description of the Missourium Theristocaulodon,”’ pp. 9, Leviathan missouriensis Koch Op. cit. (in title). Corrected form of Leviathan Missourii but changed to Missourtwm Theristocaulodon in 1843 (see preceding item). Dinotherium Australe Owen Ann. Mag. Nat. Hist., XI, pp. 329-332, figs. 1, 2. Type loc.: Darling Downs, Australia. See Owen, 1848, ibid., pp. 7-12, especially p. 9, figs. 2 and 3, for first description of femur and molar but without name. See also Owen, 1844, ibid., XIV, p. 268 (Dinotherium australe = Diprotodon australis). Mastodon australis Owen Ibid., XIV, p. 271, figs. 1 and 2 on p. 269. Type loc.: Specimen brought by native to Count Strzlecki in Australia, from cave further in the interior than the ossiferous caves of the Wellington valley. Probably a South American specimen accidentally misplaced with Australian speci- mens by this traveler (Count Strzlecki). See Falconer, 1857, table opposite p. 319, and 1868, II, pp. 271-276; also Jack and Etheridge, 1892, p. 688. Mastodon arborense Koch ‘Die Riesenthiere der Urwelt,”’ p. 18. Apparently erroneously ascribed to von Meyer. See von Meyer and Géppert, 1848, p. 705: “|Mastodon| Arborense {nusquam Myr.| Koch Riesenthiere 18.’ Also p. 706: “Mastodon Arborense (Mey.) Koch ferr. typ. ?pro] =Mastodon Arvernensis.”’ Mastodon rugatum Koch Op. cit., p. 20. See also Giebel, 1847, p. 202. Dinotherium angustidens Koch Op. cit., p. 41. Type loe.: Compubay [Cambay(?), ef. pp. 85 and 90 of present Memoir], India. See von Meyer and Géppert, 1848, pp. 424, 425: “Dinotherium. . . angustidens Koch = Mastodon angustidens Cuv.” Reference in Present Memoir Mastodon americanus Mastodon americanus Mastodon americanus Cuvieronius brasili- ensis Mammonteus primi- genius americanus Mastodon americanus Mastodon americanus DiproTropON DIpROTODON Mastodon americanus Mastodon americanus Deinotherium sp.(?) NOMENCLATURE OF THE PROBOSCIDEA Year Name Author 1845 Elephas kamensis de Blainville ‘‘Ostéographie,” p. 202. De Blainville cites this species as Hlephas Kamenskii or kamensis (see Elephas Kamen- ski Fischer, 1829, above, this list). 1845 Elephas africanus priscus de Blainville Op. cit., p. 205. European specimens considered by some palzontologists to be referable to Loxodonta africana, by others as of doubtful determination. Reviewed by Pomel, 1895.1, p. 20. 1845 Elephas ohioticus de Blainville Op. cit., p. 261. Hab.: Ohio River. 1845 Dinotherium intermedium de Blainville Op. cit., Atlas, Pl. m1. Hab.: France. 1845 *H. [Hlephas| primigenius sibiricus de Blainville Op. cit., Atlas, Pl. 11. Hab.: Siberia. See also Depéret and Mayet, 1923, pp. 183-201. 1845 *H. [Elephas] primigenius germanicus de Blainville Op. cit., Atlas, Pl. m1. Hab.: Germany. 1845 *E. [Hlephas] indicus ceylanicus de Blainville Op. cit., Atlas, Pl. mi. Hab.: India, Ceylon. Living form. 1845 *E. [Hlephas] indicus bengalensis de Blainville Op. cit., Atlas, Pl. mr. Hab.: India, chiefly Bengal and Assam. Living form. Compare Falconer and Cautley’s figures (1846 [1847, Pl. xuit]) of E. indicus (Dauntela var.) and #. indicus (Mukna var.), also Falconer (1867, p. 57) in which he presents a de- tailed comparison of the measurements between the Mukna and Dauntela varieties of Elephas indicus. He adds (p. 58): ‘‘The plates of teeth in the Mukna variety slope greatly backwards and are excessively and finely crimped; those of Dauntela are much less crimped.” 1845 *H. [Hlephas| primigenius meridionalis de Blainville Op. cit., Atlas, Pl. 11. 1845 Dinotherium |gig.| majus de Blainville ‘Ostéographie, Du Dinotherium,” p. 60. See Weinsheimer, 1883, p. 210. 1845 Dinotheriwm |gig.| medium de Blainville Loc. cit. See Weinsheimer, 1883, p. 210. 1845 Dinotherium [gig.| minus de Blainville Loc. cit. See Weinsheimer, 1883, p. 210. 1845 Dinotherium indicum Falconer Quart. Journ. Geol. Soc. London, I, p. 361. Type loe.: Perim Island, India. Type fig.: Falconer, ibid., Pl. x1v, figs. 1, la. Falconer, 1845, pp. 370, 371: ‘The Dinotherium of Eppelsheim is known to range through a very wide difference of size, dependent on sexual or individual peculiarities, and several nominal species, chiefly founded upon this character, have been described by authors. But Dr. Kaup informs me, that he now admits but two species, D. gigantewm and D. Kenigi, as he regards all the rest, such as D. Cuvieri, D. Bavaricum, D. proavum, &c., to be merely dwarfed varieties, or females of D. giganteum. M. De Blainville has arrived at nearly the same conclusion in his Ostéographie. It would be unsafe, therefore, to found any opinion regarding the Indian fossil merely on a difference of size.” 1845 Elephas Hysudricus Falconer and Cautley ‘‘Fauna Antiqua Sivalensis,’’ Pl. 1, fig. 3a (type), fig. 1846 3b (paratype). Hab.: Siwalik Hills, India. Type description: Falconer and Cautley, 1846, ‘‘“Fauna Antiqua Sivalensis,”’ letterpress, p. 41. 1845 E. [Elephas| planifrons Falconer and Cautley Op. cit., Pl. 11, fig. 5a (leetotype), fig. 5b (cotype). 1846 Hab.: Siwalik Hills, India. Type description: Falconer and Cautley, 1846, op. cit., letterpress, p. 38. 1845 Elephas insignis Falconer and Cautley Op. cit., Pl. 11, fig. 6a (lectotype), fig. 6b (cotype). Hab.: 1846 Siwalik Hills, India. Type description: Falconer and Cautley, 1846, op. cit., letterpress, pp. 37, 38. “*These perhaps may be regarded as geographic designations rather than as subspecies.”’ 1391 Reference in Present Memoir Mammonteus primi- genius Mastodon americanus Deinotherium inter- medium Mammonteus primi- genius Elephas indicus ceylanicus Elephas indicus bengalensis Deinotherium gigan- tewm Deinotherium gigan- teum Deinotherium gigan- teum Deinotherium indicum Hypselephas hysudricus Archidiskodon plani- frons Stegodon insignis- ganesa 1392 Year 1845 1846 1846 1846 1846 1847 1857 1867 1868 1846 1846 1846 1846 1847 1857 1867 1847 1847 OSBORN: THE PROBOSCIDEA Author Elephas ganesa Falconer and Cautley Op. cit., Pl. m, fig. 7a. Hab.: Siwalik Hills, India. Type description: Falconer and Cautley, 1846, op. c7t., letterpress, p. 45. Name (In Falconer and Cautley, 1846, op. cit., letterpress, p. 13.) Falconer attributed Hlephas minimus to Nesti, 1808, but the present author was unable to find this name in either of Nesti’s articles of 1808 or 1825. Elephas minimus Nesti Elephas Namadicus Faleoner and Cautley Op. cit., letterpress, p. 45. Hab.: Valley of the Nerbudda River, India. Type figs.: Faleoner and Cautley, 1847, op. cit., Pls. xir.a, and x11.B, figs. 1 and 3. Supplementary description: Falconer, 1867, ‘Description of the Plates in the Fauna Antiqua Sivalensis,” p. 15. FE. [Elephas| priscus? Falconer and Cautley Op. cit., letterpress, p. 45 (name only); also Pl. xrv, figs. 7, 7a, 7b. KE. [Elephas] (Loxod.) priscus (Goldf.) Falconer, 1857, Quart. Journ. Geol. Soc. London, XIII, pp. 345, 346, and table opp. page 319. Type loc.: Gray’s Thurrock, England. Type description: Falconer, 1867, ‘“De- scription of the Plates in the Fauna Antiqua Sivalensis,” p. 21. Type fig.: Falconer and Cautley, 1847, op. cit., Pl. xtv, figs. 7, 7a, 7b. Supplementary description: Falconer, 1868, ‘‘Palszeontological Memoirs,”’ II, p. 94, and PI. vil, as Elephas (Loxod.) priscus. Name preoccupied by Goldfuss, 1821 (see above, this list). E. |Elephas| bombifrons Falconer and Cautley Op. cit., letterpress, p. 46. Hab.: Siwalik Hills, India. Lectotype fig.: Falconer and Cautley, 1847, op. cit., Pl. xxv1; cotype figs.: Pls. xxv, XXVIII. E. {Elephas] cliftii Falconer and Cautley Op. cit., letterpress, p. 47. Type loc.: Near Yenang- yaung, Irrawaddy River, Burma. Type fig.: Clift, 1828, Trans. Geol. Soe. London, (2), II, Pt. III, Pl. xxxrx, fig. 6, as M. elephantoides. Mastodon brevirostre Gervais and de Serres Ann. Sci. Nat., (3), V, p. 268. Type loc.: Montpel- lier, Hérault, France. Cotype figs.: Gervais, 1859, ‘‘Zoologie et Paléontol- ogie Frangaises,”’ Deuxiéme ed., Pl. 1, fig. 3, Pl. mt, fig. 7 (same as in First Edition of 1848-1852). Until it is positively determined by further research that the ‘“M.’’ brevirostris of Mont- pellier is identical with the “MW.” arvernensis of Auvergne, it seems best to retain this form as a subspecies of Mastodon | Anancus] arvernensis. Elephas minimus Giebel Neues Jahrb. Min., p. 459. burg, northern Germany. See Giebel in Isis, 1845, Heft VII, p. 483, Heft XII, p. 905, also in Fauna der Vorwelt, 1847, I, Abth. I, p. 211, for description but without name. Type loc.: Seveckenberg near Quedlin- Elephas antiquus Falconer and Cautley ‘‘Fauna Antiqua Sivalensis,” Pl. x1m.p. E. (Eueleph.) antiquus Falconer, 1857, Quart Journ. Geol. Soc., London, XIII, Synop. Tab. opp. p. 319. Hab.: Locality not recorded, but undoubtedly England. Type description: Falconer, 1867, ‘Description of the Plates in the Fauna Antiqua Sivalensis,” p. 18. Type fig.: Falconer and Cautley, 1847, “Fauna Antiqua Sivalensis,” Pl. x1r.p, figs. 4, 4a. Elephas antiquus misnamed EH. meridionalis on plate (see Falconer, 1867, p. 18, and 1868, I, p. 438, together with legend), but corrected by Falconer in copy of the ‘Fauna Antiqua Sivalensis’”’ belonging to the British Museum. See also Falconer, 1868, II, pp. 176-188. Mastodon perimensis Falconer and Cautley Op. cit., Pl. xxx1, figs. 9, 9a. Type loc.: Perim Island, India. Type description: Falconer, 1867, ‘Description of the Plates in the Fauna Antiqua Sivalensis,”’ p. 44. Type fig.: Falconer and Cautley, 1847, op. cit., Pl. xxx1, figs. 9, 9a. Lydekker chose as type Brit. Mus. M.2882 (Pls. xxxvint and xxxrx of the “Fauna Antiqua Sivalensis’’), regarded as the paratype in the present Memoir. Elephas Indicus (Dauntela var.) Faleoner and Cautley Op. cit., Pl. xu, fig. xxut.a. Hab.: India. Type description: Falconer, 1867, op. cit., p. 57; 1868, op. cit., I, p. 477. Living form. Reference in Present Memoir Stegodon insignis- ganesa Palzoloxodon namadi- CUS - Hesperoloxodon antiquus Stegodon bombifrons Stegodon elephant- oides (=cliftiz) Anancus arvernensis brevirostris Hesperoloxodon antiquus Anancus perimensis Elephas indicus var. Dauntela Year 1847 1847 1847 1847 1848 1848 1850 1851 1851 1852 1854 1855 1856 1862 1867 NOMENCLATURE OF THE PROBOSCIDEA Name Author Elephas Indicus (Mukna var.) Falconer and Cautley Op. cit., Pl. xin, fig. xxur.sp. Hab.: India. Type description: Falconer, 1867, loc. cit., 1868, loc. cit. Living form. Mastodon vellavus Aymard Bull. Soe. géol. France, (2), IV, p. 414. Type loe.: Velay, France. No record of figure. While the author stated on p. 193, Vol. I, of the present Memoir that he believed both “M.” vellavus and “M.”’ Vialetii to be referable to ““M.”’ [Zygolophodon| borsoni, he thought it best nevertheless to retain these as subspecific names (p.632). Compare also Falconer, 1868, II, p. 20. Mastodon Vialetii Aymard Jbid., p. 415. Type loc.: Vialette, France. No record of figure. See note above under Mastodon vellavus Aymard, 1847. Elephas sumatranus Temminck “Coup d’CEil Général,” II, p. 91. Type loc.: District of Palem- bang, Sumatra. Type skeletons in Leiden Museum. No record of original type figure (see figure 1182 of present Memoir). Living form. Mastodon Cuvieri Pomel Bull. Soc. géol. France, (2), V, p. 258. Hab.: Gers and l’Orléanais, France. Synonym of Mastodon angustidens Cuv. (fide Trouessart, 1897, p. 700). Mastodon Buffonis Pomel Loc. cit. Type loc.: Auvergne, Perrier, France. Synonym of Mastodon borsoni Hays (fide Trouessart, 1897, p. 705). Elephas affinis Wichwald ‘‘Paleont. Rossii,’ p. 179 (fide Sherborn, 1922, ‘Index Animalium,” p. 124). See also Eichwald, 1853, ‘“‘Lethaea Rossica,” p. 350, Pl. x1, fig. 36. Type loc.: Vicinity of Taganrogh, Russia, near river Mjousse. Original in Mus. Inst. des Mines, Leningrad. Mastodon Simorrense Lartet France. Synonym of Mastodon angustidens Cuv. (fide Trouessart, 1897, p. 700). “Notice sur La Colline de Sansan,” p. 24. Type loc.: Simorre, Mastodon Gaujaci Lartet Op. cit., p. 27. Type loe.: Lombez, France. Synonym of Mastodon angustidens Cuv. (fide Trouessart, 1897, p. 700). Mastodon humboldtius Warren ‘The Mastodon Giganteus of North America,” p. 126. Hab.: South America. Warren states that Cuvier, de Blainville, and Owen regarded this species as referable to angustidens. Elephas Rupertianus Richardson ‘‘Zoology of Voyage of H.M.S. Herald,” pp. 101, 102, 141. Type loe.: Swan River, basin of Lake Winnipeg, Canada. Anancus macroplus Aymard (In Dorlhac, Ann. Soc. Agric. Puy, XIX, for 1854, p. 507). Type loe.: Mt. Coupet, near Puy, France. Synonym of M. arvernensis (fide Lartet, 1859, p. 493). Depéret, 1885, p. 159: “Le nom d’Anancus macroplus donné par M. Aymard aux molaires des sujets adultes du bassin du Puy, trés distinctes des molaires de lait décrites par Croizet et Jobert, doit également disparaitre devant le nom plus ancien d’arvernensis.”’ See also Depéret, 1890, pp. 62, 66. Mastodon pentelicus Gaudry and Lartet Compt. Rend. Acad. Sci., XLIII, p. 273 (name only). Type loe.: Pikermi, Greece. Type description: Lartet, 1859, Bull. Soc. géol. France, (2), XVI, p. 497; Gaudry, 1862, ‘‘Animaux Fossiles,”’ p. 142. Type figs.: Gaudry, 1862-1867, op. cit., Pl. xxit, figs. 1-3, also Pl. xxur. The name Choerolophodon is provisionally retained as a subgenus of Trilophodon for the species M. pentelicus, although the type of the species 7. pentelicus is close to Tetralophodon longirostris. 1393 Reference in Present Memoir Elephas indicus var. Mukna Zygolophodon borsoni vellavus Zygolophodon borsont vialetir Elephas indicus sumatranus Trilophodon angus- tidens cuvieri Zygolophodon borsont buffonis Trilophodon angusti- dens Trilophodon angusti- dens gaujact Cuvieronius humboldtii Mastodon americanus rupertianus Anancus arvernensis Trilophodon (Choero- lophodon) pentelicus 1857-1859 1857 1857 1857-1868 1858 1858 1858 1859-62 OSBORN: THE PROBOSCIDEA Author E. {Elephas| giganteus Aymard (In Falconer, Quart. Journ. Geol. Soc. London, XII, p. 321; and Faleoner, 1868, ‘‘Paleeontological Memoirs,” II, p. 20, footnote.) Name M. (Triloph.) Pandionis Faleoner Quart. Journ. Geol. Soc. London, XIII, p. 317 and Synop. Tab. opp. p. 319. Type loc.: Deccan, India [error, probably Larkana Dis- trict of Sind (fide Hopwood, letter, Feb. 10, 1932)]. Supplementary de- scription: Falconer, 1868, ‘Paleontological Memoirs,” I, p. 124. Type fig.: Falconer, 1868, op. cit., Pl. xxxrv, figs. 6, 7. M. (Triloph.) Pyrenaicus Lartet (Ex MS. Lartet, in Falconer, Quart. Journ. Geol. Soc. London, XIII, Synop. Tab. opp. p. 319.) M. [Mastodon] pyrenaicus Lartet, 1859, Bull. Soc. géol. France, (2), XVI, p. 513. Type loc.: Near Ile-en-Dodon, (Haute-Garonne), France. Type fig.: Lartet, 1859, zbid., Pl. xv, fig. 4. E. [Elephas] (Eueleph.) Armeniacus Faleoner Ibid., Synop. Tab. opp. p. 319. Type loc.: Near Khanoos, Province of Erzerum, Armenia. Supplementary description: Fal- coner, 1863, Nat. Hist. Rev., III, pp. 74,75. Type fig.: Falconer, 1863, zbid., Plo mm, fig:)2: E. {Elephas| (Loxod.) priscus (Goldf.) Falconer Ibid., pp. 345, 346, Synop. Tab. opp. p. 319; 1868, ‘““Paleontological Memoirs,” II, p. 94, as Elephas (Loxod.) priscus. Type loe.: Gray’s Thurrock, England. See above, H. [Elephas] priscus? Falconer and Cautley, 1846, 1847. Name preoccupied by Goldfuss, 1821 (see above, this list). E. [Elephas| (Eueleph.) antiquus Faleoner See under Elephas antiquus Falconer, 1847, above, this list. E. [Elephas] (Eueleph.) Columbi Falconer Quart. Journ. Geol. Soc. London, XIII, Synop. Tab. opp. p. 319. Type loc.: Brunswick canal, near Darien, Georgia. Supple- mentary description: Falconer, 1863, Nat. Hist. Rev., III, pp. 43-52, 114. Type fig.: Falconer, 1863, ibéd., p. 114, Pl. 1; 1868, ‘‘Paleontological Memoirs,” II, Pl. x, fig. 1. Neotype loc.: Phosphate Beds near Charleston, S. C. Mastodon atticus Gaudry and Lartet (In Wagner, Abh. bayer. Akad. Wiss., VIII, Abth. I, Cl. II, p. 140.) Type loe.: Pikermi, Greece. Gaudry, 1862, p. 142: ‘“C’est par une inadvertance de copie que Wagner a employé le nom d’atticus; car il nous attribue ce nom, et le seul que nous ayons proposé est celui de pentelicus.”” {| Mastodon dissimilis Jourdan (MS. 1840) Ann. Soe. Imp. Agric. Lyon, (3), I, p. Ixxxiv. Type loc.: Sadéne Basin, France. Mastodon mirificus Leidy Proc. Acad. Nat. Sci. Phila., X, pp. 10, 28. Type loc.: “Loup Fork of Platte River,’’ Nebraska (Leidy, 1873, Rept. U.S. Geol. Surv. Terr., I, p. 330); possibly near Seneca, Thomas County, Nebraska (fide Hay, 1924, Publ. Carnegie Inst. Wash., No. 322A, p. 100); “‘Pawnee Loup Branch of Platte River, Middle Loup, probably Hooker Co.” (fide Lugn and Schultz, 1934, Neb. State Mus., Bull. 41, I, p. 372). Type fig.: Leidy, 1869, Journ. Acad. Nat. Sci. Phila., (2), VII, Pl. xxv, figs. 1, 2. Cf. Rhabdobunus mirificus Hay, 1914, below, this list. Elephas imperator Leidy Ibid., pp. 10, 29. Type loc.: Loup Fork of Platte River, Nebraska (Leidy, 1869, Journ. Acad. Nat. Sci. Phila., (2), VII, p. 254; and Hay, 1914, lowa Geol. Surv., Ann. Rept. for 1912, XXIII, pp. 421, 422), possibly Seneca, Thomas County, Nebraska (see Hay, 1924, Publ. Carnegie Inst. Wash., No. 322A, p. 100); ‘Pawnee Loup Branch of Platte River = Middle Loup, proba- bly Hooker Co. [Nebraska]”’ (fide Lugn and Schultz, 1934, Neb. State Mus., Bull. 41, I, p. 373). Type fig.: Leidy, 1869, Journ. Acad. Nat. Sci. Phila., (2), VII, Pl. xxv, fig. 3. Osborn’s neotype: 1922, Amer. Mus. Novitates, No. 41, p. 4, fig. 4, from Guadalajara, Mexico. Elephas terianus Owen Rept. Brit. Assoc. Adv. Sci., 28th meeting, p. Ixxxvi (name); and Blake, 1861, Geologist, IV, p. 470 (name). Type loc.: San Felipe de Austin, Brazos River, Texas. Type description: Blake, 1862, Geologist, V, p. 58. Type fig.: Blake, 1862, zbzd., Pl. tv. Reference in Present Memoir Archidiskodon meridi- onalis(?) Trilophodon pandionis Zygolophodon pyrenai- cus Parelephas armeniacus Hesperoloxodon an- tiquus Parelephas columbi Turicius atticus Anancus arvernensis dissimilis Stegomastodon mirificus Archidiskodon impera- tor Parelephas columbi Year 1861 1861 NOMENCLATURE OF THE PROBOSCIDEA Author Dinotherium levius Jourdan Compt. Rend. Acad. Sci., LILI, p. 1011. Type loe.: Grive Saint- Alban, Isére, France. Name Elephas intermedius Jourdan Tbid., p. 1013. Type loc.: Near Lyons, Rhone Valley, France. No figure recorded, but see Lortet and Chantre, 1872 [1876], Arch. Mus. hist. nat. Lyon, I, frontispiece, plate of referred skeleton (Fig. 944 of present Memoir), also referred molars in the Muséum de Ville, Lyons (Fig. 943 of present Memoir). Very similar to Elephas trogontherti Pohlig (fide Depéret letter, August 26, 1921). 1862-1868 Hlephas Melitensis Falconer ‘The Parthenon,” p. 780. See Falconer, 1868, ‘‘Palgont. Mem., 1867 1867 1868 1868 1868 1868 1868 1868 1869 1870 II, pp. 292, 299, 307, 308. Type loc.: Zebbug Cave, Malta. Type fig.: Falconer, 1868, op. cit., Pl. x1, figs. 1, la. Elephas falconeri Busk Trans. Zool. Soc. London, VI, Pt. V, p. 251. Type loc.: Zebbug Cave, Malta. Type fig.: Busk, zbid., Pls. x~rx, L, LI (not figured in present Memoir). See Falconer, 1868, II, p. 292, footnote, where it is stated by the editor that this and other Maltese specimens had been identified by Falconer as early as July, 1860. Mastodon virgatidens von Meyer Paleontogr., XVII, p. 61. Type loc.: Near Fulda, northeast of Frankfort, Germany. Type fig.: von Meyer, zbid., Taf. 1v, figs. 1-5. Mastodon Andaranus Faleoner ‘“‘Paleont. Mem.,’” I, p. 124, footnote. Type loc.: Deccan, India. See Vol. I, p. 267, of the present Memoir. Dinotherium Perimense Falconer Op. cit., 1, p. 415 (name only). Type loc.: Perim Island, India. Dinotherium Pentapotamicum Falconer Op. cit., I, p. 5. See Dinotherium pentapotamize Lydekker, 1876, below. (In Falconer, op. cit., Il, pp. 104, 105). See note under Elephas minimus Nesti, 1846, above, this list. Elephas minutus Nesti Trilophodon Ohioticus Faleoner Op. cit., I, pp. 176 (footnote), 204. Hab.: North America. Mastodon americanus Leidy Proc. Acad. Nat. Sci. Phila., XX, p. 175. Hab.: North America. See Leidy, 1869, Journ. Acad. Nat. Sei. Phila., (2), VII, pp. 240, 395. Leidy first used the term Mastodon americanus in the 1868 article, as the equivalent of M. ohioticus and M. giganteus of authors. Mastodon obscurus Leidy Journ. Acad. Nat. Sci. Phila., (2), VII, pp. 244, 396. Type loc.: Near Greensburgh (Greensboro), Caroline County, Maryland. Type fig.: Leidy, ibid., Pl. xxvui, fig. 13. Merrill (1907, Bull. U. S. Nat. Mus., No. 53, p. 45) regarded a third superior molar, r.M3, from Tarboro, North Carolina, as the type of Mastodon obscurus. The present author refers it to Ocalientinus (Serridentinus) obliquidens of this Memoir (see Vol. I, p. 286). Stegodon sinensis Owen Quart. Journ. Geol. Soe. London, XXVI, Pt. I, p. 417. Type loc.: Alleged to be from marly beds vicinity of Shanghai, China. Type fig.: Owen, ibid., Pl. xxvui, figs. 1-3. Stegodon orientalis Owen Ibid., p. 421. Type loc.: (?)Cave near city of Chungkingfoo, Province of Szechuan, China. Type fig.: Owen, zbid., Pl. xxvut, figs. 1-4. Elephas mnaidrz Adams ‘‘Notes of a Naturalist in Nile Valley and Malta,” p. 223. Type loc.: Mnaidra Gap, Malta. Type fig.: Adams, op. cit., Pl. 1, figs. 2, 2a. Supple- mentary description as Elephas mnaidriensis: Adams, 1874, Trans. Zool. Soc. London, IX, Pt. I, p. 116; with figures of the type, Pl. vu, figs. 2, 2a; of the paratype, Pl. vu, fig. 1. 1395 Reference in Present Memoir Deinotherium levius Parelephas intermedius Palzxoloxodon meliten- sis Palxoloxodon falconeri Turicius virgatidens Trilophodon pandionis Deinotherium indicum Mastodon americanus Mastodon americanus Trilophodon obscurus Stegodon sinensis Stegodon orientalis Palzxolorodon mnaidri- ensis 1396 Year 1870 1871 1872 1872 1875 1875-1923 1876 1877 1877 OSBORN: THE PROBOSCIDEA Name Author Elephas Cornaliae Aradas Atti Accad. Gioenia sci. nat. Catania, (8), IV, p. 235. Type loe.: Catania, near the monastery of Santa Chiara, Sicily. Type figs.: Aradas, abid., figs. 1, 2. Mastodon shepardi Leidy Proc. Acad. Nat. Sci. Phila., XXII, p. 98. Type loc.: Dry Creek, Stanislaus County, California. Type fig.: Leidy, 1873, Rept. U.S. Geol. Surv. Terr., I, Pl. xx1, figs. 3, 4. Elephas Indianapolis Foster Proc. Amer. Assoc. Ady. Sei., August, 1872, p. 259 (name only). Same specimen from Indiana for which the name Hlephas Mississippiensis was subse- quently proposed. See next item. Elephas Mississippiensis Foster Nature, VI, p. 443. Hab.: Indiana, exact locality unrecorded. Name changed from Elephas Indianapolis (see preceding item), without description or figure. Mastodon proavus Cope “Synopsis of New Vertebrata from Tertiary of Colorado,” p. 10. Type loe.: Pawnee Buttes, Pawnee Creek, Weld County, Colorado. Type fig.: Cope, 1889, Amer. Naturalist, X XIII, No. 268, p. 202, fig. 6, as Tetrabelodon angustidens proavus. Mastodon productus Cope Proc. Acad. Nat. Sci. Phila., XX VI, pp. 221, 222. Type loc.: Santa Fé marls, New Mexico. Type figs.: Cope, 1877, Rept. U.S. Geogr. Surv. West of 100th Meridian (Wheeler), IV, Pt. II, Pls. uxx, figs. 1-3, Lxx1, fig. 3. Neotype fig.: Frick, 1926, Bull. Amer, Mus. Nat. Hist., LVI, Art. II, fig. 7A. Elephas Ausonius Major (MS. labels). Name in Verri, 1886, Boll. Soc. geol. Ital., V, p. 453. Type loc.: San Romano, Val d’Arno inf., Italy. Type fig.: Depéret and Mayet, 1923, ‘‘Les Eléphants Pliocénes,”’ Pl. x, figs. 1, 2. See Weithofer, 1890, p. 194, footnote, and p. 206; also Depéret and Mayet, 1923, p. 162. Dinotherium pentapotamiz Lydekker Mem. Geol. Surv. India, Paleont. Indica, (X), I, Pt. I, pp. 72, 73. (Fale. Ex MS., fide Lydekker, 1885, “Catalogue of the Remains of Siwalik Vertebrata. . . Geological Department of the Indian Museum, Calcutta,” p. 104.) Type loe.: Near Attock, Indus valley, India. Type fig.: Lydekker, 1876, zbid., pl. rx, figs. 1, 2. Lydekker, 1876, pp. 72, 73: ‘Subsequently, the same naturalist [Falconer] identified two other molar teeth from near Attock (“Pal. Mem.,’ vol. I, p. 414) as belonging to the same genus [Deinotherium); in the note on these specimens in the ‘Paleontological Memoirs’ no specific name was assigned to them, though they were considered to be of too small a size to have belonged to D. (perimense) indicum; these specimens are now in the Indian Museum; they are ticketed with labels in Falconer’s handwriting bearing the name of D. pentapotamiz, which name I have accordingly adopted. .. The specimen is distinguished by its much smaller size from any of the European species; from the first premolar of D. giganteum and D. cuvieri, it is distinguished by the following points: the antero-posterior valley is very much deeper and wider in the Indian form, rendering thereby the inner tubercles more completely conical; the posterior tubercle is mammilliform in the Indian form, whereas it is elongated transversely in the European form.” See Falconer, 1868, II, p. 5, footnote by Editor, in which he states that ‘‘A specimen of the third lower premolar of this species, from the ‘Red Marl’ at Noorpoor, found in Dr. Palconer’s collection, is labelled in his hand-writing, ‘Dinotherium Pentapotamicum, Fale.’ ”’ Mastodon (Trilophodon) Falconeri Lydekker Rec. Geol. Surv. India, X, Pt. II, p. 88. Type loe.: Potwir district, Punjab, India. Supplementary description: Lydekker, 1880, Mem. Geol. Surv. India, Paleont. Indica, (X), I, Pt. V, p. 206. Type fig.: Lydekker, 1880, zbid., Pl. xxx1m1, figs. 1, 4. C. |Cenobasileus] tremontigerus Cope Proc. Amer. Phil. Soc., XVI, pp. 584, 585. Type loce.: Probably Texas. No figure recorded. Genus and species withdrawn by Cope in 1889 (1889.2, p. 207). Reference in Present Memoir Loxodonta cornalie Rhynchotherium shepardi Parelephas(?) missis- sippiensis(?) Serridentinus proavus Serridentinus productus Hesperoloxodon antiqu- us ausonius Deinotherium indicum T etralophodon (Lydek- keria) falconerv Year 1878 1880 1880 1882 1883 1883 1883 1883 NOMENCLATURE OF THE PROBOSCIDEA Author Tetralophodon campester Cope Proce. Amer. Phil. Soc., XVII, p. 225. Type loc.: Republican River beds, Sappa Creek, Rawlins County, Kansas. Type figs.: Cope, 1889, Amer. Naturalist, XXIII, Pls. rx, x; also Cope and Matthew, 1915, ‘“‘Hither- to Unpublished Plates of Tertiary Mammalia and Permian Vertebrata,” Pls. Coe Cody Chedih Codi, Name Mastodon affinis Jourdan (Ex MS. 1859, labels in Lyons Museum), in Lortet and Chantre, Arch. Mus. hist. nat. Lyon, II, p. 308. Type loc.: (?) Elephas primigenius comune Issel Ann. Mus. Civ. Storia Nat. Genova, (1), XIV, p. 153 (as Elephas primigenius). In Zuffardi, 1913, Palewont. ital., XIX, p. 136. Type loc.: Camporosso, near Ventimiglia, Italy. Type figs.: Issel, 7bid., figs. 1-4. Elephas atlanticus Pomel Bull. Soc. géol. France, (3), VII, p. 51. Type loc.: Ternifine, near Mascara, Algeria. Cotype fig.: Pomel, 1895, ‘‘Paléontologie Monographies, No.6. Les Eléphants Quaternaires,’”’ Carte Géol. L’ Algérie, Pl. vit, figs. 1, 2. Dinotherium sindiense Lydekker Mem. Geol. Surv. India, Paleont. Indica, (X), I, Pt. V, p. 196. Type loe.: Sind, India. Type fig.: Lydekker, zbid., Pl. xxxt, fig. 4. See Lydekker, 1879, Rec. Geol. Surv. India, XII, Pt. 1, p. 43, for first notice, but with- out name. Elephas (Euelephas) antiquus var. nana Acconci AttiSoc. Toscana Sci. Nat., V, Fasc. I, pp. 146- 150. Type loc.: Cavern near Monti Pisani, Cucigliana, Tuscany, Italy. Type fig.: Acconci, zbid., Tav. rv, figs. 6, 7. Notelephas australis Owen Proc. Roy. Soc. London, XX XIII, p. 448; 1883, Phil. Trans. Roy. Soe. London, CLX XIII, Pt. III, p. 777. Type loc.: Drift deposit of ravine in district of Darling Downs, about 60 miles from Moreton Bay, Queensland, Australia. Type fig.: Owen, 1883, ibid., Pl. 11. Jack and Etheridge, 1892, p. 683: ‘‘Notoelephas [Notelephas of Owen, 1882] australis Owen. . .This genus and species were founded on portions of a tusk indicating a mammal larger than Diprotodon, in fact the largest fossil mammal yet foreshadowed amongst the extinct Australian forms. Sir Richard Owen appears to regard it as a Proboscidian Placental. . . He does not appear to apprehend any connection between this tusk and the molar tooth [Mastodon australis Owen, 1844] formerly described by him, and forming a portion of the late Count P. E. de Strzelecki’s Collection. Prof. Owen remarks that this molar is too large to be associated with the tusk, supposing the latter to have come from the upper Jaw of a full-grown individual of its species. . . Grave doubts have been expressed by several Writers as to the probability of this tooth as an Australian fossil, and it would perhaps be better to expunge it from the list.’ Mastodon Zaddachi Jentzsch Schrift. phys.-dkonom. Ges. Konigsb., Jahrg. XXIII, Abth. 2, p. 202. Type loc.: Thorn, West Prussia. Type fig.: Jentzsch, ibed., Taf. v, figs. 6a, 6b. Leptodon minor Gunn Geol. Mag., Dec. II, N. S8., X, p. 458. Type loc.: Forest bed, Norfolk, England. Type fig.: New Edition of Gunn’s ‘Sketch of Geology of Norfolk, Pl. 1, letter I” (fide Gunn, 1883, loc. cit.). This edition does not seem to have been published (ef. ““Memorials of John Gunn,” 1891, p. v). Leptodon giganteus Gunn Loe. cit. Type loc.; Forest bed, Mundesley, Norfolk, England. De- scription and type figure: Gunn, 1891, ‘“Memorials of John Gunn,” Pl. rv, fig. 2. Same as H#. Gunnii Lartet, 1883, and Elephas giganteus intermedius Gunn, 1891 (see below, this list). E. Gunnii Lartet (In Gunn, 1883, loc. cit.) Type loc.: Forest bed, Mundesley, Norfolk. Type ramus No. 361, Gunn Coll., Norwich Castle Mus. 1703. Description and type fig.: Gunn, 1891, “Memorials of John Gunn,”’ Pl. rv, fig. 2. It will be observed in this quaint description of 1883 that Gunn designates as the type of E. Gunnii Lartet the very same specimen and figure selected as the type of Leptodon giganteus. A revision of these types and specific names applied to the collection of Gunn may be found in F. Leney’s ““Type-specimens in the Norwich Castle Museum,” 1902, Geol. Mag., p. 169. 1397 Reference in Present Memoir Tetralophodon campes- ter Zygolophodon borsoni affinis Mammonteus primi- genius Palzoloxodon atlanti- Cus Deinotherium sindi- ense(?) Hesperoloxodon antiqu- us NaNUs Diprotodon australis Zygolophodon borsoni zaddachi Hesperoloxodon antiquus Hesperoloxodon antiquus Hesperoloxodon antiquus 1398 Year 1883 1884 1884 1884 1884 1885 1885 1887 1886 1886 1886 1896 1887 1887 OSBORN: THE PROBOSCIDEA Name Author Elephas Lamarmorae Major Kosmos, XIII, Jahrg. VII, p.6. Type loe.: Morimentu b. Gonnesa, Sardinia. No figures published. Elephas africanus fossilis Thomas, Philippe Mém. Soe. géol. France, (3), III, No. 2, p. 46. Cited by Trouessart, 1897-1899, p. 708, as found in Algeria. Mastodon (Trilophodon) angustidens Cuy. var. palzindicus Lydekker Mem. Geol. Surv. India, Paleont. Indica, (X), III, Pt. I, p. 19. Type loe.: Near Kamlial, northern Punjab, India. Type fig.: Lydekker, zbzd., Pl. rv, fig. 3. M. [Mastodon] euhypodon Cope Amer. Naturalist, XVIII, p. 525. Type loc.: Trail Canon, south fork of Driftwood Creek, Hitchcock County, Nebraska. Type figs.: Cope, 1889, ibid., XXIII, p. 203, fig. 7, and Pl. xu (erroneous drawing) figured as Tetrabelodon euhypodon. M.|Mastodon| serridens Cope Loc. cit. Type loc.: Texas, probably Clarendon formation. Type fig.: Cope, 1889, ibid., p. 205, fig. 8, figured as Tetrabelodon serridens. Dibelodon tropicus Cope Proc. Amer. Phil. Soe., XXII, p. 7. Type loc.: State of Michoacan, Valley of Mexico. Type fig.: von Meyer, 1867, Paleontogr., XVII, Pl. v1, as M. humboldtii. Uhde Coll. See discussion of this species (Dibelodon tropicus Cope) by Freudenberg, 1922, pp. 128, 129, in which he reproduces the original type figure of Cope from von Meyer and places it at the summit of an ascending series ending with Mastodon oligobunis progressa Freuden- berg. Mastodon perimensis var. sinensis Koken Geol. u. Pal. Abh., III, Heft II, p. 34. Yunnan, China. Type fig.: Koken, zbzd., Pl. xm, fig. 1. Type loe.: Elephas trogontherti Pohlig Zeitschr. deutsch. geol. Ges., XX XVII, Heft IV, p. 1027. Elephas (primigenius bezw. meridionalis) trogontherii Pohlig, 1887, Sitz.-Ber. niederrhein. Ges. Bonn, p. 274. Elephas (primigenius) trogontherti Pohlig, 1887 [1888?], Zeitschr. deutsch. geol. Ges., XX XIX, Heft IV, p. 799. Type loc.: Siissenborn, near Weimar, northern Germany. ‘Type figs.: Pohlig, 1888, Nova Acta Leop. Carol., LITT, p. 193, fig. 79, and p. 195, fig. 82. Cf. Hlephas intermedius Jourdan, 1861, above, this list. Mastodon cautleyi Lydekker Mem. Geol. Surv. India, Paleont. Indica, (X), III, pp. xiv-xix. Type loc.: Perim Island, India. Leetotype fig.: Lydekker, zbid., p. xv, fig. 6. Cotype figs.: Lydekker, 1880, zbid., (X), I, Pt. V, Pl. xu [subsequently referred by the present author to Tetralophodon punjabiensis|; 1884, ibid., (X), III, Pt. V, Pl. xvi, fig. 2; 1886, zbzd., (X), III, p. xv, fig. 5; Falconer and Cautley, 1847, “Fauna Antiqua Sivalensis,”’ Pl. xu, figs. 2, 2a, 3, 3a. Mastodon punjabiensis Lydekker ‘‘Cat. Foss. Mamm. Brit. Mus.,” p. 60. Type loc.: Siwalik Hills, Punjab, India. Lectotype fig.: Lydekker, 1880, Mem. Geol. Surv. India, Paleont. Indica, (X), I, Pt. V, Pl. xxi. Originally referred to Mastodon (Tetralophodon) perimensis. Mastodon (Trilophodon) floridanus Leidy Proc. Acad. Nat. Sci. Phila., XX XVIII, p. 12. Type loc.: Alachua clays, Mixson’s bone bed, near Williston, Levy County, Florida. Type fig.: Leidy and Lucas, 1896, Trans. Wagner Free Inst. Sci., IY Veh, Vain eracy ps B. |Elephas| antiquus var. minor Pohlig Verh. natur. Vereins preuss. Rhein., Jahrg. 44, p. 115. Type loc.: Seville, Spain. Stegodon trigonocephalus Martin Sammlung. Geolog. Reichs-Museums, Leiden, 1* Serie, Beitraige z. Geolog. Ost-Asiens u. Australiens, IV, Heft II, pp. 27, 36. Type loe.: Probably vicinity of Surakarta, Java. Type figs.: Martin, ib7d., Tab. 11, figs. 1, la, and Tab. m1, fig. 1. Reference in Present Memoir Palzxoloxodon lamar- morae Palzxoloxodon atlanti- cus Trilophodon paleindi- cus Blickotherium euhypo- don Serridentinus serridens Cordillerion tropicus Tetralophodon (Lydek- keria) sinensis Parelephas trogontheri Stegolophodon cautleyt Tetralophodon punjab- 1ENSLS Ocalientinus (Serriden- tinus) floridanus Stegodon trigonocepha- lus Year NOMENCLATURE OF THE PROBOSCIDEA Name Author 1887 [1888?] Hlephas (primigenius) trogontherti Pohlig See Elephas trogontherii Pohlig, 1885, 1887, above, 1888 1888 1888 1888 1888 1889 1889 1889 1889 1889 1889 1890 1890 1890 1891 this list. E. [Elephas| (primigenius) Leith-Adamsi Pohlig Nova Acta Leop. Carol., LIII, No. 1, pp. 229, 232. Type loc.: Dornap (Fuhlrott), Thuringia, Germany. Type fig.: Pohlig, zbid., p. 229, fig. 101 c-d. Mastodon bonaerensis Moreno ‘Informe Prelim. progresos Mus. La Plata,’’ pp. 17, 18 (nomen nudum). See Ameghino, 1889, Acta Acad. Nac. Cien, Cordoba, VI, p. 641. Type loc.: Arrecifes, Province of Buenos Aires, Argentina. Not figured by Moreno, but Eduardo Carette figured it in his article “Los Proboscideos Fésiles Argentinos,’’ 1919, Lam. x1, fig. 2, as D. [Dibelodon|] platensis. Mastodon argentinus Ameghino ‘‘Rapidas Diagnosis,” p. 7. Type loc.: Valley of Tarija River, northern part of Argentina, exact locality not recorded. Type fig.: Ameghino, 1889, Actas Acad. Nac. Cien. Cordoba, VI, p. 644. Mastodon platensis Ameghino Loc. cit. Type loc.: San Nicolas de los Arroyos, Province of Buenos Aires, Argentina. Type fig.: Ameghino, 1889, zbzd., p. 641. Mastodon superbus Ameghino Loc. cit. Type loc.: Pergamino, Province of Buenos Aires, Argentina. Type fig.: Ameghino, 1889, zbid., p. 647. Mastodon rectus Ameghino Actas Acad. Nac. Cien. Cérdoba, VI, p. 648. Type loc.: From the Ensenada, near La Plata, Argentina. Type fig.: Ameghino, zbid., p. 648. Elephas (primigenius) Leith-Adamsi Pohlig var. minor Viv. Foss.,” p. 711). Search in Pohlig’s contributions of 1889 does not disclose a definition of this variety, and a letter from Doctor Pohlig (Sept. 10, 1924) does not authenticate var. minor as a sub- specifie term (cf. p. 1150 of the present Memoir). (In Trouessart, 1897-1899, “Cat. Mamm. Tetrabelodon brevidens Cope Amer. Naturalist, XXIII, pp. 198-202. Type loc.: Smith River, Meagher County, Montana. Type fig.: Cope, 7zbid., p. 201, fig. 5. E. |Elephas] primigenius americanus Cope Ibid., pp. 207, 209. Hab.: North America. Elephas primigenius columbi Cope Ibid., pp. 208, 209. Type loc.: ‘Orange sand,” city of Dallas, northeastern Texas. Type figs.: Cope, zbid., Pl. xrv and text fig. 9. Elephas lyrodon Weithofer Jahrb. Geol. Reichsanst., XX XIX, pp. 79, 80. Type loc.: Val d’Arno supérieur, Italy. Type figs.: Weithofer, 1890, Beitr. Pal. Osterr.- Ung., VIII, Taf. m1, fig. 2, Taf. 1v, fig. 2, Taf. v, fig. 1. Compare observations of Pohlig (1891, pp. 314, 334). Stegodon Mindanensis Naumann Zeitschr. deutsch. geol. Ges., XLII, Heft I, pp. 166, 167. Type loc.: Mindanao, Philippine Islands. Type fig.: Naumann, 1887, Abh. Ber. k. Zool.-Anthrop.-Ethnog. Mus. Dresden, No. 6, Taf. I, figs. 1 and 2, as S. trigonocephalus. Stegodon Airdwana Martin Verh. Kon. Akad. Wetensch. Afdeel. Natuurk., Amsterdam, X XVIII, p. 4. Type loc.: Alas-Tuwa, Trinil, Java. Type figs.: Martin, 2bzd., Tab. 1, figs. 1 and 2, Tab. 11, figs. 3 and 4. Mastodon rugosidens Leidy Proc. Acad. Nat. Sei. Phila., XLII, p. 184. Type loc.: Santee beds, Beaufort County, South Carolina. No type figure recorded (see Fig. 115 of present Memoir drawn from cast, Amer. Mus. 14445). Original in Academy of Natural Sciences, Philadelphia. E.. [Elephas] primigenius Blum. var. hydruntinus Botti Boll. Soc. geol. Ital., IX, p. 709. Type loe.: La Grotta Ossifera di Cardamone, Terra d’Otranto, Italy. Type fig.: Botti, zbid., Tav.¥xxv1, figs. 1, la, 2, 2a. 1399 Reference in Present Memoir Mammonteus(?) primi- genius leith-adamsi Cuvieronius bonaeren- sis (=superbus) Notiomastodon argen- linus Cuvieronius platensis Cuvieronius superbus Cuvieronius rectus Rhynchotherium brevidens Archidiskodon meri- dionalis (female) Stegodon (Archidisko- don?) mindanensis Stegodon airdwana Mastodon americanus rugosidens Mammonteus primi- genius hydruntinus 1400 Year 1891 1891 1891 1892 1892 1892 1893 1893 1893 1893 1893 1894 1895 1896 1913 1897 1897 OSBORN: THE PROBOSCIDEA Author E. | Elephas| giganteus intermedius Gunn Same as Leptodon giganteus Gunn, 1883, subsequently named by Lartet (in Gunn, 1883) EB. gunnii (see above, this list). Name E. |Elephas| Nestii Pohlig Nova Acta Acad. Leop. Carol., LVII, pp. 303, 304. Elephas antiquus Nestii n.f., Pohlig, ibid., p. 465. Type loc.: Happisborough, Forest bed (Norfolk), Walton (Essex), Southwold (Suffolk), England. Cotype figs.: None. See Figs. 941, 1041 of present Memoir. Mastodon maderianus Ameghino Revista Argent. Hist. Nat., I, p. 248. Type loc.: Puerto Madero in Buenos Aires, Argentina. D. {Dinotherium| gigantissimum G. Stefaineseu Bull. Geol. Soc. Amer., III, pp. 81, 82. Type loe.: Gaiceana, Tecuciti, Rumania. See Stefiineseu, 1878, pp. 101-104, giving an account of discovery, also 1895, p. 178, Tab. tv and v (referred molars). Mastodon successor Cope Proc. Acad. Nat. Sei. Phila., XLIV, pp. 227, 228. Type loc.: Blanco beds, Texas. Type figs.: Cope, 1893, 4th Ann. Rept. Geol. Surv. Texas, Pl. XVI, figs. 1, la, Pl. xvi, fig. 1, as Dibelodon tropicus. D. {Dibelodon| precursor Cope Amer. Naturalist, XX VI, p. 1059 (name only). Type loc.: Mt. Blanco, Llano Estacado, Texas. Type description: Cope, 1893, 4th Ann. Rept. Geol. Surv. Texas, pp. 64, 65. Type figs.: Cope, 1893, op. cit., Pls. XVIII, XIX. T. |Tetrabelodon| serridens cimarronis Cope 4th Ann. Rept. Geol. Surv. Texas, pp. 18-20. Type loe.: East of Llano Estacado, Texas, north of south fork of Red River. Type fig.: Cope, op. cit., Pl. 111, figs. 2 and 3. Mastodon oligobunis Cope Op. cit., p. 59. Type loe.: Tequixquiac, Valley of Mexico. Type fig.: After Villada, 1903, Ann. Mus. Nac. Mexico, VII, Lam. vit (as Dibelodon Shepardi, Cope). M. |Mastodon] chilensis Philippi Zeitschr. deutsch. geol. Ges., XLV, p. 88. Lake Tagua-Tagua, Chile. Type fig.: Philippi, zbzd., p. 89, fig. 4. Type fig.: Philippi, Mastodon bolivianus Philippi Tbid., p. 89. Type loc.: Ulloma, Bolivia. ibid., p. 89, fig. 2. Mastodon Sahendi Pavlow (MS. labels Hofmuseum in Wien), Mém. Acad. Imp. Sci. St. Pétersb., (8), I, No. 3, p. 16. Type loc.: Maragha, Persia. See Kittl, 1887, p. 320, for associated fauna of Maragha. Paléontologie Monographies, No. 6, Carte Géol. L’ Algérie, pp. 32, 39. Type fig.: Elephas jolensis Pomel Hab.: Algerian seacoast, downward from Krober-Roumia. Pomel, op. cit., Pl. v, figs. 3 and 4. Elephas meridionalis antiquitatis Portis ‘‘Contribuzioni alla Storia fisica del Bacino di Roma, . .” Torino, II, p. 272. Portis’ original description of 1896 was not seen by the present author. In the Rivista Italiana di Paleontologia, II, of the same year, the name appears in a phylogenetic chart on page 331. Zuffardi in Palewontologica Italica, 1913, XIX, p. 155, includes this subspecies in the synonymy of Elephas antiquus Fale. var. trogontherioides. Mastodon angustidens Cuy. mut. ase. pygmxus Depéret Bull. Soc. géol. France, (3), XXV, p. 519. Type loc.: Kabylie, near Isserville, Algeria. Type fig.: Depéret, zbid., PI. XIX, figs. 1-3. Blephas platyrhynchus Graells Mem. Real. Acad. Cien. Exactas, Fis. nat., Madrid, XVII, p. 569. Type loe.: San Isidro del Campo, near Madrid, Spain. Type fig.: Graells, ibid., Lam. xvin, figs. 9a, 10. Type loc.: Near Reference in Present Memoir Hesperoloxodon an- tiquus Parelephas(?) trogon- therii nestii Cuvieronius (?)maderi- anus Deinotherium gigantis- stmum Stegomastodon successor Serbelodon(?) precursor Serridentinus serridens cimarronis Cordillerion oligobunis Cuvieronius chilensis Cordillerion bolivianus Tetralophodon sp. Palzxoloxodon jolensis Phiomia pygmxus Hesperoloxodon antiqu- us platyrhynchus Year 1897 1899 1900 1900 1900 1900 1901 1901 1902 1902 1902 1903 1903 1904 1904 1905 1905 NOMENCLATURE OF THE PROBOSCIDEA Name Author Mastodon aureliense, Cuv. (In Graells, bid., p. 573.) Hab.: Madrid. Mastodon angustidens var. latidens Lankester Geol. Mag., Dec. 1V, N.S., VI, pp. 289-292, PI. x1. Type loc.: Suffolk Crag, England. Figured asa trilophodont Mastodon in Quart. Journ. Geol. Soc., 1870, Pl. 34, figs. 1-4. Elephas maximus sumatranus Lydekker ‘“‘Great and Small Game of India . . . ,” p. 11. Living form. See Hlephas sumatranus Temminck, 1847, above, this list; also Lydekker, 1916, p. 83. Elephas cyclotis Matschie Sitz.-Ber. Ges. naturf. Freunde Berlin, No. 8, p. 194. Type loe.: Mwelle district, S. Cameroon. Type fig.: Heck, 1899, ‘‘Lebende Bilder aus dem Reiche der Tiere, Berlin,” Pl. exivi (original not seen by the present author). Formerly in Berlin Zoological Garden. Living form. Elephas (Loxodonta) oxyotis Matschie Tbid., p. 196. Type loe.: Upper Atbara, Sudan. Living form. Elephas (Loxodonta) knochenhauert Matschie Ibid., p. 197. Type loe.: Barikiwa, Tanganyika Territory. Living form. Palzxomastodon Beadnelli Andrews Zoologist, (4), V, August 15, pp. 318, 319 (name only); Tageblatt V Internat. Zool.-Congr., Berlin, No. 6, August 16, p. 4; Geol. Mag., Dec. IV., N.S., VIII, September, p. 401. Type loc.: Faytm, Egypt. Type fig.: Andrews, 1901, Geol. Mag., Dec. IV, N.8., VIII, p. 401, fig. 1, A, B. Meritherium Lyonsi Andrews Tageblatt V Internat. Zool.-Congr., Berlin, No. 6, August 16, p. 4; Geol. Mag., Dee. IV, N. S., VIII, September, pp. 403, 404. Type loc.: Faytim, Egypt. Type fig.: Andrews, 1901, Geol. Mag., Dec. IV, N. S., VIII, p. 404, fig. 2. Phiomia serridens Andrews and Beadnell “A Preliminary Note on Some New Mammals from the Upper Eocene of Egypt,” p. 3. Type loc.: Fayim, Egypt. Type figs.: Andrews and Beadnell, loc. cvt., figs. 1 and 2. Meritherium gracile Andrews Geol. Mag., Dec. IV, N. 8., IX, p. 292. Type loc.: Faytm, Egypt. Type figs.: Andrews, 1906, “‘A Descriptive Catalogue of the Tertiary Vertebrata of the Fayim, Egypt,” p. 127, and Pl. xvi, figs. 1 and 2. Mastodon tarijensis Ameghino An. Mus. Nac. Buenos Aires, VIII (Ser. 3*, I), p. 2 (name only). Hab.: Valley of Tarija, Bolivia. Mastodon Lydekkeri Schlosser Abh. bayer. Akad., II Cl., XXII, Abth. 1, pp. 46, 47. Type loe.: North China. Type fig.: Schlosser, zbed., Taf. xrv, fig. 8. Elephas cypriotes Bate Proc. Roy. Soc. London, LX-XI, pp. 498-500. Type loc.: Cave, Kerynia Hills, Cyprus. Cotype figs.: Bate, 1904, Phil. Trans. Roy. Soe. London, @X@VLIAB A El sisxexaeexexate Meritherium trigodon Andrews Geol. Mag., Dec. V, N.S., 1, p. 112. Type loc.: Faytm, Egypt. Type fig.: Andrews, 1906, ‘‘A Descriptive Catalogue of the Tertiary Verte- brata of the Fayim, Egypt,” Pl. rx, fig. 5. Supplementary description: Andrews, 1906, op. cit., p. 128, name changed to trigonodon. Palzomastodon minor Andrews Ibid.,p. 115. Type loc.: Fayim, Egypt. Type figs.: Andrews, 1906, ‘“‘A Descriptive Catalogue of the Tertiary Vertebrata of the Fayam, Egypt,”’ Pl. xiv, figs. 1, la, text fig. 50D. Supplementary description: Andrews, 1906, op. cit., p. 168. Palzomastodon minus Andrews Ibid., I, p. 562. Error, the author had reference to Palezomastodon [= Phiomia] minor. Palzomastodon parvus Andrews Loc. cit. Type loc.: Fayim, Egypt. Type figs.: Andrews, 1906, “A Descriptive Catalogue of the Tertiary Vertebrata of the Faytim, Egypt,” p. 148, text fig. 50C, and p. 163, text fig. 55. 1401 Reference in Present Memoir | Not determined by the present author | Zygolophodon sp. Elephas indicus sumatranus Loxodonta capensis cyclotis Loxodonta africana oxyotis Loxodonta africana knochenhaueri Palzomastodon beadnelli Meritherium lyonsi Phiomia serridens Meritheriwm gracile Cordillerion tarijensis Serridentinus lydekkeri Palzoloxodon cypriotes Meritherium trigodon Phiomia minor Phiomia minor Palxomastodon parvus 1402 Year 1905 1906 1906 1906 1907 1907 1907 1907 1907 1907 1907 1907 1907 1907 1907 1908 1908 OSBORN: THE PROBOSCIDEA Author Palzomastodon wintoni Andrews Ibid., p. 563. Type loc.: Faytim, Egypt. Type fig.: Andrews, 1906, ‘‘A Descriptive Catalogue of the Tertiary Vertebrata of the Faydm, Egypt,” p. 157, text fig. 53. Originally referred to Palzomastodon; transferred by Matsumoto (1922, p. 3) to Phiomia. Name Meritherium trigonodon Andrews Op. cit., p. 128. See Meritherium trigodon Andrews, 1904, above. Elephas africanus albertensis Lydekker Field (London), CVII, p. 1089. Type loc.: South end of Lake Albert, Africa. Type fig.: Lydekker, 1907, Proc. Zool. Soc. London, text fig. 121. Living form. Elephas africanus pumilio Noack Zool. Anz., XXIX, pp. 631-633. Type loc.: French Congo. Type fig.: Hornaday, 1905, Bull. New York Zool. Soe., October, pp. 237, 238. Living form. Elephas africanus toxotis Lydekker Proc. Zool. Soc. London, p. 385. Type loc.: Mossel Bay, western Cape Colony. Type fig.: Lydekker, zbid., text fig. 106. Living form. Elephas africanus selousi Lydekker TIbid., p. 387. Type loc.: Mashonaland, Rhodesia. Type fig.: Lydekker, zbid., text fig. 108. Living form. Elephas africanus peeli Lydekker Ibid., p. 393. Type loc.: Aberdare Mts., Kenya Colony. Type fig.: Lydekker, zbid., text fig. 114. Living form. Elephas africanus cavendishi Lydekker Ibid., p. 395. Type loc.: Lake Rudolf district. Type fig.: Lydekker, zbid., text fig. 115. Living form. Elephas africanus orleansi Lydekker Tbid., p. 398. Type loc.: North Somaliland. Type fig.: Lydekker, zbid., text fig. 118. Living form. Elephas africanus rothschildi Lydekker Ibid., p. 399. Type loc.: French Sudan. Lydekker, ibzd., text fig. 119. Living form. Lydekker states that he takes as type the statuette of “Jumbo” in the British Musuem. Professor Osborn (1931.846, p. 21) referred “Jumbo” to the subspecies Loxodonta africana oxyotis, which would make rothschildi a synonym of oxyotis. Type fig.: E. (Elephas] a. [africanus] cottoni Lydekker Tbid., II, p. 783. Type loc.: Northeastern Congo. Type fig.: Lydekker, zbzd., text fig. 111. Living form. Elephas maximus zeylanicus Lydekker ‘‘Game Animals of India,” p. 15. Living form. Elephas creticus Bate Proc. Zool. Soc. London, pp. 238-250. Type loc.: Near Cape Maleka, Crete. Cotype figs.: Bate, ibzd., Pl. xn, figs. 1-3, Pl. x11, figs. 1, 2. Paleomastodon Barroisi Pontier Ann. Soc. géol. du Nord, XXXVI, pp. 150, 151. Type loc.: Faytim, Egypt. Type figs.: Pontier, ibzd., text figs. 1 and 2. Synonym, in part (fide Matsumoto, 1922) of Phiomia wintoni and P. minor (ef. Vol. I, p. 61, this Memoir, where the present author confirms Matsumoto’s reference). Blephas (Loxedon) zulu Scott 38d Rept. Geol. Surv. (?)Natal and Zululand, pp. 259-262. Type loe.: Zululand, southeast coast of Africa. Type figs.: Scott, op. cit., Pl. xvi, fig. 6, and Pl. xvii, fig. 1. Dinotherium naricum Pilgrim Rec. Geol. Surv. India, XX XVII, Pt. I, p. 156. Name abandoned by the author and the subspecifie term gajense substituted in 1912, pp. 16 and 17. See below under Dinotherium indicum Lyd. var. gajense Pilgrim, 1912. Tetrabelodon crepusculi Pilgrim Tbid., p. 157. See Hemimastodon crepusculi Pilgrim, 1912, below, this list. Reference in Present Memoir Phiomia wintoni Loxodonta africana albertensis Loxodonta africana pumilio Loxodonta africana toxotis Loxodonta africana selousi Loxodonta africana peel Loxodonta africana cavendishi Loxodonta africana orleansi Loxodonta africana oxyotis Loxodonta africana cottoni Elephas indicus ceylanicus Palzxoloxodon creticus Loxodonta zulu Deinotherium indicum gajense Year 1908 1908 1908 1909 1909 1911 1911 1911 1912 1912 1912 1912 1912 1912 1913 NOMENCLATURE OF THE PROBOSCIDEA Name Author Stegodon javanoganesa Dubois Tijdschr. Nederl. Aardr. Genoots Amsterdam, (2), XX VB, No. 6, p. 1245. Von Koenigswald, 1933.1, p. 105: ‘‘Im grossen Ganzen scheint Stegodon trigonocephalus im Habitus von Stegodon ganesa nicht sehr verschieden gewesen zu sein. Dubois dachte sogar daran, die javanische Art nur als eine Varietat der indischen zu betrachten; ganz klar wird man iiber seine Auffassung nicht, da er die Art einmal als Stegodon javanoganesa, (1908, pg. 1245), einige Seiten weiter als Stegodon ganesa var. javanicus bezeichnet (1908, pg. 1257). Stegodon Ganesa var. javanicus Dubois Tbid., p. 1257. Type loe.: Kendeng-Schichten, Trinil, Java. Figure not found by the present author. Elephas hysudrindicus Dubois Loc. cit. Type loc.: Kendeng-Schichten, Java. Type fig.: First published figure of cotypes is in the present Memoir (Fig. 1160). Elephas Wiisti Pavlow Annuaire Géol. Min. Russie, XI, pp. 171-174. Type loc.: Tiraspol (gouv. Kherson), southern Russia. Cotype figs.: Pavlow, ibid., Pl. v, figs. 1 and 2; 1910, Nouv. Mém. Soc. Imp. Nat. Moscou, XVII, Livr. 2, Pl. 1, figs. 1-12 (Figs. 10 and 12 being the same as 1909, PI. v, figs. 1 and 2). Gomphotherium conodon Cook Amer. Journ. Sci., XXVIII, p. 183. Type loc.: Near Agate, Sioux County, Nebraska. Type fig.: Cook, zbid., p. 183. Tetrabelodon dinotherioides Andrews Geol. Mag., Dec. V, N. S., VI, p. 347. Type loc.: “Loup Fork beds,” northwestern Kansas. Type fig.: Andrews, ibid., p. 348. Elephas primus Schlesinger Monatsbl. Ver. Landeskunde v. Niederéster., V, p. 244. Meritherium Andrewsi Schlosser Beitr. Pal. Geol. Osterr.-Ung., XXIV, p. 130. Type loc.: Faytim, Egypt. Type figs.: Andrews, 1906, ‘“‘A Descriptive Catalogue of the Tertiary Vertebrata of the Fayim, Egypt,” Pls. vim, rx. Dinotherium hobleyi Andrews Abstract, Proe. Zool. Soe. London, (May), p. 35; Proc. Zool. Soc. London, (December), p. 943. Type loe.: Near Karungu, east side of Victoria Nyanza, Africa. Type fig.: Andrews, ibéd., Pl. xivim, figs. 1, la. Dinotherium indicum Lyd. var. gajense Pilgrim Mem. Geol. Sury. India, Palewont. Indica, N.5., IV, Mem. 2, p. 16. Type loc.: Gaj beds of Bugti Hills, Baluchistan. Type fig.: Pilgrim, zbid., Pl. rv, figs. 1-4. Same type specimen as Dinotherium naéricum Pilgrim, 1908, above, this list. Hemimastodon crepusculi Pilgrim Ibid., p. 17. Type loc.: Gaj zone of Kumbhi, Bugti Hills, Baluchistan. Type fig.: Pilgrim, zb7d., Pl. 1v, fig. 5; see also PI. tv, figs. 6-9, and PI. m1, fig. 4. Same as T'etrabelodon crepusculi Pilgrim, 1908, above, this list. Elephas primigenius Fraasi Dietrich Jahresh. Ver. Naturk. Wiirttemb., LX VIII, pp. 42-106. Type loc.: Steinheim a.d. Murr, Germany. Type figs.: Dietrich, cbed., Taf. rand 11, also text figs. 2, 4, 11-14, 16-21, 24-26. Mastodon arvernensis Croiz. et Job. var. progressor Khomenko Annuaire Géol. Min. Russie, XTV, Livr. 6, pp. 159-165. Type loc.: Near Gavanosy, district of Ismail, southern Bessarabia, Russia. Type fig.: Khomenko, zb7d., PI. 1. Mastodon arvernensis Croiz. et Job. var. conservativus Khomenko Ibid., p. 165. Synonym of Mastodon arvernensis Croiz. et Job. var. progressor Khomenko, 1912, above. Elephas antiquus var. insularis Soergel Paleontogr., LX, p.1. Type loc.: Carini, Sicily. Mastodon angustidens var. austro-germanica Wegner Palwontogr., LX, pp. 255-263. Type loc.: Near Oppeln, eastern Germany. Type fig.: Wagner, bed., Taf. xv, fig. 2. 1403 Reference in Present Memoir Stegodon airdwana [or S. trigonocephalus| Palzxoloxodon hysudrin- dicus Parelephas wiisti Not a probosecidean, but Dinohyus petersoni Holland Trilophodon dinotheri- oides {Not determined by the present author] Meritherium andrewsi Deinotherium hobleyi Deinotherium indicum gajense SuINA Mammonteus primi- genius fraast Anancus arvernensis progressor NOMEN NUDUM Trilophodon angusti- dens var. austro- germanicus 1404 Year 1913 1913 1913 1913 1913 1913 1914 1914 1914 1914 1914 1914 1914 1915 1915 OSBORN: THE PROBOSCIDEA Name Author Tetrabelodon corrugatus Pilgrim Ree. Geol. Surv. India, XLIII, Pt. IV, p. 293. Type loc.: Hasnot, India. Type fig.: Lydekker, 1880, Mem. Geol. Surv. India, Palzont. Indica, (X), I, Pt. V, Pl. xxxv, fig. 4, as Mastodon (Trilophodon) pandionis; see also Pl. xxxv1, fig. 1. Mastodon hasnoti Pilgrim Loc. cit. Type loc.: Near Bhimbar, northwest of Jammu, India. Type fig.: Lydekker, 1880, ibid., Pl. xutv, fig. 3, as Mastodon (Tetralophodon) stvalensis. Mastodon stegodontoides Pilgrim Ibid., p. 294. Type loc.: Lehri, Punjab, India. Type fig.: Lydekker, 1880, ibid., Pl. xxx1x, as Mastodon (Tetralophodon) latidens. Tetrabelodon macrognathus Pilgrim Tbid., p. 309. Type loc.: Near Chinji, India. No published type figure found by the present author. See figure 219 of the present Memoir. Tetrabelodon angustidens var. chinjiensis Pilgrim Ibid., p. 316 (name). Holotype of Osborn (Amer. Mus. 19421) found two miles west of Chinji Bungalow, India (see Fig. 218 of the present Memoir). Elephas antiquus Fale. var. trogontherioides Zuffardi Paleont. Ital., XIX, pp. 180, 155. Type loc.: Piedmont, Italy; lectotype from Nizza della Paglia (Astésan); cotypes from near San Paolo de Villafranca. Figures: Zuffardi, ib¢d., Tav. rx, figs. 3a-6b. Elephas primigenius Blum. var. trogontherti Pohl. Zuffardi Ibid., p. 167. Cf. Elephas primigenius mutation astensis Depéret and Mayet, 1923, below, this list. Elephas africanus Fransseni Schouteden Rev. Zool. Africaine, III, Fase. 2, p. 396. Type loc.: M’Paa near Bongo, northwest of Lake Leopold II. Type fig.: Schouteden, ibid., Pl. x1, figs. 1,2. Living form. Elephas maximus hirsutus Lydekker Abstract, Proc. Zool. Soc. London, I, p. 20 (name only). Type loc.: Kuala Pila district of the Negri Sembilan province, Malay Peninsula. Type fig.: Lydekker, 1914, Proc. Zool. Soc. London, I, text fig. 1, p. 285; 1916, “Catalogue Ungulate Mammals in British Museum,”’ text fig. 25, p. 84. Formerly in the Gardens of the Zoological Society of London. Tetrabelodon willistoni Barbour Univ. Studies, Univ. Neb., XIV, No. 2, pp. 192-194. Type loc.: Devil’s Guleh, Niobrara River, Brown County, Nebraska. Type figs.: Barbour, zb7d., Pls. v, vi, vim. Eubelodon morrilli Barbour Ibid., pp. 194-197. Type loc.: Devil’s Gulch, Niobrara River, Brown County, Nebraska. Type figs.: Barbour, zbzd., Pls. rx—x11. Tetrabelodon lulli Barbour Neb. Geol. Surv., IV, Pt. 14, p. 217. On this same page, in a footnote, the subgenus Megabelodon was suggested. Type loc.: Exposures bordering Snake River, Cherry County, Nebraska. Type figs.: Barbour, zbid., Pls. ivi. See also Barbour, 1917.1, p. 512. Mammut progenium Hay Iowa Geol. Surv., Ann. Rept. for 1912, XXIII, pp. 368-373. Type loc.: Cox gravel pit, Missouri Valley, Harrison County, Iowa. Type fig.: Hay, ibid., Pl. xxiv, figs. 1 and 2 (specimen first described and figured by Calvin, 1911, Bull. Geol. Soe. Amer., XXII, p. 213, Pls. xx, xx1, under the name Mastodon americanus). Rhabdobunus mirificus Hay ITbid., p. 374. Pohlig in 1912 chose Mastodon mirificus Leidy, 1858 (see above, this list), as the genotype of Stegomastodon (see Generic List above, p. 1379). In 1914 Hay made Leidy’s M. mirificus the type of his new genus Rhabdobunus, which, however, is invalid because preoccupied by Pohlig’s Stegomastodon (cf. Hay, 1930, p. 633). Elephas hysudricus primitivus Soergel Centralb. Min. Geol. Pal., No. 8, p. 250, No. 9, p. 283. Dinotherium styriacum Hilber Mitt. Naturwiss. Ver. Steiermark, LI, pp. 113-117. Type loc.: Oberdorf bei Weiz, Styria. Type fig.: Hilber, zb¢d., Taf. 1, fig. 1. Reference in Present Memoir Synconolophus cor- rugatus Synconolophus hasnoti Stegolophodon stegodon- toides Trilophodon macro- gnathus Trilophodon chinjiensis Parelephas trogontheri- oides Loxodonta africana fransseni Elephas indicus hirsu- tus [of doubtful val- idity—see footnotes on page 1332 of Chap- ter XX above.—EKd- itor.] Trilophodon (Geno- mastodon) willistoni Eubelodon morrilli Megabelodon lulli Mastodon progenius Stegomastodon mi- rificus {Not determined by the present author] [Not determined by the present author] Year 1915 1915 1918 1915 1915 1916 1916 1916 1917 1917 1917 1917 1917 1918 1918 1918 1918 NOMENCLATURE OF THE PROBOSCIDEA Author Elephas hayi Barbour County, Nebraska. Type figs.: 134, fig. 5d. Name Amer. Journ. Sci., (4), XL, pp. 129-134. Type loc.: Crete, Saline Hay, ibid., p. 130, fig. 1; p. 133, fig. 3; p. Elephas aurore Matsumoto Scientific Gazette, Tokyo, III, No. 5, pp. 8308-315 (Japanese only) ; 1918, Sci. Rept. To6hoku Imp. Univ., (2), Geol., III, No. 2, p. 52. Type loc.: Mt. Tomuro, Kaga, Japan. Type fig.: Matsumoto, 1918, zbzd., Pl. xx, figs. ios Subsequently (1924, Journ. Geol. Soc. Tokyo, XXXI, No. 371, pp. 256, 257, 262) made the genotype of Parastegodon. Mastodon pirayuiensis Gez ‘‘Generalidades sobre paleontologia argentina: El Mastodon platen- sis—Amegh. de Corrientes,”’ 2* edicion, p. 35. Mastodon “ligoniferus’’ Cope and Matthew ‘Hitherto Unpublished Plates of Tertiary Mam- malia and Permian Vertebrata,” Amer. Mus. Monograph Series No. 2. Type loe.: Black Hills, South Dakota. Type figs.: Cope and Matthew, op. cit., Pl. exxty, figs. 1 and 2. Elephas maximus maximus Lydekker ‘Cat. Ungulate Mammals, British Museum,’ V, p. 82. Arch. Biontol., IV, Heft I, p. 22. Type loc.: Oldoway-Tuffe, Elephas antiquus Recki Dietrich Lectotype fig.: Serengetisteppe, northern Tanganyika Territory, Africa. Dietrich, ibid., Taf. 1, fig. 2; cotypes, Taf. t-vur (in part). Tetrabelodon osborni Barbour Amer. Journ. Sci., (4), XLI, pp. 522-529. Type loc.: Near Bristow, Boyd County, Nebraska. Type figs.: Barbour, ibid., p. 523, fig. 1; p. 524, fig. 2; p. 526, fig. 3; p. 528, fig. 4. Mastodon (Bunolophodon) angustidens Cuv. {. subtapiroidea Schlesinger Denk. Naturhist. Hof- mus., I, Geol.-Pal. Reihe I, pp. 30, 31, 35, 37. Type loc.: Wies, near Eibis- wald (Styria), Austria. Cotype figs.: Schlesinger, 7bid., Taf. m1, fig. 2; Taf. Iv, fig. 1; Taf. vu, fig. 3; Taf. vin, figs. 1 and 2, and p. 31, text fig. 3. Mastodon (Bunolophodon) longirostre Kaup forma sublatidens Schlesinger Jbid., pp. 101, 102. Type loc.: Near Teschen (Schlesien), Austria. Type fig.: Schlesinger, 7bid., Taf. xvut, fig. 2. Mastodon (Bunolophodon) grandincisivum Schlesinger Ibid., p. 119. Type loc.: Maragha, Persia. Type fig.: Schlesinger, zbid., Taf. xxxtv, figs. land 2. Paratypes: Taf. xv, figs. 1 and 2, Mannersdorf near Angern, Austria. Gomphotherium gratum Hay Bull. Univ. Texas, No. LX XI, pp. 18-21. Type loc.: Pittbridge, Burleson County, Texas, on the Brazos River. Type figs.: Hay, zbid., Pl. m1, figs. 3 and 4, PI. rv, figs. 1 and 2. Gomphotherium elegans Hay Ibid., pp. 21, 22. Type loc.: McPherson, Kansas. Type fig.: Hay, 1917, Proc. U.S. Nat. Mus., LITI, No. 2198, Pl. xxv1, figs. 1 and 2. Mastodon angustidens, Cuvier var. libyca Fourtau Ministry of Finance, Egypt, Surv. Dept., pp. 84-89. Type loe.: Moghara, northern Egypt. Type fig.: Fourtau, op. cit., text fig. 58. [Mastodon angustidens| mut. Pontileviensis Mayet (In Fourtau, op. cit., p. 88—name only.) Hab.: Chevilly, Pontlevoy, France. Cotype figs.: Mayet, 1908, Ann. Univ. Lyon, Nouv. Sér. I,—Sci., Méd., Fasc. 24, Pls. vu, figs. 5, 6, x1, fig. 2 (as Mastodon angustidens). See Vol. I, p. 283 and fig. 230 of the present Memoir. Mastodon Spenceri Fourtau Op. cit., pp. 89-91. Type loc.: Moghara Desert, northern Egypt. Type figs.: Fourtau, op. cit., p. 89, fig. 60; p. 90, fig. 61. Rhynchotherium tlascale Osborn Bull. Geol. Soc. Amer., XXIX, pp. 134, 135. Type loc.: Tlascala, Valley of Mexico. Type fig.: Vol. I, fig. 448, of present Memoir. See Rhynchotherium brownt Osborn, 1936, below, this list. 1405 Reference in Present Memoir Archidiskodon hayi Stegodon aurore Cuvieronius pirayuien- sis Trilophodon ligoniferus Elephas indicus ben- galensis Palzoloxodon recki Trilophodon (Genomas- todon) osborni Serridentinus subtapi- roideus Stegolophodon sub- latidens Tetralophodon grandin- cistvUs Cordillerion gratum Tetralophodon elegans Trilophodon angusti- dens libycus Trilophodon ponti- leviensis Rhynchotherium(?) spencer Rhynchotheriwm tlascale 1406 Year 1919 1922 1921 1921 1921 1921 1921 1922 1921 1922 1922 1922 1922 1922 1922 1922 1922 OSBORN: THE PROBOSCIDEA Name Author Mastodon (Mammut) americanus Penn. forma praetypica Schlesinger Mitt. Geol. Ges. Wien, X1, p. 142. Cotype loc.: Szabadka (= Maria-Theresiopel), Batta Erd, Rakos- keresztur, Szentlérinez, and Ajnacsk6, Hungary. Original figures: Schle- singer, zbid., 1919, Taf. vi, figs. 2-4. Supplementary description and figures: Schlesinger, 1922, Geol. Hungarica, Ed. Sep., II, Fase. 1, pp. 115, 116, 227— 230, Pls. xtv—xrx. Trilophodon giganteus Osborn Amer. Mus. Novitates, No. 1, pp. 6-10. Type loc.: Eastview, near Dallas, Gregory County, South Dakota. Type fig.: Osborn, ibid., p. 9, fig. 4C. Tetralophodon barbouri Osborn Ibid., pp. 9, 10. Type loc.: Cambridge, Furnas County, Nebraska. Type fig.: Osborn, 1924, Amer. Mus. Novitates, No. 154, fig. 1. See also figs. 308F, 329D, and 337 of the present Memoir. Mastodon matthewi Osborn Amer. Mus. Novitates, No. 10, pp. 2-6. Type loc.: Snake Creek, Sioux County, Nebraska. Type fig.: Osborn, ibid., p. 3, fig. LA (Fig. 98A of the present Memoir). Osborn (1922.564, p. 4) referred this species to Miomastodon matthewi; finally (1926.706, p. 1) he made it the type of his new genus Pliomastodon. Mastodon merriami Osborn Ibid., pp. 4-6. Type loe.: Thousand Creek, Humboldt County, Nevada. Type figs.: Osborn, zbid., p. 3, fig. 1B, and p. 5, fig. 2. Mastodon tapiroides americanus Schlesinger (In Osborn, Amer. Mus. Novitates, No. 10, p. 2; fig. 1, D and D1, p. 3.) Schlesinger, 1922, Geol. Hungarica, II, Fasc. 1, pp. 224-227, Taf. xu, figs. 6 and 7, and Taf. xrv, figs. 1-4. Type loc.: Tasnad, Usztat6 Komitat, Hungary. Trilophodon (Tetrabelodon) shepardi edensis Frick Bull. Dept. Geol. Univ. Calif., XII, No. 5, pp. 405-409. Type loc.: Mt. Eden Hot Springs, San Bernardino County, southern California. Type figs.: Frick, zbid., p. 406, figs. 160-165. In the original description of T’rilophodon (Tetrabelodon) shepardi edensis Frick, 1921, the author described material which ultimately proved to belong to two different genera, namely, Rhynchotherium, and Dibelodon (=Cordillerion). The molars (Frick, 1921, figs. 160-165) are referable to Rhynchotherium shepardi edense (see Frick, 1926, Bull. Amer. Mus. Nat. Hist., LVI, Art. II, pp. 169-176); the premaxille# and tusks (Frick, 1921, Pl. 1, figs. 1, 2) to Cordillerion edensis, 1936 (see below, this list). Dibelodon edensis Osborn (in part) Amer. Mus. Novitates, No. 49, p. 2. Synonym of Cordillerion edensis, 1936 (see below, this list). Miomastodon matthewi Osborn Ibid., p. 4. See note under Mastodon matthewi Osborn, 1921, above, this list. Loxodonta griqua Haughton ‘Trans. Geol. Soc. 8. Africa, XXIV, pp. 11-13. Type loc.: Griqua- land West, Africa. Type fig.: Haughton, 7b7d., PI. 1, figs. 1, 2. Palzomastodon intermedius Matsumoto Amer. Mus. Novitates, No. 51, p. 2. Type loc.: North of Lake Qurun, Fayim, Egypt. Type fig.: Matsumoto, zbzd., text fig. 1. Phiomia osborni Matsumoto Tbid., pp. 3, 4. Type loc.: Alexandria Trail, Fayaim, Egypt. Type fig.: Matsumoto, zbid., p. 4, fig. 3. Mastodon engelswiesensis Klihn ‘Die badischen Mastodonten,” pp. x, 30. Type loc.: Engels- wies, Baden, Germany. Type fig.: Klahn, op. cit., p. 30, fig. 10. Mastodon steinheimensis Kliihn Op. cit., pp. x, xi, 35, 76. Type loc.: Steinheim, Baden, Ger- many. Type fig.: Fraas, 1870, Jahresh. Ver. naturk. Wiirttemb., XX VI, Hefte II, III, Taf. v, fig. 1 (as Mastodon arvernensis). Mastodon gigantorostris Klihn Op. cit., pp. xi, 48, 50, 87, 131. Type loe.: Bermersheim, Rhein- hessen, Germany. Reference in Present Memoir Pliomastodon ameri- canus praetypica Trilophodon giganteus Morrillia barbouri. Pliomastodon matthewi Miomastodon merriami Miomastodon tapiroides americanus Rhynchothertum shepardi edense Metarchidiskodon griqua Palzomastodon inter- medius Phiomia osborni Trilophodon engels- WIeSENSIS Trilophodon steinheim- ensis Tetralophodon giganto- rostris 1922 1922 1922 1922 1922 1922 1922 1922 1922 1922 1922 1922 1922 1922 1923 1923 NOMENCLATURE OF THE PROBOSCIDEA Author Mastodon esselbornensis Klahn Op. cit., pp. xi, 50, 73. Type loe.: Esselborn and Westhofen, Rheinhessen, Germany. Type fig.: Kléahn, op. cit., p. 92, fig. 24. Name Mastodon wahlheimensis Klihn Op. cit., pp. xi, 76. Type loc.: Wahlheim and Esselborn, Rheinhessen, Germany. Cotype figs.: Klihn, op. cit., p. 77, fig. 17; p. 94, fig. 25. Mastodon minutoarvernensis Klaihn Germany. Op. cit., pp. xu, 102. Type loc.: Herbolzheim, Baden, Mastodon gigantarvernensis Klihn Loc. cit. Type loe.: Herbolzheim, Baden, Germany. Trilophodon hicksi Cook Proc. Colo. Mus. Nat. Hist., 1V, No. 1, p. 5. Type loc.: Near Wray, Yuma County, Colorado. Type figs.: Cook, zbid., p. 9, figs. 2,3; p. 10, fig. IS joys WL rae Trilophodon paladentatus Cook Ibid., p. 6. Type loc.: Near Wray, Yuma County, Colorado. Type figs.: Cook, ibid., p. 9, fig. 1; p. 18, fig. 5; p. 14, fig. 6. Elephas jeffersonizi Osborn Amer. Mus. Novitates, No. 41, pp. 11-16. Type loc.: Near Jones- boro, Indiana. Type fig.: Osborn, zbid., p. 11, fig. 10. Elephas roosevelti Hay Proc. Biol. Soc. Washington, XX XV, pp. 100, 101 (description without figure). Typeloe.: Ashland, Cass County, Illinois. See figure 968, p. 1096, above, this Memoir. Mastodon oligobunis var. antiquissima Freudenberg Geol. u. Pal. Abh., XIV (XVIII), Heft III, pp. 118-120. Type loc.: Valley of Amajaque, Hidalgo, Mexico. Type fig.: Freudenberg, zbid., Taf. rx, figs. 3, 3a. Mastodon oligobunis var. Felicis Freudenberg Jbid., pp. 120-123. Type loc.: Puebla, Mexico. Type fig.: Freudenberg, zbid., Taf. x, figs. 1, la. Mastodon oligobunis var. intermedia Freudenberg I[bid., pp. exact locality not recorded. i 72 123-126. Type loc.: Mexico, Type fig.: Freudenberg, zbid., Taf. x1, figs. M. |Mastodon| oligobunis var. progressa Freudenberg Jbid., p. 127. Type loe.: Cannada [ean- yon] de Aculeingo, Mexico. Type fig.: Freudenberg, ¢b¢d., Taf. x1v, figs. 1, 2. El. [Elephas| Columbi var. Felicis Freudenberg Jbid., pp. 147-152. Type loc.: Tecamachalco, Puebla, Mexico. Type fig.: Freudenberg, zbid., Taf. xv1, fig. 4. El. |Elephas) Columbi var. silvestris Freudenberg T[bid., pp. 152, 153. Type loc.: Ejutla, Oaxaca, Mexico. Type fig.: Freudenberg, zbid., p. 146, fig. 19. El. [Elephas| Columbi var. Falconeri Freudenberg Jbid., pp. 153-160. Type loc.: Tequixquiac. Valley of Mexico. Paratype fig.: Freudenberg, zbzd., p. 154, fig. 21 (original- ly figured by Villada, 1903, Ann. Mus. Nac. Mexico, VII, Lam. vin, as El. primigenius Blum.) El. [Elephas| Columbi var. imperator Leidy, Freudenberg Jbid., pp. 160-171. Typical example from Spokam Bar, near Helena, Montana. Fig.: Freudenberg, zbid., p. 55, fig. 22. E. [Elephas| meridionalis mutation cromerensis Depéret and Mayet Ann. Univ. Lyon, Nouv. Ser., I,-Sci., Méd., Fase. 48, Deuxiéme Partie, pp. 150, 152, 157. Type loc.: Kessingland, Suffolk, England. Type fig.: Depéret and Mayet, zbzd., Pl. rx, fig. 1. EK. [Elephas | primigenius mutation astensis Depéret and Mayet Tbid., pp. 183, 184. Type loc.: San Paolo de Villafranca, northern Italy. Type fig.: Depéret and Mayet, ibid., Pl. xt, fig. 5 (original in Musée du Palais Carignan, Turin). Paratype fig.: bzd., Pl. x1, fig. 6 (original in Mus. Géol., Turin); figured by Zuffardh, 1913, Paleont. ital., XIX, Tav. vi [xm], fig. 2a, and Tav. v [x1] fig. 8a, as Elephas primigenius Blum var. trogontheri? Pohl. 1407 Reference in Present Memoir Trilophodon esselborn- ensis Turicius wahlheimensis Anancus minutoarvern- ensis Anancus gigantarvern- ensis Amebelodon (Trilopho- don) hicksi Amebelodon (Trilopho- don) paladentatus Parelephas jeffersonii Parelephas jeffersonii Cordillerion oligobunis antiquissimus Cordillerion(?) oligo- bunis felicis Cordillerion(?) oligo- bunis intermedius Cordillerion oligobunis progressus Parelephas columbi felicis Archidiskodon impera- tor silvestris Archidiskodon impera- tor falconeri Archidiskodon impera- tor Archidiskodon meridi- onalis cromerensis Mammonteus primi- genius astensis 1408 Year 1923 1923 1923 1923 1923 1923 1923 1924 1924 1924 OSBORN: THE PROBOSCIDEA Name Author Meritherium ancestrale Petronievies Ann. Mag. Nat. Hist., (9), XII, p. 57. Type loe.: Faydim, Egypt. Type fig.: Petronievies, ibid., p. 56, fig. 1. Anancus brazosius Hay Pan-Amer. Geol., XX XIX, pp. 112-114. Type loc.: Brazos River, near San Felipe, Texas. Type fig.: Hay, zbzd., Pl. vin, figs. 1, 2. Serridentinus simplicidens Osborn Amer. Mus. Novitates, No. 99, p. 2. Type loc.: Near Pierce, Polk County, Florida. Type fig.: Figure 231 of the present Memoir. Trilophodon progressus Osborn Ibid., p. 3. Type loe.: Driftwood Creek, Hitchcock County, Nebraska. Type fig.: Cope, 1889, Amer. Naturalist, X XIII, p. 202, Pl. x1, first as Tetrabelodon angustidens proavus, then as Tetrabelodon proavus (see Figs. 360 and 361 of the present Memoir; also p. 403, text). Tetralophodon precampester Osborn Loc. cit. Type loc.: Harlan County, Nebraska. Type figs.: Figures 308E, 329K, and 339 of the present Memoir. Loc. cit. Type loc.: Eight miles west of Pawnee Buttes, Rhynchotherium rectidens Osborn Type figs.: Figures 461 and 481 of the present Weld County, Colorado. Memoir. Rhynchotheriwm falconeri Osborn Ibid., pp. 3, 4. Type loe.: Mt. Blanco, Llano Estacado, Texas. Type fig.: Figure 468 of present Memoir (originally figured as “‘Tetrabelodon shepardii Leidy” by Cope, 1893, 4th Ann. Rept. Geol. Surv. Texas, Pl. xv). Elephas washingtonivi Osborn Tbid., p.4. Typeloc.: Pine Creek, Whitman County, Washington. Type figs.: Figures 972, 975, 893, B, B1, of the present Memoir. Harpagonotherium canadense Fischer de Waldheim (In Sherborn, ‘Index Animalium,” 1924, Pt. V, p. 1022.) Probably same as Harpagmotherium canadense Vischer, 1808, above, this list. Serridentinus mongoliensis Osborn Amer. Mus. Novitates, No. 148, pp. 1-8. Type loc.: Loh, near camp, Mongolia. Type fig.: Osborn, zbid., p. 2, fig. 1. Elephas antiquus rumanus 8. Stefainescu. Compt. Rend. Acad. Sci. (Paris), CLX XIX, p. 1418. Type loe.: Tulucesti (Covurlui), Rumania. Type fig.: Athanasiu, 1912 [1915], Annuar. Inst. Geol. Romaniei, VI, Pl. xvu, fig. 4 (as Hlephas cfr. meridionalis). Refigured by Stefanescu, 1927, ‘30 Notes sur l’Organisation des Molaires et sur la Phylogénie des Eléphants et des Mastodontes’’ (see Fig. 857 of the present Memoir). Elephas antiquus germanicus 8. Stefainescu Loc. cit. Type loe.: Tanganu (Ilfov), Rumania. Type figs.: Stefainescu, 1927, op. cit., Figs. A, B (see Fig. 1089 of the present Memoir). Elephas namadicus naumanni Makiyama Chikyi—The Globe, I, p. 381 (in Japanese; fide Matsumoto, original not available to present author); 1924, Mem. College Sci. Kyéto Imp. Univ., (B), I, No. 2, pp. 255-264 (in English). Type loc.: Sahamma, T6t6mi Province, Japan. Type figs.: Makiyama, Chikyd, I, PI. vit; Mem. College Sci. Ky6to Imp. Univ., (B), I, Pls. x1r—xv, and xv1, fig. 1. The following is an excerpt from Dr. Makiyama’s article on “Japonic Proboscidea,”’ Mem. Coll. Sci. Kyéto Imp. Univ., Ser. B, Vol. XIV, No. 1, Art. 1, p. 40, giving certain of the results of his recent studies (May, 1938): ‘This is the most common fossil elephant in Japan. The name of subspecies was given to the stenocoronine form [which] came from Sahanma near Hamamatu in Sizuoka-ken (T6t6mi) covering the Naumann’s 2. namadicus from Yokosuka as it be different from the type #. namadicus Falconer & Cautley, 1846 which iseurycoronine. After that time, I have had a number of chances to make interesting observation about the different forms of teeth, and now I have a thought that all the Japonic forms are equally separated from the Indian #. namadicus. The names listed are the synonyms: E. namadicus namadi Pohlig, 1893 of Makiyama, 1924 Parelephas protomammonteus Matsumoto, 1924 Loxodonta (Palaeoloxodon) namadica yabei Matsumoto, 1929 Blephas (Palaeoloxodon) namadicus setoensis Makiyama, 1929 E. indicus buski Matsumoto, 1929 Euelephas trogontherii of Matsumoto, 1924, and Makiyama, 1924 Palaevloxodon yokohamanus Tokunaga, 1934)” Also see note below under ‘1929 Hlephas (Palxolozodon) namadicus setoensis Maki- yama.” Reference in Present Memoir Meritherium ancestrale Trilophodon (?Tetra- lophodon) brazosius Trilophodon simplici- dens Serridentinus progres- sus Morrillia barbourt Rhynchotherium rectidens Rhynchotherium falconeri Parelephas washing- tonit Serridentinus mongo- liensis Archidiskodon plani- frons rumanus Hesperoloxodon antiqu- Us germanicus Palzxoloxodon namadi- cus naumannt Year 1924 1924 1924 1924 1924 1924 1924 1924 1924 1924 1924 1924 NOMENCLATURE OF THE PROBOSCIDEA Author Elephas namadicus namadi Makiyama Chikyu, I, p. 381 (in Japanese); Mem. College Sci. Kyoto Imp. Univ., (B), I, No. 2, pp. 263, 264 (in English). Type loe.: Dredged off island of Shédo, Sanuki Province, Japan. Type figs.: Makiyama, Chiky4, I, Pl. v1; Mem. College Sci. Kyéto Imp. Univ., (B), I, Pl. xv1, fig. 2. See note above under ‘1924 Elephas namadicus naumanni,” also note below under “1929 Elephas (Palzolorodon) namadicus setoensis Makiyama.”’ Name Euelephas protomammonteus Matsumoto Journ. Geol. Soc. Tokyo, XX XI, p. 262 (in Japanese) ; 1926, Sci. Rept. Todhoku Imp. Univ., (2), Geol., X, No. 2, pp. 43-50 (in English). Type loc.: Nagahama, Town of Minato, Kimitsu District, Province of Kazusa, Japan. Type figs.: Matsumoto, 1926, ibid., Pl. xv, figs. 1, 2 (as Parelephas protomammonteus); Pl. x1x, figs. 1-3 (the paratype, from same loeality as type). See note above, 1924, under Hlephas namadicus naumanni Makiyama. Loxodonta (Palxoloxodon) tokunagai Mastumoto Jbid., p. 267 (in Japanese); 1929, Sci. Rept. Tohoku Imp. Univ., (2), Geol., XIII, No. 1, pp. 7-11 (in English). Type loc.: Soyama, Gokayama, Hira-mura Higashi-Tonami Dist., Province of Etcha, Japan. Type figure not found by present author. Referred to Archidiskodon tokunagai by Teilhard de Chardin and Trassaert in 1937 (Pal. Sin, (C), XIII, Fase. 1, p. 44). Stegodon orientalis shodoénsis Matsumoto . Ibid., pp. 333-335 (in Japanese). ‘Type loc.: Island of Mitsugo (Mitsugo-Shima) and island of Shédo, Inland Sea, Japan. Type fig.: See Makiyama, 1938, Mem. Coll. Sci. Kyoto Imp. Univ., (B), XIV, No. 1. p. 18, fig. 7 (holotype palate). Hemimastodon annectens Matsumoto I[bid., pp. 401, 405 (in Japanese). Type loc.: Banjobora, Kaminogé-mura, Kani District, Province of Mino, Japan. Type figs.: Matsumoto, 1926, Sci. Rept. Tohoku Imp. Univ., (2), Geol., X, No. 1, Pls. I, figs. 1 and 2, 11, figs. 1-3. Trilophodon sendaicus Matsumoto [bid., pp. 402,408 (in Japanese). Type loc.: Kitayama, near Sendai, Province of Rikuzen, Japan. Type figs.: Matsumoto, 1926, Sci. Rept. Tohoku Imp. Univ., (2), Geol., X, No. 1, Pls. 1m and tv. Elephas africanus mogambicus Frade Bull. Soc. Portugaise Sci. nat., UX, Fase. 3, pp. 131, 133. Hab.: Maputo, Mozambique. Type fig.: Frade, ibid., text fig. 5. Female formerly in the Jardin de Zoologique, Lisbon. Loxodonta africana Zukowskyi Strand (In Zukowsky, Arch. Naturges., XC, Abth. A, Heft I, p. 68.) Hab.: Kaoko District, southwest Africa. Living form. Parelephas jeffersonii progressus Osborn Amer. Mus. Novitates, No. 152, pp. 1, 4, 7. Type loe.: Zanesville, Muskingum County, Ohio. Type figs.: Osborn, 1922, Amer. Mus. Novitates, No. 41, p. 13, fig. 11, and p. 14, fig. 12 (as paratypes of Elephas jeffersonit). Mammonteus primigenius compressus Osborn Ibid., pp. 5-7. Type loc.: Rochester, Indiana. Type figs.: Osborn, 1922, Amer. Mus. Novitates, No. 41, p. 8, fig. 8 (as Elephas primigenius); 1924, Amer. Mus. Novitates, No. 152, p. 6, fig. 2. Paratype fig.: Osborn, 1922, Amer. Mus. Novitates, No. 41, p. 7, fig. 7 (from Eschscholtz Bay, Alaska). Stegomastodon teranus Osborn Amer. Mus. Novitates, No. 154, p. 3. Type loc.: Blanco for- mation, Llano Estacado, Texas. Type figs.: Figures 639, 642, 644A, 645D, of the present Memoir. Stegomastodon aftoniz Osborn Loc. cit. Type loc.: About two miles east of Akron, Plymouth County, lowa. Type r.M? originally figured as Mammut mirificum by Calvin, 1909, Bull. Geol. Soc. Amer., XX, Pl. xxvir; refigured as Rhabdobunus mirificus by Hay, 1914, lowa Geol. Sury., XXIII, Pl. u. Type 1.M’, see figure 650 of the present Memoir. 1409 Reference in Present Memoir Palzoloxodon proto- mammonteus Palexoloxodon tokuna- gar Stegodon orientalis shodoénsis Serridentinus annectens Trilophodon sendaicus Loxodonta africana mo¢gambica Loxodonta africana zukowskyt Parelephas progressus Mammonteus primi- genius compressus Stegomastodon texanus Stegomastodon aftonie 1410 Year 1924 1926 1924 1925 1925 1925 1925 1926 1926 1926 1926 1926 1926 1926 1926 1926 1926 1926 OSBORN: THE PROBOSCIDEA Name Author Stegomastodon arizone Gidley (In Osborn, Amer. Mus. Novitates, No. 154, p. 4—from Gidley MS.); Gidley, 1926, U.S. Geol. Surv., Profess. Paper 140—B, p. 86. Type loc.: Curtis Flats, Cochise County, Arizona. Type figs.: Gidley, zbid., Pls. XXXII-Xxxv, Pl. xxxvitl, figs. 1-3, Pl. xxxrx, figs. 1-3. Stegomastodon nebrascensis Osborn Ibid., p. 5. Type loc.: Snake Creek, Sioux County, western Nebraska. Type fig.: Osborn, zbid., p. 4, fig. 2 B. Mammonteus primigenius americanus Osborn Proc. Amer. Phil. Soc., LXIV, No. 1, p. 33. See E. americanus DeKay, 1842, above, from near Rochester, New York. Elephas scotti Barbour Neb. State Mus., Bull. 2, I, pp. 21-24. Type loc.: Five miles south of Staplehurst, Seward County, Nebraska. Type figs.: Barbour, zbid., text figs. 7-10. {On page 1025, of the present Memoir, appears the following statement by Pro- fessor Osborn: ‘Awaiting further evidence, Osborn is inclined to regard the type of ‘Elephas scott’ as representing a young individual of Archidiskodon imperator.”” Neverthe- less he treated this species under the heading Archidiskodon imperator scotti.] Tetrabelodon abeli Barbour Neb. State Mus., Bull. 9, I, pp. 91-94. Type loc.: Devil’s Gulch, Brown County, Nebraska. Type figs.: Barbour, zbid., pp. 92, 93, figs. 52-56. Elephas maibeni Barbour Neb. State Mus., Bull. 10, I, pp. 95-118. Type loe.: About sixteen miles north of Curtis, Lincoln County, Nebraska. Type figs.: Barbour, zbid., text figs. 58-60, 63-70, 72, 74, 76-87. In 1926 described by Barbour as Archidiskodon maibeni. Archidiskodon maibeni Barbour Neb. State Mus. Bull. 11, I. See EHlephas maibeni Barbour, 1925, preceding item. Mammut francisi Hay Journ. Wash. Acad. Sci., XVI, No. 2, pp. 35-39. Type loc.: Brazos River, Pittbridge, Burleson County, Texas. Type figs.: Hay, zbzd., p. 36, fig. 3; p. 37, fig. 4. Mammut oregonense Hay Ibid., pp. 39, 40. Type loc.: Rye Valley, Dixie Creek, Baker County, Oregon. Type figs.: Hay, zbd., p. 36, figs. 1, 2. Mastodon americanus plicatus Osborn Amer. Mus. Novitates, No. 238, p.1. Type loc.: Walnut, Bureau County, Illinois. Type fig.: Osborn, ibzd., p. 2, fig. 1. Zygolophodon pyrenaicus aurelianensis Osborn Ibid., p. 2. Hab.: Chevilly(?), Cassegrain, Avaray, Franee. No type figure available to the present author. Turicius turicensis simorrensis Osborn Ibid., pp. 3, 4. Type loe.: Near Simorre (Gers), France. Type figs.: Lartet, 1859, Bull. Soc. géol. France, (2), XVI, Pl. xv, fig. 3 (as Mastodon tapiroides); Osborn, 1926, ibid, p. 4, fig. 2. Serridentinus brown Osborn Tbid., p. 4. Type loc.: Near Chinji Bungalow, India. Type figs.: Osborn, ibid., p. 5, fig. 3; p. 6, fig. 4. Serridentinus republicanus Osborn Ibid., p. 6. Type loc.: Republican River formation, north- western Kansas. Type figs.: Osborn, zbid., p. 7, fig. 5; p. 8, fig. 6. Serridentinus obliquidens Osborn Tbid., p. 9. Type loc.: Phosphate beds near Charleston, South Carolina. Type fig.: Osborn, zbid., p. 9, fig. 7. Serridentinus anguirivalis Osborn Tbid., p. 10. Type loc.: South-central Sioux County, western Nebraska. Type fig.: Osborn, zb7d., p. 10, fig. 8. Serridentinus brewsterensis Osborn J[bid., p. 11. Type loe.: Brewster, Polk County, Florida. Type fig.: Osborn, ibed., p. 11, fig. 9. Reference in Present Memoir Stegomastodon arizone Serridentinus (Ocalien- tinus?) nebrascensis Mammonteus primi- genius americanus Archidiskodon impera- tor scotti Trilophodon abeli Archidiskodon impera- tor maibent ?Rhynchotherium francisi Mastodon oregonensis Mastodon americanus plicatus Zygolophodon pyrenai- cus aurelianensis Turicius turicensis stmorrensis Serridentinus brownt Ocalientinus (Serriden- tinus) republicanus Ocalientinus (Serriden- tinus) obliquidens Serridentinus anguiri- valis Serridentinus brewster- ensis Year 1926 1926 1926 1926 1926 1926 1926 1926 1926 1926 1926 1926 1927 NOMENCLATURE OF THE PROBOSCIDEA Author Serridentinus guatemalensis Osborn Ibid., p. 12. Type loc.: Chinautla, Guatemala, Central America. Type fig.: Osborn, zb7d., p. 12, fig. 10. Name Rhynchotherium anguirivalis Osborn Ibid., p. 18. Type loe.: Sioux County, western Nebraska, Quarry 3 (Oleott Hill). Type fig.: Osborn, zbd., p. 13, fig. 11. Anancus falconert Osborn Tbid., pp. 13, 14. Type loc.: Red or Norwich Crag of Suffolk, England. Type fig.: Originally described and figured by Falconer as Masto- don (Tetralophodon) arvernensis (1857, Quart. Journ. Geol. Soe. London, XIII, Pl. xu, figs. 3, 4; 1868, ‘“Palaeont. Mem.,’’ IT, Pl. rv, figs. 3, 4). Osborn, ibid., p. 14, fig. 12. Parelephas protomammonteus (Matsumoto) typicus Matsumoto Sci. Rept. Tohoku Imp. Univ., (2), Geol., X, No. 2, pp. 43-48. Originally named Euelephas protomammonteus by Matsumoto in 1924, p. 262 (publish- edin Japanese). See above, this list. {According to Shikama (1937, p. 164) this species is synonymous with the species antiquus.—Colbert-Simpson, letter, July 12, 1939.—Editor.] Parelephas protomammonteus proximus, mut. nov. Matsumoto Jbid., pp. 48-50. Type loe.: Isone, Kokubo, Onuki-mura, Kimitsu District, Province of Kazusa, Japan. Type fig.: Matsumoto, zbid., Pl. xxrv, figs. 1, 2. [If protomammonteus, according to Shikama, is a synonym of antiquus (see preceding note under Parelephas protomammonteus typicus Masumoto), then Shikama was justified inemending the name of Matsumoto’s second species to read Parelephas proximus proxi- mus.—Colbert-Simpson, letter, July 12, 193°.—Editor.] Elephas eellsi Hay Journ. Wash. Acad. Sci., XVI, No. 6, pp. 154-159. Type loe.: Port Williams, Clallam County, Washington. Type figs.: Hay, ibid., p. 156, figs. 1, 2. Anancus orartus Hay Proc. U. 8. Nat. Mus., LX VIII (1927), No. 2625, Art. XXIV, pp. 8-14. Type loc.: West bank of Aransas River, San Patricio County, near town of Sinton, Texas. Type figs.: Hay, zbzd., Pl. 1, fig. 1; Pl. m1, fig. 1; Pl. rv, fig. 1; Pl. v, figs. 1-3. Anancus defloccatus Hay Tbid., pp. 14-16, 18. Type loc.: West bank of Aransas River, San Patricio County, near town of Sinton, Texas. Type figs.: Hay, zb7d., Pl. vr, and Pl. vim, fig. 1. Anancus bensonensis Gidley U.S. Geol. Surv., Profess. Paper 140—-B, pp. 85, 86. Type loc.: Near Benson, Cochise County, west side of San Pedro Valley, Arizona. Type fig.: Gidley, zbed., Pl. xxxn. Trilophodon (Serridentinus) pojoaquensis Frick Bull. Amer. Mus. Nat. Hist., LVI, Art. II, pp. 125, 142-150, 161, 162-165. Type loc.: About twenty-four miles north of Santa I'é, Santa Fé County, near Pojuaque, New Mexico. ‘Type figs.: Frick, 2bid., p. 148, fig. 26, and p. 144, fig. 22A. Amer. Mus. 21115 (type skull). The skull (f°.: A. M. 21125), originally considered by Frick as a paratype of Triloph- odon (Serridentinus) pojoaquensis, proved on subsequent examination to be referable to his new genus and species Ocalientinus ojocaliensis. ?Trilophodon (Serridentinus) leidii Frick Tbid., pp. 137, 140, 141, 155, 169. Type loe.: Alachua clays, Mixson’s bone bed, near Williston, Levy County, Florida. Type figs.: Leidy and Lueas, 1896, Trans. Wagner Free Inst. Sci., IV, Pl. tv, figs. 9-11; Frick, 1926, zbid., figs. 20A,B. ?Rhynchotherium (Dibelodon) edensis Frick Ibid., pp. 169-176. See note under Trilophodon (Tetrabelodon) shepardi edensis Frick, 1921, above, this list. E. [Elephas] primigenius Matsumotoi Dietrich Neues Jahrb. Min., I, Abth. B, Referate, p. 314. [The specimen considered as the type of Elephas primigenius matsumotot by Dietrich, 1927, is regarded by Shikama (1937, pp. 164, 165) as belonging to Parelephas proximus (see above, this list, under Parelephas protomammonteus proximus mut. nov. Matsumoto, 1926); therefore the former of these names becomes a synonym of the latter.—Colbert-Simpson, letter, July 12, 1989.—Hditor.] 1411 Roference in Present Memoir Serridentinus qguatemal- ensis Rhynchotherium angui- rivale Anancus falconeri Palzxoloxodon proto- mammonteus Palxoloxodon proto- mammonteus proxi- mus Parelephas(?) eellsi Cordillerion orarius Cordillerion defloccatus Cordillerion bensonen- Sis Trilophodon pojoaquen- sis Ocalientinus (Serri- dentinus) floridanus leidit Rhynchotherium shepardi edense [Not determined by the present author] 1927 1927 1927 1928 1928 1928 1928 1928 1928 1928 OSBORN: THE PROBOSCIDEA Name Author Amebelodon fricki Barbour Neb. State Mus., Bull. 13, I, pp. 131-134. Type loc.: Freedom, Frontier County, Nebraska. Type figs.: Barbour, 7b7d., text figs. 89-91. Archidiskodon transvaalensis Dart Supplement, Nature, No. 3032, pp. 41-48. Type loc.: Near Bloemhof, South Africa. Type figs.: Dart, ibid., p. 47, fig. 6 (right); p. 48, fig. 7 (left). Archidiskodon Sheppardi Dart Loc. cit.. Type loc.: Near Bloemhof, South Africa. Type figs.: Dart, ibid., p. 47, fig. 6 (left); p. 48, fig. 7 (right). Elephas indicus Buski Matsumoto Sci. Rept. Tohoku Imp. Univ., (2), Geol., X, No. 3, pp. 57, 58. Type loe.: Ninohe District, Province of Mutsu, Japan. Type fig.: Matsumoto, zbid., Pl. xxvu, figs. 2, 3. Leith-Adamsia siwalikiensis Matsumoto Japanese Journ. Geol. and Geog., V, No. 4, Art. XII. Type loc.: India. Type figs.: Falconer and Cautley, 1845, “Fauna Antiqua Sivalensis,” Pl. x1, fig. 4; 1847, Pl. x1v, fig. 8 (as Elephas planifrons). Elephas primigenius Blumenbach var. n. pachyganalis Schroeder Jahrb. preuss. geol. Landes- anstalt, XLVIII, pp. 701, 702, 718. Type loc.: Rixdorf, Germany. Type fig.: Schroeder, zbid., Taf. xxxv1, figs. la, 1b. Trilophodon (Serridentinus?) inopinatus Borissiak and Beliaeva Bull. Acad. Sei. U.S. 8. R., Cl. Sci., Phys.-Math., pp. 241-252. Type loc.: Jilanéik beds of the Turgai region, U.S.S8.R. Type figs.: Borissiak and Belfaeva, zbid., Pls. 1, 11. Archidiskodon subplanifrons Osborn Nature, CX XI, No. 3052, pp. 672, 673. Type loc.: Sydney- on-Vaal, Vaal River, South Africa. Type fig.: Osborn, zbzd., p. 672, fig. 1. Archidiskodon broomi Osborn Ibid., pp. 672, 673. Type loe.: The Bend, Vaal River, near Kimberley, South Africa. Type fig.: Osborn, zbid., p. 672, fig. 2. Elephas exilis Stock and Furlong Science, N. 8., LX VIII, No. 1754, p. 140. Type loc.: Santa Rosa Island, California. Type fig.: Stock, 1935, Sci. Monthly, XLI, September, p. 210, fig. 6 (see also Figs. 920 and 921 of the present Memoir). Elephas haroldcooki Hay Proc. Colo. Mus. Nat. Hist., VIII, No. 2, Pt. I, p. 33. Type loc.: Frederick, Oklahoma. Type figs.: Hay and Cook, 1930, zbid., IX, No. 2, Plime. 1) Ply, fig: 1, Pls: xa and xn. Trilophodon phippsi Cook Proc. Colo. Mus. Nat. Hist., VIII, No. 4, pp. 37-48. Type loc.: Near Ainsworth, Brown County, Nebraska. Type figs.: Cook, zbzd., Pls. am. {In a footnote on page 328 of Volume I of the present Memoir Trilophodon phippsi is included among the species referable to Megabelodon, the‘‘Spoonbill Mastodonts.” While on page 707 Professor Osborn mentions Megabelodon lulli, M. cruziensis, and M. joraki, we fail to find anywhere the use of the term Megabelodon phippsi. However, it was evidently his intention to assign phippsi to Megabelodon because of its tuskless mandible, the dis- tinetive character of this genus. Therefore on page 738 it has been listed under Megabelo- don with the three above-mentioned species.—Editor. } Serridentinus fricki Peterson Mem. Carnegie Mus., XI, No. 2, pp. 111-121. Type loc.: Northern flanks of Douglas Mountain on Weller Ranch, Moffat County, Colorado. Type figs.: Peterson, zbid., Pls. x1—x1v, and text figs. 21,22; also restoration by A. Avinoff (Pl. xv). Loxodonta africana angolensis Frade “Titulos e trabalhos cientificos (Curriculum vitae),’”’ p. 15. Type loe.: Region of Cunene, southern Angola, Africa. Living form. Mastodon andium Kraglievichii Berro ‘Un nuevo Mastodon en la R. O. del Uruguay.” Fide Cabrera, 1929, Rev. Mus. La Plata, XXXII, pp. 119, 141; not seen by the present author. Mastodonte de Alangasi. See ‘1931 Bunolophodon postremus Spillmann,’” below, this list. Reference in Present Memoir Amebelodon fricki Palzoloxodon trans- vaalensis Palxoloxodon sheppardi ? Palxoloxodon buski Archidiskodon plani- frons [Not determined by the present author] Trilophodon inopinatus Archidiskodon sub- planifrons Archidiskodon broomi Archidiskodon exilis Archidiskodon harold- cooki Megabelodon phippsi Trilophodon frickv Loxodonta africana angolensis Cordillerion andium kraglievichir NOMENCLATURE OF THE PROBOSCIDEA Year Name Author 1928 Platybelodon Danovi Borissiak Ann. Soc. Paléont. Russie, VII, pp. 105-120. Type loc.: Kuban region, North Caucasus, Russia. Type fig.: Borissiak, ibéd., Taf. vim, figs. 1-4. 1928 Tetrabelodon (Bunolophodon) Ayora Spillmann El Ecuador Comercial, Afio VI, No. 57. Type 1931 loc.: Quebrada [canyon] of Chalang, near Punin, southeast of Riobamba, Province of Chimborazo, Ecuador. Type fig.: Spillmann, 7bid., fig. 2 (see also figures 548 and 549 of the present Memoir). Supplementary description: Spillmann, 1931, “Die Séugetiere Ecuadors im Wandel der Zeit,’’ Erster Teil, p. 67. 1929 Elephas (Palzoloxodon) namadicus setoensis Makiyama Chikyi—The Globe, XII, No. 5, pp. 364, 365 (in Japanese). Type loc.: Seto, Japan. [Dr. Jir6 Makiyama in a letter to the editor, dated Kyoto, August 26, 1937, stated that the name Elephas (Palxoloxodon) namadicus setoensis Makiyama appears on page 365 of his article of 1929 in Chikyd, without diagnosis or figure, but by “designation points, as an example, to the specimen given in his paper ‘Notes on Fossil Elephant from Sahamma, Totomi’ (Mem. Coll. Sci. Kyoto Imp. Univ., Ser. B, Vol. I, No. 2, 1924, pp. 263, 264, PI. xvI, fig. 2, as Elephas namadicus namadi).”’ This was an outline in English of the note subsequently given by him in Japanese in the publication Chiky—The Globe, XII, No. 5, pp. 364, 365, November, 1929, with the translated title “Scientific Names of Fossils and the International Rules of Zoological Nomenclature,” pp. 358-366. Doctor Makiyama further stated that during the intervening years he had changed his opinion somewhat and that he hoped to publish the results of his studies in the near future. This article has since appeared under the title ‘“JJaponie Proboscidea,’’ Mus. Coll. Sci. Kyéto Imp. Univ., Ser B, Vol. XIV, No. 1, Art. 1, May, 1938.—Kditor. | 1929 Amebelodon grangeri Osborn Nat. Hist., XXIX, No. 1, pp. 12-16. Type loc.: Tairum Nor 1931 Basin, Mongolia. Type fig.: Osborn, zbid., p. 15 (plate). Supplementary description: Osborn and Granger, 1931, Amer. Mus. Novitates, No. 470, pp. 6, 7, text figs. 1 and 3A, Al. Transferred to genus Platybelodon, owing to fundamental differences in the structure of the tusks, e. g., the presence of dentinal rod-cones in Platybelodon (characteristic of grangeri), but which are absent in Amebelodon. 1929 Torynobelodon loomisi Barbour Neb. State Mus., Bull. 16, I, pp. 147-150. Type loc.: Sand Canyon, two and a half miles southwest of Republican City, Harlan County, Nebraska. Type figs.: Barbour, zb7d., text figs. 98-100. 1929 Trilophodon angustidens gaillardi Osborn Amer. Mus. Novitates, No. 393, p. 1. Type loc.: Villefranche d’Astaraec (Gers), France. Type fig.: Osborn, ibid., p. 1, fig. 1. 1929 Serridentinus bifoliatus Osborn Ibid., p. 2. Type loc.: Brewster, Polk County, Florida. Type fig.: Osborn, ibid., p. 2, fig. 2. 1929 Serridentinus hasnotensis Osborn Loc. cit. Type loc.: Near Hasnot, India. Cotype figs.: Osborn, zbid., p. 3, fig. 3. 1929 Serridentinus metachinjiensis Osborn Ibid., pp. 4, 5. Type loc.: One mile northwest of Chinji Bungalow, India. Type fig.: Osborn, zbid., p. 3, fig. 4. 1929 Serridentinus chinjiensis Osborn Ibid., p. 5. Type loc.: One mile and a half west of Chinji Bungalow, India. Type fig.: Osborn, zbid., p. 4, fig. 5. 1929 Serridentinus prochinjiensis Osborn Jbid., p. 6. Type loe.: Two miles west of Chinji Bungalow, India. Type fig.: Osborn, zbzd., p. 5, fig. 6. 1929 Serridentinus florescens Osborn Ibid., p. 6. Type loc.: Kholobolehi Nor region, Mongolia, five to eight miles north of camp. Type fig.: Osborn, zb7d., p. 7, fig. 7. 1929 Rhynchotherium paredensis Osborn Ibid., pp. 6-8. Type loe.: Mt. Eden Hot Springs, San Bernardino County, California. Type fig.: Osborn, zbid., p. 7, fig. 8. Originally figured by Frick, 1926, Bull. Amer. Mus. Nat. Hist., LVI, Art. II, figs. 2,8, 9 as (?)Rhynchotheriwm (Dibelodon) edensis. In the present Memoir (Vol. I, p. 496, also fig. 474) R. paredensis is regarded as a synonym of R. shepardi edense. 1413 Reference in Present Memoir Platybelodon danovi Cuvieronius ayore [Not determined by the present author] Platybelodon grangeri Torynobelodon loomisi Trilophodon angusti- dens gaillardi Ocalientinus (Serri- dentinus) bifoliatus Serridentinus hasnoten- Sis Serridentinus metachin- jiensis Serridentinus chinjien- sts Serridentinus prochin- quensis Ocalientinus (Serri- dentinus) florescens Rhynchotherium shepardi edense 1414 Year 1929 1929 1929 1929 1929 1929 1929 1929 1929 1929 1929 1929 1929 1929 1929 1929 OSBORN: THE PROBOSCIDEA Author Rhynchotherium chinjiensis Osborn Ibid., p. 8. Type loc.: Two miles west of Chinji Bungalow, India. Type fig.: Osborn, zbid., p. 8, fig. 9. Name Synconolophus dhokpathanensis Osborn Ibid., pp. 10-12. Type loe.: Three miles west of Dhok Pathan, India. Type fig.: Osborn, ibid., p. 9, fig. 10. Synconolophus ptychodus Osborn Ibid., p. 12. Type loc.: Four miles west of Chinji Bungalow, India. Type fig.: Osborn, ibid., p. 11, fig. 12. Synconolophus propathanensis Osborn Ibid., pp. 12, 18. Type loc.: Three miles east of Dhok Pathan, India. Type fig.: Osborn, ibid., p. 18, fig. 13. Stegolophodon nathotensis Osborn Ibid., pp. 13-15. Type loc.: Near Nathot, India. Type fig.: Osborn, ibid., p. 14, fig. 14. Stegolophodon cautleyi progressus Osborn Ibid., p. 15. Type loc.: Twelve miles east of Chinji Bungalow, India. Type fig.: Osborn, ibid., p. 14, fig. 15. Stegodon insignis birmanicus Osborn Ibid., pp. 15,16. Type loc.: Mingoon, opposite Mandalay, Burma. Type fig.: Osborn, zbid., p. 16, fig. 16. Stegodon orientalis grangeri Osborn Ibid., pp. 16, 17. Type loe.: Yenchingkou, near Wanhsien, Province of Szechuan, China. Type fig.: Osborn, zbid., p. 16, fig. 16. Stegodon pinjorensis Osborn Ibid., p. 18. Type loc.: Three miles north of Siswan, India. Type fig.: Osborn, zbid., p. 17, fig. 17. Archidiskodon sonoriensis Osborn TIbid., p. 18. Type loc.: One mile east of Arizpe, northern Sonora, Mexico, on the Sonora River. Type fig.: Osborn, zbzd., p. 18, fig. 18. Parelephas floridanus Osborn Ibid., p. 20. Type loc.: Manatee County, Florida, two miles south of Bradenton. Type fig.: Osborn, zb7d., p. 19, fig. 19. Parelephas columbi cayennensis Osborn Ibid., pp. 20, 21. Type loc.: Cayenne, French Gui- ana, South America. Type fig.: Osborn, zbid., p. 21, fig. 20. Elephas platycephalus Osborn — Ibid., pp. 21, 22. Type loe.: Near Siswan, bed of Amilee Creek, Simla Hills, India. Type fig.: Osborn, ibid., p. 22, fig. 21. Elephas platycephalus angustidens Osborn Ibid., pp. 22, 23. Type loc.: Three miles west of Chandigarh, Siwalik Hills, India. Type fig.: Osborn, zbd., p. 23, fig. 22. Loxodonta (Palxoloxodon) namadica (Yabei) Matsumoto Sci. Rept. Tohoku Imp. Univ., (2), Geol., XIII, No. 1, pp. 4, 5. Type loc.: Inland Sea, Japan. Type figs.: Matsumoto, zbzd., Pl. 1, fig. 2, Pl. tv. Lor. (Pal.) Tokunagai junior, mut. nov., Matsumoto Tbid., p. 10. Type loc.: Japan, precise locality unknown. Type fig.: Matsumoto, zbid., Pl. vir, figs. 1, 2. Notiomastodon ornatus Cabrera Rey. Mus. La Plata, XXXII, p. 91. Type loe.: Monte Her- moso, Province of Buenos Aires, Argentina. Type figs.: Cabrera, zbid., p. 91, fig. 2; p. 93, fig. 4. Archidiskodon vanalpheni Dart So. Afr. Journ. Sci., XX VI, p. 704. Type loc.: Sydney-on-Vaal, South Africa. Type figs.: Dart, zbid., p. 704, figs. 8, 9. Archidiskodon milletti Dart Tbid., pp. 706-708. Type loc.: Sydney-on-Vaal, South Africa. Type figs.: Dart, zbid., p. 706, figs. 10, 11. Archidiskodon loxodontoides Dart Ibid., pp. 709-711. Type loce.: Sydney-on-Vaal, South Africa. Type fig.: Dart. zbid., p. 709, fig. 18. Reference in Present Memoir Rhynchotheriwm chingiense Synconolophus dhok- pathanensis Synconolophus ply- chodus Synconolophus pro- pathanensis Stegolophodon nathoten- sis Stegolophodon cautleyi progressus Stegodon insignis birmanicus Stegodon orientalis grangert Stegodon pinjorensis Archidiskodon sonori- ensis Parelephas floridanus Parelephas columbi LAYENNENSIS Platelephas platyceph- alus Hypselephas hysudricus Palxoloxodon namadi- cus yabei Palzxoloxodon (?Archi- diskodon) tokunagat mut. Junior Notiomastodon ornatus Archidiskodon vanal- phenti Archidiskodon milletti Archidiskodon loxodon- toides 1929 1929 1929 1929 1929 1929 1930 1930 1930 1930 1930 1930 1930 1931 1931 NOMENCLATURE OF THE PROBOSCIDEA Author Archidiskodon andrewsi Dart Ibid., pp. 711-713. Type loc.: Gong-Gong, Vaal River, South Africa. Type fig.: Dart, ibid., p. 711, fig. 14. Name Archidiskodon hanekomi Dart Ibid., pp. 713-715. Type loc.: Delport’s Hope, Vaal River, South Africa. Type figs.: Dart, zbd., p. 713, figs. 15, 16. Archidiskodon yorki Dart Ibid., pp. 715, 717. near Christiana, South Africa. Type loc.: Vanasswegenshoek—Bloemheuvel, Type fig.: Dart, ibid., p. 717, fig. 19. Pilgrimia yorki Dart Ibid., pp. 719, 720. Type loc.: Below Christiana, Vaal River, South Africa. Type figs.: Dart, zbzd., p. 719, figs. 20, 21. Pilgrimia wilmani Dart Tbid., pp. 720-722. Type loc.: Below Christiana, Vaal River, South Africa. Type fig.: Dart, ibid., p. 721, fig. 22. Pilgrimia kuhni Dart Tbid., pp. 723, 724. Type loc.: Pniel Estate, South Africa. Type fig.: Dart, zbid., p. 725, fig. 24. Loxodonta prima Dart Ibid., pp. 724-726. Type loc.: Pilandsberg, Transvaal, South Africa. Type figs.: Dart, zbid., p. 725, figs. 25, 26. Loxodonta africana var. obliqua Dart Ibid., pp. 726-728. Type loe.: Valley of Steelport River, tributary of Oliphants River, northeast Transvaal, South Africa. Type figs.: Dart, cbid., p. 726, figs. 27, 28. Gomphotherium? emmonsi Hay “Second Bib. and Cat. of Fossil Vertebrata of North America,” II, p. 636. Type loe.: Halifax County, North Carolina. Type figs.: Emmons, 1858, “Rept. North-Carolina Geol. Surv.,’”’ p. 199, fig. 23; Emmons, 1860, “Manual of Geology,” p. 218, fig. 186. Gomphotherium serpentirivale Hay Op. cit., p. 639. Amebelodon sinclairi Barbour Neb. State Mus., Bull. 17, I, pp. 155-158. Type loc.: Freedom, Frontier County, Nebraska. Type fig.: Barbour, zbid., fig. 101. Pliomastodon sellardsi Simpson Bull. Amer. Mus. Nat. Hist., LIX, Art. III, pp. 203-206. Type loe.: Brewster, Florida. Type figs.: Simpson, ibid., p. 204, fig. 30; p. 205, fig. 31. Gomphotherium priestleyi Hay and Cook Proc. Colo. Mus. Nat. Hist., IX, pp. 31, 32. Type loc.: Near Frederick, Tillman County, Oklahoma. Type fig.: Hay and Cook, zbid., Pl. xm, figs. 1, 2. Prodinotherium hungaricum Ehik Geol. Hungarica, Ser. Palewont., Fase. 6, pp. 1-14. Type loc.: Kotyhaza (Dep. Nograd), Hungary. Type figs.: Ehik, cbid., Pl. 1, figs. 1-3; Pls. 11-Iv. Bull. Dept. Geol. Univ. Calif., XIX, No. 16, pp. 335-341. Type figs. : Pliomastodon verillarius Matthew Type loe.: Southeast of Coalinga, Fresno County, California. Matthew, zbzd., Pl. xL1—x.tv, text figs. 1, 2. Bunolophodon postremus Spillmann ‘Die Saiugetiere Ecuadors im Wandel der Zeit,” Erster Teil, pp. 73-107. Type loc.: Quebrada [canyon] of Cachihuayco, near Alangasi, Province of Pichincha, east of Quito, Ecuador. Type figs.: Spillmann, 1929, Natur und Museum, LIX, Heft II, text figs. 3, 4, and associated skeleton, fig. 1. Material first described in “El Ecuador Comercial,’’ 1928, Ato VI, No. 57, p. 75, as “Mastodonte de Alangasi.”’ Teleobunomastodon Revilliod, 1931, bolivianus Philippi, 1893. Revilliod, Mém. Soc. Paléont. Suisse, LI, pp. 1-21. Type loc.: Exact locality unknown, probably from the vicinity of Calacoto, valley of the Rio Desaguadero, Bolivia, south of Corocoro, or from Concordia. Type figs.: Revilliod, zbid., Pls. 1, 11, and fig. 3, p. 9. 1415 Reference in Present Memoir Palzxoloxodon(?) an- drewsi Palzxoloxodon hanekomi Archidiskodon yorki Palzxoloxodon yorki Palzxoloxodon wilmani Palzoloxodon kuhni Loxodonta prima Loxodonta africana var. obliqua Ocalientinus emmonsi NOMEN NUDUM Amebelodon sinclairi Pliomastodon sellardsi Stegomastodon priest- leyi Deinothertum hungari- cum Pliomastodon vecxil- larius Cuvieronius postremus Cordillerion (?)bolivi- anus 1416 Year 1931 1931 1931 1931 1931 1932 1932 1932 1932 1932 1932 1932 1933 1933 OSBORN: THE PROBOSCIDEA Author Torynobelodon barnumbrowni Barbour Neb. State Mus., Bull. 22, I, pp. 191-198. Type loc.: Snake River, Cherry County, southwest of Valentine, Nebraska. Type figs.: Barbour, zbrd., figs. 123, 124. Name Mastodon moodiei Barbour Neb. State Mus., Bull. 24, I, pp. 203-210. Type loc.: West Blue River, southwest of Milford, Seward County, Nebraska. Type figs.: Barbour, ibid., figs. 130, 131, 132b. Parelephas protomammonteus (Matsumoto) matsumotot Saheki Japanese Journ. Geol. and Geog., VIII, No. 3, pp. 125-129. Type loc.: Mishima, Kimitsu district, Chiba Prefecture, Province of Kazusa, Japan. Type fig.: Saheki, zbzd., Pl. xv, figs. 1-3. |Parelephas protomammonteus matsumotot Saheki, 1931, is invalid, since matsumotot in this case is a homonym of matsumotot Dietrich, 1927 (see above, this list, under Z. [Hlephas| primigenius Matsumotot Dietrich, 1927). If the material described by Saheki is distinct, as he thought it was, it is material at present without a name.—Colbert-Simpson, letter, July 12, 1939.—Editor. | Palxoloxodon antiquus italicus Osborn Amer. Mus. Novitates, No. 460, pp. 1-24. Type loc.: Pignataro Interamna, near Cassino, Italy. Type figs.: Osborn, ibid., figs. 1, 4-16. Palxoloxodon antiquus (andrewsi?) Osborn Tbid., pp. 1, 19, 21, 23. Type loc.: Upnor on the banks of the Medway, Kent, England. Type fig.: Osborn, zbzd., p. 21, fig. 15; also Andrews and Cooper, 1928, ‘On a Specimen of Elephas antiquus from Upnor,” Pls. 1-vi, text figs. 1-5, and figs. 1079-1082 and 1084 of the present Memoir. Pilgrimia archidiskodontoides Haughton Trans. Roy. Soc. So. Africa, XXI, Pt. I, pp. 4-8. Type loe.: Sydney-on-Vaal Breakwater, bed of Vaal River, South Africa. Type figs.: Haughton, zbzd., Pls. m1. Pilgrimia subantiqua Haughton Ibid., pp. 8-10. Type loe.: Delport’s Hope, half a mile from Vaal River, South Africa. Type fig.: Haughton, zbzd., Pl. rv, figs. 1, 2. Serridentinus gobiensis Osborn and Granger Amer. Mus. Novitates, No. 537, pp. 11-13. Type loc.: About forty miles southeast of Iren Dabasu, Inner Mongolia. Type fig. : Osborn and Granger, zbid., p. 12, fig. 8. Archidiskodon meridionalis nebrascensis Osborn Proc. Colo. Mus. Nat. Hist., XI, No. 1, pp. 1-3. Type loc.: Near Angus, Nuckolls County, Nebraska. Type figs.: Osborn, ibid., figs. 1, 2. Trilophodon cooperi Osborn Amer. Mus. Novitates, No. 585, pp. 1-6. Type loc.: Dera Bugti, Baluchistan. Type fig.: Osborn, ibid., p. 2, fig. 1; paratype, p. 3, fig. 2 (deseribed by Forster Cooper, 1922, Proc. Zool. Soc. London, p. 610, as referable to Bunolophodon angustidens). Tetralophodon bumiajuensis van der Maarel “Contribution to the Knowledge of the Fossil Mam- malian Fauna of Java,’ pp. 2, 3, 108-121. Type loc.: Bumiaju, central Java. Type figs.: van der Maarel, op. cit., Pls. vin, rx, x, and text figs. 17-21. Stegodon bondolensis van der Maarel Op. cit., pp. 158-164. Type loc.: Bondol near Kuwung, District Randublatung, Regency Blora, Residency Rembang, Java. Type figs.: van der Maarel, op. cit., Pl. xrv, figs. 1, 4, 5, and text figs. 24, 25. Blickotherium blicki: Frick Type loe.: Near Tapasuma, Gracios, Honduras. figs. 3, 4. Bull. Amer. Mus. Nat. Hist., LIX, Art. LX, pp. 509, 515, 527-531. Type figs.: Frick, zbed., Aybelodon hondurensis Frick Ibid., pp. 527, 528, 582. Type loc.: Near Tapasuma, Gracios, Hon- duras. Type figs.: Frick, zbid., figs. 5, 18, 18. Reference in Present Memoir Torynobelodon barnum- brown Mastodon moodiet [Not determined by the present author] Hesperoloxodon antiqu- us italicus |Hesperoloxodon antiqu- us (fide Simpson, see p. 1222 above)] Palzxoloxodon archi- diskodontoides Loxodonta subantiqua Serridentinus gobiensis Archidiskodon meridi- onalis nebrascensis Trilophodon cooperi Tetralophodon bumia- Juensis Stegodon bondolensis Blickotherium blicki Aybelodon hondurensis Year 1933 1933 1933 1933 1933 1933 1933 1933 1933 1933 1933 1933 1933 1933 1933 1934 1934 1934 NOMENCLATURE OF THE PROBOSCIDEA Name Author Serridentinus filholi Frick Tbid., pp. 509, 535. Type loc.: Gers, France. The mandible (Mus. d’Hist. Nat. Paris AC 2058, 2062) of this species was referred to in the article by Frick, 1926, Bull. Amer. Mus. Nat. Hist., LVI, Art. II, pp. 177, 178, but the name Serridentinus filholi was not assigned until 1933. Trilophodon cruziensis Frick Ibid., pp. 505, 579. Type loe.: Santa Cruz, New Mexico. Type figs.: Frick, zbid., figs. 10, 12A, 17, 23A, 25. Ocalientinus ojocaliensis Frick Tbid., pp. 509, 576,579. Type loe.: Ojo Caliente, New Mexico. Type figs.: Frick, ibid., figs. 7, 18. Trobelodon taoensis Frick Ibid., pp. 505, 580. Type loe.: Santa Cruz, New Mexico. Type figs. : Frick, 7bid., figs. 2, 13, 18. Tatabelodon riograndensis Frick Ibid., pp. 505, 581. Type loc.: Battleship Mountain, New Mexico. Type figs.: Frick, zbid., figs. 6, 13. (?)Amebelodon joraki Frick Tbid., pp. 505, 582. Type loc.: Santa Cruz, New Mexico. Type fig.: Frick, ibid., fig. 18. This type (a tuskless mandible) was first referred by the present author (Vol. I, p. 326, this Memoir) to the genus T’rilophodon, but finally in the Appendix to Volume I, pp. 707, 738, he definitely assigned it to Megabelodon. Serbelodon barbourensis Frick Ibid., pp. 506, 592, 594. Type loc.: Christmas quarry, near Ains- worth, Nebraska. Type figs.: Frick, b7d., figs. 14, 16, 27. Tatabelodon gregorii Frick Ibid., pp. 506, 597. Type loc.: Vicinity of Ainsworth, Nebraska. Type figs.: Frick, ibzd., figs. 13, 27A. (?) Trilophodon barstonis Frick Tbid., pp. 506, 607. Type loc.: Mohave Desert, California. Type figs.: Frick, 7zbid., figs. 33, 36. Mastodon raki Frick Ibid., pp. 506, 630. Type loc.: Hot Springs, New Mexico. Type figs.: Frick, zbid., figs. 25A, 29A. Mastodon americanus alaskensis Frick TIbid., pp. 506, 631. Type loc.: Vicinity of Fairbanks, Alaska. Type fig.: Frick, zbzd., fig. 29A. Elephas primigenius alaskensis Osborn (In Frick, zbzd., pp. 631, 632.) Type loc.: Vicinity of Fairbanks, Alaska. Type figs.: Osborn, this Memoir, Vol. II, figs. 1025, 1026. Serbelodon burnhami Osborn Amer. Mus. Novitates, No. 639, pp. 1-5. Type loc.: Near Ricardo, San Bernardino County, California. Type figs.: Osborn, zb7d., figs. 1, 2. Stegodon trigonocephalus praecursor von Koenigswald Wetenschappelijke Mededeelingen, Dienst Mijnbouw Nederl.-Indié, I Teil, No. 23, pp. 104, 105. Type loe.: Bumiaju, Java. Type fig.: von Koenigswald, zbzd., Taf. xxvn, fig. 2. Cryptomastodon martini von Koenigswald J[bid., pp. 111-119, Taf. xxvmt, figs. 1-3, and text figs. 8, 9. D. |Dinotherium| Bozasi Arambourg Compt. Rend. Soc. géol. France, No. 6, pp. 86, 87. Type loe.: Valley of the Omo, Abyssinia. Type fig.: Arambourg, 1935, Bull. Soe. géol. France, (5), IV, Pl. xvi (mandible). Supplementary description: Arambourg, op. cit., pp. 305-310. Mastodon grangeri Barbour Neb. State Mus., Bull. 35, I, pp. 287-290. Type loc.: Pender, Thurston County, Nebraska. Type fig.: Barbour, zbid., p. 289, fig. 170. Palzoloxodon yokohamanus Tokunaga Journ. Geog. (Tokyo), XLVI, No. 546, pp. 363-371 (in Japanese). Type loc.: Mouth of Tsurumi-gawa, Yokohama, Japan. Type fig.: Tokunaga, zbid., Pl. vit, figs. 1, 2. 1417 Reference in Present Memoir Serridentinus filholi Megabelodon cruziensis Ocalientinus ojocalien- SUS Trobelodon taoensis Trilophodon (Tatabelo- don) riograndensis Megabelodon joraki Serbelodon barbourensis Trilophodon (Tatabelo- don) gregorit Serridentinus barstonis Mastodon raki Mastodon americanus alaskensis Mammonteus primi- genius alaskensis Serbelodon burnhami Stegodon trigonoceph- alus praecursor ?2SIRENIAN { Not determined by the present author] . Mastodon grangeri [Not determined by the present author] 1418 Year 1934 1934 1935 1935 1935 1935 1935 1935 1935 1935 1935 1935 1935 1936 1936 OSBORN: THE PROBOSCIDEA Name Author Parastegodon? kwantoensis Tokunaga Tbid., pp. 365-369 (in Japanese). Type loec.: Kakio, Kanagawa Prefecture, Japan. Type fig.: Tokunaga, zbid., Pl. rx, figs. 1-3. Archidiskodon proplanifrons Osborn Amer. Mus. Novitates, No. 741, pp. 10-12. Type loc.: Gong-Gong, near the Vaal River, South Africa. Type fig.: Osborn, zbid., p. 5, fig. 2. Palzoloxodon priscus var. bosei Chakravarti Proc. 22nd Indian Sci. Congress, Calcutta, p. 209. Type loc.: Parkalta near Jammu, India. Type fig.: Bose, 1929, Quart. Journ. Geol. Min. and Metallurg. Soc. India, II, No. 3, Pl. v, fig. 10, as Stego- don bombifrons. Supplementary description: Chakravarti, 1937, Quart. Journ. Geol. Min. and Metallurg. Soc. India, IX, No. 2, pp. 39-42, Pl. v1 (figured as Palxoloxodon sp.). Gnathabelodon thorpet Barbour and Sternberg Neb. State Mus., Bull. 42, I, pp. 395-404. Type loc.: Near Ogallah, Trego County, western Kansas. Type figs.: Barbour and Sternberg, zbid., figs. 187-191. Stegodon yiishensis Young Pal. Sinica, (C), IX, Fase. 2, pp. 26-28. Type loc.: Yiishe, China. Type fig.: Young, zbzd., Pl. v, figs. 1, la. Stegodon zdanskyi Hopwood (In Young, ibid., p. 28); Hopwood, Pal. Sinica, (C), IX, Fase. 3, p. 75. Type loe.: Unknown. Type fig.: Hopwood, zbid., Pl. vu, fig. 5. Stegodon officinalis Hopwood (In Young, ibid., Fase. 2, pp. 27,30.) Hopwood, zbid., Fase. 3, p. 73. Type loe.: (?)Szechuan, China. Type fig.: Hopwood, zbid., Pl. vir, fig. 3. Trilophodon connerus Hopwood Ibid., p. 14. Type loc.: Kansu, Sining Fu, SW 20 li, Shui Ch’iian, P’u, SE 5 li, Tiao Kou, China. Type fig.: Hopwood, ibid., Pl. v, figs. 1, 2. Trilophodon wimani Hopwood Ibid., p. 19. Type loc.: Pa P’an Shan, China. Type fig.: Hopwood, ibid., Pl. v, fig. 3. Trilophodon spectabilis Hopwood Tbid., p.30. Type loc.: Said to have come from Sian, China. Type fig.: Hopwood, zbid., Pl. v1, fig. 2. Tetralophodon exoletus Hopwood Ibid., p.35. Type loc.: Shansi, China. Type fig.: Hopwood, ibid., Pl. v1, fig. 3. Pentalophodon sinensis Hopwood Ibid., p. 57. Type loe.: Yi She Hsien, Shansi, China. Type fig.: Hopwood, ibid., Pl. vu, fig. 2. Parastegodon sugiyamai Tokunaga Proc. Imp. Acad. Tokyo, XI, p. 434. Type loc.: Iruhi, in Saida village, Shikoku, Japan. Type fig.: Tokunaga, ibid., p. 433, text fig. {Not determined by the present author, but regarded by Dr. E. H. Colbert as referable to Stegodon rather than to Parastegodon (which was considered by Professor Osborn as possibly equal to Archidiskodon Pohlig or to a progressive Stegodon). Consequently this species is described in the Stegodon chapter, pp. 899 and 900 above.—Kditor. ] Mastodon atavus Borissiak ‘Travaux de l'Institut Paléozoologique de l’Académie des Sciences de VU.S.S.R., V, pp. 171-234. Type loc.: Dschilantschik River, Turgai, Russia. Type figs.: Borissiak, zb¢d., Taf. 1-v, vu, and Taf. v1, vir (in part), also text figs. 1-16. Pal. Inst. No. 2280, Leningrad. Pliomastodon nevadanus Stock Publ. Carnegie Instn. Wash., No. 473, p. 37. Type loe.: Thous- and Creek basin, about four miles northwest of the Hot Spring and on east side of Railroad Ridge, Humboldt County, northwestern Nevada. Type fig.: Stock, 7bid., Pl. 1 (incomplete skull representing most of the palate, second and third superior molars of each side, and a complete right tusk—Calif. Inst. Tech. Coll. Vert. Pal. No. 1922). Reference in Present Memoir {Not determined by the present author] Archidiskodon pro- planifrons [Not determined by the present author] Gnathabelodon thor pei [Not determined by the present author] Stegodon zdanskyi Stegodon officinalis Trilophodon connexus Serridentinus wimani Trilophodon spectabilis Tetralophodon exoletus Anancus sinensis [ Stegodon(?) sugiyamaz | {Not determined by the present author] [Not determined by the present author| Year 1936 1936 1936 1936 1936 1936 1936 1936 1936 1936 1936 1936 1936 NOMENCLATURE OF THE PROBOSCIDEA Name Author Palzoloxodon aomoriensis Tokunaga Journ. Geog. (Tokyo), XLVIII, No. 564, February, pp. 67-70 (in Japanese). Type loc.: Tenjinbayashi, on the Shichinohe-gawa, near entrance to town of Shichinohe, Kamikita-gun, Aomori Prefecture, Japan. Type fig.: Tokunaga, zbéd., Pl. 1. Supplementary description: Tokunaga and Takai, 1936, Journ. Geol. Soe. Japan, XLIII, No. 511, April 20, pp. 254-258 (in English). Deinotherium hopwoodi Osborn This Memoir, I, p. 117. Type loc.: Olduvai, near southeast shore of Lake Victoria, Tanganyika Territory, Africa. Type fig.: Osborn, op. cit., p. 104, fig. 68a. Regarded by Doctor Hopwood as a synonym of Dinotherium Bozasi Arambourg, 1934. See above, this list. Trilophodon hasnotensis Osborn Op. cit., p. 279. Type loe.: Near Hasnot, India. Type fig.: Osborn, op. cit., p. 454, fig. 417. Tetralophodon fricki Osborn Op. cit., p. 375. Type loc.: Near Clarendon, northern Texas. Type figs.: Frick, 1933, Bull. Amer. Mus. Nat. Hist., LIX, Art. IX, fig. 12 (lower) and fig. 23B (upper). Rhynchotherium brownt Osborn Op. cit., p. 494. Type loc.: San José de Pimas, Sonora, Mexico. Type fig.: Osborn, 1921, Amer. Mus. Novitates, No. 1, fig. 2C, as neotype of Rhynchotherium tlascale. See also this Memotr, I, fig. 467. On locating the type cast of Rhynchotherium tlascalz in the Geneva Museum, the neo- type mandible was found to be quite different; consequently the present author made it the type of a new species, Rhynchothertum brownt. Cordillerion edensis Osborn This Memoir, I, p. 560. Type loc.: Mt. Eden Hot Springs, San Bernardino County, California. Type fig.: Frick, 1921, Bull. Dept. Geol. Univ. Calif., XII, No. 5, Pl. u (as Trilophodon shepardi edensis); Osborn, 1922, Amer. Mus. Novitates, No. 49, fig. 1, Al, A2 (as Dibelodon edensis). See also this Memoir, I, figs. 522, 523. Cf. notes under Trilophodon (Tetrabelodon) shepardi edensis Frick, 1921, and Dibelodon edensis Osborn, 1922, above, this list. Anancus properimensis Osborn This Memoir, I, p. 647. Type loc.: Near Chinji Bungalow, India. Type figs.: Osborn, op. cit., figs. 609, 613. Pentaluphodon falconert Osborn Op. cit., p. 653. Type loc.: Siwalik Hills, India. Type figs.: Falconer and Cautley, 1847, ‘Fauna Antiqua Sivalensis,” Pls. xxxir, and XXXII, figs. 1, 2 (as Mastodon sivalensis). Miomastodon depereti Osborn Op. cit., p. 693. Type loc.: Chevilly, France. Type fig.: Mayet, 1908, Ann. Univ. Lyon, Nouv. Sér., I, Sci., Méd., Fasc. 24, Pl. vu, fig. 3 (as Mastodon angustidens). Mastodon pavlowi Osborn Op. cit., p. 694. Type loc.: Pestchana, Podolia, Russia. Type fig.: Pavlow, 1894, Mém. Acad. Imp. Sci. St. Pétersb., (8), I, No. 3, Pl. 1, figs. 1, 2,3 (as Mastodon ohioticus). Mastodon acutidens Osborn Op. cit., p.696. Type loc.: Rochester, Indiana. Type figs.: Osborn, op. cit., figs. 131, Al—-A4, 135, 656, and PI. 1, L. Stegolophodon lydekkeri Osborn Op. cit., p. 700. Type loc.: Vicinity of Bruni, northwest coast of Borneo. Type fig.: Lydekker, 1885, Proce. Zool. Soc. London, Pl. xivut (as Mastodon latidens); Lydekker, 1886, ““Cat. Foss. Mamm. Brit. Mus.,” fig. 19 (as M. latidens). Stegomastodon primitivus Osborn Op. cit., p.726. Type loc.: Northeast of Ainsworth, Nebraska. Type figs.: Osborn, op. cit., figs. 674, 675. 1419 Reference in Present Memoir [Not determined by the present author] Deinotherium hopwoodi Trilophodon hasnoten- sis Tetralophodon fricki Rhynchotherium browni Cordillerion edensis Anancus properimensis Pentalophodon falconeri Miomastodon depereti Mastodon pavlowi Mastodon acutidens Stegolophodon lydekkeri Stegomastodon primiti- vUS 1420 Year 1936 1937 1937 1937 1937 1937 1937 1938 1938 1939 1939 OSBORN: THE PROBOSCIDEA Name Author Parastegodon akashiensis Takai Proe. Imp. Acad. Tokyo, XII, No. 1, pp. 19-21. Type loc.: “Shore of the cliffy coast,” west of Nishiyagi, Okubo-mura, Akashi-gun, Hyogo Prefecture, Japan. Type figs.: Takai, ¢bd., p. 20, figs. 1, 2. {Both Mr. Fuyuji Takai and Mr. Tokio Shikama simultaneously studied the same fossil material from the Akasi District, neither being aware of the description of the other. Mr. Takai chose a molar as his type, naming the species Parastegodon akashiensis, while Mr. Shikama extended his studies on this material, including two skulls with molars, a lower jaw with molars, and some other isolated teeth from the same locality, choosing as his type one of the skulls, and naming the species Paraslegodon nipponicus in his manuscript. Meanwhile Mr. Takai had announced his species in a lecture delivered at the meeting of the Palzontological Society of Japan, November 30, 1935. While this announcement did not constitute priority of description, Mr. Shikama very generously withdrew his unpublished manuscript name in favor of Mr. Takai’s species name Parastegodon akashiensis. (Cf. footnote by H. Yabe, in Takai, 1936, zbid., p. 19).—Editor.; Pentalophodon cuneatus Teilhard de Chardin and Trassaert Pal. Siniea, (C), XIII, Fase. 1, p. 11. Type loe.: Southeastern Shansi (Yushé Basin), China. Type fig.: Teilhard de Chardin and Trassaert, 7bzd., Pl. 1, fig. 4. Mastodon intermedius Teilhard de Chardin and Trassaert bid., p. 22. Type loc.: Southeastern Shansi (Yushé Basin), China. Type fig.: Teilhard de Chardin and Tras- saert, 7bed., Pl. 111, fig. 2 a-c. Stegodon licenti Teilhard de Chardin and Trassaert Jbid., p. 27. Type loc.: Southeastern Shansi (Yushé Basin), China. Type figs.: Teilhard de Chardin and Tras- saert, 2bzd., Pl. vin, figs. la, 1b, 2, and text fig. 3. Parastegodon infrequens Shikama Japanese Journ. Geol. Geogr., XIV, pp. 127-131. Type loe.: Near Akasi (precise locality unknown). Type fig.: Shikama, 7bid., Pl. rx. Type: Anterior portion of left ramus with Me zn situ. Collection of Taki- kawa Middle School in Kobe. Parelephas proximus uehataensis Shikama Tbid., p. 165. If the name Parelephas protomammonteus typicus is synonymous with antiquus, then Shikama was justified in giving a new name, Parelephas proximus uehataensis, to include secondary specimens of P. protomammonteus typicus plus those of EH. primigenius matsu- motot.—Colbert-Simpson, letter, July 12, 1939. (See notes under Parelephas protomam- monteus typicus Matsumoto, 1926, and #. [Elephas| primigenius Matsumotoi Dietrich, 1927, above, this list.) E. |Elephas| antiquus mut. ruthenensis Astre Bull. Soc. Hist. Nat. Toulouse, LXXI, p. 30. Type loec.: Salles-la-Source (Aveyron), France. Type fig.: Astre, zbid., Pl. 1 (incomplete molar), figured as “‘Hlephas antiquus, de Salles-la-Source.”’ Bunolophodon yokotii Makiyama Mem. Coll. Sci. Kyoto Imp. Univ. (B), XIV, No. 1, pp. 12-14. Type loc.: Upper Banko Sandstone of the Meisen series, at Senkaibo in the Meisen district, North Kankyo-do, Japan. Type figs.: Makiyama, ibid., text figs. 5a and 5b. Stegodon shodoensis akashiensis (Takai, 1936) Makiyama Jbid., pp. 21-27. Cotype loc.: Eiga- sima near Akasi, and under the sea off Hayasi-zaki, Japan. Cotype figs.: Makiyama, 7bid., figs. 10-12. Palxoloxodon darti Cooke and Clark Trans. Roy. Soe. So. Africa, XX VII, Pt. 3, pp. 296-302. Type loc.: Victoria Falls, northern Rhodesia. Type figs.: Cooke and Clark, ibid., Pls. x11 and xu. Archidiskodon paramammonteus Matsumoto Zool. Mag. (Tokyo), LI, No. 10, p. 704 (in Japan- ese), p. 716 (English résumé). Type loc.: Nagahama, Minato Town, Province of Kazusa, Japan. Type fig.: Matsumoto, 7bid., fig. 3. Type: Fragment of molar. Reference in Present Memoir {Not determined by the present author] [Not determined by the present author] [Not determined by the present author] [Not determined by the present author] [Not determined by the present author] [Not determined by the present author] [Not determined by the present author] [Not determined by the present author] [Not determined by the present author] [Not determined by the present author] { Not determined by the present author] CHAPTER XXII THE GEOLOGIC SUCCESSION OF THE PROBOSCIDEA By Epwin H. CoLBerr I. INTRODUCTION. Il. ArFRica. NI OV 09 Introduction. The Eocene and Oligocene of North Africa. The Miocene of North Africa. The Miocene of Central and East Africa. The Pleistocene of North Africa. The Pleistocene of Central and East Africa. The Pleistocene of South Africa. IJ. THe Ortenv. Goa SCIEN SONS Introduction. The Miocene of Baluchistan and Sind. The Siwalik Series (Miocene—Pleistocene) of North India. The Pleistocene of Central India. The Pleistocene of Ceylon. The Pleistocene of Burma. The Pleistocene of South China. The Pleistocene of Indo-China. The Pleistocene of the East Indies (Java, Borneo, Philippines). IV. Europe. virtually no text had been written. CI COON COS Introduction. The Lower Miocene: Burdigalian. The Middle Miocene: Helvetian and Tortonian- Vindobonian. The Upper Miocene: Sarmatian. The Lower Pliocene: Pontian. The Middle Pliocene: Plaisancian. The Upper Pliocene: Astian. The Pleistocene. V. Asia. SSN Con ol tee Introduction. The Miocene of Mongolia and Central Asia. The Miocene of North China. The Pliocene of Mongolia. The Pliocene of North China. The Pleistocene of North China. The Miocene to Pleistocene of Japan. VI. NortH AMERICA. COM COUN — Se) a Introduction. The Upper Miocene: Barstovian. The Lower Pliocene: Clarendonian. The Middle Phocene: Hemphillian. The Upper Pliocene: Blancan. The Pliocene of Mexico. Proboscideans from undetermined levels in the Mio- cene and Pliocene. North American Tertiary horizons containing fragmentary proboscidean remains. The Pleistocene. The Pleistocene of Mexico. VII. CENTRAL AND SoutTH AMERICA. CU CN Introduction. The Pliocene of Central America. The Pleistocene of Argentina. The Pleistocene of the Andean valleys. The Pleistocene of Brazil and French Guiana. I. INTRODUCTION At the time of Professor Osborn’s death, in 1935, certain portions of Volume II of the Proboscidea Memoir were left uncompleted. As the editorial work on the second volume progressed, it became particularly evident that the chapter on the stratigraphic relationships of Proboscidea-bearing beds throughout the world would necessitate a considerable amount of work to bring it to completion, for although voluminous notes had been assembled, representing a great deal of work on the part of Professor Osborn and the several persons assisting him, Therefore it became apparent that a complete text for this chapter would have to be prepared, and for various reasons this task fell to the lot of the present author. For several years preceding Professor Osborn’s death I had been associated with him as his research assistant on the Proboscidea Monograph and during that association an appreciable amount of my time had been devoted to the assembling of data as to the geologic succession and occurrences of the numerous species of fossil probo- 1421 1422 OSBORN: THE PROBOSCIDEA scideans. Thus I had the opportunity to become familiar with Professor Osborn’s views as to geologic ages and correlations, and to learn, to some extent, the manner in which he planned to write and present this chapter on the geologic succession of the fossil Proboscidea. A detailed presentation of the geologic succession, such as Professor Osborn would have written, is not now considered practicable. The original author is not here to write this chapter or to supervise its writing, and it is not considered feasible to imitate his style or his method of presentation. Therefore GEOLOGICAL RELATIONSHIPS OF AFRICAN? PROBOSCIDEA Silas Sslz., 2 this chapter will be more or less in the iS a Ss si SS z : . . oye 3 gE zi |e 3 2 form of a running narrative, describing Berenice Sz\|zh 3 : 6 = Sue 2 the distribution, development, and corre- ul S 5 ’ ST I eR ERS Soa iel GRC We A Nee eel De Res 5 2 . 5 ci} && 3 3» lation of the various Proboscidea-bearing a9 2 ESA 9 5 = = 33) |2_ bo $ 223] horizons of the world. Charts and faunal Se eee |e lis a paar poke ey |. ; ; ae 4 cBe 20 5 bs 5a-3. < 35 | lists will be included only where it is &|avse woluololv| sl > EN xX ~S ie : 3257 1581212 Tsl2t2 > «|S Eee 9 2221 thought necessary to clarify the text. Fa ES =} Beh Sl SNS IS | =|8 2 yl es 2 Bz 8/32] 9 |e] 2] | P= |= fs azil2 gleal Sle S Rae peeaes = ago: Bee esl z E|E E 38.3) Fo] 2 iA 2 =. es In writing this chapter, an attempt 222. tS OS = | ais) Pek Beal Were BS OeS/1o eS] c g I= of Fi 5 a wl 2f*s| 2) Ss] ala) es] 5) 5/28, | Selessi cc! s Ee #52] will be made to set forth the various view- Z| S658| *145/2/5/E/e/4 |ES2 |Fessos) oa] 8 g|.25 9 5~ 8 — 3) rome’ | =) te x = ee a 4 . Ole cse| ISS ElElS| 2] 2 less olaee|Se2| &e| S §|2-292£| points—often very divergent—regarding FESS Spee] ss] 5] ef] Hl eee sleet 34/2 a] o vu fine £ soa O}2 380) ELESIF IEE |Al|A ey6 Ge osler |o 0] wo Olek MOE : : : =| bee) 2 pecabrebetucreaeds “SSESSTES Se| = Siar sad the geologic age and the relationships of SESel a ai FE eelAa—|E 75/2 a li aS aes : : : a6 e228 5-33 © cfs <2 .¢&]| the horizons being considered. Inall cases, a ==0n Io x Xe tog ® ce Dew os gy ’ j i Sie 09 § o4 9 6.0 o% 23-22] Professor Osborn’s views will be stressed a 004 pase Uy qizos wie S fse@e| fees 2% | )325233| (when kn and his opini ill b w pox S59 5S SRP Oe s (when known), and his opinions will be Elephas indicus | «*4A7POZNVG Elephas.: indicus M1 OG OLG SER Bae ann re ane FISSURES |\Stegodon Stegodon ganesa, Stegodon.. |. orientalis Stegodon Pee ‘orientalis’ |Palaeoloxodon halus}| miAidanensis , Palaeoloxodon = namadicus, | namadicus Hadi : Sapaees est HAIL. “snama (PHIL) : ig. _prdécursor ' 4 Stegod = | Stegolophodon elephantoides: | a apheda insignis. - ajuensis ORNEO) SE Seoey: n.trigonoceéphalus préécurso! us ons Palaéoloxodon} f pips Siegoden e PEE SnOCENE pens idiskadon planifrons| ephantoides, TIT DIOELAM Hupselephas hysudricus Seeder oe Steggdon Sfticinal is, as sinensis a LOWER MIDDLE MIOCENE |MIOCENE |PLIOCENE|PLIOCENE athanensis, corrugatus, ptyc odus, hasnoti li -Anancus_ periimensis WAGRI Dinotherium sindiense, indicum, pesiepaenr ise y hs sronader mers palaeindicus, macrognathus, chinjiensis CHINE. iSerridentinis aerotenatst metachinjiensis, browni, chinjiensis, prochinjiensis “Synconolophus_ptychodus BAe proper negate eqolo odon caulleut roqressus, ser nat Gtonste i) indicum gajense & MIDDLE erium inotherium sindiense CAMLIAL rilophodon pandionis, macrognathus £ uv a ° i) 19) Hemimastodon crepuscu Dinoth MIDDLE 1_ Doubtful : occurrence in the Pleistocene DF Levett Bradley a z Ld al ° ei | - a ty Z|. ea O Of =| Fig. 1221 In 1912, as the result of more detailed and considered studies of the Bugti fauna, Pilgrim’ came to the con- clusion that it should be correlated with the Gaj series, of Aquitanian-Burdigalian age, rather than with the Nan beds. In his discussion of the Bugti fauna, Pilgrim shows first that it is separated by considerable differences from the Lower Siwalik fauna of Sind and the Punjab. He therefore concludes that there is a great hiatus between these faunas, a conclusion that is substantiated by the stratigraphic evidence. Continuing, with a comparison between the Bugti fauna and like faunas in Europe and northern Africa, Pilgrim finally concludes that “it seems improbable that the bone beds of the Bugti hills are younger than uppermost aquitanian.”’ ‘Pilgrim, G. E., 1908. Rec. Geol. Surv. India, XX XVII, Pt. 2, pp. 141, 143. *Pilgrim, G. E., 1912. Pal. Indica, N.S.,1V, Mem. No. 2, pp. 2, 5. GEOLOGIC SUCCESSION: THE ORIENT 1441 Both Pilgrim and Forster Cooper, as the result of their independent studies on the fossils of the Bugti beds, decided that these remains represent a unit fauna of essentially Lower Miocene age. In preparing preliminary manuscript and notes for this chapter, Professor Osborn came to the conclusion that the fossiliferous beds at Dera Bugti and at Chur Lando might be assigned to two stratigraphic horizons. This determination was based on the supposed presence of Baluchitheriwm at Chur Lando and its absence at Dera Bugti, denoting in his opinion the fact that the beds at the former locality were older than those at the latter place. “Upper level: Dera Bugti bone beds; no remains of Paraceratherium or Baluchitherium. Lower level: Chur-Lando bone beds; Baluchitheriwm or Paraceratherium life zone.” (Osborn, H. F., Notes.) In his most recent notes, Professor Osborn marked the ““Chur-Lando bone bed”’ as of Lower Miocene age and the ‘Dera Bugti bone bed”’ as of Lower to Middle Miocene age.? Since Professor Osborn’s supposition of two zones or levels in the Bugti beds was quite contrary to the publish- ed opinions of both Pilgrim and Forster Cooper, it was thought advisable at the time to secure statements from these authorities as to the probabilities of such a division. I take the liberty of quoting from their letters, written in reply to the request. Personally I am not convinced of the necessity for erecting a stage for the Chur Lando bone bed with Baluchitherium, separate from the rest of the Bugti stage of the Gaj, but in any case it seems impossible that any part of the Bugti stage can be newer than the Burdigalian. This is proved by the fact that 4000 feet or more of Murrees north of the Salt Range intervenes between the Kamlial stage (Helvetian) and the Fatehjang stage which contains Baluchitherium, Anthracotherium cf. bugtiense, Brachyodus cf. africanus, Hemimeryx, Palaeochoerus pascoei and Brachypotherium fatehjangense. Thus even if the Kumbhi and Gandoi beds are the equivalent of a stage in the Murree a considerable way above the Fatehjang, they must still fall into the Burdigalian. If the Chur Lando bone bed is earlier than the oyster beds with which presumably the Kumbhi fauna is associated, there is much more to be said for Professor Osborn’s view, but this seems hardly likely, nor apparently does Cooper suggest anything in support of this conjecture. Baluchithertum may of course have died out suddenly before the bulk of the Bugti beds was deposited, but it seems to me as reasonable that the Chur Lando bone bed was a sort of cemetery for that particular species, and may have been contem- poraneous with the Kumbhi beds. If one is determined to invent stratigraphical possibilities, then I should say that the Chur Lando bone bed may be of any age later than the Stampian, that an unconformity separates it from the Kumbhi and Khajuri beds, which through the as- sociated oysters must be at any rate Burdigalian, probably Lower Burdigalian. At the same time I do not see why Baluchi- theritum bugtiense should not be Burdigalian just as much as Cadurcotherium indicum, since the latter genus elsewhere occurs in the Oligocene. Cooper alone is in a position to estimate the stratigraphical value of the Chur Lando bone bed, and if he thinks that both it and the Kumbhi beds are later than the oyster beds then both Professor Osborn’s stages must lie in the Burdi- galian. (Pilgrim, G. E., September, 1931. Personal communication to E. H. Colbert.) A good number of my specimens were picked up on the ground washed out by rains so that their precise level must remain in doubt. Personally I cannot feel that there are sufficient grounds for establishing two horizons, and it is pretty clear that Paracera- therium bugtiense is not confined to the Churlando deposit as a number of large lower molars were found scattered in other parts. This is sufficiently clear when you remember that Pilgrim was the first to describe the species and that the Churlando bone bed was not discovered and opened wp until I went to Baluchistan some time later. No one of Pilgrim’s specimens therefore can be considered as coming strictly from this bed. (Cooper, C. Forster, 1932. Personal communication to KE. H. Colbert.) PROBOSCIDEANS: Trilophodon pandionis Falconer Trilophodon angustidens palxindicus (Lydekker) Trilophodon angustidens (Cuvier) Trilophodon coopert Osborn Dinotherium indicum gajense Pilgrim Hemimastodon crepusculi Pilgrim [Surna of Osborn | 'In Volume I of this Monograph (page 275, caption to figure 221) the ““Chur-Lando” horizon is designated as being of Upper Oligocene age. Professor Osborn was inclined at times to regard this correlation as the correct one, thereby making the ‘Dera Bugti’” beds of Lower Miocene age. In this discussion his later views, as presented above, are followed. 1442 OSBORN: THE PROBOSCIDEA Srnp.—Any remarks as to the Bugti beds in Baluchistan apply equally well to the contemporaneous de- posits of Sind. Pilgrim! discussed the Lower Miocene of Sind as follows. I have elsewhere remarked upon the close similarity to the Bugti bone beds presented by certain unfossiliferous sands and conglomerates at Bhagathoro in Lower Sind. These also rest upon Lower Nari limestones and were referred by Blanford to the Upper Nari. Mr. Vredenburg, however, considers that a shell bed, which overlies them, is Upper Gaj. There seems, therefore, little doubt that these, like the Bugti beds, represent a fresh water facies of the Gaj. ... Generally, these beds do not contain more than mere fragments of vertebrates, but near Fatehjang, Anthracotherium bugtiense, Brachyodus cf. africanus, Teleoceras fatehjangense, a species very closely allied to Teleoceras blanfordi, and a species of Hemimeryx have been found. Across the Indus, in the Kohat district, there occurs in similar beds a ribbed Unio, which may be the same as one of the curious ribbed species found in the Gaj series of the Bugti hills. 3. THE SIWALIK SERIES (MIOCENE—PLEISTOCENE) OF NORTH INDIA PunJsaB, NORTHWESTERN INp1IA.—In the northern Punjab district, along the Siwalik Hills and in the Salt Range are found tremendously thick deposits of continental sediments, constituting the Siwalik Series, the most important mammal-bearing beds in the Oriental region and likewise, one of the most important mammal-bearing series in the world. Here are exposed more than twenty thousand feet of sediments, deposited as a more or less unbroken sequence ranging from the Miocene through the Lower or Middle Pleistocene periods. Since the Siwalik Series is a continuous sequence of sediments, it will be convenient to discuss the various divisions of the series together. The Siwalik Series has been divided in its larger aspects into three main divisions or groups, namely, the Lower Siwalik, the Middle Siwalik, and the Upper Siwalik beds. Each of these three principal groups of the Series has again been divided into formations or zones, as follows. Boulder Conglomerate (Tawi of Lewis) Upper Siwalik zee \(Patrot piers) Middle Siwalik {Oho Pathan agri ea Ching Lower Siwalik Karnal The establishment of this detailed sequence of deposits is the result of studies that began with the pioneer work of Hugh Falconer, a century ago, and which have continued up to the present time. Numerous authors have contributed to the Siwalik problem, notably Falconer, Cautley, Lydekker, Pilgrim, Matthew, Colbert, Lewis, and de Terra and Teilhard. Naturally opinions have differed as to the proper correlation of the several zones or formations within the Siwalik Series, but generally speaking it may be said that the lowest Siwalik horizons are of Miocene age, while the highest ones are located within the Pleistocene. The intervening beds cover the period of time between the Miocene and the Pleistocene. Proboscideans appear in the Kamlial formation and they form a very important portion of each fauna through all of the sueceeding beds. Lithologically the Siwaliks show the results of alluvial sediments accumulating in a region closely adjacent to a rapidly uplifting mountain mass. The Lower Siwaliks are sands and clays, with occasional beds of heavier material. These deposits, particularly those of the Chinji zone, are bright red in color and contain what Pilgrim has called “pseudo-conglomerates’’—bands of concretionary beds. Passing up into the Middle Siwaliks, the ‘Pilgrim, G. E., 1912. Pal. Indica, N.S., IV, Mem. No. 2, Due. GEOLOGIC SUCCESSION: THE ORIENT 1443 sandstones become somewhat coarser, with clay beds less frequent. These Middle Siwalik beds are light gray in color—not red—and they often contain large amounts of relatively unweathered minerals. In the Upper Siwaliks there are sands and clays, often heavily cemented, and capping the series is a very heavy conglomerate made up of extremely coarse gravels and large stones. These sediments pass upward from the lower to the higher beds with but few apparent breaks. Recent work has shown that there is a distinct angular unconformity between the Dhok Pathan horizon and the overlying Tatrot zone, while the Upper Siwalik Pinjor zone is separated from the capping Boulder Conglomerate by another unconformity. Although fossils range throughout the extent of the Siwalik Series, three main faunas characterize the sequence. These are the Lower, Middle, and Upper faunas, characteristic of the three groups to which these names are applied. The Lower Siwalik fauna is typically developed in the Chinji formation, the Middle Siwalik fauna in the Dhok Pathan formation, and the Upper Siwalik fauna in the Pinjor formation. The fossils of the Kamlial formation are genetically related to those of the Chinji, while the Nagri fossils are transitory between the Chinji and Dhok Pathan faunas. The Tatrot fossils may be either transitional between the Dhok Pathan and the Pinjor or identical with the latter fauna. Falconer, who knew only the Upper Siwalik fauna, regarded it as of Miocene age. Lydekker, who described a greater portion of the Middle Siwalik fauna, placed the two faunas with which he was acquainted in the Pliocene. It remained for Dr. Pilgrim to discover the Lower Siwalik fauna, and to show the true relationships of these faunas to each other and to point out their probable ages. After many years of intensive study of the Siwalik mammals, Pilgrim came to the conclusion that the Lower Siwaliks, Kamlial and Chinji, are of Middle Miocene age (Helvetian and Tortonian, respectively), while the Nagri horizon is to be placed in the Sarmatian or Upper Miocene. Furthermore, he argued that the Dhok Pathan should most properly be placed in the Pontian, and the Tatrot and Pinjor in the middle and upper portions of the Pliocene, respectively. The Boulder Conglomerate was placed by Pilgrim in the Lower Pleistocene. Pilgrim’s views,! which have been recently epitomized in a short paper on the correlation of the Siwaliks (1934), may be shown to advantage as follows. Lower Pleistocene Boulder Conglomerate Upper Pliocene (Val d’Arno) Pinjor Middle Pliocene (Montpellier) Tatrot Lower Pliocene—Pontian Dhok Pathan Upper Miocene—Sarmatian Nagri : f ‘Tortonian Chinji Middle Miocene reivetian Kamlial Pilgrim’s views as to the correlation of the Siwaliks were based on his comparison of the several Siwalik faunas with what seemed to be their equivalents in the European sequence. The outstanding characteristics of the Siwalik faunas, on this basis, would be the close resemblance of the Chinji fauna to the typical Tortonian fauna of La Grive-St.-Alban, similarly the resemblance of the Dhok Pathan fauna to the wide-spread Pontian faunas of Pikermi, Samos, and Maragha, and finally the close comparison between the Pinjor fauna and the so-called Upper Pliocene fauna of Val d’Arno. 'Pilgrim, G. E., 1934. Amer. Mus. Novitates, No. 704. 1444 OSBORN: THE PROBOSCIDEA In 1929 Dr. Matthew,! having studied the Siwalik fossils in London and Calcutta, attempted to modify Pilgrim’s correlation by stepping the whole Siwalik Series up in the geologic column from the position assigned to it by Pilgrim. In doing this Matthew was guided by two main considerations. 1. The appearance of new invading elements in a fauna is a safer guide to its correlation than the disappearance of old elements or the average composition of the fauna as a whole. The appearance of these new elements must be interpreted in the light of what is known of their origin and dispersal. When this is as directly recorded and fully documented as it is in the case of Tertiary Equide or Camelide, the evidence appears not open to any effective challenge. But more often the appearance of new elements in a fauna may be explained in several ways, the relative probability of which is not easy to test. 2. India and the Oriental region generally are today characterized by the survival of many primitive types of mammals as well as by the absence, scarcity, or recent appearance of some of the most progressive and specialized mammals. It compares in these respects with West Africa and tropical America. While it does not necessarily follow that this was true during the later Tertiary, yet it should be so considered until evidence proves the contrary; and so far from proving the contrary I believe that all of the evidence conforms with this assumption and much of it is difficult to explain in any other way. Matthew argued that since Hipparion in India first appears in the Chinji beds, these deposits cannot be older than the oldest Hipparion-bearing beds of North America, which latter mark the first appearance of Hipparion as a direct descendant of the North American Merychippus. In other words, Hipparion arose in North America and subsequently migrated to the Old World—consequently its appearance in the eastern hemisphere must of necessity be later than its first appearance in North America. And Matthew regarded the appearance of Hip- parion as indicative of the faunal change that marked the transition from the Miocene into the Pliocene. Likewise, Matthew postulated that Hquus arising in North America at the beginning of the Pleistocene, did not reach Eurasia until after its first appearance in the New World. Consequently, the Upper Siwalik beds con- taining Hquus would of necessity be of Pleistocene age. In this respect, it might be said that Matthew placed the Val d’Arno and Villafranchian faunas of Europe in the Lower Pleistocene because of the presence of Equus in these horizons—a decision differing from the traditional European practice of assigning this stage to the Upper Pliocene. It might be said that Matthew’s correlation of the Siwaliks was not based on the evidence of the Equide alone. He showed that the Chinji giraffes are comparable to the Pontian giraffes, while the Dhok Pathan giraffes are more advanced than any giraffes of the Pontian. And in the Upper Siwaliks, he cited the appearance of the camel as additional evidence of the Pleistocene age of these beds. Therefore, he argued, the general resemblances of the Siwalik faunas show their homotaxial but not their correlative identities. These are relict faunas, in which the influx of new types from the outside furnish the real clews as to their age. Pilgrim, in 1931, answered Matthew’s argument as follows: Matthew (1929) has recently sought to replace my correlation of the Dhok Pathan stage with the Pontian by another which, to judge from his diagram on p. 441, puts both it, as well as the earlier Nagri stage, later than Pikermi; assumes a gap, which does not exist, between the Middle and Lower Siwalik, and makes the Chinji and Kamlial stages start in the Vindobonian and end at an horizon which is the equivalent of Pikermi. His argument is mainly based on the first occurrence of Hipparion in India at the top of the Chinji stage, but he considers that the remainder of the fauna, including the Carnivora, support it. Apparently, while admitting in part the occurrence in the Dhok Pathan of species allied to those of Pikermi, he regards these as relics of an earlier age. . . If this is so, we have the choice of alternatives: (1) either such forms must have migrated from the Holarctic region in Pikermi times and lingered on in India to a much later epoch; or (2) such migration did not take place until post-Pikermi times. If we adopt the first alternative we ought to find that the Nagri fauna and that of the uppermost Chinji, which by hypothesis are the equivalent of Pikermi, contain Pikermi species or species at a similar stage of development; but if the second, neither the Nagri nor Chinji fauna ought to contain any immediately ancestral types of the Pontian fauna of the Holarctic region. Actually, however, neither is true of the Carnivora, nor it may be said of other mammalian orders. We do ‘Matthew, W. D., 1929. Bull. Amer. Mus. Nat. Hist., LVI, pp. 442, 443. *Pilgrim, G. E., 1931. Catalogue of the Pontian Carnivora of Europe, Brit. Mus. (Nat. Hist.), pp. 151, 152. GEOLOGIC SUCCESSION: THE ORIENT 1445 not find Pikermi species at any earlier stage than the Dhok Pathan (the Perim Island beds I now regard as belonging to the Dhok Pathan stage); while the Nagri and, still more so, the Chinji stages contain species that are quite definitely ancestral to (or at any rate more primitive than) those of the Dhok Pathan and the Pontian of Europe alike. Matthew’s argument from Hipparion is more difficult to answer. If Hipparion originated in North America in Pontian times it obviously could not have reached India before the Pontian. But his premises lack absolute proof. First, although badly preserved equid teeth do occur at the top of the Chinji, are these certainly Hipparion and what stage of development do they represent? Secondly, did Hipparion originate in North America and not in Central Asia? Thirdly, if it did, is the age of the Valentine, in which Hipparion first appears in America, demonstrably contemporaneous with and not earlier than, say, the Sebastopol fauna of the Black Sea region in Europe? The discovery of a fauna which contains the immediate ancestors of the hitherto described ‘Hipparion fauna’ of China will settle the question definitely. Personally, I anticipate that such a fauna will be comparable to some extent with that of the Chinji stage of India. In 1935 the present author, as the result of protracted studies on the Siwalik collection in the American Museum of Natural History, came to a conclusion essentially similar to Dr. Matthew’s views as to the age of the Siwaliks, in short, that the Chinji fauna is equivalent to the “Valentine” or transitional Miocene-Phocene of North America and is slightly pre-Pontian, or Pontian, in age, that the Dhok Pathan fauna is definitely post-Pon- tian and that the Pinjor fauna is definitely of Lower Pleistocene age. There were a few slight differences between my interpretations and those of Matthew, namely, the raising of the Chinji into the Lower Phocene and the con- sideration of the Siwaliks as a continuous series rather than as separated by two major breaks, the view taken by Matthew. Consequently the correlation adopted by the present author is as follows: Mowerbleistocenes 4. 0. sa408 4.8 {Boulder Conglomerate \ Pinjor Transitional Tatrot Middle to Upper Pliocene.......... {Dhok Pathan (Nagri Lower Pliocene Chinji Upper Miocene Kamlial Subsequently (1937) G. E. Lewis! published still another opinion as to the ages of the several Siwalik faunas. On the basis of new evidence, especially his stratigraphic field studies, he adopted the following correlation of the beds forming the Indian sequence. Middle Pleistocene Tawi (new name for Boulder Conglomerate) Lower Pleistocene Tatrot (including Tatrot and Pinjor) Upper Pliocene Break Middle Pliocene Dhok Pathan Lower Pliocene Nagri Upper Miocene Chinji Middle Miocene Kamlial_ This correlation differs from those of Matthew and of Colbert by placing the Lower Siwaliks even lower than was granted by these two authors, but not so far down as they were placed by Pilgrim, and by extending the Upper Siwaliks higher into the Pleistocene than had previously been done. Lewis’ views as to the age of the Chinji beds are based to a great extent on the supposed Upper Miocene appearance of Hipparion in North America, in the Mint Canyon beds of the Pacific coast. He points out that this Hipparion mohavense is very close to the Siwalik Hipparion, that it is probably close to the ancestor of the Asiatic species, and that the Mint Canyon formation in which it appears is topped by marine beds carrying Miocene invertebrates. Lewis, G. E., 1937. Amer. Journ. Sci., (5), XX XIII, p. 197. 1446 OSBORN: THE PROBOSCIDEA As Lewis has remarked, it is probable that Hipparion mohavense is as close or closer to the Asiatic Hipparion than are any other North American species of this genus, while Merychippus suman is very likely ancestral to these Hipparion types. Consequently Merychippus sumani and Hipparion mohavense are the forms that should be compared with the Asiatic Hipparion. The real crux of Lewis’ argument is the question as to whether the Mint Canyon formation is of Upper Miocene age. This is the interpretation given by Maxson, who described the mammalian fauna from these beds. It should be pointed out here, however, that Stirton (1933)! has disputed the Miocene age of the Mint Canyon, linking this formation with the Ricardo of Lower Pliocene age, rather than with the Barstow of Upper Miocene age—as was done by Maxson, and placing it above the Barstow but below the Ricardo in the geologic column. One important point in this discussion of the Mint Canyon is, of course, the age of the overlying marine beds. It should be noted that the invertebrate fossils are not well preserved, and that Woodring’s correlation of the de- posits with the Cierbo formation was more or less tentative. Stirton places the Mint Canyon assemblage as equivalent in age to the Cierbo and regards both of these formations as being about as close in their time relation- ships to the Ricardo as they are to the Barstow. Lewis regards the Nagri as the Pontian equivalent in the Siwalik Series, and places the Dhok Pathan in the Middle Pliocene, as did Matthew and Colbert. Finally Lewis shows a great stratigraphic break between the Dhok Pathan and the Upper Siwaliks, a fact of which the present author was unaware when his work was carried forward. The Tatrot, which the present author considered as possibly transitional between the Upper Pliocene and the Lower Pleistocene, is regarded by Lewis as an equivalent of the Pinjor horizon. Lewis chooses the name Tatrot for this level, a choice that may not be general- Likewise, Lewis’ designation of the Boulder ) ly followed, due to the long-established use of the term ‘‘Pinjor.’ Conglomerate as ‘“Tawi”’ may not be generally accepted. In 1936 de Terra and Teilhard placed the Tatrot and Pinjor in the Lower Pleistocene and the Boulder Con- glomerate in the Middle Pleistocene, as did Lewis. Since these two authors were concerned only with the Pleisto- cene of India, their conclusions are not discussed here. In several recent papers (1939, 1940)*, Pilgrim has defended his views as to the correlation of the Siwaliks, and has offered rebuttals to the arguments of Matthew, Colbert, de Terra and Teilhard, and Lewis. While admitting as proven fact the North American origins of Hipparion and Equus, Pilgrim nevertheless main- tains that the general characters of the Chinji and Dhok Pathan faunas prove their pre-Pontian and Pontian relationships, respectively. This author holds that Matthew’s and Colbert’s idea that the Siwalik faunas are homotaxially similar to but correlatively later than comparable faunas in Europe is untenable. Therefore, in order to explain the presence of Hipparion in the Chinji and to retain this formation at a pre-Pontian level, Pilgrim suggests that the various Lower Pliocene formations of North America may be actually older than has hitherto been admitted by American palzontologists. Pilgrim disagrees with Lewis’ procedure of making the Dhok Pathan an equivalent of Roussillon and Montpel- lier and placing these latter in the Plaisancian. As he shows by his arguments, these European faunas are of Astian age, while the Plaisancian represents a general faunal gap between the Lower and the Upper Pliocene in 'Stirton, R. A., 1933. Amer. Journ. Sci., (5), X XVI, p. 570. *Pilgrim, G. E., and A. T. Hopwood, 1939. Ree. Geol. Surv. India, LX XIII, Pt. 4. Pilgrim, 1939.1. Pal. Indica, N. S., XXVI; 1940.1, Geol. Mag., LXXVII, pp. 1-27. GEOLOGIC SUCCESSION: THE ORIENT 1447 Europe. Pilgrim further maintains that the Tatrot is a truly distinct horizon, not closely related to the Pinjor as considered by de Terra and Teilhard, or identical with it as claimed by Lewis. In the light of these recent discussions, Pilgrim presents his latest correlation of the Siwalik Series as follows: Pleistocene......... Post-Cromerian Boulder Conglomerate Villafranchian Pinjor HOGENes ye sats eee Astian Tatrot Pontian Dhok Pathan ‘ Sarmatian Nagri MHOCENE? =r. =: =: ; Tortonian Chinji Tortonian Kamlial In conclusion, it may be said that the Siwalik Series of India represent a period of deposition ranging from the middle or upper part of the Miocene well into the Pleistocene. The lowest Siwalik horizon, the Kamlial formation or zone, is definitely of pre-Pontian age. Above this is the Chinji formation, carrying Hipparion, which may be correlated as either Pontian or as representing a period of time immediately preceding the Pontian, according to the manner in which the origin and migrations of Hipparion from North America are interpreted. Following the Chinji is the Nagri formation, transitional between the underlying Chinji and the overlying Dhok Pathan. The Dhok Pathan is Pontian or post-Pontian, according to the manner in which the Siwalik faunas are interpreted in relation to Eurasiatic and North American faunas. Between the Dhok Pathan and the succeeding Upper Siwaliks is a break, representing an Upper Pliocene interval during which no sediments were deposited. The Upper Siwaliks are composed of two horizons of Lower Pleistocene age (Tatrot plus Pinjor) separated by an erosional break from the uppermost Boulder Conglomerate (Tawi of Lewis), possibly of Middle Pleistocene age. Prerim IstAanp.—The fauna from Perim Island was originally considered by Dr. Pilgrim to be equivalent to the Lower Siwalik fauna. Subsequently, however, he changed his opinion and placed the Perim Island assemblage in the Dhok Pathan stage. This latter procedure would seem to express correctly the true relationships of the Perim Island beds. 4. THE PLEISTOCENE OF CENTRAL INDIA NARBADA-GODAVARI ALLUVIUM.—Extensive deposits of Pleistocene age are exposed in Central India in the Narbada and the Godavari valleys. These sediments are approximately contemporaneous with each other (Pilgrim, 1905) and they represent, at least in part, the final stages of the Pleistocene in India. The alluvial deposits of the Narbada and the Godavari rivers have been known for many years, and from time to time fossil mammals, definitely related to the Upper Siwalik fauna, have been found in them. Consequent- ly various papers have been published in which these sediments and their contained fossils are discussed. One of the latest contributions is that of de Terra and Teilhard (1936), in which the problem of the Upper Siwalik and later Pleistocene deposits of India is considered. These authors find that in northern India the Boulder Conglomerate stage is followed by a long erosion inter- val, after which there comes the Potwar silt, a fine deposit, partly of fluviatile, partly of eolian origin, of Upper Pleistocene age. The Potwar silt is succeeded by another long erosion interval, while finally, at the top of the section, are redeposited Potwar sediments and loesses. Turning now to the Narbada valley, de Terra and Teilhard! make the following remarks as to the Narbada alluvium. 'Terra, H. de, and P. Teilhard de Chardin, 1936. Proc. Amer. Philos. Soc., LX XVI, No. 6, pp. 820-822. 1448 OSBORN: THE PROBOSCIDEA PROBOSCIDEA: | eee | Chinji | Dhok Pathan Tatrot Pinjor Dinotherium sindiense Lydekker aes Dinotherium indicum Falconer x x Dinotherium pentapotamiz Lydekker x Dinotherium angustidens Koch x Trilophodon pandionis Falconer Trilophodon angustidens palxindicus (Lydekker) Trilophodon macrognathus (Pilgrim) Trilophodon chinjiensis (Pilgrim) Trilophodon hasnotensis Osborn Tetralophodon falconeri (Lydekker) Tetralophodon punjabiensis (Lydekker) Serridentinus hasnotensis Osborn Serridentinus metachinjiensis Osborn i Serridentinus browni Osborn Serridentinus chinjiensis Osborn i. Serridentinus prochinjiensis Osborn YG P&S 24 | 2S Rhynchotherium chinjiensis Osborn Synconolophus dhokpathanensis Osborn ae Synconolophus propathanensis Osborn Synconolophus corrugatus (Pilgrim) Synconolophus ptychodus Osborn Synconolophus hasnoti (Pilgrim) a Anancus perimensis (Falconer and Cautley) Anancus properimensis Osborn Pentalophodon sivalensis (Cautley) Pentalophodon falconeri Osborn ; 7 Stegolophodon latidens (Clift) Stegolophodon cautleyi (Lydekker) fr Stegolophodon cautleyi progressus Osborn Stegolophodon nathotensis Osborn - Stegolophodon stegodontoides (Pilgrim) Stegodon bombifrons (Faleoner and Cautley) Stegodon cliftii (Falconer and Cautley) Stegodon elephantoides (Clift) Stegodon ganesa (Falconer and Cautley) Stegodon insignis (Falconer and Cautley) Stegodon pinjorensis Osborn Archidiskodon planifrons (Falconer and Cautley) , oa H ypselephas hysudricus (Falconer and Cautley) Platelephas platycephalus (Osborn) GEOLOGIC SUCCESSION: THE ORIENT 1449 The formation consists here [Narbada valley] of two different horizons, each of which begins with a basal gravel overlain by brown and pinkish or orange coloured coneretionary clays and silts. In the lower zone the conglomerate is coarser and more cemented, the clay is more intensely coloured and also richer in concretions than in the upper zone. Fossils occur chiefly near the disconformity which separates both zones. . . The ‘lower zone’ of the Narbadda Pleistocene can be equated with the Upper Siwalik ‘Boulder Conglomerate’ on faunistic, archeological and lithological grounds. The association of advanced Hlephas with Hippopotamus and large Bos suggests a stage slightly younger than the older Upper Siwaliks. In harmony with this is the appearance of an early Paleolithic culture in the basal gravel, clearly calling to one’s mind the picture of heavy accumulation of river deposits during the glacio-pluvial stage of late Siwalik times in the Punjab. It follows that on these grounds a further correlation between the ‘Upper Zone’ and the ‘Potwar silt’ becomes rather plausible. Both are separated by a long erosion interval from the underlying beds and both contain implements of Soan type. The cotton soil might then well represent the latest Pleistocene which possibly is homotaxial with the redeposited Potwar silt and the second loess in the Punjab. According to these authors, the relationships of the Narbadda alluvium might be represented in the following manner. Narbada valley Punjab Cotton soil Re-deposited Potwar and second loess | | Erosion interval | U. Pleist. | | Narbada alluvium | Potwar silt upper zone | Erosion interval M. Pleist. | Narbada alluvium Boulder conglomerate | | lower zone The mammalian fauna of the Narbada alluvium is as follows: Helarctos namadicus (Falconer and Cautley) Hippopotamus palexindicus Falconer and Cautley Stegodon insignis (Falconer and Cautley) Hippopotamus namadicus Falconer and Cautley Stegodon ganesa (Falconer and Cautley) Cervus duvaucelli Cuvier Palzxoloxodon namadicus (Falconer and Cautley) Bubalus palxindicus Falconer Rhinoceros unicornis Linnseus Leptobos fraseri Riitimeyer Equus namadicus Faleoner and Cautley Bos namadicus Faleoner Sus namadicus Pilgrim Closely related to the Narbada alluvium is the Godavari alluvium, exposed also in peninsular India, but on the eastwardly flowing drainage system. Pilgrim,! in 1905, made a study of the Godavari deposits, and below are given certain remarks that he made with regard to this formation. As compared with the alluvial deposits of the Narbada, which flows in a contrary direction, and enters the sea on the west coast of India, our knowledge of those of the God4vari is very limited, both as regards the fossil contents, and even as to the nature, thickness and superficial extent of the alluvium itself. . . Comparing the fauna of the Godavari alluvium with that of the older Narbada deposits, we shall see that the only three mammalian species found in it up to now are identical with Narbada forms. Of these H. antiquus (namadicus) Fale. et Caut., and Hippopotamus palzindicus F. et C., are quite absent from the older deposits of the Siwalik beds, while Equus namadicus F. and C. comes up from below. We are therefore justified in regarding the two series of alluvia as of approximately the same age, and any conclusion we arrive at with respect to the Narbada deposits must apply equally to those of the Godavari. PROBOSCIDEAN: See faunal list, above. 'Pilgrim, G. E., 1905. Ree. Geol. Surv. India, XXXII, Pt. 3, pp. 199, 218. 1450 OSBORN: THE PROBOSCIDEA 5. THE PLEISTOCENE OF CEYLON CryLton.— During the years 1935 and 1936 Upper Siwalik fossils were discovered in Ceylon, and described by Deraniyagala.! These fossils are extremely fragmentary but are sufficiently complete to show that they are true Upper Siwalik forms. The faunal list is as follows. Palzoloxodon Rhinoceros Hexaprotodon ?sivalensis 6. THE PLEISTOCENE OF BURMA IRRAWADDY VALLEY (IRRAWADDY SERIES).—The Cenozoic history of Burma may be summed up by saying that there was continuous subsidence accompanied by continuous deposition. Consequently the sediments have accumulated to a great thickness, with but minor breaks in the series. Of course there were variations in the rate of subsidence and deposition, causing a considerable amount of interfingering of marine and continental deposits, but on the whole since Middle Eocene times a gradual encroachment of the continental beds from the north to the south took place, thereby causing the Burmese gulf to retreat southwardly. A series of deposits represent the Tertiary in Burma, of which the Pondaung sandstone of Upper Eocene age and the Pegu beds of Oligocene to Pliocene age are the continental, mammal-bearing facies. Correlative with the upper portions of the Pegu beds are the lower phases of the Irrawaddy series. These Irrawaddy beds constitute the upper part of the sedimentary section in Burma, ranging from the Lower Pliocene up through the lower phases of the Pleistocene. Fossils are fragmentary and scarce in the Pegu beds, and it is only in the Irrawaddy series that proboscideans are found. Two faunas have been identified in the Irrawaddy beds, a lower one of Pliocene age, probably correlative with the Dhok Pathan fauna of the Siwaliks, and an upper one of Pleistocene age, correlative with the Pinjor fauna of the Siwaliks. The two faunas are separated from each other by a vertical range of about 4500 feet of sediments. There is some question as to the actual presence of proboscideans in the Lower Irrawaddy beds. Pilgrim, in 1910, listed the mastodont, Stegolophodon latidens, as belonging to the Lower Irrawaddy fauna, but this occurrence was questioned by Stamp in 1922.2 Subsequent work in Burma would seem to indicate that this animal is a member of the Upper Irrawaddy fauna; certainly a number of discoveries made in recent years establish it as definitely belonging in the Lower Pleistocene of Burma. Therefore, since its presence in the Lower Irrawaddy beds is very doubtful, it will be considered here as limited to the Pleistocene in Burma. The Upper Irrawaddy fauna is in all respects an eastward extension of the typical Upper Siwalik fauna of India. In numerous cases there is a specific identity between elements constituting the Burmese faunas and those elements in the Pinjor fauna of northwestern India. Where species are not identical in the two regions, they are so closely related as to leave little doubt as to their derivation from common ancestral types. Therefore there is no reason to regard the Upper Irrawaddy fauna as other than of Lower Pleistocene age, strictly correlative with the Pinjor fauna to the west. "Deraniyagala, P. E. P., 1935. Journ. Roy. Asiatic Soc. (Ceylon), XX XIII, No. 88; 1936, Geol. Mag. (London), LX XIII, No. 865. *Stamp, L. Dudley, 1922. Geol. Mag. (London), LIX, No. XI, pp. 481-501. GEOLOGIC SUCCESSION: THE ORIENT 1451 That there was an extension of the Upper Siwalik fauna eastwardly throughout a considerable portion of the Oriental region, is shown by the presence of characteristic Siwalik types in various East Indian Islands, to the south and east of Burma. This extension of the Upper Siwalik fauna has been designated by von Koenigswald! as the “Siva-Malayan” fauna, having its origin in northern India, pushing into Burma and reaching Java and adja- cent regions in Lower Pleistocene times. PROBOSCIDEANS: Stegolophodon latidens (Clift) Stegodon elephantoides (Clift) Stegodon insignis birmanicus Osborn Hypselephas hysudricus (Falconer and Cautley) SHAN PuiatreAu (Mocox Fissures).—The Irrawaddy faunas of Burma are found in the Irrawaddy River valley, in stream and flood-plain deposits. As contrasted with this, is the fauna found in the limestone caverns or fissures of the Shan Plateau. This fauna is distinguished by Stegodon orientalis, Palzoloxodon namadicus, Ailuro- poda baconi, and Hystrix. These are essentially the very forms that characterize the mammalian assemblages found in limestone caverns and fissures to the east of Burma, in Yunnan, Shansi, and Szechwan. There are other mammals also in the Burma caverns that show the close relationships of these deposits to the cave deposits of China, animals such as Rhinoceros, Sus, and various ruminants. But it is upon the basis of the first four forms enumerated that the affinities of this cavern fauna in various regions may be established. Of late years, particularly because of the work of Teilhard and de Terra, Pei, Bien, von Koenigswald, and others, there is a tendency to regard the cave faunas of China as belonging to a Middle Pleistocene stage of de- velopment. In Burma there is the Upper Irrawaddy fauna which seems to be definitely older than the mammals from the Mogok caves, while in China there are such assemblages as those of Nihowan and Ma-Kai, which would seem to precede the cave faunas. In short, the cave faunas represent a post-Villafranchian development through- out the Orient. So far as Burma is concerned, the Mogok fauna may be compared with the Narbada assemblage on the west, as well as with the cave faunas to the east. But an analysis of such a comparison will show that although certain Narbada elements are to be found in the Mogok fauna, the bulk of the assemblage is closely related to, if not identical with the cave faunas of China. Of course the most important correlations are with the Choukoutien fauna of North China, containing Sinanthropus, and the Trinil fauna of Java, containing Pithecanthropus. Von Koenigswald, recognizing the continuity over a broad area in the Orient of this Middle Pleistocene fauna, designated it as the ‘““Sino-Malayan” fauna. Seemingly having had its origin in the southern part of China, the Sino-Malayan fauna spread to the south to the East Indian Islands, and to the west to Burma. Consequently the two Pleistocene faunas of Burma represent invading assemblages, the Lower Pleistocene or Upper Irrawaddy fauna coming in from the west, and the Middle Pleistocene or Mogok fauna coming in from the east. PROBOSCIDEANS: Stegodon orientalis Owen Palzoloxodon namadicus (Falconer and Cautley) 'Koenigswald, G. H. Ralph von, 1938-1939. Peking Nat. Hist. Bull., XIII, Pt. 4, pp. 293-298. 1452 OSBORN: THE PROBOSCIDEA 7. THE PLEISTOCENE OF SOUTH CHINA SzECHWAN PRovINcE, Cu1na.—In southwestern China, in the province of Szechwan, are numerous pits and fissures developed in the limestone that forms the surface outcrops, and many of these pits contain rich deposits of fossil mammals. They have been worked by the Chinese since ancient times for their “dragon bones,”’ which constitute one of the staples of the Chinese drug trade. The fossils from the Szechwan pits were first described by Owen, who regarded them as of Pliocene age. Subsequently various authors, notably Koken, Schlosser, and Matsumoto, described fossils from the Szechwan pits. Finally, in 1923, Matthew and Granger! described a new and a very complete series of fossils from Szechwan (collected by the latter author). At the time, these authors retained the older viewpoint of an Upper Pliocene age for the material with which they were working. Subsequently, however, Matthew came to the conclusion that the Szechwan fauna is more properly to be placed in the Lower Pleistocene, a view that was never published, but clearly stated in manuscript notes. Perhaps the most important mammals in the Szechwan fauna are Stegodon orientalis, a large stegodont with very long grinding teeth, a gibbon, Bunopithecus, a langhur monkey, a large Hystriz, a dhole related to the modern Indian dhole, a giant panda, a gigantic tapir, Megatapirus, a chalicothere, Nestoritherium sinense, a Rhinoceros, and an extremely large gaur, Bibos geron. The generally modern character of this fauna places it as definitely of Pleistocene age, for there are too many advanced elements in it to allow its inclusion in the Pliocene. The association of Stegodon orientalis, Hystrix, and the giant panda at once establishes a strong resemblance to the Mogok fauna of Burma, discussed on a preceding page. Indeed, upon the basis of the entire faunal assemblage at Mogok and in the Szechwan fissures, there is every reason to think that they are contemporaneous, representing essentially a single fauna stretching from Burma through southern China. As has been pointed out in the discussion of the Mogok fauna, there is good reason to think that these cave faunas are of Middle Pleistocene age since they are preceded both in Burma and in China by faunas having a Villafranchian aspect. I take the liberty to quote at this place from Matthew’s unfinished manuscript on the Szechwan fauna: This is a fairly typical fauna of southeastern Asia, plus a number of species now extinct or limited in their range. Marked features are the presence of Stegodon, but no mastodons or mammoths,” the giant tapir, and a rhinoceros, but no horses, a single tooth of Chalicotheriwm. Among the Carnivora is a hyena related to the spotted hyzenas, Cyon but no Canis, a very large marten as big as the American fisher Martes pennanti, and a tiger, civet, particolored bear and true bear not very different from modern survivors. The artiodactyls include muntjac and sambhur gazelle, serow, gaur and yak, also some other species which we have not yet succeeded in identifying; they may be extinct species. The rodents are almost wholly a large bamboo rat allied to the Chinese species but as big as the Malayan. Altogether this fauna appears to be the fauna of South Chinese forests and mountain valleys, as we may suppose it to have been before civilized or semicivilized man cut down the forests, cultivated the valleys, and brought about the extinction of the larger and more specialized animals, driving the remainder of the fauna into the hills. The absence of horses, of true dogs, of mammoths and of mastodons is in marked contrast to the Pliocene fauna of North China, where all those animals had already appeared. The latter was apparently a plains fauna. Teilhard, Young and others,’ in 1933, indicated the fissure deposits of Szechwan and adjacent localities as extending over a period of time in the Lower Pleistocene that embraced both the Sanmenian and the Choukoutien deposits. In another part of this same paper, they indicated that the fissure deposits of South China might be limited to a period of time contemporaneous with the Choukoutien deposits. This is their latest opinion, and is now generally held by authorities on the Pleistocene mammals of Asia. ‘Matthew, W. D., and Walter Granger, 1923. Bull. Amer. Mus. Nat. Hist., XLVIII, Art. XVII, pp. 563-598. 2Palxoloxodon namadicus has recently been discovered in Szechwan, and described by Young. 3Black, Teilhard de Chardin, Young and Pei, 1933. Mem. Geol. Surv. China, Ser. A, No. 11, Table III, p. 158; Map III, opposite page 164. GEOLOGIC SUCCESSION: THE ORIENT 1453 The mammalian faunas from other caverns in southern China are so similar to that of the Yenchingkou pits as to need no particular consideration at this place. It is quite evident that all of these occurrences represent a single fauna that was rather widely spread during Middle Pleistocene times. Mention should be made of the caves in Kwangsi, the fauna of which has been described by Teilhard, Young, Pei and Chang, and the Hoshangtun Cave in Yunnan, described by Bien and Chia. Also there should be mention- ed the isolated occurrence of a stegodont said to have been found near Shanghai and described by Owen as Stegodon sinensis. This is probably of the same age as the other South Chinese Stegodonts. PROBOSCIDEANS: Stegodon orientalis Owen Palzxoloxodon namadicus (Falconer and Cautley) 8. THE PLEISTOCENE OF INDO-CHINA FrencH Inpo-Cuina.—There are cavern deposits in northern French Indo-China, seemingly quite similar to those of southern China, described above. Since they contain the same fauna as the Chinese caverns, they must be considered as correlative with the latter, representing a southern extension of the characteristic Middle Pleistocene cave fauna of the Orient. Mansuy,! who described the mammalian fossils from Indo-China, identified the stegodont material as belonging to Stegodon “‘cliftc”’ and Stegodon insignis. A comparison of his excellent plates with material on hand suggests the possibility that the Stegodon from Indo-China is of the species orzentalis. In fact, Mansuy regarded S. orientalis as probably synonymous with S. insignis, and this view undoubtedly influenced him in his identification of the material. PROBOSCIDEANS: Stegodon orientalis Owen.—(S. insignis according to Mansuy.) Stegodon elephantoides (Clift)—(This species? probably orientalis.) Palzoloxodon namadicus (Falconer and Cautley) 9. THE PLEISTOCENE OF THE EAST INDIES JAvA.—Since the discovery of Pithecanthropus in 1892, Java has assumed a position of great importance to the students of human prehistory and the evolution of life. Consequently numerous contributions have appeared during the course of the past forty years, discussing the phylogenetic relationships and the probable geologic age of Pithecanthropus. It has been only recently that detailed geological investigations have been carried on in Java, with a view to clarifying some of the hitherto obscure points as to the succession of upper Cenozoic sedi- ments in Java. For many years it was supposed that the beds at Trinil, in which Pithecanthropus was discovered, might be placed in the Upper Pliocene or the Lower Pleistocene, thereby allocating to Pithecanthropus the honored position of being the oldest known hominoid. But recent studies by various authors have shown that the upper Cenozoic stratigraphy of Java is much more complex than it was originally thought to be—therefore our ideas as to the age of Pithecanthropus and the associated fauna have necessarily undergone considerable revision. ’ In 1931 van Es published a detailed geologic study entitled ““The Age of Pithecanthropus,”’ in which he showed that there was a volcanic boulder breccia and beneath it a sand and conglomerate underlying the Trinil beds. Both of these older horizons he regarded as of Pleistocene age. 'Mansuy, H., 1916. Mem. Surv. Geol. Indochine, V, Fase. II, pp. 1-26. 1454 OSBORN: THE PROBOSCIDEA In 1932 van der Maarel! described a new fauna from Bumiaju, in Central Java, and came to the following conclusions as to the stratigraphic relationships of the Javanese strata: “1. The Bumiaju fauna is older than the Trinil fauna. 2. The Trinil fauna is certainly of Pleistocene age, more particularly either Lower [or] Middle Pleisto- cene, but not Upper Pleistocene. 3. Accordingly the Bumiaju fauna is of upper pliocene or lower pleistocene age.”’ In recent years von Koenigswald? has carried on a series of careful studies regarding the upper Cenozoic mammal-bearing sediments in Java, and he has distinguished a number of successive horizons within the Pleisto- cene. These and correlative horizons in India may be listed as follows: JAVA INDIA Sampoeng zone (Neolithic) Ngandong zone Potwar Trinil zone Narbadda Pleistocene Boulder Conglomerate Djetis zone Pinijor Kali Glagah zone J Tji Djoelang Tatrot Pliocene Tji Sande zone Incidentally von Koenigswald placed the Kali Glagah and the Tji Djoelang zones in the Upper Pliocene, since he regards the Pinjor fauna as a transitional Pliocene-Pleistocene assemblage and the Tatrot fauna as of Upper Pliocene affinities. The Tji Djoelang and Kali Glagah faunas contain Lower Pleistocene elements, e. g., Tetralophodon, Stego- don, Merycopotamus, and Hippopotamus, such as are found in the Tatrot and Pinjor faunas of India. While these animals may also be found in Upper Pliocene sediments, the general resemblance of the two Javanese faunas listed above to the Tatrot and Pinjor faunas of India justifies their inclusion in the Pleistocene according to the definition of the period adopted in this work. The Djetis fauna is more advanced, although it contains a few persistent remnants of the Siwalik fauna— notably Nestoritherium sivalense. There are, however, various characteristic Oriental elements in this fauna, such as Rhinoceros cf. sondaicus, tapir, Paradorurus, pangolin, and Symphalangus. Von Koenigswald was fully justified in placing the Djetis fauna fully as late as or probably later than the Pinjor assemblage of India. The Trinil fauna is directly comparable with the Choukoutien fauna of China, and both of these are now regarded as definitely post-Villafranchian in age. According to the definitions adopted in the present work the Trinil beds may therefore be regarded as approximately of Middle Pleistocene age. There are numerous elements in the Trinil fauna correlating it with the cave faunas of China, such as Stegodon, Viverra, Felis, Hystrix, Muntia- cus, Cervus, Bubalus, and Simia. Finally, the Ngandong fauna of Java is a very late Pleistocene assemblage, with a few new elements, but distinguished mainly from the older Trinil fauna by the presence of a Neanderthal type of man. 'Van der Maarel, F’. H., 1932. “Contribution to the Knowledge of the Fossil Mammalian Fauna of Java,” p. 199. *Koenigswald, G. H. R. von, 1935. Proc. Akad. Wet. Amsterdam, XXXVIITI, No. 2, pp. 188-198; 1939, Quartir, Zweiter Band, pp. 28-53. GEOLOGIC SUCCESSION: THE ORIENT 1455 The faunas of the several Javanese Pleistocene horizons are too extensive to list here, so only the probo- scideans will be enumerated. PROBOSCIDEANS: Sampoeng zone Elephas indicus Linnzeus Ngandong zone Stegodon trigonocephalus Martin Palzoloxodon cf. namadicus (Falconer and Cautley) Trinil zone ?Cryptomastodon martini von Koenigswald [=Sirenian? (Osborn)]| Stegodon trigonocephalus Martin Palzoloxodon cf. namadicus (Falconer and Cautley) Djetis zone Stegodon trigonocephalus praecursor von Koenigswald Elephas sp. Kali Glagah zone Tetralophodon bumiajuensis Maarel Stegodon trigonocephalus praecursor von Koenigswald Archidiskodon planifrons (Falconer and Cautley) Tji Djoelang zone Stegodon sp. BorneEo.—In 1885 Lydekker! described a tooth that he identified as Mastodon latidens, from the northwest coast of Borneo. The specimen forming the subject of the present notice was forwarded from Borneo to the Secretary of this Society by Mr. A. H. Everett, C.M.Z.S., who stated that it was found during the early part of the present year by a Kadayan in the jungle in the vicinity of Bruni, on the north-west coast of Borneo. Owing to the country being in a disturbed state Mr. Everett could not visit the locality to make further inquiries; but there seems no doubt that the history of the specimen is a true one. Comparing this tooth with the many teeth of Mastodon latidens from Burma and India, Lydekker came to the conclusion that the tooth from Borneo was specifically identical with the Burmese-Indian forms, and therefore he decided that the deposits in Borneo from whence this tooth was recovered might be questionably of Pliocene age. Subsequently (1936, Volume I of this Monograph, p. 700) Professor Osborn made the tooth from Borneo the type of a new species, Stegolophodon lydekkerv. Stegolophodon latidens from Burma, where the type was discovered, would seem to be restricted to the Upper Irrawaddy beds of Pleistocene age, while in India the species ranges down into the Pliocene Dhok Pathan horizon. It is an interesting fact that Lydekker emphasized the close mineralogical comparison between the Borneo tooth and those teeth of M. latidens from Burma. With this fact in mind, and remembering that the Stegolophodon latidens in Burma is found in the Pleistocene level, there would seem to be some reason for thinking that the 'Lydekker, R., 1885. Proc. Zool. Soc., London, 1885, p. 777. 1456 OSBORN: THE PROBOSCIDEA Borneo specimen might be of Pleistocene age. It should be pointed out, however, that Professor Osborn placed this specimen in the Pliocene (but with a query) as did Lydekker. PROBOSCIDEAN: sStegolophodon lydekkeri Osborn. PHILIPPINE IsSLANDS.—On the basis of published records it would seem that fossil proboscideans from the Philippine Islands are represented by a single tooth from the island of Mindanao. This specimen was described many years ago by E. Naumann, under the name of Stegodon mindanensis. Naumann compared it with Stegodon trigonocephalus of Java, pointing out the fact that there were great similarities between the species. Therefore it would seem possible that the beds containing these two forms in Java and Mindanao are geologically more or less equivalent to each other. In this connection it may be pointed out that Stegodon trigonocephalus in Java is found typically in the Ngandong beds, above the Trinil zone of probable Middle Pleistocene affinities. It is an interesting fact, however, that stegodonts closely related to S. trigonocephalus are found in lower beds in Java, ranging down to the basal Pleistocene. Therefore it would seem that the Mindanao deposits might be repre- sentative either of Lower or of Middle Pleistocene times in the Philippine region. PROBOSCIDEAN: Stegodon mindanensis Naumann. IV. EUROPE 1. INTRODUCTION It may be said that the Proboscidea entered Europe at the beginning of the Miocene—considering the Burdigalian to be the opening stage of this period. From that time until the final stages of the Pleistocene, these animals were prominent in the extinct mammalian faunas of the European region; therefore, a consideration of the Proboscidea-bearing beds of Europe is virtually a discussion of the continental stratigraphy of the Miocene, Pliocene, and Pleistocene of that region. A thorough discussion of the continental Tertiary and Quaternary sediments of Europe is indeed a large order, too much to be included within the scope of this present work, and beyond the capabilities of one who does not have an intimate first-hand knowledge of the complex relationships of these Old World mammal-bearing horizons. Therefore, it is proposed to discuss very broadly the general succession of upper Tertiary and Pleisto- cene continental deposits in Europe, giving particular attention to the more typical localities at which the several stages are best developed. In doing this the localities at which proboscidean types were discovered will, of course, be especially stressed. In dealing with the Tertiary mammalian faunas of Europe one general consideration must be kept in mind, particularly during the earlier phases of the Tertiary namely, that this was primarily a mediterranean area period—interspersed with numerous islands and peninsulas. In this respect, the European region is to be con- trasted with America and Asia, which from the end of the Cretaceous were, generally speaking, broad continental platforms, with the shallow sea borders lapping over the edges, but not in the main encroaching materially beyond their present limitations. It was not until late Tertiary times that Europe underwent a general emergence so that most of its present area was permanently lifted above sea level. Naturally, the broken-up character of the land areas of Europe in earlier Tertiary times had a distinct influence on the development of the mammalian faunas, and similarly the final emergence of this region as a continental mass also is reflected in the expression of GEOLOGIC SUCCESSION: EUROPE 1457 the assemblages of land animals. So it is that the earlier separated faunas are finally replaced by associations of mammals that become more and more cosmopolitan, with an eventual culmination in the appearance of the almost world-wide Hipparion faunas and subsequent assemblages that are so characteristic of the last phases of Tertiary times. GEOLOGICAL RELATIONSHIPS OF EUROPEAN PROBOSCIDEA, FRANCE ITALY AND ’ CENTRAL EUROPE EASTERN EUROPE| ASIA MINOR ENGLAND BELGIUM - SPAIN | ™ MEDITERRANEAN GERUSTRIA ROLAND E. MEDITERRANEA SWITZERLAND ISLANDS HUNGARY RUMANIA ISLANDS Hesperoloxodon antiquus Parelephas wiisti Palaeoloxodon esperoloxodon |Mammonteus primigenius aie ntiquus hydruntinus ; cupriofes M ammon te us primigenius platythynchus Loxodonta cornaliae es Hesperoloxodon | Pal&toloxodon (Widely distributed over Europe) Pa]4eoloxodon | mnaidriensis, antiquus germanicus creticus alconeri, melitensis, lamarmorae q 3 Hesperoloxodon antiquus ttalicus Parelephas intermedius Parelephas trogontherii Mastodon pavlowi |Parelephas P P 1S P. * I 4 i Mammonteus pgm genius anmentacus lesperoloxodon anfiquus raasi (Widely distributed over Europe) « « leith-adamsi Parelephas trogontherii Mammonteus primigenius Taasi «“ « leith-adamsi Archidiskodon meridionalis Parelephas trogontherioides Mammonteus primigenius astensis Hesperoloxodon antiquus ausonius « «« nanus 3 GUNZ | Archidiskodon| Anancus arvernensis meridionalis cromerensis Archidiskodon P antfrons rumanus Zygolophodon borsoni zaddachi Anancus gigantarvernensis ‘ minutoarvernensis Zygolophedon borsoni Anancus arvernensis “ « brevirostris “ « dissimilis “ « macroplus Zygolophodon borsoni buffonis “ « ~—vellavu “ « vialetii ana cus a Iconeri Turicius virqatidens Dinotherium Stegolophodon Sigantissimum sublatidens Dinotherium medium Dinotherium Siganteum Dinotherium proavus |Trilophodon Miomustodon tameorles « —podolicum| (Cheerolophodon) See esnus “ uralense pentelicus Anancus arvern-|Turicius atticus ensis progressor Tetralo hodon nancus grandincisivus intermedius Pliomastodon americanus Braciupics Trilophodon esselbornensis Tetralophodon longirostris « gigantorostris Platybelodon danovi Dinotherium bavaricum “ secundarium Trilophodon steinheimensis Dinotherium intermedium Dinotherium levius Zygolophodon pyrenaicus Turicius turicensis é “ engelswiesensis ma «« simorrensis “ angustidens minutus Trilophodon angustidens «“ “ austro-germanicus “ G Caine : Serridentinus subtapiroideus “ “ gai lardi Dinotherium cuvieri Zygolophodon pyrenaicus aurelianensis Turicius tapiroides Miomastodon depereti Trilophodon pontileviensis stidens cuvieri “ an Bernidentinus SA Tholi Dinotherium hungaric um DFlevett Bradley Fig. 1222 2. THE LOWER MIOCENE: BURDIGALIAN It is not the purpose in this exposition to go into the question as to whether the Aquitanian is more properly to be considered as the closing stage of the Oligocene or the opening stage of the Miocene, for this horizon precedes the appearance of proboscideans in Europe, and its consideration here would be fruitless. Suffice it to say, therefore, that the Miocene may be considered for the purposes of convenience to have opened with the Burdigalian. 1458 OSBORN: THE PROBOSCIDEA The development of Burdigalian mammals in Europe is perhaps best exemplified by the fauna of the Sables de l’Orléanais of central France. This fauna, known since the time of Cuvier, has long been regarded as typifying the Burdigalian stage in the European region, but in recent years as knowledge of the stratigraphy and palzeon- tology of these beds has become more exact, there have arisen the inevitable differences of opinion as to the exact relationships of the several horizons and their contained mammalian faunas within the Sables de l’Orléanais each to the other. The problem was stated by Mayet! in 1908 as follows. Les sables de l’Orléanais ne sont pas une formation géologique ayant la remarquable unité qu’on lui a longtemps atri- buée. A priori, il était 4 supposer que cette masse énorme d’alluvions granitiques qui couvre encore actuellement une partie du centre de la France, avait exigé un temps fort long pour étre formée, charriée, déposée. Les renseignements stratigraphiques et les données paléomammalogiques me paraissent confirmer cette induction. On peut distinguer dans le Burdigalien de l’Orléanais, dont l’expression ‘Sables de |’Orléanais’ est pour ainsi dire synonyme, plusieurs facies locaux, d’4ge trés probablement différent. It is Mayet’s contention, and in this he follows the lead of his illustrious countryman, Professor Depéret, that the Burdigalian of central France is inaugurated by the Calcaire de Montabuzard, underlying the Sables de ’Orléanais, and between these two horizons he would place the Sables de Chitenay. Thus, his Burdigalian section would be, generally speaking, as follows: Sables de la Sologne Marnes de |’Orléanais Sables de l’Orléanais Marnes du Blesois Sables de Chitenay Calecaire de Montabuzard Mayet has envisaged the Burdigalian in central France as beginning with the deposition of sediments in a vast lake, the “lac de Beauce.”” The Calcaire de Montabuzard is a local facies of this deposition, and as such contains a mammalian fauna which Mayet finds to be essentially more primitive than the typical Sables de VOrléanais fauna. The Sables de Chitenay are also regarded by Mayet as constituting a local facies of the basal Burdigalian, deposited by a large Miocene river which flowed from the Central Plateau into the lac de Beauce at about the time the deposition of the Calcaire was coming to anend. Either contemporaneous with or immediately succeeding the sands of Chitenay are the Marnes du Blesois. Then come the Sables de |’Orléanais in the strict sense of the word, with the large mammalian fauna typical of these sands. And finally at the top of the section are unfossiliferous beds, the Marnes de l’Orléanais and the Sables de la Sologne. A much different interpretation of the Burdigalian and associated faunas of central France was put forward by Stehlin in 1908.2. A decade or so before this time, Depéret had called attention to the very important fact that the fauna of the Sables de l’Orléanais contains two elements, one consisting of indigenous mammals, derived directly from the Aquitanian mammals of the same region—the other being composed of immigrant forms coming in from African, Asiatic, and American centers of origin. Stehlin elaborated on this thesis and attempted to show that the relative abundance of immigrant forms as compared with indigenous types is indicative to a certain extent of the general advancement in age of the several faunas within which they are contained. Following this line of thought, Stehlin came to the conclusion that the fauna of Chitenay is the most archaic of the Burdigalian mammalian assemblages, because it contains the smallest percentage of immigrant forms. Thus, he would designate the Chitenay fauna as lower Burdigalian, intermediate in position between the upper Aquitanian faunas, such as Saint Gerand-le-Puy and the characteristic middle Burdigalian fauna of the true Sables de l’Orléanais. ‘Mayet, L., 1908. Ann. Univ. Lyon, N.S., I, Fasc. 24, p. 313. *Stehlin, H. G., 1908. Bull. Soc. géol. France, (4), VII, p..545. GEOLOGIC SUCCESSION: EUROPE 1459 Perhaps the greatest discrepancy between the views of Stehlin and those of Depéret, Mayet and others, is his assignment of the Caleaire de Montabuzard to a position not below the Sables de l’Orléanais, but rather to an elevated place considerably above the Sables. Here again Stehlin has based his views on the fact that there is a very large immigrant element in the Montabuzard fauna, as compared with the indigenous mammals. Thus, this authority would regard the Montabuzard assemblage as being perhaps equivalent to or slightly later than the typical Sansan fauna. Stehlin admits, however, that: Si, un jour, on découvre une faunule de Mammiféres dans quelque lambeau de caleaire de l’Orléanais indubitable, ¢’est- a-dire directement recouvert par des sables fluviatiles burdigaliens fossiliféres, la question de Montabuzard sera tranchée. The localities at which Burdigalian faunules have been found in central France are numerous and are listed by Depéret, by Mayet, and by Stehlin, to which authors the reader is referred. Mayet, particularly, has given detailed discussions of the important localities at which Sables de l’Orléanais deposits containing mammalian remains have been discovered. As to the fauna of the Sables de l’Orléanais, the following forms might be accorded particular attention: Pliopithecus—according to Stehlin present in the Faluns de Pontlevoy and not in the Sables de l’Orléanais, strictly speaking. But Mayet placed this genus in the Sables, as well as in the Faluns. Certainly an advanced anthropoid, that might be of Middle rather than of Lower Miocene affinities. Amphicyon—typically Miocene. Pseudzlurus—also typically Miocene. Steneofiber—a characteristic Lower Miocene form. Proboscideans—all essentially primitive—Lower to Middle Miocene. Brachyodus—a persistent type from the Oligocene. Palzocherus—another persisting Oligocene form. Listriodon—characteristic of the Miocene. L. lockharti of the Sables is a very primitive member of the genus and might be considered as indicative of the beginning of the Miocene. 5} Palzomeryx, Amphimoschus, Procervulus, Dicrocerus—all primitive Miocene cervulines. Anchitheri'um—the earliest appearance of this genus in Europe. Macrotherivum—quite characteristic of the Miocene, and it may occur as early as the beginning of the period. Diceratherium—this genus? Aceratherium, Teleoceras—again typical Miocene forms. It might be well to mention here some important localities in addition to those discussed, namely, Chitenay, Chevilly, Eggenburg in Lower Austria, and Briittelen in Switzerland, at which latter place a shoreline facies has been discovered, with an association of marine forms. In a recent paper (1934) by Roman and Viret,' a Burdigalian fauna from La Romieu is described. This fauna is found to be contemporaneous with the upper Burdigalian of the Sables de l’Orléanais, particularly as characterized by the faunas of Baigneaux and of Chevilly. In this connection, it might be said that these authors make some interesting comments as to the characters of the upper Burdigalian fauna in general, as compared with ‘Roman, F., and J. Viret, 1934. Mem. Soe. géol. France, N. 8., IX, No. 21. 1460 OSBORN: THE PROBOSCIDEA the lower Burdigalian assemblage below, and the Helvetian fauna above. In short, the conclusions of Roman and Viret as to the characters of the upper Burdigalian fauna or faunas of Central Europe are as follows: The upper Burdigalian fauna differs from the older lower Burdigalian fauna in that: There are more of Brachyodus onoideus, Amphitragulus, Palexomeryx garsonnini, and typical Brachypotherium aurelianense. The following forms appear for the first time: Listriodon lockharti, Hemicyon, near H. gériachensis, Pseudxlurus quadridentatus, Lagomeryx of small size, Dorcatherium, Dinotherium, as characterized by the small D. cuviert, and a large Brachypotherium. The upper Burdigalian differs from the younger Helvetian in that: There have not as yet appeared such forms as Pliopithecus, Machairodus, Ursavus, Micromeryx, Potamo- therium, and above all the antelopes, which latter appear suddenly in the Middle Miocene of Sansan. Anchitherium aurelianense is present but as a small mutation—that is, it has not attained the size of the typical later Miocene Anchitherium. The same is true of Steneofiber. Palxocherus is present but not Hyotherium. The proboscideans of the Burdigalian are as follows: Dinotherium cuviert Kaup Type—Chevilly Turicius tapiroides (Cuvier) Type—Caleaire de Montabuzard Turicius turicensis (Schinz) (According to Mayet, type from Sarmatian) Dinotherium bavaricum von Meyer (According to Mayet, type from Helvetian) Miomastodon deperett Osborn Type—Chevilly Trilophodon pontileriensis Mayet and Fourtau Type—Pontlevoy Trilophodon angustidens cuviert (Pomel) Type—Gers Serridentinus filholi Frick Type—Gers Zygolophodon pyrenaicus aurelianensis Osborn Type—Chevilly Dinotherium hungaricum Ehik Type—Kotyhaza, Hungary 3. THE MIDDLE MIOCENE: HELVETIAN AND TORTONIAN = VINDOBONIAN HetvetiaAn.—The Middle Miocene of Europe, the Vindobonian, may be divided into two stages, the Helve- tian and the Tortonian, each typified by distinctive mammalian faunas. The characteristic Helvetian fauna is that of Sansan. The Sansan fauna, discovered by Lartet, and studied by this same authority and by Milne-Edwards, and later by Filhol, comes from a hill near the little village of Sansan in southern France. At this place there is a sue- cession of freshwater sediments, beginning with a clay bed at the bottom, followed by a caleareous layer, which in turn is capped by a series of marls containing freshwater molluscs and numerous fossil vertebrates—particularly mammals and birds. In contradistinction to the preceding Burdigalian fauna, as developed in the Sables de l’Orléanais, the Sansan assemblage is definitively a more modernized fauna, with no important persisting Oligocene types, which were so common in the Burdigalian faunas. Indeed, the Sansan fauna is characterized by the appearance of many mam- GEOLOGIC SUCCESSION: EUROPE 1461 malian forms indicative of the beginning of evolutionary developments that culminated in the modernized faunas of uppermost Cenozoic and Recent times. The origin of the Sansan deposits was discussed by Lartet, whose conclusions were substantiated by Filhol. Lartet had the following to say about the genesis of these beds: Toutes ces circonstances m’avient porté a penser que |’accumulation successive des fossiles organiques qui constitue le dépét de Sansan se serait effectuée au fond d’un marais, ou si l’on veut d’un petit lac o& auraient vécu les espéces aquat- iques dont on retrouve des débris; tandis que les eaux torrentielles auraient pu entrainer dans ce bas-fond, soit des osse- ments dispersés, soit les cadavres entiers des animaux terrestres établis 4 demeure ou du moins se montrant passagérement sur les terres environnantes, émergées pour un temps plus ou moins long.! L’hypothése de Lartet parait bien justifiée. I] y a eu A Sansan un lac, dont la plus grande partie a disparu, a été emportée i l’epoque du creusement des vallées. Dans ce lac venaient se déverser différents cours d’eau, susceptibles de déborder A certains moments et de couvrir probablement d’assez vastes espaces, d’ot ils emportaient les animaux y vivant au moment de leurs crues subites, pour les déposer en un point oU existait un remous.” Depéret has discussed at length the composition and relationships of the Sansan fauna, consequently the fauna in its entirety will not be listed here. But the following important genera contained within the fauna may be given some consideration. Pliopithecus—here we see the continuation of a relatively advanced anthropoid, which already has appeared in the Burdigalian fauna of the Sables de l’Orléanais and the Faluns de Pontlevoy. Insectivores—several genera, such as Hrinaceus, Sorex, Talpa, which are similar to the modern forms. Rodents—like the insectivores, genera leading to modern types. There should be mentioned particularly: Steneofiber—a characteristic Lower to Middle Miocene castoroid. This, or a related type, is found in the Lower Miocene of North America. Amphicyon—a typical Miocene canid, showing advanced characters. Hemicyon—one of the more advanced “‘bear-dogs,” distinetly later than Lower Miocene, and found at various Middle to Upper Miocene localities in Europe, Asia, and North America. Machairodus and Pseudelurus—Mhiocene Felide. Other carnivores are distinctly leading to more modern types. Anchither‘'um—the typical Miocene equid of the Eastern Hemisphere. Macrotherium—similarly, the typical Old World Miocene chalicothere. Acerather‘um—continuation of a form that first appeared in the Burdigalian. Listriodon—typical Miocene suid. Palexomeryx—a very typical element in the Miocene faunas of Europe. Trilophodon. Turicius. Of the European deposits correlative with those of Sansan, perhaps the most important are those of Engels- wies and Georgensgmiind in Germany, and Eibiswald and Goriach in Austria. Klahn (1922) describes the Engelswies deposits as chalky, brackish water sediments, probably deposited in an estuary or the mouth of a river that flowed out into the Middle Miocene mediterranean of Central Europe. These beds contain a typical Middle to Upper Miocene fauna, characterized notably by Dinothervum, Barchypo- therium, Aceratherium, Anchitherium, Macrotherium, and Hyotherium. Wlihn would correlate this fauna with the Steinheim assemblage, which would place it somewhat higher than it is regarded in the present work. 'Lartet, E., quoted by H. Filhol, 1891, pp. 5, 6. *Filhol, H., 1891, p. 6. 1462 OSBORN: THE PROBOSCIDEA The fauna from Goriach, described by various authors but particularly by Toula and by Hofmann, would seem to be a typically Middle Miocene mammalian assemblage. The presence in this fauna of characteristic Miocene genera, the lack of truly advanced Miocene forms, and the persistence of certain types from the Lower Miocene seem to favor the correlation of Gériach as approximately of Sansan age. There might be noted here such genera as Amphicyon, Dinocyon, Hemicyon, Trilophodon, Anchitherium, Steneofiber, Aceratherium, Hyo- therrum, Paleomeryx, Dicrocerus and other primitive deer, all of which relate this fauna very closely with the Sansan assemblage. The genus Cebocherus, described by Hofmann as from Goriach, is at best of doubtful refer- ence, for the material is very fragmentary. One would hardly expect this form in the Miocene. Others faunas of Helvetian age will compare with the Sansan fauna in about the same way as these, as con- sidered above. PROBOSCIDEANS: See list of Vindobonian proboscideans (p. 1464 below). TortTontan.—Of the Tortonian deposits in which fossil mammals have been found, none is perhaps better known than La Grive-Saint-Alban, Isére, in southern France. This locality has been known for many years, and has been studied extensively by Depéret, who published several large papers dealing with the fauna and the stratigraphic relationships of that deposit. Depéret has shown that the mammal-bearing beds of La Grive-Saint-Alban are in reality fissure deposits, formed during Middle Miocene times, in pits, fissures, and caves that were eroded out of Mesozoic limestones. In similar pockets at Mont-Ceindre, near Lyons, Depéret discovered such a wealth of bat remains that he was led to think that perhaps some of the deposits in this general complex or series were formed in Miocene bat caves. Some fifty years ago this author! described the typical deposits at La Grive-Saint-Alban, as follows (Depéret, 1887, p. 60): Ces fentes sont toutes ouvertes vers le haut, et pénétrent plus ou moins profondément dans l’épaisseur du terrain jurassique; la plupart cependant atteignent le niveau du sol de la carriére. Le remplissage de ces fentes, s’est opéré, il me semble, par le haut, et sous l’influence d’un simple ruissellement 4 la surface du caleaire; la faible vitesse des eaux pluviales sur ce plateau horizontal, et la lenteur du phénoméne de remplissage me paraissent suffire 4 expliquer l’absence si remarquable, dans les maté- riaux charriés, de cailloux roulés et méme de graviers, qui pourraient faire penser A un charriage opéré par un cours d’eau de quelque importance. Les seuls éléments étrangers a |’argile que j’ai observés sont des cristaux spathiques, qui paraissent résulter de l’évaporation répétée d’eaux riches en acide carbonique. Quant 4l’argile rouge, je pense qu’elle n’est ici que le simple résidu de la lente dissolution du calcaire, opérée d’une maniére continue par les eaux pluviales, pendant la longue émersion du continent mayencien, et accumulée dans les fentes par les eaux de ruissellement. A few years later (1892) Depéret? had the following to say as to the age and relationships of the fauna from La Grive-Saint-Alban. Ainsi que je l’ai déja indiqué en 1887, la présence A la Grive du Rhinoceros brachypus et du Dinotheriwm, qui manquent a Sansan, mais se retrouvent dans l’horizon un peu plus élevé de Simorre (Gers) annonce des affinités importantes entre la faune de la Grive et celle de cette derniére localité, sur laquelle nos connaissances sont malheureusement fort incomplétes. Ce dernier parallélisme est d’autant plus probable que, ainsi que je |’ai déja indiqué, les animaux de la Grive, tels que le Pliopithecus, le Machairodus Jourdani, \a Lutra Lorteti, le Cricetodon Rhodanicum, le Protragocerus Chantrei paraissent représenter dans leur ensemble un degré d’évolution légérement plus avancé que celui des espéces représentatives dans le gisement de Sansan. .. . Le gisement des sables de Steinheim (Wurtemberg) doit étre considéré comme |’équivalent exact de celui de la Grive. [in effet, sur les quarante-sept espéces de la Grive, dix-sept se trouvent aussi a Steinheim. . . . Les autres gisements de Suisse, d’Allemagne ou d’Autriche que l’on peut considérer comme a peu prés synchroniques de eelui de la Grive-Saint-Alban sont: les lignites de Gériach (Styrie); Georgensgmiind; Giinsburg et les sables a Dinotherium de Baviere; Ellg, Kapfnach; Ries (Nordlingen); Vermes (Jura Bernois); mais la faune de ces diverses localités est beaucoup plus pauvre que celle des autres gisements précités. 'Depéret, C., 1887. Arch. Mus. hist. nat. Lyon, IV, p. 60. *Depéret, C., 1892. Arch. Mus. hist. nat. Lyon, V, Pt. 2, pp. 4, 5. GEOLOGIC SUCCESSION: EUROPE 1463 Subsequently, in 1899, Gaillard! published a paper supplementing the studies of Depéret, and describing new or little known mammals from La Grive-Saint-Alban. Gaillard’s conclusions as to the age of this deposit were essentially the same as those of Depéret. Since the time that Depéret and Gaillard made their detailed studies of the fauna of La Grive, the general trend has been to substantiate and strengthen the conclusions of these authors as to the age of this mammalian assemblage. That the Helvetian and Tortonian stages are very closely related is shown by a comparison of their respective faunas. Thus, the Sansan and La Grive faunas may be compared, by genera, as follows. SANSAN La GrivrE-SAInt-ALBAN SANSAN La GRIVE-SAInT-ALBAN [SV METRMODEOIRS 9. cicioote eno SORE IOS CENCE TO ROE Pliopithecus VQUerrO is crseuy en eae seers epee ieee ieee Renee ee Viverra T a2 CGD a: 5556 BUCS SERS Oe re IRAN eee eS ae Vespertilio Herpestes Rhinolophus Vesperugo Progenetta [5 TGDTECUBS 6:6 bo tle Sra Ace cL ae eee eee Erinaceus WWhachav0dus are at ee OO ete ree Machairodus CN EUERGR 5 « 6.6 web OCEAN ORE TOE nee eee ree Galerix IP SCULRUUNULS ter cat PRR ToT oe COL CCIE Pseudelurus Lanthanotherium Aelurogale Mygale Dimylus plophodone sae ee COTA eS Oe Eee ee Trilophodon TEE 5 « © 6 6b Suge BORIS eee arene Sorex Turicius Dinotherium TDD. © 6 GUS ce NRO CIRCE Tarn ee eee are ee Talpa FV LO ONG RIOH ete a oe tie aR GIT oa OS Oda we Se Anchitherium IGIFIQ > 6 66 AHO ORD EEE eee Sciurus ACETAL eT re eeee spots SE ACC ee Aceratherium EELOD OLIMAR hee ee cose ainl 2 aisle es epee aed Br Cricetodon Macrotherium 1 PDTROOS o-'ono cers GEE Gre RCE REE aca Ra ea M yorus Chaeromonusn cnn eee oe ee Cheromorus Meriones? Cherotheriwm Arvicola? VERT ERINI Te 3 abs pnts cros Gombe OLD OE ttoe BUS ae Listriodon Steneofiber Hyotherium Lagopsis EY TNOSCHUSH seem oper eR OO Hyzemoschus Prolagus SY RYO OAT ROS Goel Oxon Oot aeeg EUS Oren © pickets © cc! Micromeryx REIL UOT Py area cyan (fle wlota aegis me Gemmle sh Hemacyon PGLLOMETY Ls ernest ecey eo Cae Moore Oe CE Palxomeryx Pseudocyon Morphelaphus Dicrocerus AOTIUOSEODS ol G-Ds 6-6 O10 BOS SOCIO ACER Orca Ter Amphicyon Strogulognathus SCORED 6 ollie & Stree ORIG REC aire Trochitis Palzxocervus Mustela Plesictis Cervus Putorius? Martes Antilope Protragocerus Proputorius Lutra From this comparative list it may be seen that there is little definitive evidence that enables one logically to separate the typical Helvetian faunas from those of Tortonian age. Indeed many of these faunas are considered as of one or the other of these two stages, by different authors, and no two authorities will correlate the Middle Miocene deposits of Central Europe in exactly the same way. Therefore there is much to be said in favor of lumping the several Middle Miocene faunas within one time division, the Vindobonian. Of the other so-called Tortonian faunas, some of the most important are those of Simorre and Villefranche d’Astarac in France, of Steinheim, Mésskirch, and Oppeln in Germany. The Steinheim fauna, preserved in an impure chalky ooze, or mud, contains the following diagnostic genera: Amphicyon, Viverra, Chalicomys, Aceratherium, Macrotherium, Anchitherium, Listriodon, Dicrocerus, and Palxo- meryx. This assemblage is generally regarded as of Middle to Upper Miocene age, equal to the La Grive fauna and perhaps younger than the typical Helvetian faunas. Kahn? speaks of it as follows: In Begleitung des unten zu beschreibenden Mast. steinheimensis deutet die Fauna auf obermiozines Alter hin. Diese ist dieselbe wie die von Kgl. Neudorf bei Oppeln und Grive-St. Alban. The fauna of Simorre is in all respects correlative with that of La Grive-Saint-Alban, and for this reason is often considered as typical of the Tortonian in Europe. The remarks as to the age and relationships of the fauna of La Grive apply equally well to that of Simorre. Gaillard, Claude, 1899. Arch. Mus. hist. nat. Lyon, VII. *Klahn, H., 1922. “Die badischen Mastodonten, etc.,” p. 33. 1464 OSBORN: THE PROBOSCIDEA Certain other European faunas, representing the final stages of the Vindobonian or Tortonian, may be con- sidered here. Of these, particular mention should be made of Giningen, Saint Gaudens, Elgg, and Monte Bamboli; Kapffnach, Giinsburg, and San Isidro. The (Eningen fauna, and flora preserved in fine-grained calcareous beds near Zurich, Switzerland, has long been famous for the excellence and diversity of the fossils comprising it. It is especially characterized by numerous plants, insects, reptilian and amphibian remains, as well as those of fossil mammals. Incidentally, it was here that the type of Andrias scheuzeri was discovered. De Lapparent' makes the following remarks concerning the (Eningen fossils: On attribue au sommet du tortonien les couches d’(!ningen, sur les bords du lae de Constance, caleaires en minces pla- quettes, extraordinairement riches en fossiles, insectes, poissons (Leuciscus), reptiles, etc. Mais c’est surtout par sa flore que le gite d’(iningen est célébre. Heer ena décrit prés de 500 espéces, parmi lesquelles bon nombre de formes européennes sont mélées A des types asiatiques, africains, australiens ou méme américains. L’assise inférieure ou couche A insectes d’Ciningen est composée d’environ 250 feuillets, o& l’on distingue jusqu’aux saisons successives: les fleurs de camphrier annongant le printemps, les fruits d’orme et de peuplier |’été, ceux de camphrier et de Diospyros approche de l’automne. . . Heer pense qu’il régnait 4 Giningen un climat analogue 4 celui de Madére, du Japon méridional et de la Géorgie, soit une moyenne annuelle de 18 4 19 degrés. The numerous faunas contemporaneous with or correlative with Giningen and Saint Gaudens were listed by Depéret in 1906. PROBOSCIDEANS: VINDOBONIAN PROBOSCIDEANS See list of Vindobonian proboscideans, below. Dinotherium levius Jourdan T. La Grive-Saint-Alban Dinotherium intermedium Blainville Type locality? France Dinotherium bavaricum Meyer H. Gmiind Dinotherium secundarium Kaup T. Simorre Zygolophodon pyrenaicus (Lartet) T. Ile-en-Dodon Zygolophodon borsoni affinis (Jourdan) Type locality? Trilophodon angustidens (Cuvier) T. Simorre Trilophodon angustidens minutus (Cuvier) Saxony Trilophodon angustidens gaujaci (Lartet) Lombez Trilophodon angustidens austro-germanicus Wegner Oppeln Trilophodon angustidens gaillardi Osborn T. Villefranche d’ Astarac Trilophodon engelswiesensis Klahn H. Engelswies Trilophodon steinheimensis Wlaibn . Steinheim Serridentinus subtapiroideus (Schlesinger) H. Wies Turicius turicensis (Schinz) T. Elgg Turicius turicensis simorrensis Osborn T. Simorre T.—Tortonian H.—Helvetian 4. THE UPPER MIOCENE: SARMATIAN The close of the Miocene and the opening of the Pliocene in Eurasia was inaugurated by a new cycle of sedimentation. This was, in brief, the beginning of the regression of Miocene seas, with a consequent inauguration of extended continental sedimentation, characterized at first by littoral deposits, and then, in a progressive fashion, 'De Lapparent, A. de, 1906. Traité de Géologie, p. 1615. GEOLOGIC SUCCESSION: EUROPE 1465 by lake beds and river channels. The change from a predominantly mediterranean and island type of landscape, typical of the European Miocene, to a truly continental type of landscape, so characteristic of the Pliocene, may be said to have begun in the Upper Miocene, progressing with increasing rapidity through the Sarmatian and finding its culmination in the Pontian. The question of the correlation of Sarmatian deposits and faunas, which are characteristically of marine, shallow-water facies, with the continental mammalian faunas of the same period is indeed difficult. This problem is chiefly concerned with the query as to just what may be considered as truly Sarmatian or equivalent mam- malian faunas. Certain deposits, such as Saint Gaudens, (Eningen, Monte Bamboli and related beds, listed by Osborn in 1910 (p. 257) as of Sarmatian age, are more properly to be considered as representing the uppermost phases of the Vindobonian or Tortonian in Europe, as shown in preceding pages of this chapter. This would seem to be the general consensus of opinion of authorities on the matter and need not be discussed at any greater length here. Yet the question of whether or not any mammalian faunas can be truly correlated with the marine Sarmatian stage is an important one, and it has been recently considered by von Koenigswald and by Tobien. Borissiak, in 1914 and 1915, described a mammalian fauna that was found at Sebastopol, on the borders of the Black Sea, in beds containing unmistakable Sarmatian molluscs. According to this author, the fossils were found as a “‘breche ossifere”’ in a small calcareous lens, intercalated within the upper zone of the middle Sarmatian of that region. Naturally, the occurrence of this fauna led the author, and numerous subsequent writers, to believe that the Sebastopol fauna was in truth a mammalian assemblage of Sarmatian age. Yet it is a very curious fact that the mammalian fauna of Sebastopol is typically Pontian in its characters. It contains Achtaria, a giraffid quite similar to Palzotragus, Tragoceras, Aceratherium, and Hipparion; all in all an assemblage that recalls Pikermi and Samos. Naturally, if this fauna is of true Sarmatian age, then it offers undoubted proof of the pre-Pontian occurrence of Hipparion—a fact of prime importance. For, if as some authors think, Hipparion appears in Eurasia before the beginning of the Pontian, the most important of the several criteria definitive of the Pontian loses much of its diagnostic value. The idea that the Sebastopol fauna is truly of Sarmatian age, or that Hipparion occurs in the Old World prior to the advent of the Pontian, has recently been disputed by von Koenigswald. This author believes that rather than a pre-Pontian appearance of what seems to be a typically Pontian fauna, the Sebastopol deposit represents a cave, or possibly a fissure deposit, in which animals of a later age have been intruded into sediments older than the period during which they lived. This explanation of the Sebastopol occurrence, if valid, goes a long way towards clearing up what has always been a stumbling block in the correlation of Eurasiatic mam- malian horizons. Von Koenigswald! says that: ‘“‘Das sarmatische Alter der Hipparion-Fauna von Sebastopol kann bei der Art der Einlagerung und der Erhaltung der Reste nicht als unbedingt erwiesen angesehen werden.”’ Von Koenigswald goes on in the same contribution to dispute the idea that Hipparion is present as a pre- Pontian form in any of the several localities where it has been reported as of an age preceding the characteristic Pontian fauna. As opposed to this view is the recent work of Tobien (1938), in which it is maintained that not only at Sebastopol, but at various other localities, particularly in eastern Europe, there are beds of true Sarmatian age, 'Koenigswald, G. H. R. von, 1931. Sonderabdruck aus dem Centralblatt f. Min., ete., Abt. B, No. 1, p. 45. *Tobien, H., 1938. Zeits. deutsch. Geol. Ges., XC, Heft 4. 1466 OSBORN: THE PROBOSCIDEA bearing Hipparion. The localities other than Sebastopol cited by Tobien are the trans-Caucasian region (Alekse- ev, 1930), Linz (Giimbel, 1894), Bavarian Flinz (Freudenberg, 1928), Odessa (Andrussow, 1905), Constantinople (Dietrich, 1933), Taraklia in Bessarabia (Khomenko, 1913, 1914), Thrace (von Arabu, 1916, 1919), Rumania (Krejci-Graf, 1932). And in addition, Tobien describes some teeth and astragali of Hipparion from (iningen (Ohningen). So the problem stands at the present time. If these localities are truly of Sarmatian age, then they represent what might be considered as the final stage of the continental Miocene in Europe, a stage during which Hipparion and other Pontian types appear, foreshadowing the tremendous spread of the Hipparion faunas of the true Pontian. If, on the other hand, these localities are of Pontian age (at least those in which Hipparion is present) then it may be said that for practical purposes there are no continental mammalian faunas that may be proven as the exact equivalents of the marine Sarmatian, and that the continental Miocene of Kurope comes to a close with such upper Vindobonian or Tortonian assemblages as those of Saint Gaudens and the like. At best the problem works down to one of definitions, and as such is a difficult one to settle satisfactorily. 5. THE LOWER PLIOCENE: PONTIAN The new faunal and sedimentation cycle that had its beginnings in the Sarmatian continued with increased momentum into the following Tertiary subdivision in Europe, the Pontian. The mediterranean sea, so character- istic of the Middle Miocene of Central Europe, became increasingly restricted, continental flood-plain and lacustrine deposits replaced marine beds, and mammalian faunas became exceedingly wide-spread and character- istic. In this connection, there need only be repeated the well-known fact that the Pontian was the time during which the characteristic Hipparion fauna spread throughout Eurasia. This fauna, which all in all was remark- ably uniform throughout the extent of its range, stretched from Spain and western Europe to China, from the shores of the Mediterranean as it was at that time into northern Europe. It was a wide-spread assemblage of mammals living in a plains or steppe environment, and showing a great preponderance of grazing perissodactyls and artiodactyls, accompanied by the carnivores that might be expected with an ecological and faunal association such as this. The presence of the Hipparion fauna in the Oriental and the Asiatic areas, and its significance with regard to the correlation of the upper Tertiary beds of these regions, has been discussed in other sections of this chapter. At the present time the discussion will be limited to the occurrence and significance of the Pontian Hipparion fauna in Europe. In a discussion of the continental Pontian faunas, the fact must be kept in mind that these mammalian assemblages mark the beginning of a new period in the history of mammalian faunas. This was the time when truly modernized types of advanced mammals made their first appearance, and particularly when immigrant forms suddenly became conspicuous in the Eurasiatic faunas, to characterize these faunas wherever they might be developed. Of course, as the name implies, the most characteristic of the immigrant genera is Hipparion. That this genus is of undoubted North American origin can no longer be questioned by anyone who has made a careful study of the succession of the Equide. Consequently, as has already been pointed out in other sections of this chapter, the appearance of Hipparion in the Old World must be subsequent to the time at which it arose in North GEOLOGIC SUCCESSION: EUROPE 1467 America—whatever that time may be. Certainly Hipparion did not become a genus distinct from its ancestor Merychippus, before the uppermost part of the Miocene in North America. And even though some evidence would seem to indicate that Hipparion might have made its first appearance in North America during Upper Miocene times, other evidence has been interpreted to indicate the fact that the genus did not appear in the New World until the beginning of the Pliocene. Turning now to a consideration of other Pontian types, we see numerous immigrant and specialized forms appearing for the first time in Eurasia. Among the carnivores it is interesting to note that true bears such as /ndarctos and Ursavus—as distinguished from the ancestral ‘‘bear-dogs,”’ such as Hemicyon—first appear in the Pontian. Likewise, the hyzenas, of Asiatic and Oriental origin, become well-established elements in the Tertiary faunas in Pontian times. True enough, these animals first appear in Upper Miocene times, in the immediately pre-Pontian deposits, but it is in the Pontian that the hyzenas become widely distributed throughout the European region. A conspicuous element in the Pontian faunas of Europe, particularly along the eastern border of the Mediter- ranean, is the aardvark, Orycteropus. The tubulidentates, though now of African distribution, did not necessarily originate or evolve on that continent; indeed, all of the fossil aardvarks of pre-Pleistocene age come from the Eurasiatic and Oriental regions, so that the group may well have originated somewhere in the north. In a different category is the gigantic hyracoid, Pliohyrax—a conspicuous member of the Pikermifauna. We know quite definitely that the hyracoids originated in Africa in Eocene times, and so the presence of a member of this group in the European Pontian implies an immigration into the fauna from a southerly source. The most conspicuous elements of the Pontian fauna, with the exception of Hipparion, are the numerous ruminants. Here is an advanced group of mammals, appearing for the most part as newly evolved elements in the mammalian assemblage. Our evidence points rather strongly to the fact that the ruminants of advanced type are relatively recent developments in the mammalian world. That is, the common ancestor of the deer, of the giraffids, and of the bovids is probably to be found in the Miocene. In the European Pontian is a very primitive giraffe, Palzotragus, not greatly different in most of its characters from the modern African okapi. With Palzotragus is Samotherium, which is nothing more nor less than an en- larged type of Palzxotragus, and Helladotherium, a gigantic giraffid belonging to a group that was to become widely spread during Upper Pliocene and Pleistocene times. It would seem probable that the origin of the giraffids might have been Oriental—that they were immigrants into Europe and into Africa, and that the modern okapi represents a persistent primitive form, pushed to the periphery of the range for this family by more specialized types developing at or near the center of origin. This was the age of the first flowering for the gazelles and the antelopes, and these animals are to be found in great profusion throughout the Eurasiatic and Oriental Pontian. There is a host of forms (recently monographed by Pilgrim) that give to the Hipparion fauna a very African appearance. It would seem, however, that these various gazelles, sheep, antelopes, and pre-cattle were of northern origin and migrated in Pontian or post-Pontian times to the southern continent. 1468 OSBORN: THE PROBOSCIDEA With these considerations in mind, what is the age of the Pontian? Among French scholars the European Pontian has been generally accepted as of Upper Miocene age, while many Germans would place the Pontian at the beginning of the Pliocene. Many years ago, as pointed out elsewhere in this chapter, Matthew suggested the desirability of making the continental Pliocene throughout the world coincident with the appearance of Hip- parion. This definition for the beginning of the Phocene is, on the whole, as satisfactory a designation as has ever been proposed. Consequently, the Pontian is here considered as of Lower Pliocene age, corresponding in Europe to those stages in Asia, the Orient, and North America at which Hipparion first becomes a definitely identifiable element in the mammalian faunas. The outstanding Pontian localities of Europe are listed below, taken from the list as presented by Pilgrim in his ‘‘Pontian Carnivora of Europe.” These localities, stretching from Spain on the west to the Black Sea region and Asia Minor on the east, are characterized and correlated by the uniformity and the similarity of their faunas. Spain Aragon, Catalonia, La Mancha France Montredon, Orignac, Cerdagne, Mont Léberon, Cucuron, Croix Rousse Germany Eppelsheim, Salmendingen, Melchingen, Trochtelfingen Sicily Gravitelli Greece Pikermi Macedonia Salonica, Veles Samos Samos Hungary Baltdvar, Polgardi, Csakvar, Baroth-kopecz Black Sea (Sebastopol), Novo Elisavetovka, Taraklia Kischinev, Grebeniki, Tschobrotschi Persia Maragha Of these, perhaps the best known from their faunas, are: Mont Léberon, Cucuron, Eppelsheim, Pikermi, Samos, and Maragha. Of course, the presence of typical Pontian faunas characterized by Hipparion at the above localities affords strong evidence as to their general unity and contemporaneity. Yet it is quite possible, as Pilgrim has pointed out, that all of these faunas may not be exactly equivalent, each to the other, in age. That is, homotaxial factors may play a certain part in the spread and delimitation of the Pontian complex in Europe—just as they most probably did (as shown in another section of this paper) in India. Indeed, certain authorities would subdivide the Pontian, particularly on the basis of the geology of the Black Sea area, into at least three zones. But in spite of these considerations it is to be remembered that the Pontian faunas represent essentially a definite phase in the evolution of mammalian faunas, and as such are for all practical purposes to be considered together as a unit, even though there may not be exact time identities between them—due to homotaxial lags dependent upon mi- grations or other causes. This whole question has been no better stated than by Pilgrim (1931), as follows: At the same time, there is no doubt that the ‘Hipparion fauna’ does occur at different levels, of which the lowest, including perhaps the Sebastopol fauna described by Borissiak [but see remarks in the discussion preceding this on the Sarmatian problem] (1915), has been referred to the Upper Sarmatian; the fauna of Taraklia (Khomenko, 1914) and Novo-Elisavetovka (Alexejew, 1916) are classed as Maeotic, or intermediate between the Sarmatian and the Pontian, while the remaining localities are truly Pontian. It is hardly conceivable that these three stages should not be represented among the so-called ‘Pontian’ deposits of the rest of Europe. It may be that increased material, and intensive study of what is already in our museums, may one day render such a correlation possible, but in my opinion the state of our knowledge does not permit of it at present. . . However this may be, there can be no doubt that we are dealing with a definite faunistic unit. Its homogeneous character is specially striking when we compare it with the Vindobonian fauna of La-Grive-Saint-Alban which preceded it or that of Monte Olivola, Rousillon and Montpellier, which followed it. The question, however, arises as to whether such a fauna, which includes both Sarmatian and Pontian elements, can strictly be called Pontian. These names were definitely applied to certain marine strata in the Black Sea region, and their application should not be either restricted or extended. Many writers, seeing this, have written of this fauna as the ‘Hipparion fauna’, but since the genus Hipparion seems to have a wider range even in Europe and still more so in Asia and North America than the fauna we are considering, that term does not meet the need, and we seem therefore bound to speak of the Sarmato-Pontian fauna. 'Pilgrim, G. E., 1931. Catalogue of the Pontian Carnivora of Europe. Brit. Mus. (Nat. Hist.), pp. 147, 148. GEOLOGIC SUCCESSION: EUROPE PONTIAN PROBOSCIDEA 1469 Trilophodon (Choerolophodon) pentelicus (Gaudry and Lartet) Pikermi Turicius atticus (Wagner) Pikermi Tetralophodon grandincisivus (Schlesinger) Maragha Tetralophodon longirostris (Kaup) Eppelsheim Dinotheriwm medium Kaup Eppelsheim Dinotherium giganteum Kaup Eppelsheim Trilophodon esselbornensis (Kahn) Esselborn Turicius wahlheimensis (Klihn) Wahlheim Tetralophodon gigantorostris (Klihn) Bermersheim Miomastodon tapiroides americanus (Schlesinger) Tasnad Pliomastodon americanus praetypica (Schlesinger) Dinotherium proavum Eichwald Batta-Erd Podolia, Russia Dinotherium podolicum Eichwald Podolia Dinotherium uralense Kichwald Ural Mts. Anancus arvernensis progressor Khomenko S. Bessarabia Anancus intermedius (Kichwald) Volhynia Platybelodon danovi Borissiak (Sarmatian) Kkuban, N. Caucasus, Chokrak Beds 6. THE MIDDLE PLIOCENE: PLAISANCIAN Following the expansion of Pontian mammals in Europe, there would seem to be a gap in the succession of mammalian faunas—perhaps the result of extensive marine inundations. This fact has been pointed out by numerous authors, notably by Depéret, who was the first student to recognize and elucidate this distinct break in the sequence of Phocene mammalian faunas in Europe. Osborn (1910)! emphasized the importance of Depéret’s conclusions, as quoted below, and more recently Pilgrim (1939)? pointed out the reality of a Middle Pliocene hiatus in the European fossil mammal faunas. As this is chiefly a marine phase, the terrestrial mammalian fauna is imperfectly known. The typical deposits are those of the lignites of Casino (Tuscany) which are correlated by Depéret with the lacustrine deposits of Autrey in the valley of the Saone, France. On the east coast of England is a marine formation, the Coralline Crag of Suffolk, containing a mastodon and a rhinoceros (Dicerorhinus). Of course this gap is not complete, for there are scattered mammalian remains known from deposits of Plaisancian age, but generally speaking the continental facies in this stage are of little importance. And it is interesting to note that in many cases the fossil mammals of Plaisancian age are found in littoral deposits, some- times associated with diagnostic marine invertebrates. All of these facts were made clear by Depéret, in his classic studies of some fifty years ago. Depéret considered the Plaisancian and the succeeding Astian as closely related, and together they constituted his ‘older Pliocene”’ which he distinguished from the “‘new Pliocene”’ or Sicilian. In the older Pliocene, according to this author, were numerous holdovers from the Pontian fauna, while the Sicilian is marked by the influx of a new and modernized fauna, quite distinct from those preceding it. According to the chronology adopted in this 1Osborn, H. F.,1910. “The Age of Mammals,” pp. 311, 312. *Pilgrim, G. E., and A. Tindell Hopwood, 1939. Rec. Geol. Surv. India, LX XIII, Pt. 4, pp. 445, 446. 3Depéret, C., 1893 (1894). Bull. Soc. géol. France, (3), XXI, pp. 529, 530. 1470 OSBORN: THE PROBOSCIDEA present work, the Pontian, of course, represents the beginning of the Pliocene, while the Plaisancian and the Astian, still showing mammalian types of Pontian origin, constitute the middle and upper phases of the Pliocene, respectively. The Sicilian, with the influx of new and modernized types, such as Equus and Archidiskodon, represents the beginning of the Pleistocene (Depéret, 1894, op. cit., pp. 529, 530): On distingue nettement dans le faciés marin du Pliocéne trois étages: le Pliocéne infériewr ou Plaisancien, dont le type est dans les argiles bleues subapennines; ensuite le Pliocéne moyen ou Astien, fondé sur les sables jaunes trés fossiliféres du pays d’ Asti ot ils surmontent avec évidence l’étage des argiles bleues. Ces deux étages marins .. . . sont étroitement unis l’un A l’autre par leur distribution géographique et méme par leur faune (qui dénote une mer assez chaude), de sorte que l’on pourrait A la rigueur, avec MM. de Rouville, de Stefani, Welsch, ete., les considérer comme les deux phases successives, la seconde plus littorale, d’un méme dépot, sur un fond de mer en voie d’exhausse- ment graduel. Depéret then goes on to show that in Italy mammalian remains are rare in the marine facies of the Plaisancian, and in the Astian as well, while to the north, especially in France, the Plaisancian deposits constitute a series of blue clays with invertebrate remains, underlying or antecedent to the mammaliferous Astian sediments. Pilgrim has recently restated Depéret’s arguments, and has particularly emphasized the Plaisancian faunal gap by demonstrating that the succeeding Astian fauna shows closer resemblances to the Sicilian fauna above it, than to the older Pontian fauna. Very little of the Roussillon and Montpellier fauna became extinct between that stage and that of the Val d’Arno, Perrier and Senéze, as compared with the multitude of families and genera which appear for the last time in the Pontian. I can see no grounds for regarding the two faunas as very different in age, while on the contrary a great zoological gap is suggested between Roussillon and the Pontian. Two littoral or marine stages, the Plaisancian and the Astian, intervene between the Villafranchian and the stage which corresponds to the mammaliferous lignites of Casino. The fauna of the Casino lignites is perhaps slightly younger than that of the Pontian. Depéret, therefore, most reasonably considered that the Roussillon fauna corresponded with the Astian, while the Plaisancian filled the faunal gap between the Pontian and Roussillon faunas. There is no mammal fauna known in Europe which can be said to correspond with certainty to the Plaisancian. [Italics my own. |] Three proboscideans have been referred very questionably to the Plaisancian. These are: Turicius virgatidens (von Meyer) Fulda, Germany Stegolophodon sublatidens (Schlesinger) Teschen, Austria Dinotherium gigantissimum Stefinescu Gaiceana, Rumania 7. THE UPPER PLIOCENE: ASTIAN The final Tertiary stage in Europe is that of the Astian, typified by the deposits of Villanova, Asti, in Italy, and characterized by the very rich mammalian faunas of Roussillon and Montpellier in France. Depéret demon- strated very ably, as already shown above, the fact that the Astian fauna or faunas are essentially modernized— much closer to the Pleistocene and Recent faunas of the Holarctic region than to the Pliocene faunas preceding this stage. This point has been strongly emphasized by Pilgrim, in a recent paper, and it was more generally stated by Osborn in 1910,! as follows: Surveying this. . . Pliocene fauna as a whole we are struck by the great predominance of animals closely related to exist- ing forms. If the living zodlogist should imagine himself in France at this period, he would see only four animals which would appear entirely novel and unknown, namely, the saber-tooth tiger, the mastodon, the hipparion, and the hysnarctos; all the rest of the fauna would seem to be a very strange commingling, or congress, of African, European, and Asiatic mammals of the present day. Not a single North American element would be observed in this assemblage, unless we except those elements of more remote migration, such as the hares, the tapirs, and possibly the hipparions and the foxes. 1Osborn, H. F.,1910. ‘The Age of Mammals,” p. 317. GEOLOGIC SUCCESSION: EUROPE 1471 One of the chief deposits of Astian age, perhaps the most important one in Europe, is that of Roussillon, fully described by Depéret in his classic monograph of 1890.!_ According to this author, after the retreat of the Plaisan- cian sea, the Roussillon basin was filled with a considerable thickness of argillaceous and silicious sandstones, and in addition caleareous muds, all having a fluviatile origin. It is in these latter beds that the characteristic Rous- sillon fauna was deposited. En revanche, ces limons sont le gisement d’une riche faune de vertébrés terrestres et fluviatiles, dont les débris charriés par les courants, se retrouvent aujourd’hui dans quelques points privilégiés, qui correspondent sans doute 4 des remous ou 4 des parties plus stagnantes du cours des anciennes rivieres. Les piéces osseuses sont presque toujours isolées, souvent brisées et méme un peu roulées; il est assez rare de rencontrer plusieurs os en connexion. Cependant diverses parties d’un méme squelette se retrouvent quelquefois dans un rayon peu étendu; méme le squelette de la Testwdo Perpiniana a pu étre déterré tout entier grace A la protection efficace de la boite osseuse, et 4 ’habitude qu’ont les Tortues de rétracter leurs membres et leur téte dans Vintérieur de la carapace. Les ossements se rencontrent dans les limons d’eau douce sur presque toute l’étendue du bassin du Roussillon; les localités les plus riches sont: Villemolaque, Trouillas, le mas Belrich dans la vallée du Réart; Thuir, Millas, le Soler, le Serrat d’en Vac- quer, la citadelle et les briqueteries des portes Canet et St-Martin, 4 Perpignan, dans la vallée de la Tét; les briqueteries de Rivesaltes dans la vallée de l’Agly. Pilgrim? (1939) has presented a very helpful review of the Roussillon fauna, particularly with regard to its relationships with European assemblages below and above it. Some of his remarks are as follows: The character of the European fauna altered considerably between the Pontian and the stage of Roussillon. Numerous families and genera have disappeared. . . . There are very few surviving genera from the Pontian, and almost all of these differ specifically. .. . On the other hand numerous species from Roussillon are identical or nearly so with Villafranchian forms. The differences which the fauna of the Villafranchian displays from that of Roussillon are mainly due to what are invading forms, which does not necessarily imply any great difference in age... . When we observe that the comparable forms are either specifically the same or differ very little from one another, we are forced to conclude that little development took place between the two levels. Very little of the Roussillon and Montpellier fauna became extinct between that stage and that of the Val’Arno, Perrier and Senéze, as compared with the multitude of fami- lies and genera which appear for the last time in the Pontian. I can see no grounds for regarding the two faunas as very differ- ent in age, while on the contrary a great zoological gap is suggested between Roussillon and the Pontian. The Astian fauna, as known from the assemblages of Roussillon and Montpellier, is large and varied and shows a composition somewhat as follows: Among the primates are Dolichopithecus, Semnopithecus, and Macacus, the first of which failed to survive into Pleistocene times. The carnivores display a considerable variety, including Vulpes, Ursus (both appearing for the first time in Europe), Agriotherium, Lutra, Viverra, Hyena, Megantereon, Epimachaerodus, and Felis. A large group of rodents, including Hystriz, and one form, Ruscinomys, which became extinct at the end of the Pliocene. The proboscidean genera, Zygolophodon and Anancus. Among the perissodactyls, Hipparion as found in the Astian represents a long-persistent survivor from Pontian times. This genus, which generally became extinct at the end of the Pliocene, did persist in some regions into the Lower Pleistocene. Also there is to be noted Tapirus and Dicerorhinus. Among the artiodactyls Potamocherus and Capreolus appear here for the first time. There are also various bovids, such as Palzoryx and Gazella. 1Depéret, C., 1890. Mém. Soe. géol. France, ITI, p. 9. *Pilgrim, G. E., and A. Tindell Hopwood, 1939, op. cit., pp. 443-445. 1472 OSBORN: THE PROBOSCIDEA There is also Orycteropus, a large species as compared with the characteristic Pontian forms—although smaller than the recent types. Of particular significance is the absence of certain diagnostic genera from the Astian faunas, notably Equus, Elephas (Archidiskodon), and bovines, such as Leptobos. It is the absence of these types that gives conclusive proof of the distinction of the Astian fauna from the succeeding Villafranchian, where these animals as immigrant forms appear for the first time in Europe. This difference was noted by Depéret, and has been emphasized by later authors. In addition to the characteristic Astian mammalian faunas of France, and to a lesser extent of Italy, there are limited occurrences of mammals of this age in other parts of Europe, notably along the eastern coast of England. Here is found the Red Crag deposit, a marine sediment containing occasional mammals. The mam- mals, when found in the Red Crag, prove to be of the same types as those found at Montpellier and Roussillon. The occurrences of the types of Astian proboscideans are given by Osborn as follows. In this connection it might be mentioned that Pilgrim (1939) denies the presence of Zygolophodon borsoni at Roussillon, although Depéret in his monographic study lists this species among the Roussillon fauna. ASTIAN PROBOSCIDEANS Zygolophodon borsoni (Hays) Villanova, Asti, Italy Zygolophodon borsoni buffonis (Pomel) Auvergne, France Zygolophodon borsoni zaddachi (Jentzsch) Thorn, Germany Zygolophodon borsoni vellavus (Aymard) Velay, France Zygolophodon borsoni vialetii (Aymard) Vialette, France Anancus arvernensis (Croizet and Jobert) Perrier, France Anancus arvernensis brevirostris (Gervais and de Serres) Montpellier, France Anancus arvernensis dissimilis (Jourdan) Sadne Basin, France Anancus arvernensis macroplus Aymard Mt. Coupet, Puy-en-Velay, France Anancus gigantarvernensis (Klaihn) Herbolzheim, Germany Anancus minutoarvernensis (Klihn) Herbolzheim, Germany Anancus falconert Osborn Suffolk, England 8. THE PLEISTOCENE There was no abrupt change from the Pliocene to the Pleistocene in Europe, a condition which was generally true for the other parts of the World. Consequently the boundary between the two epochs is difficult to define, with the result that there are manifest differences of opinion as to what event or events in geologic history may be properly regarded as significant of the opening of Pleistocene times in this region. In a general way, the Pleisto- cene is the Great Ice Age, when portions of the northern hemisphere were partially covered by several successive glaciations, and it might be supposed that the advance of the first continental glacier would serve to mark the beginning of Pleistocene history. But some of the European authorities, notably Boule, would place the first glaciation in the Upper Pliocene. Moreover a great portion of the Harth’s surface was not affected by glacial phenomena, so that other criteria must be used, no matter what may be the opinion as to the time of the first glacial advance. The sequence of pluvial and dry periods, the development of marine and river terraces and other geological phenomena have been widely studied and variously used as aids in Pleistocene chronology in the non- GEOLOGIC SUCCESSION: EUROPE 1473 glaciated regions of the Earth, yet even these evidences, useful as they are in establishing the succession of diastrophic, climatic or sedimentary events within the Pleistocene period, have not been generally satisfactory in an attack upon the problem of the beginning of the Pleistocene. Therefore the evidences of wide-spread changes in the mammals are perhaps the most significant and the most useful of the criteria studied, in establishing the open- ing of Pleistocene times. The mammals were rapidly evolving animals, able to migrate quickly over most of the Earth’s surface, so that changes in mammalian faunas necessitated by the adaptations required by the development of new and unusual environmental conditions would almost instantly (from a geological point of view) be reflected throughout wide areas. The problem has been excellently stated by Hopwood,' as follows: Apparently the boundary between the Pliocene and Pleistocene is difficult to determine on purely geological evidence, so that it is necessary to approach the problem in some other way, preferably by means of the fossil mammals since they furnish the raw material for the rest of the inquiry. There are many factors involved, but two of them are far more important than the others. First of these is the proposition that the best boundaries in time are those expressed by a change in fauna, which means in practice the incursion of new types, rather than the disappearance of old ones. The second proposition is, that the time necessary for the distribution of new types of active quadrupeds, e.g., horse or bison, over a very wide area is negligible from the geologi- cal point of view. At the beginning of the so-called Sicilian or Villafranchian time division in Europe and Asia, there was a sudden appearance of new and modernized types of mammals, part of which were the descendants of indigenous ancestors, part of which were immigrants. Notable among these new types were the modern forms of horses (Hquus), of cattle (Bos), and of elephants (Archidiskodon, Elephas and related genera). As long ago as 1911, Haug suggested that the appearance of these key types, the first an immigrant from the New World, the others of Old World origin, should mark the beginning of the Pleistocene in Europe. In recent years this view has been reiterat- ed, notably by Matthew and by Hopwood. If accepted, then the typical Sicilian or Villafranchian faunas of the European region are to be regarded as of Lower or Basal Pleistocene age, and it is the influx of these new types that marks the beginnings of the period. As opposed to this, many European authorities are inclined to regard the Villafranchian fauna, containing Equus, Bos, and Elephas, as marking the summit of the Pliocene. In the present chapter the former interpreta- tion will be favored. Granting that the opening of the Pleistocene is marked by the appearance of numerous modernized mammals, especially Hquus, Bos, and EHlephas (in the broad sense of the term), we may now consider the problem of dat- ing the sequence of events within Pleistocene times. This has occupied the attention of many students over a long period of years. In Europe the criteria most generally used for establishing dates within the Pleistocene are those of the glacial succession, as delineated especially by Penck and by Geikie, and of the sequence of marine and river terraces, as described by de Lamothe, Depéret, and others. In northern Europe there are evidences of four major glaciations, with interglacial periods between them. In the Mediterranean region there are the remains of four marine terraces, which may be traced in part around the Atlantic coast and into the North Sea. In 1919, Depéret indicated the relationships between the European glaciations and marine terraces in the following tabular form. NorTHERN HUROPE ALPS MrEpDITERRANEAN Fourth glaciation Mecklenburgian Wiirm Monastirian 18-20 meter terrace Third glaciation Polonian Riss Tyrrhenian 30 meter terrace Second glaciation Saxonian Mindel Milazzian 55-60 meter terrace First glaciation Scanian Giinz Sicilian 90-100 meter terrace ‘Hopwood, A. T., 1935. Proc. Geol. Assoc., XLVI, Pt. 1, p. 46. 1474 OSBORN: THE PROBOSCIDEA It must not be supposed that this compilation represents an exact correlation, since it is known that the marine terraces may be correlated with the river terraces, while these latter underlie the moraines of the glaciations to which they are most closely related. Therefore, it is probable that the terraces are more or less of interglacial age, and immediately precede the glaciations with which they may be roughly correlated. The important fact is that there is a fourfold division of the Pleistocene in Europe, as based either upon the development of marine and river terraces, or upon the sequence of glaciations. And each glaciation may be considered as occupying the latter portion of the general terrace stage, as shown below. Upper Monastirian IV Wiirm IV L is ower Monastirian nal Upper Tyrrhenian III Riss Lower Tyrrhenian II Upper Milazzian II Mindel Lower Milazzian I Upper Sicilian I Giinz Lower Sicilian Many students, following particularly the work of Penck, have envisaged a succession of ‘‘warm”’ and ‘‘cold” mammalian faunas in Europe that might be correlated more or less closely with the successive advances and retreats of the great ice sheets. According to Penck’s scheme, the four glaciations were more or less equally developed, but separated from each other by unequal intervals, of which the median one, the Mindel-Riss inter- glacial, was by far the most protracted, and the last one, the Riss-Wiirm, the shortest. He also distinguished the Wiirm glaciation from the others by his supposition that it terminated in a series of minor fluctuations, rather than by a single and steady decline. This idea of alternating faunas controlled by glacial phenomena has been expressed by Osborn’ in the following words: The principal contributors to the theory of northward and southward migrations and to the succession of faunas are Nehring, Woldrich (1882), and more recently Penck. In considering the distribution and migration of the mammals throughout the Glacial Period, we must constantly keep in mind the differences of latitude. . . Penck also observes that we cannot hope to trace a continuous evolution of forms during Pleistocene times, because we are not dealing with a development of one successive series in one locality, but with the cyclical alternation of a number of different faunas compelled to migrate through the alternations in the temperature and in the floras, the mammals disappearing and re- turning at intervals too brief to allow of any marked evolutionary changes. Herein lies our difficulty when we attempt to distin- guish between the tundra faunas of the late glaciations and the forest faunas of the late interglacial epochs, because the faunas return not only with the same generic but the same specific types, as especially illustrated in the case of the mammoth (#. primigenius) and the giant deer (Cervus megaceros). A considerably different picture has been presented recently (1933) by Stehlin,? who thinks that the glacial periods were not only separated each from the other by interglacial periods of varying lengths, but also that they were in themselves much more different from each other in intensity than was admitted by Penck. Thus Stehlin, basing his conclusions upon the development of the mammalian faunas, supposes that the glaciation at the begin- ning of the Pleistocene was relatively much less intense than the two glaciations coming after the long Middle Pleistocene interglacial period. (Incidentally, this author, like Boule, recognizes only three glaciations in the Pleistocene.) According to Stehlin, the maximum of glacial activity was reached towards the end of the Wiirm ‘Osborn, H. F., 1910, op. cit., pp. 388, 389. “Dubois, A., and H. G. Stehlin, 1933. Mem. Soc. Pal. Suisse, LIJ—-LIII, p. 272. GEOLOGIC SUCCESSION: EUROPE 1475 glacial period, a conception based upon the sequence of European Pleistocene faunas and which regards the “warm” or temperate types of mammals as prevailing through a great part of the Pleistocene, to be followed in “cc Wiirm and then in post-Wiirm times by an invading arctic assemblage: . la plus grand invasion des animaux arctiques dans les latitudes moyennes et méridionales ne coincide avee le développement maximal des glaciers wiirmiens, mais lui succéde. . .” This view is somewhat in accord with the recent tendency to discount to a large degree the reality of alternat- ing “cold” and “warm” faunas, and to regard the development of the Pleistocene mammals as a rather continuous process, with minor fluctuations due to the alternation of mild and severe temperature conditions. Hopwood (1939, 1940), for instance, has the following to say on this subject: At this stage it may be useful to discuss some of the points which govern the use of mammalian faunas as indices of the age of Pleistocene deposits. It should be self-evident that the composition of a fauna will, in the main, depend on the climate as well as on the environment; the literature is full of reference to ‘cold’ and ‘warm’ faunas, and also to faunas of ‘forest,’ ‘parkland,’ or ‘plains’ type ... Moreover, one cannot be absolutely certain that any particular species indicates a particular type of climate. ... When ‘warm-climate’ mammals are considered it is well to keep in mind the wide range of climate acceptable to such animals as the leopard and tiger, and to remember that even a hippopotamus can survive an English winter with a minimum of extra shelter and warmth. Except with the aid of a long series of fossils, even an approximation to the true date is all but impossible in the absence of such easily recognisable species as Equus robustus Pomel, Rhinoceros etruscus Falconer, Elephas meridionalis Nesti, or Rangifer tarandus (Linn.)"™ Geographical changes of this magnitude [uplift of Alps, etc.] might be expected to have influenced the climate and the fauna in some way or another, but this does not seem to have happened. Admittedly, much has been written about alternating ‘warm’ and ‘cold’ faunas, particularly in Germany, where the teachings of Penck were most influential, but in fact there is very little evidence to support the more extreme expositions of this view. . .2 It has been the practice among many European paleontologists to recognize three general mammalian faunas in the Pleistocene of Europe, corresponding roughly to the lower, middle, and upper phases of the period. Although, as de Lapparent has stated, the various individual animals constituting these faunas extend beyond the vague limits of each assemblage to mingle with each other, nevertheless associations of certain mammals are characteristic of the early, middle, and late portions of Pleistocene times. Thus he would recognize an early warm fauna, typified by Hlephas antiquus, a middle cool and humid period with Elephas primigenius and Rhinoceros tichorhinus, and finally a late cold fauna with the woolly mammoth and reindeer predominating. This was the general succession recognized by Osborn in his “‘Age of Mammals” in 1910, and described in a somewhat similar form recently (1935) by Boule and Piveteau.* While this conception of the succession of Pleistocene faunas in Kurope may be true in a general way, it is probably oversimplified. In the first place, it seems evident from recent work in various parts of the world that many if not most of the Pleistocene mammals appeared at an early stage in the history of the period, to continue with but little change through its extent. With the exception of certain rapidly developing types, such as the hominids and the proboscideans, there was probably but little evolution of a super-specific nature occurring within the relatively short duration of Pleistocene times. There were some extinctions, it is true, and these, as much as anything else, serve to distinguish the characters of the faunas in successive phases of the Ice Age. As regards this, it would appear that the evidence for extinctions at various times within the Pleistocene is certainly much more convincing in Eurasia than it is in the Americas. 1Hopwood, A. T., 1939. Proc. Prehist. Soc., N.S., V, Pt. 1, pp. 13, 14. *Hopwood, A. T., 1940. Proce. Geol. Assoc., LI, Pt. 1, p. 85. 3Boule, Marcellin, and Jean Piveteau, 1935. ‘Les Fossiles.”’ 1476 OSBORN: THE PROBOSCIDEA Hopwood has outlined the relationships of the appearance and disappearance within the Pleistocene in Europe as follows. Glacial IV.—Extinction of Hlephas primigenius, Rhinoceros tichorhinus, Hyena spelxa, Ursus speleus. Interglacial—Extinction of Elephas antiquus, Rhinoceros megarhinus, Felis leo spelexa. Glacial ITI.— Interglacial—First appearance of Bos primigenius, Rangifer tarandus, Rhinoceros tichorhinus, Ursus spelexus, Hyzxna spelxa. Glacial I].—Extinction of Hyena arvernensis, Canis nescherensis, Equus robustus, Equus stenonis, Rhi- noceros etruscus, Elephas meridionalis. Interglacial Glacial I.— These were some of the principal changes that took place during the development of the Pleistocene faunas in Europe, and it is mainly by looking at them that we can see the progress of Pleistocene history reflected in the assemblages of Ice Age mammals. Compared with changes such as these, the influx or egress of mammals adapted to warm or cold climates was of relatively slight importance. As Hopwood states': “Climatic variations may have favoured first one type and then the other, but neither type was completely expelled from the area.”’ The typical Basal or Lower Pleistocene proboscideans of Europe were Archidiskodon planifrons, Archidiskodon meridionalis, and Hesperoloxodon antiquus. Parelephas trogontherii and Elephas primigenius, perhaps not quite so ancient in their first appearance as the above mentioned forms, nevertheless were present in the European Pleis- tocene before the close of its lower phase. Seemingly there was an early extinction of the Archidiskodon group, specifically A. planifrons and A. meridionalis, probably in the interval marking the close of the first interglacial and the opening of the second glacial periods. Hesperoloxodon antiquus probably became extinct in middle or later Pleistocene times, perhaps near the end of the third interglacial period according to Hopwood. Finally, during or after the fourth glaciation there was an extinction of Hlephas primigenius. PROBOSCIDEANS: Mammonteus primigenius (Blumenbach) Mammonteus primigenius hydruntinus (Botti) Mammonteus primigenius fraasi (Dietrich) Mammonteus? primigenius leith-adamsi (Pohlig) Mammonteus primigenius astensis (Depéret and Mayet) Loxodonta cornaliae Aradas Hesperoloxodon antiquus (Falconer and Cautley) Hesperoloxodon antiquus germanicus (Stefanescu) Hesperoloxodon antiquus italicus Osborn Hesperoloxodon antiquus platyrhynchus (Graells) Hesperoloxodon antiquus ausonius (Major) Hesperoloxodon antiquus nanus (Acconci) Palzxoloxodon mnaidriensis (Adams) Palzoloxodon falconert (Busk) Palxoloxodon melitensis (Falconer) 1Hopwood, A. T., 1940, op. cit., p. 86. Palxoloxodon lamarmorae (Major) Palxoloxodon cypriotes (Bate) Palzxoloxodon creticus (Bate) Parelephas trogonthervt (Pohlig) Parelephas trogontherti nestii (Pohlig) Parelephas trogontherioides (Zuffardi) Parelephas intermedius (Jourdan) Parelephas wusti (Pavlow) Parelephas armeniacus (Falconer) Archidiskodon meridionalis (Nesti) Archidiskodon meridionalis cromerensis (Depéret and Mayet) Archidiskodon planifrons (Falconer and Cautley) Archidiskodon planifrons rumanus (Stefanescu) Mastodon pavlowt Osborn Anancus arvernensis (Croizet and Jobert) GEOLOGIC SUCCESSION: ASIA Vv. ASIA 1. INTRODUCTION Ast, as here used, comprises that portion of the continent north and west of the Himalaya Mountains and the Tibetan Plateau. Thus there is included within it Siberia, Mongolia, North China, Japan, and Turkestan. This separation of the more northerly and westerly sections of Asia from the Oriental Region is logical and natural, being based in large part on the modern zoogeographic realms. That part of Asia delimited above constitutes at the present time the eastern moiety of the Palearctic Realm, and the evidence would seem to be indicative of the fact that during the later stages of the Cenozoic, the Palearctic portion of Asia was zoogeographically distinct from the more southerly and easterly Oriental section, just as it is today. 2. THE MIOCENE OF MONGOLIA AND CENTRAL ASIA Moncouia: Lon Formation.—The Loh formation is exposed to the south of Uskuk Mountain in the Tsagan Nor basin of Mongolia, where it overlies the Upper Oligocene Hsanda Gol beds. The only fossils identified to date from the Loh formation are Serridentinus mongoliensis Osborn and Baluchitheritum mongoliensis Osborn. (This latter form is probably not Baluchitherium but rather a rhinoceros of the Coelodonta group.) From the evidence of these fossils, and due to the fact that the Loh rests directly on the Upper Oligocene Hsanda Gol, it would seem very probable that the Loh formation represents a Lower Miocene or Burdigalian phase in the sedimen- tary history of Mongolia. It might be pointed out here that the fossils of the Loh beds might be placed in later phases of the Miocene, particularly the Middle Miocene, with as much justification as in the Lower Miocene. Indeed, this series was designated as ‘‘Lower to Middle Miocene”’ by Professor Osborn in Volume I of this Monograph. Berkey and Morris! described the Loh formation as follows: Directly overlying the Hsanda Gol clays, about five miles south of Uskuk Mountain, there is a group of olive green clays less than one hundred feet thick. No clearly defined physical break can be seen between the two for- mations, but the upper olive clays yielded fossils which Dr. W. D. Matthew correlates with the Lower Miocene of Europe. PROBOSCIDEAN: Serridentinus mongoliensis Osborn. tSerridentinus Ocalientinus) orescens KHUNUK = Pisa OrCee aN GE Pe Eel 1S) OIC. E ANCE Serridentinus mongoliensis afavus MGI OuGsE NCE: Sa oncanere Trilophodon inopinatus cf angustidens GEOLOGICHL RELATIONSHIPS OF ASTATIC PROBOSCIDEX Mammonteus primigenius. ‘Palatoloxodon naumanni- Mamadicus. (oy setoensis-. namadicus yabei | Stegodon 2 --“orientalis,-: eo SINeNSI Ss. | Parelephas protomammonteus (EPaldtoloxodon)! Palseoloxodon: | tokunagai!-. Trilophodon.: sendaicus =~ Sernidentinus! : annectens " Trilophodon palaeindicus ‘Stegolophodon| lafidens Stegodon sp.2- DF kevett Bradley Tune Gur Formation.—The Tung Gur formation of Mongolia is typically exposed along a northeast to southwest trending escarpment in the immediate vicinity of Gur Tung Khara Usu, Inner Mongolia. This escarp- 1Berkey, C. P., and F. K. Morris, 1927. Natural History of Central Asia, II, p. 365. 1478 OSBORN: THE PROBOSCIDEA ment is part of a broad table-land situated to the northeast of the Kalgan-Urga trail and near the border between Inner and Outer Mongolia. The Tung Gur formation was described by Spock in 1929! and again by the same author in 1930.1. On the basis of geologic evidence alone Spock suggested that this formation might be of Pliocene age, which was the opinion first held by Professor Osborn as a result of his studies of Platybelodon, the most important and by far the most spectacular fossil mammal from this horizon. Soon after the Tung Gur fauna was discovered, however, P. Teilhard de Chardin suggested (verbally) that the fauna might be of Miocene age—or more specifically of pre-Pontian affinities. Teilhard’s conclusion was based upon his observations of the fossils as they were collected in the field. This correlation of the Tung Gur was subsequently verified by the detailed studies of the fauna by various authors including the present writer. Consequently, Professor Osborn came to regard the Tung Gur formation as of Upper Miocene age, thereby placing Platybelodon as an earlier stage of shovel-tusked mastodont than he had first considered it to be. (See Volume I, pages 463-466, of this Monograph.) The Tung Gur fauna is too large for detailed consideration or listing at this place. Some of the important forms constituting the fauna, however, may be briefly considered. As for the rodents, there is an Amblycastor closely related to Miocene species of the same genus occurring in North America. Among the carnivores, the giant canid or ursid, Hemicyon, may be linked with certain Miocene species referable to this genus, particularly as the Tung Gur form shows numerous relatively primitive, dog-like characters. There is a hyena of the Crocuta group, which might, on the other hand, perfectly well be of Pliocene age. The same is true of Platybelodon, since it shows many specialized characters. Perhaps one of the most significant of the Tung Gur animals is Anchitherium, definitely a Miocene type. The presence of this forest horse and the complete absence of any trace of Hipparion constitute strong evidence in favor of a Miocene age for the Tung Gur formation. The rhinoceroses, too, would seem to be of Upper Miocene affinities. A pig, Listriodon, on the whole shows affinities with Miocene species, although this is a genus that persisted from the Miocene into the Plocene in some localities. The deer, Stephanocemas and Lagomeryx, show definite relationships with comparable Upper Miocene forms from Europe and Asia. Of the bovids, one species, Ovoceras noverca is, according to Pilgrim, a small form that might have been ancestral to certain Pontian species. From this brief review it becomes evident that the Tung Gur fauna, although containing some possible Pliocene elements, is, in its general aspects, of Upper Miocene affinities, a conclusion that is greatly strengthened by the presence of Anchitherium and the absence of Hipparion in the assemblage, particularly since the absence of the latter form would seem to be real and not due to accidents of collecting. It would seem that the Tung Gur fauna, with a mixture of woodland and plains species, represents a borderland assemblage of mammals. PROBOSCIDEANS: Platybelodon grangeri (Osborn) Serridentinus gobiensis Osborn CENTRAL Asta: JILANCIK Breps, TurGAr Recion.—In a series of papers published between 1927 and the present time Borissiak has described a mammalian fauna from the Turgai region of Central Asia, discovered in the Jilancik (or Dschilantschik) beds of Miocene age. The Jilancik beds rest on a series of Oligocene deposits contain- ing Indricotherium, and in turn are succeeded by a Pliocene horizon in which Hipparion is present. At the ‘Spock, L. E., 1929. Amer. Mus. Novitates, No. 394; ibid, 1930, No. 407. GEOLOGIC SUCCESSION: ASIA 1479 present time the Jilancik fauna (excavated during a long period of years, beginning in 1914) would seem to be composed entirely of proboscideans and rhinoceroses, as follows. Trilophodon inopinatus Borissiak and Beliaeva Trilophodon cf. angustidens (Cuvier) Trilophodon atavus (Borissiak)! Brachypotherium aurelianense Nouel Aceratherium depereti Borissiak This fauna has a distinct Miocene aspect, and due to the determination of elements within it identical with or close to certain Lower Miocene species of Europe, Borissiak has referred the assemblage to the Burdigalian. With regard to the age of the Jilancik beds, the following remarks might be appropriate at this place. Trilophodon inopinatus Although Borissiak thought that this species was close to Serridentinus mongoliensis of the Loh forma- tion, Osborn placed it definitely in the genus T’rilophodon. Borissiak, as the result of his comparisons, considered the Loh formation to be virtually identical with the Jilancik beds in age. Osborn, on the other hand, compared Trilophodon inopinatus with the primitive Trilophodon cooperi, from the Bugti beds of Baluchistan. It is inter- esting to notice that both comparisons are with Lower Miocene horizons. This agrees with other evidence on the age of the Jilancik beds. Trilophodon angustidens If this identification is valid, the Jilancik beds may be correlated with the Lower Miocene or Burdigalian of Europe. Trilophodon atavus This species is very close to T. angustidens. This again is a link with the Lower Miocene of Europe. Brachypotherium aurelianense According to Borissiak, this characteristic European Burdigalian form is present in the Jilancik sediments. He has shown, however, that the Jilancik form displays certain advanced traits in the direction of the Middle Miocene B. brachypus of Europe. Aceratherium depereti Placed by Borissiak between A. lemanense and A. tetradactylum in its evolutionary stage of develo pment. This species shows certain specializations, notably the very much elongated nasals, which mark an advance beyond the typical Burdigalian forms. PROBOSCIDEANS: See faunal list, above. 3. THE MIOCENE OF NORTH CHINA Cuina.—The study of the fossil mammals of China dates from the middle of the last century, when Davidson published a short notice of some paleontological material gathered together in Shanghai by W. Lockhart. Among these first-known of the Chinese fossils was a tooth of an elephant. From that time until the present day a suc- This form has been recently described by Borissiak, under the name of Mastodon atavus. It is closely related to T'rilophodon angustidens, as Borissiak points out; consequently it is here referred to this latter genus. 1480 OSBORN: THE PROBOSCIDEA cession of students, notably Owen, Gaudry, Koken, Schlosser, and in recent years the various authors describing the collections made by the Geological Survey of China, the Paleontological Institute of Upsala University, and the American Museum of Natural History, have added increasingly important and voluminous data concerning the paleontological history of northern China and Mongolia. The history of Upper Cenozoic mammals in North China would seem to begin with the Middle or Upper Miocene and to continue almost uninterruptedly through the Pliocene into the Pleistocene. Through this stretch of geologic time two periods are marked by the expansion and the unusual abundance of fossil mammals. These are the Lower Pliocene or Pontian, in which the large, widely distributed ‘“Hipparion fauna” occurs, and the Lower and Middle Pleistocene, in which there are several faunas, notably the ““Hquus fauna” or Villafranchian assem- blages of the flood-plain deposits and the later Choukoutien cave faunas. As to the beds preceding the almost universal Hipparion clays, there is little to be said. At best, these sediments are physically but slightly differentiated from the Pontian deposits, if they are at all distinguishable. There have been described, from time to time, certain fossils that would seem to be of definitely Upper Miocene rather than of Pontian affinities, and it is on the basis of these discoveries, as much as anything, that the presence of pre-Pontian sediments in North China is inferred. Such is the case of Trilophodon connexus, which Hopwood regards as a very primitive stage in the buno- dont mastodonts, closely comparable to the Miocene Trilophodon coopert of Baluchistan. Trilophodon cooperi is a Lower Miocene species; whether, on the basis of this, T’rzlophodon connexus should be regarded as a very early Miocene form, or rather a structurally primitive species persisting into the Upper Miocene, is a question open to some doubt. It is possible that this species may represent a Sarmatian or an equivalent age in North China. On the other hand, there is no reason why the supposed pre-Pontian species in North China might not be structurally primitive forms persisting into the Lower Pliocene. In this connection it might be noted that Trilophodon wimani, considered by Hopwood as possibly of Sarmatian age, was placed by Teilhard in the Pontian. Hopwood based his conclusions on the primitive structure of Trilophodon wimani, and also on the fact that it was associated with Listriodon gigas, a pig showing affini- ties, according to Miss Pearson, with certain Miocene forms of Europe. Yet Teilhard’s conclusions as to the age of Trilophodon wimani were based on his thorough knowledge of the stratigraphic relationships of the Cenozoic deposits of China, and therefore they carry great weight. PROBOSCIDEANS: Trilophodon connexus Hopwood Trilophodon wimani Hopwood (Pontian, according to Teilhard) Andersson,! in his discussion of the Cenozoic of North China, describes the Lu Tzu Kow beds, exposed in the Pao Te Hsien area of Shansi. According to Andersson these beds, discovered by Zdansky, are of Upper Cenozoic age, but they underlie the Hipparion clay. Yet although the Lu Tzu Kou beds are exposed beneath the Hip- parion beds, there is a strong possibility that the age difference between the two series is not great. As the Hipparion beds represent the transition from Miocene to Pliocene, the Lu Tztt Kou beds could eventually be sup- posed to represent the Miocene, but there are some facts at hand which indicate that they are in age nearly related to the Hipparion beds. 1Andersson, J. G., 1923. Mem. Geol. Surv. China, Ser. A, No. 3, p. 107. GEOLOGIC SUCCESSION: ASIA 1481 4, THE PLIOCENE OF MONGOLIA KuuNUK Formation.—The Khunuk formation was described by Granger! as follows: In the Kholobolchi Nor region of Mongolia there are three areas of late deposits considered, in the field, as being of Pleis- tocene age; the geologists were of the opinion that these three exposures were of the same age, and the name ‘Khunuk’ was given to the formation. The three exposures mentioned in the Kholobolchi Nor region are respectively north, northeast, and east of the lake; the type of Serridentinus florescens comes from the northern exposure of the ‘Khunuk’ and represents the only specimen obtained in that particular locality. As Osborn has shown, it is possible that the Khunuk formation is approximately correlative with the Hung Kureh beds of the Tsagan Nor region, which latter deposits contain a small mammalian fauna of Pliocene re- lationships. The single mastodont described from the Khunuk formation is an advanced type of serridentine, which would favor the assigning of a Pliocene age to these sediments. It is not possible to say at what stage in the Pliocene the Khunuk and Hung Kureh formations should be placed, but it is very probable that they are in the main Pontian equivalents, representing a phase in the sedi- mentary history of Mongolia immediately subsequent to the period during which the very fossiliferous Tung Gur Upper Miocene sediments were deposited. PROBOSCIDEAN: Ocalientinus (Serridentinus) florescens (Osborn). 5. THE PLIOCENE OF NORTH CHINA LOWER PLIOCENE The Hipparion clays of North China form a mantle covering extensive areas of valley and plateau country, dissected by streams. The localities at which fossils have been discovered in the Hipparion clays are so numerous that they cannot be listed here, nor can comparisons between them be attempted. It might be well, however, to quote Andersson’s? description of the Hipparion clays at Chi Chia Kou in Pao Te Hsien, one of the most richly fossiliferous and best-known localities in this horizon. The Chi Chia Kou region is a plateau land dissected by an intricate system of ravines, in the bottom of which the sub- stratum of the Hipparion clay is almost everywhere visible. The basement rock is formed by the Carboniferous coal series in nearly horizontal beds. The basal layer of the Hipparion series is a conglomerate bed, at most 4 meters thick with gray matrix. Above this basal conglomerate rests the red Hipparion clay with a maximum thickness of 65 meters. In the clay there are inter- bedded gravel beds of little persistence and occasionally also lenses of sand. In certain horizons there are also irregular lime con- cretions in the clay. Round the fossil mammal bones are seen infiltrations of lime. In the 65 meters of red, mostly entirely barren clay there is a well defined bone-carrying horizon 25 meters above the bottom and 35 meters underneath the top of the deposit. The bone layer is mostly less than a meter thick and so nearly horizontal that there is not more than 5 meters variation of altitude of the bone bed throughout the whole Chi Chia Kou area. In the bone horizon there are not bones everywhere, but rather pockets or nests rich in bones separated by some meters of barren clay. . . According to Dr. Wiman and Dr. Zdansky the Hipparion fauna indicates steppe conditions, but the occurrence of Giraffine and Suid points to the existence of groups of trees and of water pools spread over this steppe. As the clays are mostly barren over wide areas and rich in mammals only in the three areas mentioned, it might be inferred that in the otherwise very dry steppe there were locally oases with trees, water pools and occasionally also sheet floods after the rains. It seems as if these genial conditions prevailed only during a short, well defined period. 1Granger, Walter, in H. F. Osborn, 1936. Volume I of this Monograph, p. 397. “Andersson, J. G., 1923, op. cit., pp. 107, 108, 110. 1482 OSBORN: THE PROBOSCIDEA PROBOSCIDEANS: (Pontian of China) Serridentinus wimani (Hopwood) Serridentinus lydekkeri (Schlosser) Trilophodon spectabilis Hopwood Tetralophodon exoletus Hopwood Tetralophodon sinensis (Koken) (This horizon?) Pentalophodon sinensis Hopwood = Anancus sinensis (fide Osborn) Pentalophodon cuneatus Teilhard and Trassaert Zygolophodon borsont (Hays) (This horizon?) Mastodon americanus (Kerr) (This genus and species?) ‘“‘Mastodon”’ sp. Mastodon intermedius Teilhard and Trassaert Stegodon licentt Teilhard and Trassaert Stegodon zdanskyi Hopwood (This horizon?) MIDDLE PLIOCENE Only within recent years has the Middle Pliocene been recognized in North China. The distinction of this stage has been difficult because it is lithologically closely related to the typical Pontian deposits, and particularly because the fauna is not well known. In their important contribution entitled ‘“The Pliocene Lacustrine Series in Central Shansi,’”’ Licent and Trassaert! describe sediments overlying the Pontian beds but underneath the San- menian or Lower Pleistocene. These authors have shown that the lacustrine sediments of Central Shansi occupy an extensive, shallow Permo-Triassic syneline. The beds representative of the Upper Cenozoic are 100 meters or more in thickness and entirely of a freshwater type. They are characterized by the “extremely deceptive recurrency of the same facies: rusty or reddish sands, green marls, etc.,’’ thereby making the differentiation of the several horizons con- tained within the series extraordinarily difficult. Nevertheless, Licent and Trassaert recognize three zones within these lacustrine sediments. The lowest zone, designated as Zone 1, is of Pontian affinities. Above it is Zone 2, of Middle Pliocene age, succeeded by Zone 3, representing the Sanmenian. In addition it is pointed out that there is very likely a fourth zone, coming between zones 2 and 3, representative of a stage between the Middle Pliocene and the Sanmenian. Zone 2. In the following zone 2, better studied in the Changtsun basin, the deposits become less coarse, and a typical lacustrine condition is prevailing: green and bluish marls, containing many bird, turtle, fish-remains, freshwater shells (Lym- nea, Planorbis, thin-shelled Unionidz) and carbonised plant-remains. A small Hipparion, the Chilotherium, a tapiroid Mastodon and the Stegodon are still present (as in zone 1). But a remarkable type of strepsiceros Antilope (ef. Antilospira T. & Y.) seems to appear for the first time, and also a special Castor (characteristic of the Ertemte fauna of Mongolia) Dipoides majori Schl. A middle Pliocene age seems to be indicated. Suggestion of a fourth sedimentary zone. The palzontological analysis of the fauna collected in the Yiinchu basin suggests that, between zone 2 and zone 8 a fourth horizon might be eventually recognised in the Pliocene deposits. First, collected by country people, from such localities as Malan, we have, embedded in a characteristic matrix of a dark red hue, canon-bones of a big Hipparion, already associated with a Bison, but without any sure trace of Equus. Furthermore, amongst the best fossils purchased in the area, are several teeth of a thick lamelled Elephant (Hlephas ef. planifrons), a form never observed in Nihowan, and yet too much advanced for being conveniently referred to the Mastodon-Stegodon fauna of zone 2. It would seem therefore that the lower part of zone 3 might have to be separated as a special unit, distinguishable from the true Nihowan beds by the absence of Horse and the presence of archaic Elephant. This point however is not yet supported by clear stratigraphical evidences. 1Licent, E., and M. Trassaert, 1935. Bull. Geol. Soc. China, XIV, No. 2, pp. 214, 216. GEOLOGIC SUCCESSION: ASIA 1483 UPPER PLIOCENE (?) Teilhard and Trassaert,! in 1937, divided the Pliocene (as they considered the extent of its upper and lower limits) into three ‘‘zones,”’ namely: (1) A lower zone comprising the Pontian Hipparion beds, (2) a middle zone of Pliocene age but subsequent to the Pontian, and (3) an upper zone comprising the Villafranchian, which these authors placed in the Upper Pliocene. In addition to the characteristic ‘Middle Pliocene,” as recognized by Teilhard and Trassaert, and as described by Licent and Trassaert (see above), there would seem to be an upper zone or subzone which is immediately antecedent to the base of the Villafranchian. Since the Villafranchian is considered as Lower Pleistocene (as will be shown below) it would seem logical to suppose that this upper portion of “Zone II’ represents the uppermost Pliocene in North China. Teilhard and Trassaert made the following comments concerning it: But, near Chinglo (N. Shansi, cf. Teilhard and Young, 1931, p. 52, fig. 15), in an horizon representing clearly Zone II (rather large Deer and Hipparion, Antilospira, no Equus . .), Teilhard and Young have collected several years ago an isolated lamella of D; decidedly referable to a primitive Elephant, and not toa Stegodont. An upper horizon (planifrons subzone) might therefore have to be recognised some day at the top of Zone II. ProBosciDEANS: (Middle and Upper Pliocene of China) Zygolophodon borsoni (Hays) Mastodon intermedius Teilhard and Trassaert Stegodon zdanskyt Hopwood Stegodon orientalis Owen (This zone?) Archidiskodon planifrons (Falconer and Cautley) (Uppermost Pliocene) 6. THE PLEISTOCENE OF NORTH CHINA The Pleistocene of China is of great importance, not only because it has yielded large and rich mammalian faunas that are directly ancestral to the modern faunas of northeastern Asia, but also because it has produced some of the most complete and the most significant remains of fossil man. For this reason, a rather complete and detailed discussion of the Quaternary beds of China will be necessary. Tue VILLAFRANCHIAN OF CuINA.—The Pleistocene succession of North China may be regarded as beginning with the Villafranchian deposits of Nihowan, and also possibly of Yushé. This assignment of the Villafranchian of China to the beginning of the Pleistocene is not entirely in accord with the views of Teilhard,? who recognizes the fact that there is a growing sentiment among vertebrate paleontologists to regard the appearance of Hquus, elephants, and cattle as truly indicative of the beginning of the Pleistocene throughout the world, but who feels that in China it is most convenient to end the Tertiary sequence with the diastrophic movements that resulted in the cutting of the gorges, subsequent to the deposition of the typical Villafranchian sediments. Consequently, he would regard the accumulation of the Choukoutien deposits and the first appearance of man as indicative of the beginning of the Pleistocene in North China. In North China, the major period of gorges cutting (Fenho stage) first placed by Bailey Willis at the base of the Malan loess, and later transferred by Barbour and myself to a pre-Nihowan stage (2), would find its true place in an intermediate position, namely between the Sanmenian (Nihowan) and the Choukoutien stages of deposition. . . . 1Teilhard de Chardin, P., and M. Trassaert, 1937. Palaeontologia Sinica, Ser. C, XIII, Fase. 1, p. 53. °*Teilhard de Chardin, P., 1937. Bull. Geol. Soc. China, XVII, No. 2, pp. 173-175. 1484 OSBORN: THE PROBOSCIDEA In North China, again, (just as in France and also in North India), the first appearance of horse (more and more generally aecepted by the paleontologists as indicating the base of the Pleistocene in the Old World) antecedes clearly the major diastro- phisms which would be the best limit between the Tertiary and the Quaternary from a geological point of view. Therefore, the advantage of keeping the Villafranchian (first ‘Hquus-beds’) in the Pliocene, instead of referring it to the Pleistocene, should be re-considered. So far as North China is concerned, the most natural base of the Quaternary is given by the appearance of Man on a modernized topography, after the Villafranchian. Yusn&.—According to Teilhard,! the basal portion of the Villafranchian of China is to be found in the Yushé basin, and consequently he has distinguished this part of the Pleistocene sequence as a separate subzone within the Villafranchian. Strangely enough, no Archidiskodon has been recorded up to now from the typical Villafranchian formation of Nihowan. In order to explain this difference (if it really does exist), we may suppose that the Nihowan beds represent only the top of the Villafranchian (‘namadicus subzone?’), the base of the Villafranchian being on the contrary only present in Yushé, and characterized perhaps there by the latest Archidiskodon (‘tokunagai subzone’?).. . In Yushé, as well as in Nihowan, the Stegodonts seem to have disappeared before the beginning of the Villafranchian. Yet, some new finds made in S. Shansi suggest that the group (represented by Stegodon orientalis) has lasted as far up as the Lower Pleistocene inclusively along the northern border of the Tsinling, as it did in Central and Southern China (Szechuan, etc.). This definition of a basal zone of the Villafranchian would seem to be in accord with the evidence put forward by Licent and Trassaert for a distinct zone between the Middle Pliocene and the Villafranchian of Central Shansi. Since the relationships of this distinct subzone or horizon in Yushé is with the overlying Villafranchian, rather than with the underlying Pliocene sediments, it would seem best to place it as the first member of the Pleistocene group of sediments in North China. PROBOSCIDEANS: Palzxoloxodon tokunagai (Matsumoto) Palxoloxodon namadicus (Falconer and Cautley) NIHOWAN (SANMENIAN).—The true Villafranchian of North China is best exemplified by the sediments and their contained fauna as exposed in Nihowan. Teilhard? has shown that originally only two distinct phases were recognized in the Upper Cenozoic of North China, namely, the Pontian Hipparion beds and the Pleistocene loess. Subsequently, as the result of more detailed work in the North China area, a series of post-Pontian, pre-Loess deposits came to light, occupying the interval between the levels originally known. But then again the first impression of the geologists was that they had to deal with a single polymorphous complex. On one hand, no internal disconformities were observed, at that time, in the newly recognised sedimentary block. And, on the other hand, the common occurrence in all these ‘post-Pontian and pre-Loessic’ deposits of closely related types of Horse, Hy- aena, Rhinoceros, Rodents and Mollusca suggested for all of them a same general age. Thus was borne the conception of the ‘Sanmenian,’ a single stratigraphical and faunistical unit, rather vaguely referred either to the Late Pliocene, or to the Lower Pleistocene. As Teilhard has shown, there is a definite faunal and physiographic break between the Nihowan and Chou- koutien phases, hitherto included in the Sanmenian. Consequently, he has felt it necessary to limit the term Sanmenian to the Nihowan deposits and their included fauna, thereby making the Choukoutien formation definitely post-Sanmenian in its relationships. In accordance with his views as to the Upper Pliocene age of the Villafranchian, Teilhard has placed the Sanmenian as the latest part of the Pliocene, relating it to the lacustrine Middle Pliocene sediments of North China, and separating it quite definitely from the overlying Choukoutien phase. It is proposed here to regard the Sanmenian as a lower phase of the Pleistocene sequence in China (preced- ed perhaps by the basal Villafranchian beds of Yushé) and related to the Choukoutien deposits. Of course, in 'Teilhard de Chardin, P., and M. Trassaert, 1937. Palaeontologia Sinica, Ser. C, XIII, Fase. 1, p. 53. *Teilhard de Chardin, P., 1937. Bull. Geol. Soc. China, XVII, No. 2, p. 170. GEOLOGIC SUCCESSION: ASIA 1485 following this line of procedure, the weight of the argument is being given to the faunal evidence, as based on the appearance of new and advanced Pleistocene genera, rather than to the physiographic evidence cited by Teilhard, as based on diastrophism and the rejuvenation of the streams. It all comes down to a choice between two lines of evidence which do not coincide, as shown by the following diagram. FauNnaL EvIDENCE PHYSIOGRAPHIC KvIDENCE Malan Loess Choukoutien | Pleistocene Break = (Gorge Cutting) Pleistocene Sanmenian (Nihowan) Li Break (Influx of advanced fauna) Pliocene Pliocene Pliocene Teilhard’s' latest views as to the Nihowan fauna have been expressed as follows: In spite of the fact that some important physiographic changes have to be supposed (and have in fact been partly traced) in Shansi between the ‘Hquus-Camelus beds’ of Nihowan and their underlying Middle Pliocene beds (containing neither Horse nor Camel), it remains that both the Nihowan and the pre-Nihowan (Middle Pliocene) series have in common decided litho- logical and faunistical affinities. Both are dominantly lacustrine in facies, and both are characterized by a same ‘sub-tropical Asiatic’ fauna. ... What I mean by this latter expression will only be clear when the description of the Yushé fauna has been given in Palzontologia Sinica. Anticipating however on the publication of these memoirs, I can already say that, from the material lately collected in SE Shansi, it appears now that, at the middle of the Pliocene, two different faunistical blocks were mixing in the forested steppes of North China: (1) A block of south-eastern affinities: abundance of Cervulids (several extinct genera of Munjack deer parallelizing in some way the living south-American deer), Rusa and Axis, associated with Stegodon. (2) A peculiar Central-Asiatic block, chiefly characterized by a harvest of beautiful, mostly strepsicere, antelopes (more than five new genera recognized so far). This was apparently a special ‘Antelopes province,’ parallelizing in many respects the modern African province, but separated from it since the base of the Pliocene. Now the important point bearing on our present subject is that, on the whole, the two above mentioned Cervine and Antilopine blocks were still thriving in North China after the appearance of such forms which (as horse, camel, bison, Huclado- ceros) characterize the Villafranchian of Europe. In spite of many important differences in their respective faunas (disap- pearance of the Stegodon, Aceratherids, Giraffids, etc.) (cf. 2), the Middle Pliocene and the Villafranchian (Nihowan) of North China are marked by the same general topography (a rather mature surface), the same climate (rather warm), and the same fundamental types of deer and antelopes. PROBOSCIDEAN: Palzxoloxodon namadicus (Falconer and Cautley). CHoUKOUTIEN.—The post-Villafranchian, pre-Loess Pleistocene deposits of North China have become of extreme importance within the last few years because of the discovery of the extinct man, Sinanthropus, at Choukoutien, southwest of Peking. The Choukoutien deposits were discovered by J. G. Andersson some twenty years ago, and as the result of many years of excavation a large and varied mammalian fauna has been taken out of these beds and described. The Choukoutien sediments are thick fillings in limestone caves or fissures, and as such need no stratigraphic description at this place. 'Teilhard de Chardin, P., 1937, op. cit., pp. 171, 172. 1486 OSBORN: THE PROBOSCIDEA Various authors have described the several groups of mammals constituting the Choukoutien fauna, so that the detailed information as to this mammalian assemblage is voluminous, to say the least. Recently Teilhard has summarized the Choukoutien mammalian fauna by showing that from the standpoint of its geological significance the following classification may be formulated. 1. Forms that were present in the Sanmenian of Nihowan (Villafranchian), but which did not persist into Choukoutien times. 2. Nihowan forms present in the Choukoutien deposits, but absent in the subsequent Malan Loess. 3. Loess forms present in Choukoutien, but absent in Nihowan. 4. Loess forms absent from Choukoutien. 5. Forms that were present in all three formations. The distribution of the significant mammalian types in the three formations enumerated above may be shown graphically in the following manner. VILLAFRANCHIAN LATE PLEISTOCENE Sanmen-Nihowan Choukoutien Malan Loess Mustela pachygnatha— Elasmotheriwum——~ Postschizother‘'um—— Hipparion—————— Eucladoceros————_—> Rusa Ochotonoides Stphneus tingi———— Machairodus Machatrodus Hyzxna cf. sinensis—————>__ Hyena cf. sinensis Equus sanmeniensis—————>_ Equus sanmeniensis Rhinoceros cf. mereckit———>_ Rhinoceros mercki Ovibovids > Ovibovids Nyctereutes > Nyctereutes ——————————> WNyctereutes Meles — Meles > Meles Camelus———————————_ Ca melus > Camelus Gazella————_————————_ Gaazella > Gazella Spirocerus > Spirocerus — Sptirocerus Bison — Bison > Bison Ovis + Ovis > Ovis Cyon + Cyon Siphneus cf. fontanieri > Siphneus ef. fontanieri Euryceros > Huryceros Bubalus———————————_ Bubalus Hyena spelxa Equus hemionus Cervus elaphus Bos primigenius GEOLOGIC SUCCESSION: ASIA 1487 From this it will be seen that the Choukoutien fauna is truly intermediate between the Villafranchian fauna of Nihowan and the late Pleistocene fauna of the Malan Loess. As to the relationships of the Choukoutien mam- malian assemblage to the faunas either above or below it, Teilhard and Young! made the following statements. Taking the facts more in detail, the Choukoutien types in many cases are distinctly different from their corresponding Nihowan or Sjara-osso-gol forms, being less primitive than the former, and less advanced to some extent than the latter. For example: Hyaena cf. sinensis, Rhinoceros cf. tichorhinus, Spirocerus wongi, Bison palaeosinensis, Ovis shantungensis, from Nihowan are more primitive than Hyaena sinensis, R. tichorhinus, Spirocerus peti, Bison sp., Ovis cf. ammon from Choukoutien. On the contrary, Hyaena sinensis, Euryceros pachyosteus, Bubalus teilhardi from Choukoutien can be held as more archaic than Hyaena spelaea, Euryceros ordosianus, Bubalus wansjocki from the Loess. The conclusion is obvious. In full accordance with the stratigraphical facts which will be set forth subsequently below in section 5, the Choukoutien fauna fits so exactly between the Late Pliocene (Nihowan) and the Upper Pleistocene (Sjara-osso- gol) ages that it is not easy to decide to which of them it stands more closely related. The Choukoutien formation has conse- quently broadly to be considered as of Lower Pleistocene age. In order to account for the change between the Nihowan and the Choukoutien faunas, a long period of time must be allowed during which some climatic changes most probably took place. The fauna is still dominantly of a palaearctic type, but with a clearer tendency to break into a special east Asiatic type. Some southern migration, possibly along the sea coast, may be sug- gested by the appearance of the Bubalus. Recently Teilhard has placed the Choukoutien deposits in the Middle Pleistocene, as a result of field work and correlative studies made throughout southeastern Asia. By following this procedure, the Choukoutien deposits are brought into line with the cave deposits of southern China and Burma, described in a preceding section of this chapter. On the basis of present evidence, this would seem to be the most logical correlation for the Choukoutien beds. There seems to be but little doubt that there was a widely spread Lower Pleistocene fauna in Asia and the Orient (Upper Siwalik, Upper Irrawaddy, Djetis, Nihowan) followed by an equally extensive Middle Pleistocene complex, which upon the mainland is commonly preserved in caves (Mogok, Hoshangtun, Trinil, Kwangsi, Szechwan, Choukoutien). Tue Logess oF NortH Cuina.—At the top of the Cenozoic succession in North China is the Loess, a character- istic and wide-spread deposit forming a prominent and easily-recognized capping layer over a large area north of the Yellow River. Fossils of an advanced type have been found in the Loess, and for this reason the deposit is generally regarded as of Upper Pleistocene age. The following remarks were made by Andersson,? in 1923, re- garding the mammalian and avian fauna of the North China Loess. The most common mammal remains in the loess are tusks and molars of an Elephas, which according to Dr. Zdansky may possibly be Hlephas namadicus, which was originally found in the Pleistocene alluvium of the Narbada valley of India, where it occurs together with two species of Hippopotamus, and several other mammals. The reappearance of this Indian species in the North China loess would be very surprising, especially in view of Richthofen’s eolian theory. However, Dr. Zdansky’s pro- visional determination has to be tested by a much closer study of the specimens. At any rate, it is beyond doubt that the com- mon loess elephant is not the Mammoth as has formerly been suggested. Among the numerous isolated finds which have been made in loess-like material the following species can with fair safety be assigned to the loess: Rhinoceros affinis simus, Ovis? sp., Hyena sp., Ursus sp. A skull of a Castorid also was obtained from undoubted loess, and this is another find which tends to weaken Richthofen’s eolian theory. The same applies to a recently found Sus sp. Only in a single case, have we ever come across what deserves the name of a small bone accumulation in the loess. This was in SE Shansi, in Yuan Chii Hsien, and the locality has been studied by Dr. Zdansky who has communicated the section fig. 39. In this place were found: Hyena, Equus, Cervus and a turtle. The deer, a forest animal, and the turtle, a water animal, are further finds which hardly agree with Richthofen’s eolian theory. Lastly we have to mention a fossil which seems to conform better with the said theory, namely the egg shells of an Ostrich, a bird described under the name Struthiolithus. .. The modern ostrich is a steppe bird, and the mere mode of occurrence of these unbroken shells, often two or more together, seems to indicate that the nests were occasionally covered by a wind-drift deposit which prevented the eggs from maturing and preserved the shells in an unbroken state. Earlier authors attributed a great thickness to the loess deposits of North China—in some cases as much as 1,500 feet. According to Andersson, however, the loess never exceeds a thickness of more than 50 or 60 meters. 1Teilhard de Chardin, P., and C. C. Young, 1933. (In Black, Teilhard, Young and Pei), Mem. Geol. Surv. China, Ser. A, No. 11, pp. 48, 49. Andersson, J. G., 1923, op. cit., p. 127. 1488 OSBORN: THE PROBOSCIDEA Andersson’s revised figures for the thickness of the loess are based on the fact that he differentiated the older beds, which the earlier students had confused with the loess and included in it. The loess of North China, named the Malan, has been divided by Andersson into two beds or horizons. The lower deposit he has called the primary loess or the true loess. The upper bed is the secondary loess or the re- deposited loess. As is indicated by the name, this secondary loess is merely material from the primary loess, reworked in very late Pleistocene times. The redeposited loess, because of the taxonomic relationships and the slight mineralization of its contained fossils, has been defined by Andersson as a post-loessic deposit. Thus it may be considered as an horizon bridging the time between the latest Pleistocene and the beginning of Recent conditions in North China. Lately Teilhard and Young have thrown new light on the problem of the post-loessic faunas of North China in their study entitled, “On the Mammalian Remains from the Archeological Site of Anyang.” These authors show that in the ancient city of Anyang, capital of the Shang dynasty during the period between 1400 B.C. and 1100 B.C., there were numerous mammals similar to the post-loessic fossils found in other parts of China. In their analysis of the Anyang fauna, Teilhard and Young list three faunal divisions, namely: 1. The wild indigenous animals, such as the raccoon-dog, bear, badger, tiger, bamboo-rat, hare, water-deer and the sika. 2. The domesticated animals, including the pig, dog, sheep, goat, cattle, water-buffalo, and the macaque. 3. Imported mammals, the whale (probably bones collected along the seacoast), the elephant, tapir, and a small bear. The presence of the elephant, which has been identified as Hlephas indicus, in North China is interesting, to say the least. The evidence would seem to show, however, that this elephant was never wild here, but rather was imported (probably in the form of tribute) by the ancient peoples of this old city. As Teilhard and Young! have remarked, ‘‘Such beds [the redeposited loess and similar deposits] are still, geologically, a ‘no man’s land’; more attention should be paid in [the] future to the redeposited loess, to peat-bogs and to subrecent cave deposits.” The difficulty in dealing with these redeposited loess and post-loessic beds is that they really do bridge the period between the Pleistocene and the present day. Consequently such beds are defined only in a most provision- almanner. As Teilhard and Young have shown, if Rhinoceros, Elephas namadicus or Elephas primigenius, Hyena, and Huryceros are present, the deposits may be called Pleistocene with some degree of assurance. On the other hand, there is no reason why some of these forms may not persist beyond the closing of the Pleistocene period— whenever that might have been. Therefore the problem resolves itself into a question of drawing an artificial line across a series of continuous events which are so close to us that it is not possible for us to gain a true perspec- tive as to their importance or their interrelationships. PROBOSCIDEANS: Palzxoloxodon namadicus (Falconer and Cautley) Loess EKlephas primigenius Blumenbach Upper Pleistocene, North China, Mongolia Elephas indicus Linn. Sub-Recent, Anyang 'Teilhard, P., and C. C. Young, 1936. Palaeontologia Sinica, Ser. C, XII, Fase. 1, p. 57. LIST OF THE FOSSIL LAND MAMMALS OF JAPAN AND KOREA Speciric NAMEs! LOCALITY PRIMATES Macacus fuscatus Bly RopENTIA Sciurid Lepus brachyurus brachyurus Tem. CARNIVORA Meles anakuma Tem. PROBOSCIDEA Hemimastodon annectens Mat. (Serridentinus) *Prostegodon latidens (Clift) (Stegolophodon) Trilophodon sendaicus Mat. (Serridentinus) *Trilophodon palxindicus Lyd. Parastegodon aurorae Mat. (Stegodon) *Stegodon bombifrons Fale. and Caut. *Stegodon clifti Fale. and Caut. *Stegodon orientalis Owen *Stegodon sinensis Owen Stegodon sp. Parelephas protomammonteus Mat. (Palxoloxodon) Parelephas trogontherii Pohlig Loxodonta tokunagai Mat. (Palzoloxodon) Loxodonta namadica naumanni Mak. 1924 (Palxoloxodon) Elephas namadicus setoensis Mak. 1929 (Palxoloxodon) Loxodonta namadica yabei Mat. Elephas indicus buski Mat.(Palxoloxodon) Mastodon sp. PERISSODACTYLA Anchitherium hypohippoides Mat. *Equus caballus Linné Palzxotapirus yagi Mat. Teleoceras pugnator Mat. Rhinoceros (Teleoceras) tokiensis Tok. Rhinoceros (Teleoceras?) kaniensis Tok. Rhinoceros (Aceratherium?) watanabei Tok. Rhinoceros (Acerathertum?) maki Tok. Rhinoceros sp. Rhinoceros shindoi Tok. Rhinoceros koreanicus Tok. Rhinoceros sp. Sus nipponicus Mat. Sus leucomystax Tem. Amphitragulus minoensis Mat. Dicrocerus tokunagai Mat. Cervavus oweni hirabayashit Tok. Cervus nippon nippon Tem. Cervus kazusensis Mat. Cervus riukiuensis Mat. *Cervus cfr. elaphus Linné Cervus sp. Cervus sp. Cervus sp. *Hlaphurus davidianus M—Edwards Muntiacus astylodon Mat. Giraffa nipponica Mat. Bubalina sp. *Bison occidentalis Lucas *Bibos geron Mat. *Common to Japan and continental Asia. 1The proboscidean names in parentheses are the generic references used by Professor Osborn. Kyushu (Buzen) Honshu (Mino) Kyushu (Buzen) Honshu (Shimotsuke), Kyushu (Buzen) Honshu (Mino) Honshu (Hitachi, Rikuzen) Honshu (Rikuzen) Honshu (Mino) Honshu (Kaga) Honshu (Kazusa) Honshu (Ise) Honshu (Kazusa, Shimotsuke, Echigo, Suo, Nagasaki), Inland Sea Inland Sea Kyushu (Hyuga) Honshu (Kazusa) Honshu (Kazusa, Mikawa, Omi, Shinano), Inland Sea. Honshu (Etchu) Honshu (Musashi, Totomi, Sagami, Hitachi, Kazusa, Mikawa, Shimosa) Inland Sea Honshu (Ugo, Noto), Inland Sea, Shikoku, Hokaido (Ishikari) Honshu (Musashi, Mino, Mutsu) Korea (Joshin) Honshu (Mino) Honshu (Hyogo, Korea (Keikido) Honshu (Mino) Honshu (Mino) Honshu (Mino) Honshu (Mino) Honshu (Suo) Korea (Kokaido) Honshu (Suo) Honshu (Shimotsuke), Kyushu (Buzen) Korea (Keikido) Taiwan (Taikei) Honshu (Ugo) Honshu (Shimotsuke), Kyushu (Buzen) Honshu (Mino) Honshu (Iwaki) Honshu (Shinano) Honshu (Musashi, Shimotsuke, Ugo), Kyushu (Buzen), Inland Sea Honshu (Kazusa) Loochoo Is. Korea (Heian-nando) Honshu (Shimoosa) Honshu (Totomi) Honshu (Sagami) Honshu (Kazusa) Loochoo Is. Honshu (Kazusa) Honshu (Omi) Inland Sea Inland Sea 1489 Miocene xX ~ xXXKKKKXK X xX X Pliocene XXX X Pleistocene x x X x Sek 282K XX XK KX xX X XK KX XK OG 1490 OSBORN: THE PROBOSCIDEA 7. THE MIOCENE TO PLEISTOCENE OF JAPAN Japan.—The fossil mammals of Japan, though for the most part very poorly preserved, have been found at numerous localities, showing that Cenozoic continental deposits, particularly those of the Upper Cenozoic, are widely distributed throughout the islands. Most of the mammal-bearing localities are on the island of Honshu, and a majority of these deposits are of Pleistocene age. In all of the Japanese deposits, whether of late Tertiary or of Pleistocene age, the proboscideans are the dominant elements in the faunas. The evidence of the fossil mammals would seem to indicate that Japan was connected with Asia during Miocene, Pliocene, and early Pleistocene times. Certain elements of the Miocene fauna of Asia reached Japan, and in subsequent times there was likewise a constant infiltration of new migrants from the mainland. This influx from Asia into Japan reached its height seemingly during the early part of the Pleistocene, for at that time numerous proboscideans as well as certain other forms found in Japan were specifically identical with mainland types. An interesting feature of the Japanese faunas is the dominance of the proboscideans, especially during Pleisto- cene times. Whether this dominance of the Proboscidea is real or illusory is a question difficult to decide on the basis of extant evidence. That is, there may be many proboscideans in the Japanese deposits because these fossils are large and more apt to be preserved than are more delicate, smaller mammalian types. Or it may be that the proboscideans actually effected a crossing from the mainland to the islands where other smaller, less adaptable mammals failed. Recently Dr. Tokunaga! published a compendium of all the fossil land mammals found in Japan and Korea. His list of genera and species, with their horizons and localities, is repeated here (p. 1489) in so far as it bears on Upper Cenozoic forms. VI. NORTH AMERICA NN TRODUCTION It would seem that the Proboscidea first reached North America from the Old World in Upper Miocene times, and from that time until the end of the Pleistocene, these animals played an important réle in the development of the mammalian faunas of the New World. Moreover, the evidence of recent discoveries shows that certain of the mammoths and mastodons did not become extinct until after man had crossed from Asia into America and was well established in his New World home. In the following pages the relationships of the various Proboscidea-bearing formations of North America will be discussed. Only those beds in which proboscideans are actually recorded will be included in the discussion, and they will be grouped by their ages, according to whether they are of Miocene, Pliocene, or Pleistocene affinities. In considering the Miocene and Pliocene beds, the several formations as named according to prevalent American usage will be described and compared. To give the discussions a certain degree of order and comprehension, the formations will be grouped according to their occurrence in the eastern portion of North America, in the Great Plains region, and in the Mountain and Pacific Coast region respectively. The treatment of the Pleistocene must of necessity be somewhat different. Here the sequence of the glacial and interglacial periods in North America in relation to the occurrence of Pleistocene proboscideans will be discussed. 1Tokunaga, Shigeyasu, 1933. Amer. Mus. Novitates, No. 627, pp. 2-4. GEOLOGIC SUCCESSION: NORTH AMERICA 1491 2. THE UPPER MIOCENE: BARSTOVIAN PAWNEE CREEK ForMATION, CoLoRADO.—Dr. Matthew, in his first description of the Pawnee Creek beds of northeastern Colorado, recognized two fossiliferous layers or zones within the formation, but at that time he pointed out the fact that the separation between these zones, either stratigraphically or faunistically, was very inconstant, so he was inclined to regard the zoning of the Pawnee Creek beds as of little importance. Later, in 1909, in 1918, and in 1924, he regarded the Pawnee Creek formation as a unit, containing a single fauna. GEOLOGICAL RELATIONSHIPS OF NORTH AMERICAN PROBOS CIDER, PACIFIC COAST INTERMONTANE PLATEAUS GREAT PLAINS AND MOUNTAINS EASTERN U.S. (WASHINGTON) Parelephas washingtonii (Mastodon americanus, Parelephas columbi, Archidiskodon imperatfor, Mammonteus primigenius,} P. rchidiskodon Archidiskodon maibeni, scoth, hayi,merid- arelephas eelsi Pareles ; ealinbittelicis ?Rhynchotherium BrneaiG ieee i Morrillia barbouri, tonal, nebrascensis reas iskodon INDIANA Widely distributed francisi Mastodon grangeri, PLEISTOCENE N. ica. sonoriensis, Stegomastodon| Mastodon mMoodiei, | Mastodon over America Mastodon ; imperator. . Mastodon Cortillecs prigstley! ttegomastodon acutidens Archidiskodon ore genens!s ime acne mele gets Wei Mastotacee, enius| Stegomastodon. Parelephas ii exilis "Pbfconeri SAN. Se Stegomastodor’ aftoniae 3 primitivus JERS TEHAMA i i ee PE. BO conaer uncertain Horizon uncertain Gs. CAROTENE ropicus,| Steqomastfodon ordillerion orarius i H ; Redeposited in Stegomastodon Searizonae |C. defloceatus qpeilephode ligpelients PLEISTOCENE > OS NEBRASKA TRILOPHODON ANGUSTIDENS, FRANCE Pe So TRILOPHODON FRICKI, cotoraoo TRILOPHODON COOPERI, BALUCHISTAN (ORAKI, NEW MEXICO GNATHABELODON THORPEI, Kansas An TRILOPHODON PHIPPS1, NEBRASKA MEGABELODON LULLI MORRILLIA BARBOUR], NEBRASKA TETRALOPHODON_PUNJABIENSIS, INDIA AMEBELODON FRICKI, NEBRASKA TETRALOPHODON (LYDEKKERIA) FALCONERI, NDIA TETRALOPHODON ELEGANS, KANSAS ALGERIA y Q TETRALOPHODON LONG/ROSTRIS, GERMANY FAYUM PHIOMIA’ OSBORNI, TETRALOPHODON CAMPESTER, KANSAS < PHIOMIA WINTONI, FaYum +7, TETRALOPHODON a YDEKKERIA) SINENSIS, CHINA a PHIOMIA SERRIDENS, Favua PHIOMIA MINOR, FAyUM 180" 165 150" 135" Jt EQUATOR EQUATOR) g 15° 30" 45° 60° 75u OOF 105° 135" 150" i L DF Levelt Bradley 935 180" Origin, Migration and Evolution of. oPhiomia, aAmebelodon, «Tetralophodon, x Morrillia. Trilophodon, aMegabelodon, ¢Gnathabelodon, Osborn 1935 PLATE XV Geologic range: Trilophodon, Lower Miocene to Middle Pliocene; Megabelodon, Mio-Pliocene and Lower Pliocene. Gnathabelodon, Middle(?) Pliocene. Phiomia, Lower Oligocene and Upper(?) Oligocene; Amebelodon, Middle Pliocene. Pleistocene. Tetralophodon, Mio-Pliocene to Middle Pliocene; Morrillia, Middle & q ¢ —_ ie =, - 7 - i Ss iat ay oe i 4: a oa ea ee ==) “AQQUYSY F3i8 2 ee te é <*4 ne a i \ i SUMMARY 1557 Genomastodon and Tatabelodon.—As regards Genomastodon and Tatabelodon, Professor Osborn was unable to find clear generic distinctions to validate these terms. Of the latter he remarks on page 325 of Volume I that the “superiorly grooved and upcurved longirostral symphyses of these types [T'atabelodon riograndensis and T. gregorii of Frick] scarcely serve for generic distinction from Trilophodon to which they appear to be related both in length of jaw and in the dimensions of the 4% ridge-crested third inferior grinding teeth.’”’ It will be noted, however, that both Genomastodon and Tatabelodon are retained as subgenera or synonyms of T'rilophodon (see Vol. I, pp. 293, 298, and 324). Micration.—The classic Trilophodon angustidens of the Middle Miocene of France is now traced to the T. angustidens libycus of North Africa through the very primitive T. pontileviensis of the Lower Miocene of France and the 7’. cooperi of the Lower Miocene of Baluchistan; it passed through the Lower and Middle Miocene stages of Europe and Asia into the extremely long-jawed American types of the Upper Miocene and the Pliocene of Dakota, Nebraska, Colorado, and New Mexico, to which different generic names have been assigned, such as Megabelodon and Genomastodon (sce migration map, Fig. 1227). Dr. Chester Stock lists Trzlophodon sp. as occurring in the Ricardo deposits of the Mohave Desert, California (see Stock, 1928.1, pp. 43-47). Recently Doctor Hopwood has described members of this subfamily from China (see p. 702 of Vol. I of this Memoir). So far as known, Trilophodon became rare, or migrated from Eurasia to America, at the close of Miocene or beginning of Pliocene time both in Europe and Asia. Superfamily: MASTODONTOIDEA Osborn, 1921. Family: BUNOMASTODONTID Osborn, 1921 (Syn. TRILOPHODONTID Simpson, 1931)! Subfamily: GNATHABELODONTIN® Barbour and Sternberg, 1935 Genus: Gnathabelodon Barbour and Sternberg, 1935 (Cf. Vol. I, Appendix, pp. 710-714, and Fig. 668; Vol. II, p. 1533, and Fig. 1227, also Pl. xv) GNATHABELODONTS, typical “‘shovel-jawed longirostrines,” of progressive bunomastodont type, with broad tuskless mandibular symphysis and very broad molars. The present author states on page 711 of Volume I that the “mandibular and dental characters widely separate Gnathabelodon from the shovel-tuskers, flat-tuskers, or any of the remaining genera of the Bunomastodontide and Serridentid and are in full accord with Barbour’s con- ception of this as representing a distinct subfamily remotely related to the Amebelodontine.””’ Gnathabelodon.—Skull very large, the width across the occipital condyles being 294 mm. Lower jaws longirostral, expanding broadly towards the extremities with an elongated and longitudinally groov- ed symphysis—a broad, birdlike, toothless bill analogous to that of the duck-billed Platypus in the Monotreme order. Superior tusks massive, rounded, and outcurved, with upturned tips; no enamel. Inferior tusks wanting, replaced by a broadly expanded, sharpened bony border, probably sheathed with hardened epidermis. Superior and inferior grinding teeth tetralophoid to pentalophoid, ptychodont, with double entotrefoils and rudimentary ectotrefoils in superior molars, double ectotrefoils on inferior molars. In general, of progressive, bunomastodont type with central conules. Intermediate molars trilophodont. In old age reduced to a single pair above and below. Cement in valleys of third superior molars. Dental formula incompletely known: I ¢3:5 M 35 Ridge-crest formula: M 2;, M 3¢;. Horizon.—Middle(?) Pliocene. Compare Vol. II, p. 1533, and figures 1224, 1227. Micration.—The genotype Gnathabelodon thorpei is the only species of this genus thus far discovered, and was found near Ogallah, Trego County, western Kansas. See footnote 2 on page 1555 of this chapter.—Hditor.| 1558 OSBORN: THE PROBOSCIDEA Superfamily: MASTODONTOIDEA Osborn, 1921. Family: BUNOMASTODONTID Osborn, 1921 (Syn. TRILOPHODONTID Simpson, 1931)! Subfamily: AMEBELODONTIN® Barbour, 1929 Genera: Amebelodon Barbour, 1927; Phiomia Andrews and Beadnell, 1902 (Cf. Vol. I, pp. 333-338, 690, 715-719, Pls.v—vu, x, x1, Fig. 277; Vol. II, p. 1533, Figs. 1220, 1224, 1227, Pl. xv) AMEBELODONTS, longirostral trilophodont mastodonts of North Africa and Central United States, differing from the typical Longirostrine in the high degree of specialization for the shoveling or digging functions of the inferior tusks, which are reinforced by internal dentinal lamine. This tusk structure precludes the analogy of Amebelodon to Platybelodon in which the inferior tusks consist of closely compacted rod-cones. The postsymphy- seal portions of the jaw and grinding teeth undergo little progressive evolution except in size; it is the anterior symphyseal and rostral region of the amebelodont shovel-tuskers that undergoes such an extraordinary special- ization, while the grinding teeth remain substantially the same. Amebelodon, typical ‘shovel-tusker’ of Nebraska, descendant of Phiomia of the Fayim. Cranium and rostrum uniformly abbreviated and reduced in size. Mandibular rostrum relatively long and slender. Lower jaws elongated (longirostral), nearly horizontal, moderately expanded towards the extremities. Superior tusks reduced in size, downcurved, outcurved, with persistent outer enamel band; inferior tusks without enamel band, composed of concentric dentinal lamine (1-9), instead of dentinal rod-cones as in Platybelodon, greatly elongated, somewhat broadened, closely appressed in the median line, tips rounded, alveolar portion plano-concave above. Mass of the Amebelodon tusk more than seventy times greater than that of the Phiomza tusk. Molars large, narrow, with double trefoils and central conules; inferior molars with five ridge-crests; cement present. From external cingulum rise many short, blunt cones. Dental formula: Incompletely known. Horizon.— Middle Pliocene. Compare Vol. II, p. 1533, and figures 1224, 1227. Phiomia, ancestral ‘Faytim shovel-tusker’ of North Africa. Of medilongirostral type; cranium low, abbreviated; palate long and narrow; symphysis long; progressive dolichopy, also progressive doli- chodonty and bunomastodonty, the latter character tending to place these animals in the direct line of ancestry of Amebelodon fricki. Superior tusks sharply pointed, downcurved, with enamel band on out- er surface. Inferior tusks horizontal, spatulate, without enamel, composed of concentric dentinal lamine (1-7); incisive alveoli elongate (P. osborni); Di, with straight inner border, convex outer border marked by crenulations or serrations, enamel confined to the tip. Molars typically bunodont, relatively long and narrow (progressive narrowing of M; prophetic of Amebelodon fricki), brachyodont, rudimentary or progressive conules; generally trilophodont, last inferior molar in Phiomia osborni subtetralophodont. Dental formula: Diz, C3 1 Dp # P= Mi? Ridge-crest formula: Dp 234 Dp 32+ Dp 43 P 3; P 42. M 13 M 23 M 33; Horizon.—Lower Oligocene (Fluvio-marine formation) and Upper(?) Oligocene. Compare Vol. I, figure 23; Vol. II, p. 1533, and figures 1220, 1224, 1227. Micration.—Up to the year 1934 the progressive species of Phiomia from the Fayim of North Africa were believed by the present author to be broadly ancestral to the Trilophodon angustidens phylum of Europe, chiefly because of the fundamental bunomastodont pattern of the Phiomia molars with ‘central conules’ conspicuous in Phiomia osbormi. Finally, however, he was of the opinion that Phiomia was the direct ancestor of Amebelodon, as stated in Volume I, p. 715: ‘‘Not only in external form but in the internal macroscopic and microscopi¢ structure of the incisive tusks (see Pl. v, A-F3) Phiomia wintoni leads into Phiomia osborni and after an immense interval of geologic time”! spanning the Upper Oligocene, the entire Miocene, and Lower Pliocene periods, the See footnote 2 on page 1555 of this chapter.—Editor.] *{“Thirty million year evolution of the lower jaw and paired inferior tusks from the Phiomia minor of the Oligocene of North Africa into the Amebelodon fricki of the Pliocene of Nebraska’”’ (cf. Osborn, 1935.937, p. 408, fig. 3; also Pl. y—with modifications—of Vol. I of the present Memoir).—Editor.]} SUMMARY 1559 Middle Pliocene A mebelodon frickt appears in Nebraska as the indubitable successor of Phiomia osborni of the Oligocene River Nile of Egypt.’”’ The abundance of Phiomia is attributable to its having a flood-plain habitat similar to that of Trzlophodon and other longirostrines, which are by far the most numerous proboscideans through- out Oligocene, Miocene, and Lower Pliocene times. Superfamily: MASTODONTOIDEA Osborn, 1921. Family: BUNOMASTODONTIDA! Osborn, 1921 (Syn. TRILOPHODONTID Simpson, 1931)! Subfamily: TrrraLOPHODONTIN van der Maarel, 1932 Genera: Tetralophodon Falconer, 1847, 1857; Morrillia Osborn, 1924. Subgenus: Lydekkeria, Osborn, 1924 (Cf. Vol. I, pp. 342-379, 690, 703-705, Pls. x, x1, and Fig. 307; Vol. II, p. 1533, Figs. 1221, 1224, 1227, Pl. xv) THE TETRALOPHODONTS, imperfectly known Medilongirostrines of Eurasia and North America, with four- crested or tetralophodont intermediate molars, single trefoils in the Miocene stage (Lydekkeria), rudimentary double trefoils in the Pliocene stage (Tetralophodon); brachyodont progressive to hypsodont, with cement and complete doubling of trefoils in Pleistocene time (Morrillia). Cranium brachycephalic, with rounded temporal region; moderate elongation of the rostrum. Palate contracted, with posterior nares directly opposite pentaloph of M*. Lower jaw medilongirostral. Superior tusks with enamel band. Inferior tusks flattened, oval, retrogressive, absent(?) in females (T. campester), without enamel band (the exception being 7’. grandincisivus). Superior and inferior tusks straight (Fig. 320). Molars progressively brachyodont to subhypsodont (Tetralophodon), to hypsodont and covered with cement (Morrillia). Intermediate molars tetralophodont. Third molars long and nar- row, with 5% to 6 ridge-crests (T'etralophodon), to 8% superior crests (Morrillia); third superior molars in T. fricki exhibit forward inclination of the crests. Central conules variable, cones and conelets relatively smooth and regular; progressive doubling of the trefoils. Third grinders only in use in old age. Dental formula: 1 *2-? Dp =* M #2 Ridge-crest formula: Dp 2?“ Dp 3° Dp 4%8 M 1% M 242 M 32282 Horizon.—Mio-Pliocene to Middle Pliocene (Tetralophodon), to Middle Pleistocene (Morrillia). Compare Vol. I, figure 307; Vol. II, p. 1533, and figures 1221, 1224, 1227, also Pl. xv. It has been erroneously assumed by some authors that animals of the Trilophodon angustidens type gave rise to animals of the Tetralophodon type by abbreviation of the jaw and by the addition of a fourth crest on the intermediate molars. This assumption is disproven by three facts: (1) The true T'rilophodon persisted after the true Tetralophodon had appeared; (2) Tetralophodon is a Medilongirostrine, whereas Trilophodon is a Hyper- longirostrine; (3) a true ancestral species of Tetralophodon (Lydekkeria), e.g., Mastodon | = Tetralophodon (Lydek- keria)| falconeri Lydekker, is recorded in the Middle Pliocene, Dhok Pathan beds, of India. Consequently the separation by Maarel in 1932 of the Tetralophodonts from the Trilophodonts under the subfamily name Tetralo- phodontinz has been adopted in the present Memoir. These medilongirostral mastodonts arose independently from an ancestral Oligocene stage which might be comprised within the genus Phiomia. This stage was less specialized than Phiomia osborni, which leads directly into Amebelodon fricki, as mentioned above under the Amebelodontine (p. 1558). The distinctions between the genus T'rilophodon, the typical genus Tetralophodon, and the genus Morrillia are entirely in progressive characters which developed from the Mio-Pliocene into Pleistocene times. The subgenus Lydekkeria Osborn, 1924, genotype Mastodon falconeri Lydekker, 1877, of supposed Middle Miocene age by Lydekker, Pilgrim, and Osborn, now proves to be of Middle Pliocene age, according to Hopwood [See footnote 2 on page 1555 of this chapter.—EHditor.] 1560 OSBORN: THE PROBOSCIDEA (see Vol. I, p. 353), and therefore a contemporary of ‘Mastodon’ [Tetralophodon| punjabiensis Lydekker, 1886. This being the case, Lydekkeria may become a synonym of Tetralophodon, and T. punjabiensis may become a synonym of 7. falconert. This leaves the Miocene ancestry of Tetralophodon uncertain, but the primitive Mio- Pliocene stage (Lydekkeria) appears to be ancestral to the Lower Pliocene Tetralophodon. Micration.—By referring to figure 1227 of the present Memoir, also to Pl. xv, it will be noted that the general trend of migration is from the west to the east. A fragmentary molar of a member of the Tetralophodon- tine (‘M.’ longirostris?) is recorded from the lacustrine beds of Smendou north of Constantine, northern Africa, Professor Osborn in 1925 in his article in NarurAL History on ““The Elephants and Mastodonts Arrive in Amer- a,’’ pp. 12 and 13, states that the ‘“‘Tetralophodonts are in all countries very rare, yet we can trace their long migration through eastern Europe into India and China, until finally they arrive in Kansas and Nebraska.”’ Since that time Tetralophodon bumiajuensis of Java has been described by van der Maarel, and the present author, in the first volume of this Memoir, described 7’. fricki from northern Texas. From the limited evidence now at hand it would appear that Tetralophodon became rare in Eurasia early in Pliocene time but lingered until Middle Pleistocene time on the ancient flood-plains of Nebraska. Superfamily: MASTODONTOIDEA Osborn, 1921. Family: BUNOMASTODONTID Osborn, 1921 (Syn. TRILOPHODONTID Simpson, 1931)! Subfamily: Nororostrin Osborn, 1921 (syn. Cordillerionine Scott, 1937) Genus: Cordillerion Osborn, 1926 (Cf. Vol. I, pp. 543-566, also Pls. x, x1, and Fig. 501; Vol. II, p. 1534, and Figs. 1224, 1225, 1227, Pl. xv1) Cordillerion.—Cranium mastodontoid, of relatively low, simple character as in the primitive Trilophodonts; rostrum elongate. Jaws broad, deep, and abbreviated (brevirostral) ; symphysis is of medi- um length. Superior incisive tusks spiral or horizontal, down- and outcurved, around which is wound more or less spirally a broad enamel band. Inferior incisive tusks wanting, probably present in juvenile stages. Grinding teeth brachyodont, never hypsodont, moderately elongate, with single trefoils, parallel in this respect with Trilophodon, central conules more or less conspicuous, cement in progressive stages. Intermediate molars with tendency to tetralophodonty (subtetralophoid) in M 2. In final phyletic stages grinding tooth action concentrated on second and third superior and inferior molars. Adaptation chiefly to a mountain habitat, grinders adapted principally to a browsing habit, tusks probably used in the uprooting of plants for food. Undoubtedly of Old World ancestry, although no member of the Notorostrine has as yet been found in the Eastern Hemisphere. Dental formula: I *** Dp #¢ M ?3 Ridge-crest formula: Dp 23 Dp 3 ie 4; M 13 M 233% M 3 cae Horizon—Pliocene and Pleistocene. Compare Vol. I, figures 501 and 680; Vol. II, p. 1534, and figures 1224, 1225, 1227, also Pl. xvi. Tue Nororostrines.—The earliest member of the subfamily Notorostrine is the South American Masto- donte des Cordiliéres of Cuvier described in 1806 and later (1824) designated by him Mastodon andium, thus superseding the name Mastodon cordillerarum of Desmarest, 1820-1822. This species was for a long time confused with the Mastodonte humboldien, also of South America, described by Cuvier in the same publication of 1806. In January of 1921—a century later—Professor Osborn reviewed (Osborn, 1921.515, p. 10) these two classic species, including also the Dibelodon | =Cordillerion| tropicus and Mastodon |= Stegomastodon| successor of Cope, designating them as the ‘“‘Brevirostrines of South America,” and in September of the same year he established ‘See footnote 2 on page 1555 of this chapter.—Editor.] Abin d CORDILLERION ANDIUM, ECUADOR ts ) RHYNCHOTHERIUM FALCONERI, Texas le ANANCUS FA AE CONERS: ENGLAND RHYNCHOTHERIUM SHEPARDI E, CALIFORNIA > Cc ley EDENSIS, CALIFORNIA ANANCUS ARVERNENSIS, FRANCE S PENTALOPHODON SIVALENSIS, 1NOIA INDIA” RHYNCHOTHERIUM BROWNI, mexico AY BELODON HONDURENSIS, HONoURAS SIND ANANCUS PERIMENSIS INDIA - A A RHYNCHOTHERIUM TLASCALAE, SYNCONOLOPHUS ~CORRUGATUS, INDIA MEXICO BLICKOTHERIUM ) BLICKI, HoNnoURAS a SYNCONOLOPHUS PROPATHANENSIS, INDIA RHYNCHOTHERIUM CHINJIENSE, iNnoia SYNCONOLOPHUS PTYCHODUS, INDIA BLICKOTHERIUM / EUHYPODON, Dal W) NEBRASKA Yann a ANANCUS PROPERIMENSIS, INDIA RHYNCHOTHERIUM SPENCERI, EGyer EQUATOR DF Levett Bradley 1936 Origin, Migration and Benen ae tell ekhynchotherium, aBlickotherium, aclybelodon, ¢Anancus, *Pentalophodon, xSynconolophus. Osborn 1935 PLATE XVI Rhynchotherium, Middle Miocene to Upper Pliocene; Blickotherium, Pliocene; Aybelodon, Anancus, Mio-Pliocene to uppermost Pliocene; Pentalophodon, Upper Pliocene; Synconolophus, Mio-Pliocene to Middle Pliocene. Geologie range: Cordillerion, Pliocene and Pleistocene. Pliocene. i PAA YE SN SUMMARY 1561 his new subfamily, the Notorostrine (1921.526, p. 330) to embrace not only the four species just mentioned but six additional species, namely, Mastodon bolivianus and M. chilensis of Philippi, and M. platensis, M. rectus, M. argentinus, and M. superbus of Ameghino, on the ground that they were profoundly separated from other buno- mastodonts by the progressive abbreviation of the jaw, and that they were the only members of the great order Proboscidea which, as far as known (1921), entered the South American continent. Professor Osborn’s researches had not then revealed the fact that other mastodonts, namely, Rhynchotherium Falconer and Notiomastodon Cabrera, had also wandered into South America as well as one species of elephant belonging to the Parelephas phylum, i.e., Parelephas columbi cayennensis Osborn, of French Guiana. Perhaps a still more important step was taken by Professor Osborn in 1923 (1923.601, p. 1) when he separated the Mastodon humboldtii group from the M. andium group, making M. humboldti the type of his genus Cuvieronius, distinguished by double trefoils on the superior and inferior molars, and by simple, rounded, outwardly and up- wardly curved superior tusks, lacking enamel band (see p. 540 of Vol. I), in contradistinction to M. andium with single trefoils, more or less conspicuous central conules in the molars, and broad, spirally enameled superior tusks, a peculiarity not observed in any other proboscidean, to which subsequently (1926.706, p. 15) he assigned the generic name Cordillerion. This subfamily of short-jawed bunomastodonts, therefore, is monophyletic, including the genus Cordillerion only. Micration.—As mentioned above, the Notorostrine probably originated in the Old World; fossil evidence of this fact, however, is lacking, for the earliest occurrence, as far as known, of the genus Cordillerion is the Upper Pliocene C. edensis Frick-Osborn, found at Mt. Eden Hot Springs, San Bernardino County, California; other discoveries are at Benson, Arizona, and near Sinton, Texas, also of the Pliocene. It would seem, therefore, that the trend of migration was southward, through Mexico, into South America, as far as Uruguay. Superfamily: MASTODONTOIDEA Osborn, 1921. Family: BUNOMASTODONTID4 Osborn, 1921 (Syn. TRILOPHODONTID Simpson, 1931)! Subfamily: RayncHorostrin Osborn, 1918, 1921 (syn. Rhynchotheriine Cabrera, 1929) Genera: Rhynchotherium Falconer, 1862, 1868; Blickothervum Frick, 1933; Aybelodon Frick, 1933 (Cf. Vol. I, pp. 475-513, 690, also Pls. x, x1, and Figs. 451, 483; Vol. II, p. 1531, and Figs. 1220, 1221, 1224, 1227, also Pl. xvi) Tur RHYNCHOROSTRINES, beak-jawed mastodonts, typified by Rhynchothertum Falconer, so named because of the sharply deflected rostrum. The subfamily Rhynchorostrine is the least known of any of the bunomasto- donts. The genotype of Rhynchotherium is the cast of a lower jaw, the original of which was found at Tlascala, Valley of Mexico. It was at first characterized by Falconer (in a letter to Lartet of Sept. 12, 1856) as resembling “very much the figure in Alcide D’Orbigny’s Voyage, described by Laurillard as M. Andium,” but he stated that the ‘‘Genoese paleontologists had provisionally named it Rhynchothercum, from the enormous development of the beak, approaching Dinotherium.”’ This letter was not published until 1868 (see Pal. Mem., Vol. II, pp. 74, 75). He further stated that he saw this cast in Genoa, an unfortunate substitution for Geneva (see Falconer, 1863, p. 56), an error to which Felix and Lenk called attention in 1891, but which was apparently unnoticed until Professor Osborn’s researches for the present Memoir led him to make inquiry of Dr. Hans Georg Stehlin, who kindly located the cast in the Muséum d’Histoire Naturelle of Geneva. Through the courtesy of the Director, Dr. Pierre Revilliod, a duplicate cast was presented to the American Museum. Professor Osborn was the first to use the specific name tlascale in 1918 (Osborn, 1918.468, table opposite p. 134); thus the genotype is Rhynchotherium tlascale. The discovery in 1911 by Dr. Barnum Brown near San See footnote 2 on page 1555 of this chapter.—Editor.] 1562 OSBORN: THE PROBOSCIDEA José de Pimas, Sonora, Mexico, of a fine specimen of a lower jaw referable to Rhynchotheriwum—in fact, made the neotype by Professor Osborn in 1921 (1921.515, pp. 5, 6), owing to the supposed loss of the cast—served to confirm the presence of this genus in Mexico. On recovery of the cast in Geneva, it was found that the type and neotype differed sufficiently to warrant the naming of a new species; consequently Professor Osborn assigned to the neo- type the name Rhynchotherium browni (see Vol. I of the present Memoir, p. 494). During the years following Faleoner’s description, specimens of Rhynchotherium were unearthed in California, Colorado, Montana, Nebraska, and Texas. The Rhynchorostrine include the fewest and the least known species of the bunomastodonts. Tracing the phylum back into its Eurasiatic or Old World ancestry, we find a primitive lower jaw with downturned symphysis and laterally compressed incisive avleoli (Fig. 455) described by Fourtau in 1918 as Mastodon spenceri from Moghara, Egypt, but provisionally referred by Professor Osborn to Rhynchothertum spenceri, which may confirm the hypothesis of the African origin of the Rhynchorostrinz. Also doubtfully referred by Professor Osborn to Rhynchotherium (R. chingiense) is a left ramus, with M2, Ms, 7m sztu, found by Dr. Barnum Brown in 1922 in the Mio-Pliocene of India, near Chinji Bungalow. Dr. D. K. Chakravarti in 1935, p. 209, states that he sees no evidence of the downturning of the symphysis, and that the ‘“‘warping of the alveolus of the tusk appears to the present author to be a case of distortion superinduced upon the specimen after its entombment”’; he suggests that ‘it should be referred to Trilophodon angustidens (Cuvier) var. chinjiensis (a name proposed by Pilgrim in 1913 for a form discovered in the Salt Range. .. ).” Rhynchotherium.—No complete cranium known. Face of medium length, maxilla partly down- turned. Mandible of medium length; symphyseal portion strongly deflected. Superior and inferior tusks, I’, I., laterally compressed, bent downwards and outwards, with broad external enamel band; inferior tusks slightly upeurved. Grinding teeth of bunomastodont type, relatively broad, simple, brachyodont, single trefoils only, inferior ‘central conules’ variable, superior ‘central conules’ absent. Molar enamel thick. Third superior grinders with three and a half to four plus ridge-crests; third inferior grinders not exceeding four and a half to five ridge-crests; functional grinders in adults reduced to M 2-M 3, finally to M 3. Intermediate grinders trilophodont. Dental formula: I =? Dp # M # Ridge-crest formula: Dp 2% Dp 33 Dp 4% M 1 M 22, M 3°%4+ Horizon.—Middle Miocene to Upper Pliocene. Compare Vol. II, p. 1531, and figures 1220, 1224, 1227, also Pl. xvi. Up to the year 1929 the Rhynchorostrine had been considered as monophyletic. The discovery, however, near Tapasuma, Honduras, by Mr. John C. Blick of the Frick Expedition of 1929, of two long slender mandibles, one adolescent, the other mature, amplified our knowledge of these rare and probably forest-living animals, with the surprising result, as described by Mr. Childs Frick in 1933 in his article on ““New Remains of Trilophodont- Tetrabelodont Mastodonts,” that two new genera were added to this subfamily, namely, Blickotherium, genotype Blickotherium blicki, and Aybelodon, genotype Aybelodon hondurensis. The distinctive feature of Aybelodon is the enamel-less incisors, differing in this respect from other members of the Rhynchorostrinz, but the profile of the type jaw reveals its rhynchotherine affinity and places it in a ‘‘distinet and unique line of succession.” Blickotherium.—No complete cranium known; see, however, figure 462 A for palate of Mastodon euhypodon Cope referred to Blickotherium by the present author. Mandible elongate, extremely slender, less deflected than in Rhynchotherium. Symphysis laterally compressed, downwardly curved. Inferior tusks upeurved, compressed laterally, broad external enamel band. Apparent presence of a replace- ment P, in the adolescent mandible. Ridge-crest formula (Frick): Dp 45; (possible replacement P,) M 1s M 2;;, Ridge-crest formula (Osborn): M 1; M 2s M 33, Horizon.—Pliocene. Compare Vol. II, p. 1531, and figures 1224, 1227, also Pl. xvr. SUMMARY 1563 Aybelodon.—Cranium unknown. Mandibular rami of long, relatively slender type; vertical ramus, condyle and coronoid process low. Symphysis strong, very elongate, laterally and vertically compressed, upcurved, extremely narrow, transverse exceeding vertical diameter, as contrasted with Blickotheriwm blickt. Inferior tusks large, broad, closely appressed, upeurved, without trace of enamel band. Superior tusk (referred), possibly representing a distal section of upper left side, exhibits tendency toward inward and forward curvature, the presumed outer surface with remnants of enamel band. First and second inferior molars small; third inferior molars relatively broad and short; very thick enamel. Second inferior molars probably trilophoid; third inferior molars fully tetralophoid with rudimentary pentalophid. Ridge-crest formula: M 2;; M 335 Horizon.—Pliocene. Compare Vol. II, p. 1531, and figures 1224, 1227, also Pl. xv. In summarizing the persistent primitive and progressive characters of the Rhynchorostrines, we may mention the following: Persistent Primitive Characters.—(1) Grinding teeth persistently brachyodont or short crowned; (2) no evidence of hypsodonty or deposition of cement; (3) enamel unusually thick; (4) simple external trefoil spurs on inferior molars and internal trefoil spurs on superior molars, no trace of doubling of the trefoil spurs; (5) inferior ‘central conules’ variable, superior ‘central conules’ absent. Progressive Characters.—Superior tusks progressive, elongated, and laterally flattened, with broad persistent enamel band. Inferior tusks progressive, compressed laterally, elongated, upturned, with persistent lateral enamel band. The tusks of Aybelodon hondurensis, however, are entirely without enamel. Mraration.—From the Miocene of ?Egypt, through the Mio-Pliocene of ?India, to the Middle to Upper Miocene of Montana, Upper Miocene-Upper Pliocene of California, Nebraska, Texas, Colorado, Mexico, and Honduras. See especially figure 483 of Volume I of the present Memoir, and the accompanying PI. xvt. Superfamily: MASTODONTOIDEA Osborn, 1921. Family: BUNOMASTODONTID Osborn, 1921 (Syn. TRILOPHODONTID& Simpson, 1931)! Subfamily: BrevirosTrRIN& Osborn, 1918 (syn. Pentalophodontine Scott, 1937) Genera: Anancus Aymard, 1855, 1859; Pentalophodon Falconer, 1857, 1865; Synconolophus Osborn, 1929 (Cf. Vol. I, pp. 617-665, 690, Pls. x, x1, and Fig. 590; Vol. II, p. 1534, and figures 1221-1223, 1227, also Pl. xv1) Tur BREVIROSTRINES.—We now come to one of the mastodont divisions that never reached America, as far as is known at present, namely, the Brevirostrine, a subfamily provisionally named by Professor Osborn in 1918 (1918.468, p. 136) and fully defined by him in 1921 (1921.515, p. 11). In Volume I, p. 627, of the present Memoir, he summarizes the characters of this subfamily, as follows: ‘“These short-jawed bunomastodonts are readily distinguished from all others by four principal characters: First, the skull and jaws short, as in the elephants; second, the grinding teeth finally reduced to one above and one below, as also in the elephants and in the Stegodonts; third, perhaps most distinctive and unique, the alternation of the main internal and external cones which are placed diagonally instead of transversely to the long axis of the crowns, a character first observed by Cautley in describing his species Mastodon sivalensis also by Falconer (1846, p. 50) in comparing the Mastodon sivalensis of the Siwaliks with M. latidens, and later (1868, II, pp. 29-31) with the Mastodon arvernensis of the British coast Upper Pliocene; the fourth distinctive character is the plication or folding (Synconolophus) of the enamel borders of the molar ridges, giving the hippopotamus- and piglike character termed ‘chcerodont’ by Schlesinger, the 'ISee footnote 2 on page 1555 of this chapter.—Hditor.] 1564 OSBORN: THE PROBOSCIDEA grinding action of the teeth being obviously as in pigs, and the hippopotami. The grinders, originally brachy- odont, become subhypsodont; thus the single superior and inferior molar teeth become highly effective masti- cating organs, labyrinthodont in pattern and serving effectually in the comminution of food.”’ It appears that the Brevirostrine include three phyla, namely, Anancus Aymard, 1855, 1859, Pentalophodon Falconer, 1857, 1865, and Synconolophus Osborn, 1929. Stegomastodon Pohlig, 1912, at first regarded as belonging to the Brevirostrinz, was removed by Professor Osborn to his new family Humboldtide, because of a “fundamental resemblace of the grinding teeth to those of ‘Mastodon’ humboldtit.”’ Anancus, ‘straight-tusked brevirostrine’ of southern Eurasia. Aymard based his genus on Anancus macroplus, 1855, a generic and specific name applied to bones and teeth of an adult mastodont similar to the species described and figured by Croizet and Jobert as Mastodon arvernensis from Auvergne, based on the milk dentition. Anancus macroplus is therefore regarded as synonymous or identical with A. arvernensis, which latter becomes the genotype. The term Anancus signifies without bend, probably in allusion to the straight tusks or upper incisors. This phylum is purely Eurasiatic. A. arvernensis and A. falconert may possibly be derived from A. perimensis or A. properimensis of India. Anancus.—Cranium approaching true elephantine type, with brachycephaly, hypsicephaly, and fore- and-aft abbreviation; relatively primitive elongation of the rostrum in front of the orbits, an adaptation to the presence of two grinding teeth in use at the same time, namely, M 2-M 3; frontal region relative- ly narrow or laterally compressed; summit also narrow, since the temporal fosse are divided by a relatively narrow frontoparietal plate as compared with the very broad frontoparietal plate in Pentalo- phodon falconeri; occipital condyle very prominent and pedunculate—a unique feature in the Probo- scidea. Anterior portion of symphyseal region abbreviate. Superior tusks straight, elongate. No in- inferior tusks. True molars brachyodont to subhypsodont; cones simple, smooth, compressed fore and aft. Single external median conules in inferior molars; internal median conules in superior molars. Deciduous molars (premolars) grooved, ptychoid, or plicated, externally. No trefoils. Crowns brachyo- dont, cones erect (Anancus macroplus and related species); crowns subhypsodont, strongly inclined and alternating (A. falconeri). Internal proversion of superior and inferior ridge-crests. Intermediate molars tetralophodont. Dental formula: I *?-° Dp #4 M +3 Ridge-crest formula: Dp 22 Dp 3 +2+ Dp 4 “4 M 14 M 24:42 M 3 33 Horizon.—Mio-Pliocene to uppermost Pliocene. Compare Vol. II, p. 1534, and figures 1221, 1222, 1227, also Pl. xvt. Pentalophodon, ‘five-crested straight-tusker’ of the Siwaliks. Falconer in 1857 characterized this genus as having the intermediate molars divided upon a quinary ridge formula, evidently with Mastodon sivalensis of Cautley in mind at the time, for later (1865, p. 262) he clearly designated it as the genotype. His descriptions and definitions are also based on specimens of superior and inferior molars figured in plates Xxxv1, Xxxvui of the “Fauna Antiqua Sivalensis,” as well as on a cranium figured in plate xxx1n, fig. 2, with altogether different and much more progressive grinding teeth. A close comparison of the molars shows that this referred cranium belongs to a much more progressive species, if not genus, than the type molar of ‘Mastodon’ sivalensis, which Professor Osborn in Volume I, p. 653, of the present Memoir made the type of a new species, namely, Pentalo- phodon falconert. This Indian form (Pentalophodon sivalensis) has a line of southern Asiatic ancestry of its own, branching from Anancus perimensis and distinct from the west European A. arvernensis. Pentalophodon.—Cranium lofty, hypsicephalic, brachyopic. Occipital condyles elevated. Sym- physis abbreviate. Incisive tusks unknown. Intermediate molars pentalophodont. Molars subhypso- dont; cones elevated, strongly alternating, inclined forwards in the lower molars; superior and inferior ridge-crests extending obliquely forwards from the external to the internal faces, that is, internal proversion; typically composed of three main irregular cones set widely apart (P. sivalensis), closely SUMMARY 1565 compacted (P. falconeri); crown progressively chcerodont with twenty-two aristogenes in M; (P. sivalensis), twenty-seven in M® (P. falconert); enamel smooth, not ptychoid; no ecto- and entotre- foils; central conules present in two anterior valleys. Small amount of cement sometimes present. Dental formula: I °° Dp =} M {2 Ridge-crest formula: Dp 4; ?M 1, M 24%2 M 35324 ni Horizon.—Upper Pliocene. Compare Vol. I, p. 648; Vol. II, p. 1534, Figs. 1221, 1223, 1227, and Sacvi. Synconolophus, ‘syncone-crested brevirostrine’ of the Siwaliks. Falconer’s plates or descriptions do not seem to show grinding teeth referable to this peculiar genus, but Lydekker in his memoir of 1880 on “Siwalik and Narbada Proboscidia”’ figures examples of teeth certainly referable to Synconolophus. The warping or dislocation of the cones and crests in the superior and inferior molars is much more extreme than in Anancus or in Stegomasto- don, also the multiplication of separate intermediate conelets, originally springing from the trefoils, is quite distinct from the regular single or double trefoils of Stegomastodon. The whole surface of the crown becomes crowded with a labyrinthine pattern of cones, conelets, and more or less separate intermediate conelets, with warped or dislocated ridge-crests in apparent disorder, giving the appearance best expressed in the Greek generic term Synconolophus. The genotype is Synconolophus dhokpathanensis, based on a giant cranium found by Dr. Barnum Brown in 1922 in the Dhok Pathan horizon of India. The three-ridged second molar, M?’, suggested comparison with T'rilophodon, but the broad four-ridged third molar, M7’, differed from that of Trilophodon and also from that of Tetralophodon punjabiensis which occurs in the same geologic horizon. It was not until perfect superior molars were found (see Fig. 631—paratype), also from the same horizon a pair of referred superior and inferior molars (Fig. 632), that the structural and phyletic key to this very aberrant genus could be determined (ef. Vol. I, pp. 654-665). Synconolophus.—Cranium platycephalic, dolichopic, brachycranial (Vol. I, p. 649) [dolichocranial? —see p. 347], somewhat more elongate than in Anancus or Pentalophodon. Postnarial opening far back of grinders, a generic distinction from Tetralophodon. Mandibular symphysis broad, trough shaped, concave superiorly (S. propathanensis), indicating a large, fleshy lower lip. Superior tusks large, up- turned, and out-turned, without enamel. Inferior tusks reduced or absent (S. propathanensis). Milk tusks with enamel. Grinders with ridge-crests strongly arched or dislocated; valleys blocked with warp- ed, compacted conelets and conules, giving on wear astrongly chcerodont, ptychoid, crowded, compressed, enamel-bordered appearance; internal proversion. No trefoils. Cement strongly developed in S. dhokpath- anensis and S. propathanensis. Cones smooth to deeply grooved. Intermediate molars with trilophodont ridge formula and small rudimentary half ridge-crests, representing pro-protoloph and tetartoloph. Dental formula: Di *?° I “* Dp =¢ M #2 Ridge-crest formula: Dp 4° M 1*°* M 2%3# M 3¢+% Hor1zon.—Mio-Pliocene to Middle Pliocene. Compare Vol. IT, p. 1534, and figures 1221, 1227, also Pl. XVI. GEOGRAPHIC RANGE.—(Vol. I, p. 629): ‘Armed with very powerful upper tusks, the enamel bands of which early disappear, propelled by massive limbs supported on a powerful frame, these animals appear to have been adapted to browsing on the succulent vegetation of the warm-temperate zones, chiefly of the 40th and 30th parallels, from France and the southeast coast of England, through northern Italy, then through India, but never reaching North America, so far as we know at present. Analogous but unrelated are the species of Stegomastodon.”’ A step further must be taken in the migration of these animals, owing to the determination by Dr. A. Tindell Hopwood (1935.1, pp. 57-60) of molars from Shansi, China, as belonging to Pentalophodon sinensis. Professor Osborn accepted Doctor Hopwood’s reference of these molars to a member of the Brevirostrine, but regarded them as referable rather to Anancus than to the more progressive Pentalophodon which has a greater number of ridge-crests than are present in the Chinese molars (see Appendix to Vol. I, p. 722). 1566 OSBORN: THE PROBOSCIDEA Superfamily: MASTODONTOIDEA Osborn, 1921. Family: HUMBOLDTID Osborn, 1935, 1936 (Syn. STEGOMASTODONTID Scott, 1937) Subfamily: HumBouptina Osborn, 1934, 1936 (syn. Stegomastodontine Scott, 1937) Genera: Cuvieronius Osborn, 1923; Hubelodon Barbour, 1914; Stegomastodon Pohlig, 1912 (Compare Vol. I, Chap. XII, Chap. XIII, pp. 667-684, and Appendix, pp. 722-728, also Pls. x, x1, and figures 501, 590; Vol. II, p. 1535, and figures 1224, 1225, 1227, also Pl. xvi1) THE HumBoLpTINEs, ‘hyperptychoid Humboldtine mastodonts’ of the southern United States and South America. The decision of Professor Osborn in 1934 (Osborn, 1934.924, p. 183, fig. 2) to dismember his subfamily Notorostrine (which included the Mastodon |Cordillerion| andium and M. |Cuvieronius| humboldtii groups, as well as the Hubelodon morrilli of Barbour) was occasioned by his recognition of the differences in the crania and dentition of the members of these groups. For example, the Cuvieronius skull converges toward that of Hle- phas and of Mastodon americanus, while the Cordillerion skull retains the relatively low, simple character of the primitive Trilophodonts; furthermore, in Cuvieronius hwmboldti the superior tusks are straight or upcurved without enamel band, and the superior and inferior molars are broad with typically double trefoils and central conules either inconspicuous or wanting; whereas in Cordillerion andium a broad enamel band winds more or less spirally around the elongated superior tusks and the molars are narrow with single trefoils on the internal side of the upper molars and on the external side of the lower molars, as in other bunomastodonts. In the same article (1934, p. 183) Professor Osborn definitely placed Eubelodon morrilli in the direct ancestral line of the Humboldtines or Humboldtine (fig. 2, our Pl. x), and accepted Dr. Cabrera’s inclusion of Leidy’s Mastodon |= Stegomastodon| mirificus, 1858, as well as other Stegomastodonts, in the Humboldtinz, which sub- family is defined on pages 575 and 615 of Volume I of the present Memoir. Eubelodon.—From 1914, when Doctor Barbour described Hubelodon morrilli, until 1929, the type specimen was the single representative of this genus. The discovery of a superb cranium by the Frick Expedition of 1929, at Horsethief Canyon, near Ainsworth, Nebraska, and referred by Mr. Childs Frick in 1933 to this species, shed additional light on the dentition of Hubelodon, as both the second and third superior molars are present, the latter beautifully displaying the coronal pattern of 44 ridge-crests with double trefoils, at once reminding one of Cuvier’s type of ‘Mastodon’ humboldtii. Characters of Eubelodon, ‘ancestral Humboldtine mastodont’ of Nebraska. Cranial and facial region primitive, subequal, less elevated than in Cuvieronius humboldtii. Occipital condyle low com- pared with elevated condyle and occipital crest of Cuvieronius superbus. Rostrum and inferior mandible elongate, somewhat downturned and pointed, with no trace of alveolus for tusks. Superior tusks relatively short, simple, rounded, out-turned, tips chisel shaped, slight downcurvature, conserving the primitive downcurvature of the paired superior incisors of all primitive mastodonts, divergent, without enamel band. No inferior tusks. Grinders broad with double trefoils, four ridge-crests and talon. Superior molars with strong entotrefoils, incipient ectotrefoils; indications of cement; broad internal cingulum; outer side more convex, inner side more plane or concave. Cingulum less distinct in inferior molars. Pelvis excessively broad; humerus robust; inferior in height and width to Warren Mastodon (Mastodon americanus). Ridge-crest formula: M 2} M 342 Horizon.—Middle(?) Pliocene. Compare Vol. II, p. 1535, and figures 1224, 1227, also Pl. xvit. Cuvieronius.—In giving the history of the genotype of Cuvieronius, namely, the Mastodonte humboldien Cuvier, 1806 [= Mastodon humboldtii Cuvier in Desmarest, 1818, Mastodon humboldii Cuvier, 1824], Professor CUVIERONIUS SUPERBUS, CUVIERONIUS PLATENSIS, A CUVIERONIUS HUMBOLDTII, CHILE ARGENTINA Cs CUVIERONIUS AYORAE, STEGOMASTODON ARIZONAE, ona STEGOMASTODON SUCCESSOR, TEXAS EUBELODON MORRILLI, NEBRASKA STEGOMASTODON AFTONIAE, IOWA arcentina CUVIERONIUS POSTREMUS, ecvu4oor ; STEGOMASTODON MIRIFICUS, hihs NEBRASKA Saeiae STEGOMASTODON PRIMITIVUS, NEBRASKA ARIZONA STEGOMASTODON TEXANUS, TEXAS 60} 45) J 15 | & MEN IRS | | -; ; | t+) + = se = YY EQUATOR $ EQUATOR I T i Hee al | | 180 16: (35 150" 105° 30W. DF Levelt Bradley 1935 = 120 30" 60" 90°E 105 120° 150" Origin, Migration and £volutio eEubelodon, aCuvieronius, * Stegomastodon. PLATE XVII i OF Osborn 1935 Geologic range: Eubelodon, Middle(?) Pliocene; Cuvieronius, Pleistocene, Upper Pleistocene (C. postremus); Stegomastodon, Upper Pliocene and Lower to Middle Pleistocene. iia. Lae - - i SUMMARY 1567 Osborn states in Volume I of the present Memoir, p. 575, as follows: “The weight of Cuvier’s authority has been so great that since 1806 the generic name Mastodon was applied by all authors to humboldtzi, until Cope proposed (1884.2, p. 2) the ill-fated generic name Dibelodon, signifying two-tusker, which he based upon the four-tusker Mastodon shepardi of Leidy. Cope’s practice of including Mastodon andiwm and M. humboldtii also within the genus Dibelodon was followed by Lull and Osborn until Osborn discovered Cope’s error and proposed the generic name Cweronius (1923.601). Meanwhile all the European authors continued to use Cuvier’s name Mastodon, including Boule in his important Memoir of 1920.” Characters of Cuvieronius, ‘Humboldtine mastodont’ of the Pampean and Andean regions of South America. Cranium elevated, facial region prominent, expanded, with air-cell chambers as in Mastodon and Elephas. Rostrum of inferior mandible relatively abbreviate. Tusks relatively large, simple, rounded, upturned and out-turned or straight, without enamel band; relatively short, robust, upturned in primitive species (C. ayore and C. postremus). No inferior tusks. Molars broad with typically double trefoils; single trefoils in primitive species (C. ayorx, C. postremus). Third superior and inferior mo- lars tetralophodont to pentalophodont. Intermediate molars with rudimentary tetartoloph. Superior ridge-crests transverse or retroverted internally; inferior ridge-crests proverted internally. Superior and inferior crowns either plane or curved on external side. Central conules inconspicuous or absent. Ridge-crest formula: M 13t M 28% M 342°: Horizon.—Pleistocene. Upper Pleistocene (C. postremus). Compare Vol. II, p. 1535, and figures 225227, also. Pl. xvit. Stegomastodon.—This genus was first thought by Professor Osborn to be the American representative of his subfamily Brevirostrine, but it has recently been determined that the migration of the true Brevirostrines (Anancus, Pentalophodon, and Synconolophus) did not extend to the North American continent. We are still without knowledge of the Asiatic and possibly African forbears of Stegomastodon. The Stegomastodon mirificus group now proves to be closely related in its molar-tooth structure to the Cuvieronius humboldtii group of the subfamily Humboldtine. This observation was first made by Dr. Angel Cabrera in 1929 and was confirmed by Professor Osborn (see Vol. I, p. 612, this Memoir) after a further “close comparison of the fundamental grinding- tooth pattern in Stegomastodon,’”’ which disclosed the fact that ‘‘both the superior and inferior ridge-crests of the true Brevirostrines of Eurasia are proverted, whereas in the Humboldtines of America the inferior ridge-crests are proverted and the superior ridge-crests are either retroverted or transverse (Cwvieronius) or centroverted (Stegomastodon).”’ Characters of Stegomastodon, Pohlig’s ‘roof-toothed Humboldtine mastodont’ of the southwestern United States. Cranium abbreviate; palate elongate, postnarial opening far back of grinders (S. tecanus). Mandibular rostrum progressively abbreviated. Superior tusks short, massive, upcurved, outcurved, without enamel band. No inferior tusks. Tusks in S. primitivus first projected directly out from the alveoli after which they were simply outcurved and then by inward rotation the tips were finally incurved at the extremities. This peculiar tusk structure may remove it from the direct ancestry of the other species of Stegomastodon and place it in a line of its own. Juvenile tusks straight. Molars brachyodont to subhypsodont; cones slightly alternating, deeply plicated, ptychoid, folded, giving a labyrinthine pattern to the worn grinding surfaces; double to quadruple to multiple trefoils in both superior and inferior molars, completely blocking the valleys; enamel thick; no central conules; external pericones block the spaces between the five main crests (S. texanus ref.). Superior ridge-crests centroverted; inferior ridge-crests proverted internally as in Anancus, Pentalophodon, and Snyconolo- phus. Cement in transverse valleys. Intermediate molars with trilophodont to tetralophodont ridge formula, that is, 3% ridge-crests. Ridge-crest formula: Dp 43 M 13% M 23% M 3feovs-sie ts Horizon.—Upper Pliocene and Lower to Middle Pleistocene. Compare Vol. II, p. 1535, and figures 1224, 1227, also Pl. xvit. 1568 OSBORN: THE PROBOSCIDEA The final step taken by Professor Osborn in 1935 was the creation of a new family, the Humboldtide (see Osborn, 1935.937, fig. 2, forname only; 1936, Vol. I, p. 722, for definition), to embrace the Hwbelodon-Cuvieronius- Stegomastodon group, or Humboldtine. Micration.—(Vol. I, p. 611): “Putting together the important resemblances between specimens of Eubelo- don morrilli of Nebraska and the relatively little-known primitive remains of tusks and grinding teeth from the Inferior Pampean of Argentina, we reach the very important conclusion that EH. morrilli is a Middle Pliocene ancestral stage in the subfamily Humboldtine on its way from Eurasia through North America to South America. In the Andean region there appear the primitive short-limbed forms Cuvieronius ayore and C. postremus, and the primitive typical C. hwmboldti of Bolivia, (?)Ecuador, and Chile, quadrupeds of smaller size and possibly of mountain habitat. Possibly a direct descendant of Hubelodon is the straight-tusked Cuvieronius rectus of the Inferior Pampean of the Argentine. Finally there appears in the Pampean the giant Cuweronius swperbus and the more specialized C. platensis as distinguished by Cabrera.’”’ On a subsequent page (p. 685) Professor Osborn states that we may be certain that at some future time we may trace Hubelodon back to northern or central Asia and ultimately perhaps to Africa. Superfamily: MASTODONTOIDEA Osborn, 1921. Family: SERRIDENTID Osborn, 1935, 1936 Subfamily: SERRIDENTIN# Osborn, 1921 Genera: Serridentinus Osborn, 1923; Ocalientinus, Serbelodon, and Trobelodon Frick, 1933 (Cf. Vol. I, pp. 380-473, 729, also Pls. x, x1, and Figs. 344, 373; Vol. II, p. 1535, and figures 1221-1225, 1227, also Pl. xvi) Perhaps in no other subfamily has there been such confusion in the determination of species as in the Ser- ridentine, especially in the differentiation of the Trilophodont and Serridentine unworn molars, in fact, for a century these animals have been confused with T'rilophodon. The salient features which clearly separate the subfamily Serridentinze from the Longirostrine (or Trilophodonts) are: First, the central conules observed in all species of T’rilophodon are entirely absent in the Serridentine; second, these central conules are functionally replaced in the Serridentine by crests or spurs which arise on the sides of the superior internal cones and on the inferior external cones; and third, these internal and external crests subdivide into two, three, four to six smaller conelets, which produce a serrated aspect, hence the term ‘serrate-toothed mastodonts’ or Serridentine. By referring to Professor Osborn’s first papers, it will be noted (Osborn, 1921.515, p. 8) that he designated as the ‘‘T. |Trilophodon] serridens Phylum” four species, namely, Mastodon serridens Cope, Tetrabelodon serridens cimarronis Cope, Dibelodon precursor Cope, and questionably Mastodon turicense Schinz. Again in his definition of the new subfamily Serridentinze (Osborn, 1921.526, p. 330) he stated that the “‘Serridentine apparently spring from M. turicense Schinz, 1824, of the Middle Miocene of France and Switzerland—a rare animal, probably because a forest dweller,” and calls attention to the distinguishing spur or crest on the sides of the superior and inferior grinders of the Serridentinz. Professor Osborn’s final conclusions are given in Volume I, p. 382, of the present Memoir: ‘“‘One of the most interesting results of the prolonged research in preparation for this Memoir is the discovery of two clearly distinguishable phyla of mastodonts (Turicius and Serridentinus) which by all previous investigators both in Europe and America had been confused with the phylum Trilophodon. . . In the present chapter [Chap. X] it is shown that the grinding teeth, e.g., of Mastodon (B.) subtapiroidea of the Lower Miocene lignites. . . widely differ from those of Trilophodon and strongly resemble those of Serridentinus. . . [see Figs. 350, 351, 352]; it is possible that Twricius [genotype Mastodon turicense Schinz] and Serridentinus arise from a similar stock not unlike Serridentinus subtapiroideus. The evolution of the true Turicius phylum, however, is entirely 3 NOTIOMASTODON ORNATUS, ARGENTINA V6) 1 OCALIENTINUS FLORESCENS, moncotra “ ; (| 7 4 : J nee SERBELODON BURNHAM1I, NOTIOMASTODON ARGENTINUS, CALIFORNIA SERRIDENTINUS Z| ee Ul GUATEMALENSIS, GUATEMALA oy OCALIENTINUS J OBLIQUIDENS, TORYNOBELODON SOUTH CAROLINA BARNUMBROWNI, NEBRASKA SERRIDENTINUS ‘“ANGUIRIVALIS, AEG TORYNOBELODON LOOMISI, SERBELODON REBAR! 3 BARBOURENSIS, NEBRASKA » {| OCALIENTINUS FLORIDANUS, FLoRioa db SERRIDENTINUS SERRIDENS, Z SERRIDENTINUS PRODUCTUS, Y, ; 7 c NEW MEXICO ~ ~ (2S : SERRIDENT, BR 4 7N NI TROBELODON TAOENSIS, Ah) ES ME Be é ) NEW MEXICO SERRIDENTINUS SRS GGT ere FD NE SES: ) NEW MEXICO | PLATYBELODON GRANGERI, moncotsa SERRIDENTINUS GOBIENSIS, INNER MONGOLIA PLATYBELODON DANOVI, , : NORTH CAUCASUS SERRIDENTINUS SUBTAPIROIDEUS, SERRIDENTINUS MONGOLIENSIS, MONGOLIA AUSTRIA AEP) SERRIDENTINUS ANNECTENS, JsaPan >\ - SUMMARY 1569 distinct, leading into sharply crested forms such as T. virgatidens, while the Serridentinus phylum is (A) increasing- ly Serridentine, ending in Ocalientinus (Ser.) floridanus, or (B) more crested, ending in S. guatemalensis.”’ While, therefore, there is a strong resemblance in the Miocene stages between species of Serridentinus and species of Turicius, there is very little resemblance between final species in these two phyla which are very widely divergent. The Serridentine are polyphyletic, embracing four genera—Serridentinus Osborn, 1923, and the Ocalientinus, Serbelodon, and Trobelodon of Frick, 1933. Professor Osborn’s researches led him in 1935 (Osborn, 1935.937, fig. 2—see also Vol. I, p. 729, of the present Memoir) definitely to separate these ‘serrate-toothed mastodonts’ (the Serridentine), also the Platybelodontine (Platybelodon, Torynobelodon) and the Notiomastodontine (Notiomasto- don) from all other mastodonts, thereby constituting a very large and highly diversified new family, the Serri- dentide, which at the present time includes no less than forty species (Vol. I, p. 741), united by the common character of the serrated crests arising from the ectoconelets in the lower molars and from the entoconelets in the upper molars, also by the enamel bands on the superior tusks, but generically separated by the widely radiating adaptations of the inferior tusks—horizontal oval in Serridentinus, oval, greatly enlarged in Serbelodon, flattened into a pair of broad shovels in Platybelodon, abbreviated and rounded in Torynobelodon, and entirely wanting, with brevirostral mandible, in Notiomastodon. Hasits.—Members of the Serridentide, especially the true Serridentinus, rarely occur in the same fossil beds with members of the Bunomastodontide, hence it is inferred that they had different feeding habits. The more or less perfectly serrated crests on the inner side of the upper molars and on the outer side of the lower molars point to leaf-cutting and browsing habits, especially true of the typical Serridentines, which, it is thought, may have frequented forests and were principally browsers, while the typical flat-tuskers (Platybelodon), the cutting-tuskers (Serbelodon), and the sub-shovel tuskers (Trobelodon) frequented shallow lakes and river borders. Characters of Serridentinus, typical ‘serrate-toothed serridentine,’ ‘prod-tusker’ or ‘scaptobelodont,’ of Eurasia and North America. Cranium low, dolichocephalic; undoubtedly rounded above like the skulls of Trilophodon and Tetralophodon; the chief difference between the skull of Serridentinus and that of Trilophodon is the abbreviation of the rostrum, maxillo-premaxillary above, and symphyseal portion of the mandible below. The symphysis in species of American Serridentinus is decurved and contains two large tusks of flattened oval section. Jaws moderately elongate (medilongirostral), relatively stout, tending to decurvature. Superior tusks laterally compressed, downcurved and outcurved, retaining broad lateral enamel band. Inferior tusks without enamel band, large, horizontally flattened, closely appressed, and never greatly elongated as in Trilophodon. Intermediate molars trilophodont from the beginning to the end of the phylum as in the Trilophodon phylum. Grinding teeth relatively broader than in Trilophodon, primitively brachyodont progressive to subhypsodont; prominent serrated trefoil spurs or crests arise from the anterior and posterior faces of the entocones in the superior molars, from the ectocones in the inferior molars; median conules absent in the valleys, unlike Trilophodon in which they are present. Three superior and inferior grinding teeth (M 1—M 3) in use at the same time; functional retention and usage of upper and lower second and third molars, in contrast to the Longirostrine (Trilophodon) and Tetralophodontine (Tetralophodon) in which the mastication is finally concentrated on the third upper and lower molars. Internal cingulum more or less crenulate on superior molars, extend- ing as far back as the antero-internal border of third crest of the third molars; rudimentary external cingulum only in valley between first and second crests (S. productus). Dental formula: I ¢2° Dp #¢ P #§ M23 Ridge-crest formula: Dp 23; Dp 33 Dp 43 P 3z P 42+ M 13 M 23 M 33%. Hori1zon.—Lower Miocene to Upper Pliocene. Compare Vol. II, p. 1535, and figures 1221-1224, 1227, also Pl. xvii. Characters of Ocalientinus, ‘multiserrate-crested serridentine,’ ‘prod-tusker’ or ‘scaptobelodont,’ of the United States and Mongolia, directly ancestral to ‘Serridentinus’ republicanus of Kansas and to ‘S.’ floridanus of Florida. Cranium tall, narrow. Mandible elongate, symphysis extremely narrow with deep 1570 OSBORN: THE PROBOSCIDEA superior longitudinal channel. Superior tusks as in Serridentinus, with latero-inferior enamel band. Inferior tusks without enamel, upcurved, relatively short. Molars with roundly blunted cones and conelets, trefoil spurs dominant, florescent. Relative breadth of molars correlated with foreshortening of cranium and heightening of occiput. Both superior and inferior ridge-crests sloping forward, slowly increasing in height. Ridge-crest formula: Dp 4% P 3? P 4s M 15 M 28 M 3,4, Horizon.—Miocene to Upper Pliocene (O. obliquidens redeposited in Lower Pleistocene). Compare Vol. II, p. 1536, and figures 1223, 1224, 1227, also Pl. xvim. Characters of Serbelodon, Frick’s ‘sub-shovel-tusked serridentine’ of the western United States. Cranium imperfectly known, probably more elongate than that of Trobelodon. Mandible extremely abbreviated and broadened, the length equaling that of Serridentinus productus. Symphysis short, symphyseal rostrum extremely short and relatively broad. Superior tusks heavy, downwardly and outwardly curved, with strong enamel band. Inferior tusks concavo-convex, relatively short, flattened, heavy, spatulate; distinguished from Platybelodon by total absence of dentinal rod-cones, the interior being composed of dentinal laminz. Molars relatively long and narrow with serridentine pattern— superior molars with entoconelet spurs, inferior molars with ectoconelet spurs, but without serrated borders. Ridge-crest formula: M 2% M 3+ Hor1zon.—Lower Pliocene, Upper Pliocene. Compare Vol. II, p. 1536, and figures 1224, 1227, also Pl. xv. Characters of Trobelodon, Frick’s ‘brevidentate serridentine’ of New Mexico, of ‘sub-shovel-tusk’ type. Cranium broad or brachycephalic. Mandible relatively decurved. Symphysis relatively elongate, narrow, slightly upcurved. Superior tusks with moderate enamel band; inferior tusks moderately heavy, biconvex, pointed, absence of dentinal rod-cones. Third superior molars short and broad, with blunt cones; expanded entotrefoils as in Serridentinus; third inferior molars also short and broad, with ex- panded ectotrefoils—trefoil structure of primitive serridentine type. Ridge-crest formula: M 23 M 3% Horizon.—Mio-Pliocene. Compare Vol. II, p. 1537, and figures 1224, 1227, also Pl. xvi. Micration.—The following is the geographic distribution of species of the Serridentine, the trend of mi- gration being eastward from southwestern Europe to North America (see Pl. xvim, also Figs. 373 and 1227): France, Austria, India, Mongolia, China, Japan, California, Nebraska, Kansas, Colorado, New Mexico, Texas, Guatemala, and eastward in the United States to North and South Carolina and Florida. Superfamily: MASTODONTOIDEA Osborn, 1921. Family: SERRIDENTID Osborn, 1935, 1936 Subfamily: PLATYBELODONTIN# Borissiak, 1928 Genera: Platybelodon Borissiak, 1928, 1929; Torynobelodon Barbour, 1929 (Cf. Vol. I, pp. 338, 339, 459-472, 729, 730, also Pls. x, x1, and Figs. 344, 373; Vol. II, p. 1537, and figures 1222, 1223, 1227, also Pl. xvitt) The subfamily Platybelodontinze was founded by Borissiak in 1928 on his Platybelodon danovi of the Chokrak beds, Kuban district, North Caucasus, the chief distinguishing character being the long and wide spoon-shaped symphysis with wide, flat incisors. Prior to this description, and wholly unknown to Professor Borissiak, a speci- men found in Nebraska had been described by Dr. E. H. Barbour as Amebelodon fricki in 1927 and made the type of his family Amebelodontide (subsequently, 1929, modified to Amebelodontine). Professor Osborn assumed (1931) that the Amebelodontide Barbour, 1927, preoccupied the Platybelodontine Borissiak, 1928, especially as both possessed large shovel tusks supposedly penetrated by dentinal tubules or rod-cones; also at the time he was not aware of the highly progressive serridentine pattern of the Platybelodon molars, in widest contrast to the trilophodont pattern of the Amebelodon grinders, or of the fact that the second superior and inferior molars possessed four ridge-crests, unlike Trilophodon and members of the subfamily Serridentinz with three ridge-crests. SUMMARY 1571 As stated above under the subfamily Amebelodontine (p. 1558), the structure of the incisive tusks of A mebelo- don and of its ancestral form Phiomia is that of progressively concentric dentinal lamine, an observation made in 1934, 1935, when the inferior tusks of Phiomia were sectioned and carefully compared with those of Amebelodon. Likewise the inferior tusks of Platybelodon were sectioned, revealing compact interior dentinal rod-cones instead of concentric dentinal lamine; also the tusk of the American related genus Torynobelodon was found to be composed of rod-cones, resulting in the reference of this genus to the Platybelodontine. Quoting Dr. Edwin H. Colbert in Volume I, p. 715, of the present Memoir, “It would seem possible, then, that the presence or absence of these rods distinguishes the two subfamilies Platybelodontinze and Amebelodontinz respectively .. . It would seem likely, also, that in the Platybelodontine the tusks are square across the ends, whereas in the Amebelodontinz the ends of the tusks are rounded. . . The origin of the rods constitutes a perplexing question. To me [Colbert] they appear to be derived from the lamellar ivory; perhaps they represent a secondary breaking down of the concentric cones of dentine that form the tusk. Perhaps they are structures developed in the pulp cavity independently of the lamellar ivory. .. A magnified cross-section of a dentinal rod of Torynobelodon loomisi shows that it is built up of concentric layers of dentine—presumably unvascular orthodentine. An examination of the tusk of 7’. loomisi shows that the rods occupy the central portion of the tusk, and that they are surrounded by typical ivory. The rods are quite variable as to size, ranging from one to five millimeters in di- ameter. Many of the rods branch. There are numerous nodules or egg-shaped excrescences on the sides of various rods, and on some rods the external surface is granular. The rods are of varying lengths. The rods are ‘bundled’ together, so to speak, in the tusk, but they are not compressed; consequently they are round and not hexagonal. The interstices between the rods were seemingly empty; if there was a filling of dentine it has been destroyed during the process of fossilization. In many places the rods seem to grade into the lamellar ivory.” Characters of Platybelodon, Borissiak’s typical ‘flat-tusker’ of Mongolia. Cranium broad, low, dolichopic. Mandible with long and wide spoon- or shovel-shaped symphysis; extreme constriction and narrowing of mandibular groove (according to the theory of Dr. William K. Gregory, 1934, the lower sur- face of the tongue occupies this narrow groove which broadens out into the deeply hollowed suprarostral concavity). Low and massive rami (type). Superior tusks small, rounded, relatively short and slightly outturned and downturned; inferior tusks wide and flat, with regularly and sharply defined borders, com- posed of numerous and quite closely compacted dentinal rod-cones. Upper surface without enamel, with irregular longitudinal grooves, lower surface protected by enamel [?dentine] longitudinally striated (fide Borissiak). Anterior serration of the entocones in the superior molars and of the ectocones in the inferior molars, as distinguished from the double or anterior and posterior serration of the corresponding ento- and ectocones in the Serridentinus and Ocalientinus molars. Fourth superior and inferior deciduous premolars and second molars with four ridge-crests instead of three ridge-crests as in Trilophodon, Amebelodon, and members of the subfamily Serridentine, resembling Tetralophodon in this respect. Dentalitormula: Wi-.-5 | --, Opa — 4 Ml Ridge-crest formula (based on complete collection of P. grangeri in American Museum): Dp 1; Dp 25Jaecwon Dp 35:55 Dp 455 P 3 Saat P43 M15) M 224, M 3534, Horizon.—Upper Miocene. Compare Vol. II, p. 1537, and figures 1223, 1227, also Pl. xvi. Characters of Torynobelodon, Barbour’s ‘dredge-tusker’ of Nebraska. Cranial characters not yet described by Doctor Barbour. Mandible thin, ladle shaped, with low ascending rami; abbreviated as compared with Platybelodon grangeri; symphysis relatively narrow with relatively broad bar between rostrum and rami. Tusks short, broad, ladle or spoon shaped, upcurved, roughly and deeply corrugated, composed of dentinal rod-cones. Dental formula incompletely known: I 53 M =; Ridge-crest formula incompletely known: M 2s M 3 ;, Horizon.—Pliocene. Compare Vol. II, p. 1537, and figures 1224, 1227, also Pl. xvmt. 1572 OSBORN: THE PROBOSCIDEA Micration.—Up to the present time members of this subfamily have been found in the North Caucasus and in the Gobi of Mongolia. A left lower incisor from Texas, in the Colorado Museum, may be referable to Platybelodon; also Torynobelodon, which appears to be a collateral branch, and somewhat less perfect mechanically, of the Platybelodonts of Central Asia, comes from the Pliocene of Nebraska. Superfamily: MASTODONTOIDEA Osborn, 1921. Family: SERRIDENTID Osborn, 1935, 1936 Subfamily: NorroMastopDONTIN« Osborn, 1936 Genus: Notiomastodon Cabrera, 1929 (Compare Vol. I, pp. 541, 590-592, 691, 730, 731; also Pls. x-xu, and Fig. 501; Vol. II, p. 1537, and figures 1225, 1227, also Pl. xv) The third subfamily of the Serridentide is the Notiomastodontinz Osborn, based on the Notiomastodon ornatus of Cabrera, 1929. The reasons set forth by the present author for his reference of this subfamily and genus to the Serridentide will be found on page 730 of Volume I of this Memoir, and are briefly as follows: The inferior molars have prominent serrated ridges arising from the ectoconelets as in Serridentines, and the adjacent mesoconelets are reduced or wanting, in contrast to Cuvieronius and Cordillerion of South America, in which the trefoil spurs arise from the mesoconelets; it is radically different from the fundamental molar pattern of any of the Bunomastodontide, the trefoil structure of which invariably rises from the mesoconelet or is in close connection with it through the central conules. Also evidencing the Serridentine relationship are the laterally compressed, downcurved superior tusks, with broad external enamel band, all trace of which band is absent in the contemporary Cuvieronius of the Pampean region. Characters of Notiomastodon, Cabrera’s ‘short-jawed serridentine’ of the Pampean. Mandible abbreviated, typically stout, lacking inferior tusks; symphysis strongly abbreviated with a transversely concave groove for the tongue; coronoid small and condyle elevated. Superior tusks down- and out- turned, with broad enamel band as in Serridentines and Trilophodonts. Inferior molars bunolophodont, with prominent serrated ridges arising from the ectoconelets as in the Serridentines; mesoconelets re- duced or wanting. Double series of trefoils on wear (genotype). Ridge-crest formula: M 1, M 2; M 3ivaq Horizon.—Pliocene and Lower Pleistocene(?)._ Compare Vol. I, Pls. x and x1; Vol. II, p. 1537, and figures 1225, 1227, also Pl. xvi. Micration.—The Notiomastodonts seem to have been confined to South America, as the genotype, Notio- mastodon ornatus from Buenos Aires, and a tusk from the valley of the Tarija River, described by Ameghino in 1888 as Mastodon argentinus, referred by the present author to Notiomastodon, are the only specimens recorded up to the present time. Superfamily: MASTODONTOIDEA Osborn, 1921. Family: MASTODONTIDA® Girard, 1852 (syn. in part MAMMUTID& Cabrera, 1929) Subfamily: PaLasoMASTODONTINA Osborn, 1936 Genus: Palsxomastodon Andrews, 1901 (Cf. Vol. I, pp. 138, 139, 143-149, 689, 691, 692, also Pls. 1, 1, x, x1, and Figs. 24, 86, 123a; Vol. II, p. 1529, and figure 1220, also Pl. xrx) The theory that Meritherium gave rise to Palzomastodon, and that Palzomastodon gave rise to Mastodon angustidens, generally so interpreted from Dr. C. W. Andrews’ Memoir of 1906, was not accepted by Professor Osborn, who, in 1909, even went so far as to separate Maritherium from direct relationship to the Proboscidea. This opinion, however, he afterward (1921) modified, in fact, he definitely regarded Meritherium as constituting es ee ACUTIDENS, var, INDIANA hs “ant MAS TODO. eS N. AMERICA MASTODON _PAVLOW!, 1 od Te PLIOMASTODON “VEXILLARIUS, CALIFORNIA ZYGOLOPHODON BORSONI, 1TALy Har TURICIUS TIRGA TIDENS, GERMANY STEGOLOPHODON STEGODONTOIDES, INDIA a STEGOLOPHODON LYDEKKERI, BORNEO ase y Z A\ STEGOLOPHODON LATIDENS, BURMA L « ae STEGOLOPHODON CAUTLEY! PROGRESSUS, INDIA MIOMASTODON TAI TAPIRG IDES Atel AMERICANUS, HUNGARY FAYUM a MIOMASTODON MERRIAFII, NEVADA Saas 2 é e , of, en ee PALAEOMASTODON_INTERMEDIUS, FAYOM Lo { / a wat MIOMASTODON DEPERETI, FRANCE PALAEOMASTODON BEADNELLI, TURICIUS TURICENSIS, SWITZERLAND PALAEOMASTODON PARVUS, FAYUM EQUATOR Origin, Migration and Evolution of « Pal&tomastodon, Ai Panmastiodiom. sPigniasedon * Mastodon, aTuricius, x Zygolophodon, ¢Stegolophodon. Osborn 1935 PLATE XIX Geologic range: Palzomastodon, Lower Oligocene. Miomastodon, Lower to Middle Miocene, and Lower Pliocene; Pliomastodon, Lower to late Pliocene; Mastodon, early to late Pleistocene. Turicius, Lower Miocene to Middle(?) Pliocene; Zygolophodon, Lower Miocene to Upper Pliocene. Stegolophodon, Mio-Pliocene to Upper(?) Pliocene [Lower Pleistocene?]. SUMMARY 1573 one of the great primary stocks of the Proboscidea, namely, the Mceritherioidea. Doctor Andrews also (1909), while still regarding Maitherium as a proboscidean, reconsidered the possibility of its not being in the direct line of ancestry of Palzomastodon; and Doctor Matsumoto (1924), while placing Meritherium in a side line of its own, concluded that Palzomastodon appeared nearly to correspond to a theoretical ancestry of the Zygolophodon-Mastodon phylum, thus suggesting that it was related to the subfamily which the present author designated as the Mastodontine. This theory was immediately approved by Professor Osborn, although it must not be overlooked that in 1923 he had clearly separated the true Mastodon americanus from the Zygolophodon of Vacek, at which time he described a new subfamily, the Zygolophodontine. In 1934 Professor Osborn became convinced that Palzomastodon should be removed from direct ancestry to the true Mastodon, because he observed that the progressive third superior and inferior molars of P. intermedius exhibited the presence of proto- and metaconules, thus blocking the median sulcus characteristic of all the Masto- dontine. On page 139 of Volume I of the present Memoir we find the following: ‘In the carefully drawn Plates I, 1, m1, and tv, illustrating the evolution of the grinding teeth in Mastodon, Zygolophodon, Turicius, and Stegolo- phodon, a median longitudinal sulcus separates the external and internal cones both in the superior and inferior molars. This demonstrates that the ancestral proboscidean molar was tetrabunodont, as in Meeritherium, not hexabunodont, as in Palzomastodon. .. By close comparison of all the figures of the upper and lower grinding teeth of Palzomastodon, Miomastodon, Pliomastodon, and Mastodon, it is observed: (1) That the molar crowns in Palzomastodon are mainly tetrabunodont, i.e., each protoloph (superior) and each protolophid (inferior) is com- posed of a main external bunoid cone and a main internal bunoid cone; in the superior molars (Fig. 94D) where the conules persist the main crown is hexabunodont. The presence of proto- and metaconules blocking the median sulcus forbids the ancestry of Palzomastodon to Mastodon. (2) The vestigial intermediate protoconules and metaconules are observed in the hexabunodont superior molars of Palzomastodon intermedius, thus the crested upper grinders are hexabunodont or six coned, whereas the lower grinders are subtrilophodont (Fig. 93, M®-Ms,, Fig. 94). This primitive condition of the cones connects Palzomastodon with its undiscovered sexituber- cular-quadritubercular ungulate ancestors; the conules observed in the third superior molar, M?, of P. intermedius (Figs. 93 and 89) are not seen in M? of the same species (Fig. 92); the conules are vestigial or disappearing struct- ures.”’ Accordingly Professor Osborn withdrew Palzomastodon from the subfamily Mastodontine and created a new subfamily, Paleomastodontine, for its reception (see Vol. I, Appendix, p. 691). Palzomastodon, the ‘ancient mastodon’ of North Africa, Eocene-Oligocene. Skull imperfectly known. Palate relatively short and broad. Mandible elongated anteroposteriorly (medium jawed) mandibular symphysis relatively short as compared with Phiomia. Superior tusks unknown. Inferior tusks rounded and relatively abbreviated. Molars brachyodont, of bunolophodont rather than of buno- mastodont type, progressive in size and evolution from tetrabunodont cones (Palzomastodon parvus) to hexabunodont cones (P. intermedius, P. beadnellc); bilophodont superior molars, subtrilophodont in- ferior molars, in other words, the tritoloph and tritolophid are in a formative stage, tritoloph rudimen- tary even in M*. Intermediate proto- and metaconules progressively developing to perfect the transverse crests, thus closing the median longitudinal sulcus characteristic of all the Mastodontinz. Molars exhibit- ing central conules; trefoils rudimentary or absent. Basal cingula neither very strong nor very rough. Dental formula: 1,455 Ct Dp; PM? Ridge-crest formula: P 23 P 31% P 43 M 172 M 2 #* M 3% Horizon.—Lower Oligocene (Fluvio-marine formation). Compare Vol. II, p. 1529, and figure 1220; also Pl. xrx. Hasirs AND Hasirat.—Probably forest loving with transversely secant molar crests for cutting herbage. Up to the present time, however, specimens of Palzomastodon have been found in the Fayim region only. 1574 OSBORN: THE PROBOSCIDEA Superfamily: MASTODONTOIDEA Osborn, 1921. Family: MASTODONTID Girard, 1852 (Syn. in part MAMMUTIDA Cabrera, 1929) Subfamily: Masropontina Brandt, 1869—Osborn, 1910 Genera: Mastodon Cuvier, 1806-1817; Miomastodon Osborn, 1922; Pliomastodon Osborn, 1926 (Cf. Vol. I, pp. 132, 1388, 150-190, 690, 693-697; also Pls. 1, x, x1, and Figs. 86, 123a, 123b; Vol. II, p. 1530, and figures 1222, 1224, 1227, also Pl. x1x) In Professor Osborn’s article in NaTrurRAL History of 1925 (Osborn, 1925.637), we find on page 10 the follow- ing statement: “The true mastodons of our American forests appear to have arisen from the diminutive Palxo- mastodon of the primordial river Nile. The reason these animals have left no trace of their 10,000-mile and 2,000,000-year journey from the Nile region to the forests bordering the Ohio and the Hudson rivers is that fossilization of forest-living fauna has always been rare. The ancestral Palzomastodon of the Nile region is itself very rare... not even fossilized teeth of this race were scattered in Europe to show the route.”’ During the ten-year interval from 1925 to 1935, Professor Osborn changed his views regarding Palzomastodon being the ancestor of the true Mastodon, as expressed in the foregoing paragraphs (p. 1573) on the Palzesomastodon- tine and also on pages 138 and 139 of Volume I of the present Memoir: ‘‘While we are certain that the true Pal- zomastodon possesses a longer jaw but a broader and shorter skull than its Lower Oligocene contemporary Phiomia, while the superior grinding teeth are comparatively short and broad, and while in the crowns of the superior and inferior grinding teeth the proto- and metaconules forbid the ancestral relationship of Palzomastodon to Plio- mastodon, Miomastodon, and Mastodon, we must await further knowledge of the cranium and of the cutting teeth, also of the incisors of Palzomastodon, before we can form a positive opinion on this very important and interesting question.” Again, in the Appendix to Volume I, p. 692, he voices uncertainty as to the ancestral forms of the Mastodontinze which embraced the four genera, Palezomastodon, Miomastodon, Pliomastodon, Mastodon, until he withdrew Palzomastodon, making it the type of a new subfamily Paleomastodontine: ‘Despite the extreme rarity of the remains of forest-loving and browsing mastodonts, exemplified by the rarity of Palzomastodon as compared with Phiomia in the Fayim deposits, several discoveries of isolated grinding teeth have been made in the Tertiary deposits of Europe and North America which are now recognized as belonging to successors of still unknown true Mastodontine of Africa and ancestors of the true Mastodontine of North America.”’ The contrasting characters, so far as known, of the three genera now constituting the Mastodontine, are as follows: Mastodon, ‘typical mastodon’ of Eurasiatic and North American forests. Cranium brachycephalic, brachyopic. Mandible and symphysis abbreviated; progressive reduction of rostrum. Superior tusks large, rounded, upturned, with slightly indicated annular ring growths; inferior tusks straight, cylindri- cal, variable in old age; no enamel. Marked sexual disparity in female tusks. Grinders relatively elon- gate, bilobate, with strong median sulcus between inner and outer lobes; no conules; rudimentary tre- foil spurs on superior inner lobes, on inferior outer lobes; summits of lobes simple or crowned with small conelets; extreme binary fission of summits of crests in Mastodon acutidens, in which as many as twenty-one conelets may be counted; progressive plication of the surface enamel. Ridge-crests lopho- dont, progressively elevated, subhypsodont. Progressive development of third crest (tritoloph and -lophid) in intermediate molars; progressive development of fifth crest rudiment (pentalophid) in third inferior molars. Permanent premolars suppressed, except P'-P,, which form in the jaw but do not erupt. Cingulum slightly stronger on inner side than on outer side. Dental formula: I 0? Dp =% P ¢:°:t (vestigial) M +3 0-0-4 Ridge-crest formula: Dp 2% Dp 33 Dp 43 M 13 M 23 M 3%% 4M Horizon.—Early to late Pleistocene. Compare Vol. II, p. 1530, figures 1222, 1224, 1227, and Pl. xix. SUMMARY — 1575 Pliomastodon, ‘Pliocene mastodon’ of Eurasia and North America, an ancestral stage of Mastodon. Distinctive, low browed, flat crested; abbreviated postnarial region and broadly divergent premaxillary region; approximation of condyles to superior molars (M°), extreme backward recession of anterior nares and shortening of lower jaw, probably also an unusual width of the occipital crest, suggesting an ex- ceptionally large development of the proboscis (characters based on Pliomastodon vexillarius Matthew, 1930—see Vol. I, pp. 161-163). Superior tusks suboval to rounded in section, upturned, enamel band vestigial or wanting; a “‘narrow strip of thin enamel” within the alveolar base of the tusk in P. vexillarius. Inferior tusks probably stout (as inferred from the alveoli—about 50 mm. in diameter near the posterior end—in P. sellardsi Simpson, 1930). Molars intermediate in hypsodonty between Miomastodon and Mastodon; a distinctive feature is the posterior narrowing of the crown of the third superior molars (P. matthew). Ridge-crests with rudimentary trefoil spur extending from entocones of superior molars and from ectocones of inferior molars; expanded ectotrefoils (P. matthewi). No median conules. Persis- tent longitudinal sulcus; also persistent four conelets. 0-2-0 Dental formula incompletely known: I &2° Dp? P? M 43 Ridge-crest formula: P 43; M 1; M 23 M 3,44, Horizon.—Lower to late Pliocene. Compare Vol. II, p. 1530, and figures 1222, 1224, 1227, also Pl. TK, Miomastodon, ‘Miocene mastodon’ of Eurasia and North America. Skull and skeleton unknown. Mandible and symphysis moderately elongated; rostrum laterally compressed, abbreviate as compared with Trilophodon or Serridentinus. Superior tusks downturned, broadly oval in vertical section; broad enamel band on concave external surface (Miomastodon merriam?), on convex external surface (M. tapiroides americanus); inferior tusks rodlike, horizontal, vertically oval. Molars brachyodont to subhypsodont; internal and external lobes or cones of each loph separated by median or longitudinal sulcus; summit of each lobe or cone double or bifid, rounded; mesial expansion of rudimentary trefoils in ento- and ectoconelets, upper and lower; central conules absent; retarded binary fission of cones and conelets in proto- and metalophs; conelets never exceeding four in each loph or transverse ridge-crest. Persistent median sulcus. Dental formula incompletely known: I *3% Dp? P? M +2 Ridge-crest formula: P 42; M 13; M 23 M 3,4; Horizon.—Lower to Middle Miocene, Lower Pliocene. Compare Vol. II, p. 1530, and figures 1222, 1224, 1227, also Pl. xrx. Mieration.—(Cf. Vol. I, pp. 132, 133): “Springing from unknown ancestors of the Eocene-Oligocene of North Africa, the true Mastodontine appear in the Lower Miocene Miomastodon |M. depereti| of western Eurasia [France] and soon find their way eastward into North America, appearing in the Middle Miocene species of Nevada, Miomastodon merriami, and in the Lower Pliocene species of western Nebraska, Pliomastodon matthew, meanwhile leaving behind in Austria-Hungary their relatives, Miomastodon tapiroides americanus and Pliomastodon americanus praetypica, which give rise to the rare true Mastodon of southern Russia, referred to ‘Mastodon ohioti- cus’ by Pavlow [and by the present author to Mastodon pavilowi]. Rarely found in northern and western North America, they multiply rapidly in the favorable forests of the middle and eastern United States in the typical form Mastodon americanus.” Fossil remains are found even as far northeast as Nova Scotia and New Brunswick, as evidenced by a recent (1936) discovery at Hillsboro, N. B., and identified by Director William MacIntosh of the New Brunswick Museum as Mastodon americanus. Superfamily: MASTODONTOIDEA Osborn, 1921. Family: MASTODONTID Girard, 1852 (Syn. in part MAMMUTID& Cabrera, 1929) Subfamily: ZYGOLOPHODONTIN& Osborn, 1923 Genera: Zygolophodon Vacek, 1877; Turicius Osborn, 1926 (Cf. Vol. I, pp. 191-223, 282, 690, 697-699, also Pls. 11, 111, x, x1, and Figs. 137, 159; Vol. II, p. 1530, and figures 1222, 1227, also Pl. x1x) When in 1923 Professor Osborn created his subfamily Zygolophodontine, he had not observed the pronounced differences in the molar structure of species of Zygolophodon Vacek, 1877, which later (1926.706, p. 3) actuated him 1576 OSBORN: THE PROBOSCIDEA to describe a new genus, Turicius. Especially noticeable were (1) the strong subdivision of the main cones of certain species (Zygolophodon) into from four to six subequal conelets, whereas in other species (Turicius) the cones subdivided into from five to nine conelets, and even into twenty-five (see Pl. 11, pp. 184, 135); also (2) the presence of strong ‘trefoil spurs’ on the supero-internal and infero-external cones of the latter (Turiczus) in contrast to the absence (or vestiges) of these spurs in the former (Zygolophodon). The species included within Turicius show several resemblances to those included within the genus Serridentinus (e.g., the trefoil spurs just mentioned), but in many other characters Turicius is clearly separable from Serridentinus and more closely related to Zygolo- phodon. The history of the Zygolophodontine will be found in detail in Volume I, Chapter VII, covering the period from 1770 to 1936, in which is set forth the grounds on which this subfamily is separable from the Mastodontine, to which it is most nearly related. In brief, the median longitudinal sulcus (Fig. 89) of Mastodon americanus disappears in Zygolophodon and transversely perfected crests evolve; in Mastodon the main cones feebly subdivide at the summits into two conelets,’ progressive to four (Mzomastodon), whereas in Zygolophodon there are four to six main conelets, and from five to nine and even twenty-five in certain specimens of T'uricius. In other words, from the principal characters of the type grinding teeth we observe (cf. Vol. I, p. 199) “that the universal generic character both of Zygolophodon and of Turicius is the absence or reduction of the longitudinal sulcus, fissure, or commisure, which Hays pointed out as the chief character distinguishing his type (Fig. 154) of M. [Zygolophodon| borsoni from M. americanus. The absence or reduction of this sulcus distinguishes all the ascending species of Zygolophodon and of Turicius, from early Miocene to late Pliocene or early Pleistocene time, from all the ascending or ancestral stages of Mastodon in the corresponding period. We also observe that the fission of the cones into ‘conelets’ is rapidly progressive and distinct in Zygolophodon and in Turicius, whereas it is decidedly retarded in Mastodon. |The final stage of evolution, however, in the Mastodon acutidens molar (Fig. 656) converges towards the final evolution stage in the zygolophodont molar (Turicius virgatidens—Fig. 168).| In many other features... the progressive dental and skeletal evolution of Zygolophodon parallels that of Mastodon. ...(1) the abbreviation of the jaws, (2) the suppression of the premolar dental succession, (3) the reduction of the lower incisive tusks, (4) the progressive abbreviation of the cranium. ... From the fact that the Zygolophodon molars are relatively broader and shorter than those of Mastodon, it is probable that the cranium of Zygolophodon was more brachycephalic than that of Mastodon.” The collateral relationship of the Zygolophodontinz to primitive species of Stegodonts (Stegolophodontine, Stegodontinz) of Asia is treated in Volume I, p. 195, and in Chapter XIV of the present volume, p. 819. Such relationship was first adumbrated by Schlosser as early as 1903, and was followed in 1917 by the suggestion of Schlesinger that primitive European species may have given rise to primitive species of Stegodonts of Asia to which he gave the new generic name Stegolophodon—both of which suggestions Professor Osborn was inclined to favor. An excellent columnar presentation of the outstanding characters of the grinding teeth of the Mastodon, Zygolophodon, Turicius, and Stegolophodon phyla is given on page 211 of Volume I, supplemented by Plates 1-1v (pp. 134-135) demonstrating the molar evolution. Reference is made in the last column to a description of the collateral ancestor of Stegodon in Chapter XV; this should read Chapter XIV, as in revising the present volume it was found desirable to rearrange the chapters somewhat (see pp. 819-822 above). Professor Osborn made a further suggestion on page 202 of Volume I, to the effect that, owing to certain strong resemblances (e.g., supero-internal and infero-external trefoil spurs) between the grinding teeth of Turicius and Serridentinus, despite the entirely different lower tusks in the two genera, there might be an affinity of the See, however, Maslodon acutidens Osborn (described on pp. 696, 697 of Vol. I of the present Memoir) in which as many as twenty-one conelets may be counted on certain of the lophs.—Editor.| SUMMARY 1577 genus Turicius to Serridentinus which a fuller knowledge would demonstrate. Among the distinguishing charac- ters, in addition to the differences in the tusks, he mentions (p. 223) the development of a pure transverse crest in the molars of Turicius, as against the development of corresponding conelets and crests in the American Serridentinus in which the cones and conelets do not become connate and remain quite distinct at the summit, also the progressive strengthening of the trefoil spurs in Serridentinus, which become more and more prominent and conspicuous in progressive stages, surmounted with small conelets, whereas in progressive stages of Turiczus, like 7. atticus and T. virgatidens, the trefoil spurs are reduced to fine ridges accented with numerous small conelets. See, however, his final conclusions on pages 1568, 1569 above, namely, that the Twrictus phylum is entirely distinct. A condensed summary of the contrasting characters in the two genera Zygolophodon and Turicius is as follows: Zygolophodon, typical ‘yoke-crested mastodont’ of Eurasiatic Miocene to Pleistocene. Cranium and skeleton relatively unknown, probably similar to Mastodon americanus. Symphysis more abbreviated than in Turicius. Superior tusks rounded, enamel band disappearing early, before Lower Miocene time. Inferior tusks reducing early; no enamel (Z. pyrenaicus). Grinders permanently blunt, brachyodont (Z. borsoni), not becoming subhypsodont. Ridge-crests or lophs directly transverse, not arched, each loph divided into four to six distinct subequal conelets. Conules or tubercles in median valleys disappear- ing in early stages (vestigial conules in anterior valley of Z. pyrenaicus). Trefoil spurs vestigial or absent. Median longitudinal sulcus disappearing, primitive, vestigial. Fifth inferior crest, i.e., pentalophid, slowly progressive. Intermediate molars trilophodont (fide Schlesinger). Suppression of the premolar dental succession. 2-4 7 1-3 Dental formula: I $33 Dp = M 7 Ridge-crest formula: Dp 2; Dp 3; Dp 43 M 13 M 23: M 3¢%4 Horizon.—Lower Miocene to Upper Pliocene. Compare Vol. II, p. 1530, and figures 1222, 1223, 1227, also Pl. xix. Turicius, sharp-crested ‘Zurich mastodont’ of the European Miocene and Pliocene. Cranium and skeleton relatively unknown. Symphysis progressively pointed and reduced in length, horizontal not decurved. Superior tusks oval in section, rodlike, with sharply defined enamel band except in Turicius virgatidens. Inferior tusks without enamel, straight, suboval in section, undergoing progressive reduction. Molar ridge-crests sharply transverse and elevated, with conelets progressive from four to twenty-five. Median longitudinal sulcus vestigial in early stages, completely disappearing in progressive stages. No median conules. Progressively strong trefoil spurs on superior internal cones and on inferior external cones. Conelets increasingly connate at summit, rising into sharp, subhypsodont transverse crests. Postero-inferior molars with four well-developed crests, the tetartolophid slowly progressive, the pentalo- phid rudimentary but progressive in the higher stages. Gradual repression of the premolar dental suc- session, as in Mastodon americanus and Zygolophodon borsont. Dental formula: Di = I = Dp = Mi Ridge-crest formula: Dp 2°; Dp 323; Dp 43 M 13 M 23? M 333, Horizon.—Lower Miocene to Middle(?) Pliocene. Compare Vol. II, p. 1536, and figures 1220, 1222, 1227, also Pl. xx. Micration.—The types of Zygolophodon and Turicius, as known up to the present time, occur in southern and central Europe. As will be seen by referring to figure 159 and Plate xix, a referred specimen of T'uriczus comes from North Africa, also, according to Matsumoto, one of Zygolophodon from Japan. Recently (cf. pp. 698 and 699 of the present Memoir) Doctor Hopwood has described a juvenile mandible regarded by him as belonging to Mastodon americanus but according to Professor Osborn referable to T'uricius, also a molar of Zygolo- phodon borsoni, both from China. Zygolophodon borsoni is relatively abundant in Siberia. Again consulting figure 159 and Pl. xrx, it would seem that the trend of migration was from North Africa, through southern and central Europe eastward to Siberia, China, and Japan. The Zygolophodontine apparently never reached America. 1578 OSBORN: THE PROBOSCIDEA Superfamily: MASTODONTOIDEA Osborn, 1921. Family: MASTODONTID: Girard, 1852 (Syn. in part MAMMUTIDA Cabrera, 1929) Subfamily: STEGOLOPHODONTIN® Osborn, 1936 Genus: Stegolophodon Schlesinger, 1917 (Cf. Vol. I, p. 700, Pls. tv, x1 and Fig. 661; Vol. II, Chap. XIV, p. 837, also p. 1531, and Figs. 691, 1221, 1223, 1228, Pl. xix) The subfamily Stegolophodontinz, type Stegolophodon Schlesinger, 1917, was defined by Professor Osborn as “‘transitional in molar tooth structure between the Mastodontoidea and the Stegodontoidea’’—the latter a superfamily first created by him to embrace both Stegolophodon and Stegodon (see Osborn, 1935.97, fig. 2). On further study, however, the median sulcus characteristic of the Mastodontine was found to persist in the anterior ridge-crests of Stegolophodon, thus pointing toward a very remote relationship between the Mastodontoidea and the Elephantoidea—indeed, a comparison shows that the most primitive elephantoid molar known, that of Archidiskodon proplanifrons of South Africa (subfamily Mammontine), is, in section, very like the molar teeth of a Mastodon (subfamily Mastodontinz), that is, without trefoils or central conules blocking the valleys. Professor Osborn, therefore, became convinced that Stegolophodon should be removed from the Elephantoidea and placed as a terminal member of the Mastodontoidea, leaving the genus Stegodon as the sole generic representative of the superfamily Stegodontoidea. The reader will find a detailed account of the phylogeny of the Stegolophodonts and Stegodonts in Chapter XIV of the present volume, wherein they are treated together according to Professor Osborn’s original views, as unfortunately he had not reached this chapter in his final revision. We may summarize the distinctive characters of Stegolophodon, as follows: Stegolophodon, ‘roof-crested pro-stegodont’ parallel and collateral in evolution with the true Stegodon of eastern and southern Eurasia. Cranium low, dolichocephalic, suggesting the primitive type of skull and tusks seen in Trilophodon and Serridentinus. Mandibular symphysis not certainly known, but probably short and tuskless. Superior tusks straight with broad enamel band. Inferior tusks undiscovered. Lophs as in Mastodon and Zygolophodon, tendency to form from four to six trans- versely arranged cones and conelets (conelets somewhat irregular) and to consolidate into ridge-crests; molar pattern transitional between the Zygolophodon type and the Stegodon type; no median conules and no trefoil spurs. Closure of enamel in base of transverse valleys, that is, V-shaped; enamel thick; cement usually absent. Dental formula: I °?-°Dp #4 M 22 Ridge-crest formula (based largely on Stegolophodon latidens) : Dp 2; Dp 33 Dp 44 M 1*%4 M 2 #5 M38 248+ Horizon.—Mio-Pliocene to Upper (?) Pliocene [Lower Pleistocene?]. Compare Vol. II, p. 1531, and figures 1221, 1223, 1228, also Pl. x1x. ORIGIN AND Micration.—It was doubtfully suggested by Schlosser in 1903 (1903, p. 191) that west European species, originally described as Mastodon turicensis |M. tapiroides, M. pyrenaicus, ete., separated as Zygolophodon by Vacek in 1877 and by Osborn in 1926 as Turicius (M. turicensis, M. tapiroides) and Zygolophodon (M. pyrenai- cus)|, may have given rise to primitive Asiatic species [such as Stegolophodon cautleyi|; also that M. turicensis |= Turicius tapiroides| of the Lower Miocene of Europe may have given rise to the M. [Stegolophodon|] latidens of the Lower Pliocene |?Lower Pleistocene] of Asia from which in turn sprang off the true Stegodonts, such as Stegodon insignis. Professor Osborn’s final opinion regarding the relationships of the Stegolophodontine (see pp. 197 and 819-822 above) would seem to be in favor of Schlosser’s suggestion of 1903, also that of Schlesinger of 1917, that primitive species of Europe [Mastodon (Stegolophodon) sublatidens| may have given rise to primitive 4 STEGODON A/RAWANA JAVA STEGODON TRIGONOCEPHALU* UAVA STEGODON PINIORENS/S S/SWAN, INDIA STEGODON ORIENTALIS GRANGERI SZECHUAN, CHINA STEGODON /NS/GNIS STEGODON GANESA S/WALIKS, /ND/IA S/WALIKS, INO/IA \y STEGODON BOMBIFRONS S/IWAL/IKS, INOIA EQUATOR Origin, Migration and Evolution of Stegodon PLATE XX Geologic range: Stegodon, Lower Pliocene [?Lower Pleistocene, see p. 824 above], Middle Pliocene to Upper Pleistocene. Since this map was prepared the range of Stegodon has been extended into Africa, as far south as Kaiso, on Lake Albert. SUMMARY 1579 species of Asia to which Schlesinger has given the generic name Stegolophodon (cf. Vol. I, pp. 195, 197, and Pls. II-Iv, also p. 700). The Stegolophodonts are not known to have reached America. Fossil remains have been found in Austria, in Perim Island, the Punjab and the Siwaliks of India, also in Burma, Japan, and Borneo. Their possible line of migration in indicated in figure 1228 and on the accompanying Plate xrx. STEGODONTOIDEA Superfamily: STEGODONTOIDEA Osborn, 1935, 1936. Family: STEGODONTIDA Young-Hopwood, 1935 Subfamily: Srecopontinz Osborn, 1918, 1921 Genus: Stegodon Falconer and Cautley, 1847, 1857 (Cf. Vol. I, Pl. x1, and Chap. XTV of the present volume, pp. 807, 853, also Figs. 686-688, 691, 1221, 1223, 1228, and Pl. xx) As fully stated in Chapter XIV above, Professor Osborn separated the true Stegodonts from the Elephanto- idea, giving them a new superfamily name, Stegodontoidea, which includes the genus Stegodon only of the sub- family Stegodontine, the genus Stegolophodon having been removed by him to the Mastodontoidea, subfamily Stegolophodontinz. The family Stegodontide, first mentioned by Dr. C. C. Young in 1935 and fully defined by Dr. A. Tindell Hopwood later in the same year, embraced both Stegodon and Stegolophodon. In the present Memoir this family is logically included in the superfamily Stegodontoidea, although Professor Osborn never indicated such reference other than in his chart of 1935 (Osborn, 1935.937, fig. 2), and even in this instance he also included both genera, Stegodon and Stegolophodon. The separation of the Stegodontoids from the Elephan- toids was based chiefly on the fact that in section the molars of the former showed the valleys separating the adjacent ridges to be closed or V-shaped at the bottom and those of the latter to be open or U-shaped. Another determining factor was the extremely short face of the Stegodontoids which, in his opinion, could not have given rise to the longer face of the Elephantoids. With this classification in mind, the following characters have been compiled pertaining solely to the Stegodontinz, genus Stegodon. Stegodon, so named because of the resemblance of the toothed ridges of the grinding teeth to a series of roof-gables; more primitive than the true elephants. Skull and tusks do not lead into either the Elephantine or the Mammontine types. Cranium brachycephalic, brachyopic, of mastodontoid (Stegodon bombifrons), to extremely abbreviated, female? (S. insignis), more elongated, male? (S. ganesa), more triangular (S. trigonocephalus) form; rostrum elongated to support the tusks; palate short, depressed well below the occipital condyles (bathycephalic); narial openings elevated. Jaws greatly abbreviated. Superior tusks horizontal or subhorizontal in direction, parallel, and slightly upcurved, without trace of enamel band, attaining great dimensions (S. ganesa). Inferior tusks disappearing very early. Grinding teeth short crowned; progressive heightening of ridge-crests (brachyodont to subhyp- sodont), the distinctive feature being the rapid multiplication of transverse crests by the addition of crests posteriorly; increasing curvature of the occlusal surface; enamel valleys closed or V-shaped at the bottom instead of U-shaped as in the elephantoids, filling with cement. Particularly interesting and significant is the transformation of the original cones by binary or rarely by ternary fission into conelets (maximum 20+ in S. airdwana). Ridge-crests intermediate between Stegolophodon and Archidiskodon planifrons types. Probably browsers rather than grazers. Dental formula: Di *** I =" Dp = M = Ridge-crest formula: Dp 2°* Dp 3&3 Dp 423; M 1%2 M 2&2 M 3224 Horizon.— Lower Pliocene [?Lower Pleistocene, see p. 824 above], Middle Pliocene to Upper Pleisto- cene. Compare this chapter, p. 1539, and figure 1228. OrIGIN AND Micration.—Professor Osborn’s views on the origin of the true Stegodonts are given on page 25 of Volume I of the present Memoir, as follows: ‘It has been assumed by practically all palzeontologists that the Elephants were descended from the Stegodonts. . . . The Stegodonts were of independent origin and formed an 1580 OSBORN: THE PROBOSCIDEA independent parallel branch terminating in the highly specialized Elephas [Stegodon| aurore Mastumoto from the Upper(?) Pliocene of Mt. Tomuro, Japan, now . . . separated by Osborn as the Stegodontoidea superfam. nov.” Also on page 853 of the present volume, where the generic characters of Stegodon are enumerated, occurs the following, agreeing in substance with the above conclusions: ‘‘Phylum parallel to that of the true Archidiskodon and Elephas, not directly ancestral, readily distinguished by cranial and dental characters.” It will be noted on page 819 above that Schlosser suggested the possible derivation of the true Stegodonts from certain Miocene species of western Europe since referred by the present author to the Zygolophodontine, that is to say, these may have given rise to the Lower Pliocene [?Lower Pleistocene] M. [Stegolophodon| latidens of Asia, from which in turn sprang off the true Stegodonts, such as Stegodon insignis. Professor Osborn regarded this phylogenetic problem as of such importance that he stated on page 197 of Volume I that it would be treated more fully in Volume II, under the Stegodontinz [of the Stegodontoidea superfam. nov.]. It is with regret that we are unable to carry out his intention in this respect, as he left no record of his final views on the subject other than is given above. The accompanying migration map (PI. xx), therefore, omits the hypothetical European point of departure and indicates the Siwaliks of India as the center of dispersal, from which radiate three distinct routes, namely, to Burma and Java, to central China, and to the northern part of China onward to Japan. Up to the present time Stegodon has not been found in the Western Hemisphere. See Chapter X XII, p. 1436 above, where Dr. Edwin H. Colbert cites from a letter of January 14, 1938, from Dr. A. Tindell Hopwood, in which the range of Stegodon is extended into Africa as far south as Kaiso on the eastern shore of Lake Albert. ELEPHANTOIDEA It will be recalled that Professor Osborn long held the opinion that Africa was the center of adaptive radia- tion—or the “‘Shomeland’’— of the Proboscidea. With this thought ever in mind, he pursued his researches over a period of nearly thirty-five years, and, happily, the results, so far as can be determined at the present time, have confirmed his views, as will be seen from the following citation from his last paper on the subject, ‘“The Ancestral Tree of the Proboscidea. Discovery, Evolution, Migration and Extinction over a 50,000,000 Year Period,” 1935.937, p. 410: In the Elephantoidea we have discovered a brilliant example of aristogenic growth and extensive migration in the evolution of Archidiskodon proplanifrons, a very primitive stage found in the Vaal River gravels of South Africa, into the Archidiskodon maibeni of Nebraska. Archidiskodon proplanifrons has a molar crown pattern like that of a mastodont, with an enamel length of 690 mm.; Archidiskodon maibeni has eighteen tall ridge plates and an estimated enamel length of 9000 mm. This is the first time that the evolution of the elephantoid molar from a theoretic mastodont prototype has ever been actually demonstrated .. . The 15,000 mile migration of Archidiskodon, from the Vaal River of South Africa through intermediate stages in North Africa, France, Italy, Britain, India, then the long geographic break to the Niobrara River in Nebraska where Leidy discovered these animals in 1858, is the most remarkable trek of any species of plant or animal thus far discovered in the life history of the earth. The by-product is the fact that these stages give us an unprecedented opportunity of measuring precisely the actual rate of evo- lution between Upper Pliocene and Middle Pleistocene time. The enamel length increasing from 690 mm. to an estimated 9000 mm. affords an evolution of approximately 15 mm. of enamel length per thousand years. In this introduction to the subfamily members of the Elephantoidea it may not be amiss to give the present author’s definitions of alloiometrons and aristogenes, as well as an outline of his ‘‘elephant enamel or ganometric method” of measuring Pleistocene time, particularly in view of the fact that reference has been made in the fore- going quotation to aristogenic and ganometric usage in his researches. Compare page 1582, also figure 300 of Vol. I, and figures 1231, 1239, 1240 of this chapter. “‘Aristogenes are new adaptive units originating directly in the geneplasm and slowly evolving into important functional service.” Hence aristogenesis (first known under the term “definite variation,” then as “‘rectigradation’’) SUMMARY 1581 ‘9s a creative process from the geneplasm of entirely new germinal biomechanisms; the process is continuous, gradual, direct, definite in the direction of future adaptation.” In time contrast to alloiometrons (that is, changes of proportion or intensity which may be expressed in measurements and indices, and which appear to be immediate and more or less temporal adaptive reactions to new habits), aristogenes are secular, appearing very slowly in the course of long periods of geologic time. Lines of ordinal, family, generic and specific descent may be distinguished by the potentiality of certain new geneplasmic aristogenes. (Modified from Osborn, 1934.922, pp. 202, 210.) NEBRASKA UPPER PLEISTOCENE High-crowned giant Archidiskodon 18 lofty ridge-plates. Enamel length = mm APCHIDISKODON (iMPERATOR) MAIBENI NEBRASKA 4068MM, 13/4)0" 1931 NEBRASKA LOWER PLEISTOCENE Medium-crowned Archidiskodon PLEISTOCENE OF AMERICA /4 ridge-plates. Enamel length = mm ARCHIDISKODON MERIDIONALIS NEBRASCENSIS 3672mm2‘n" 1933 S. NEBRASKA FRANCE LOWER PLEISTOCENE Medium-crowned elephant molars /3-/4 ridge-plates Enamel length=2824 mm ARCHIDISKODON MERIDIONALIS 372mm. 12/2" 1931 SFRANCE DURFORT FRANCE LOWER PLEISTOCENE Zow-crowned elephant molars LOWER PLEISTOCENE OF FRANCE mm /3 ridge-plates. Enamel length= ARCHIDISKODON PLANIFRONS R 3745 mm,\2/3n2" e 1931 FRANCE CHAGNY INDIA. UPPER PLIOCENE Molars of primitive elephant type TYPE ARCHIDISKODON PLANIFRONS 193) INDIA SIWAUKS 1980mM,6'6’s (0-12 ridge-plates. Enamel length = 15/5 mm. SOUTH AFRICA. MIDDLE PLIOCENE TYPE Molars transitional from mastedon to elephant ARCHIDISKODON SUBPLANIFRONS SOUTH AFRICA PT SEG 6 ridge-plates. Enamel length=650mm. “Tes SOUTH AFRICA. MIDDLE PLIOCENE Mastodon-like molars PLIOCENE. AFRICA_INDIA TYPE ARCHIDISKODON PROPLANIFRONS 1450mm 4'9"e SOUTH AFRICA Sta ridge-plates. Enamel length= 690mm. ARCHIDISKODON: THREE-MILLION-YEAR EVOLUTION AND MIGRATION SOUTH AFRICA TO AMERICA AND MEXICO HFO. 1935 Fig. 1239. Geneplasmic evolution of the archaic-toothed mammoths during a three-million-year period so far as known to April, 1935. Twenty-seven years of continuous exploration and research yielded all the ascending stages in the geneplasm of this archaic-toothed mammoth. After Osborn, 1938.1, fig. 11. For example, compare Osborn, op. cit., p. 221: “In the elephantoid division the low transverse ridge-crest is perfected in the upper Pliocene of the African ancestral elephants (Archidiskodon). The unique aristogenic Allometric Adaptations of the Limbs Osborn - Gregory. 1929 FEMUR Shortening with increasing speed Lengthening with 27.1% of total length increasing weight 48.6% of total length Lengthening with increasing Speed 41.1% of total length Shortening with ALLOMETRONS: Adaptive Speed and Weight Proportions (Left) Horse. Cursorial: Pes 41% of total length: Femur 27-1% of total length (Right) Elephant-Graviportal: « 17/% « — # ” « FB6% « “ Analogous ALLOMETRONS Speed and Weight evolve in Mammals and Keptiles Fig. 1240. (Left) Hquus: Alloiometrons of speed; (Right) Hlephas: Alloiometrons of weight. After Osborn, 1938.1, fig. 6. Abbreviating femur of the horse, 27.1 % of total length. Elongating femur of the elephant, 48.6 % of total length. Tibia of the horse relatively constant, 31.7 % of total length. Tibia of the elephant relatively constant, 34.3 % of total length. Elongating pes of the horse, 41.1 % of total length. Abbreviating pes of the elephant, 17.1 % of total length. Similar cursorial alloiometrons evolve in all quadrupeds attaining speed, irrespective of phyletic relationship. Similar graviportal alloiometrons evolve in quadrupeds irrespective of mammalian (Proboscidea) or reptilian (Sauro- poda) affinity. 1582 OSBORN: THE PROBOSCIDEA potentiality of the transverse ridge-crests to convert pairs of cones by transverse binary fission into transverse ridges is followed by the elevation of these ridge-crests into the elephantine ridge-plates, as first manifested in the roof-toothed Stegodon of southern Eurasia. It is parallelee in the distinct Archidiskodon ridge-plated stock of the upper Pliocene of South Africa. This dark continent gave rise to the world dominant elephantoid division of the Proboscidea . . . Aristogenesis combined with alloiometric extension is carried to the biomechanical extreme in the divergently adaptive grinders of the three mammoths [Archidiskodon, Parelephas, Mammonteus|. The contrasts in the total length of the enamel foldings of the gigantic Archidiskodon (8,000 mm), of the gigantic Parelephas (10,000 mm), of the relatively small Mammonteus (6,000 mm) are coordinated with the relative intensities of their struggle for existence.’’ To summarize, in Professor Osborn’s own words (op. cit., p. 234): In biomechanical evolution there are two distinct processes. The one, long known, consists in the alloiometric modification of existing adaptations as in changes of proportion and of function. The other, discovered in course of researches on the phylo- geny of the horses, titanotheres and proboscideans, consists in the gradual geneplasmic origin of new and distinct adaptations; it is to the latter originative and creative process that the term Aristogenesis is applied. Both processes become part of the hereditary equipment of the organism. We may now proceed to epitomize the three subfamilies embraced within the superfamily Elephantoidea, family Elephantide, namely, the Mammontine, Loxodontine, and Elephantine, in uniformity with the fifteen subfamilies of the Mastodontoidea, and the single subfamily each of the Moeritherioidea, Deinotherioidea, and Stegodontoidea. Superfamily: ELEPHANTOIDEA Osborn, 1921. Family: ELEPHANTID Gray, 1821 Subfamily: Mammontina Osborn, 1921! Genera: Archidiskodon Pohlig, 1885, 1888; Metarchidiskodon Osborn, 1934; Parelephas Osborn, 1924; Mammonteus Camper-Osborn, 1788-1924! (Cf. the present volume, Chapters XVI, XVII, and XVIII, also Pls. xx1 and xx, and Figs. 815, 822, 933, 1006, 1221-1225, 1228) The name Mammontinze Osborn, 1921, was substituted for Euelephantinze Osborn, 1918, owing to the fact that the genus Hwuelephas Falconer, 1857, was invalid (see Chap. XIX, p. 1175 above). Originally this subfamily, aside from the mammoths, doubtfully included Elephas hysudricus of India, subsequently found to belong to the Elephantine, genus Hypselephas Osborn, because of profound cranial and dental differences. A definition of this subfamily reads as follows on page 32 of Volume I: Subfamily Mammontine (Mammoths)—Osborn, 1921.515, p. 1. Of close original affinity to the Elephantinae, including (a) the southern mammoths Archidiskodon planifrons and A. meridionalis of southern Eurasia, A. ¢mperator of North America, all with broad-plated teeth and few crests; also (b) the northern mammoths which apparently include Parelephas trogontherti of western Europe, P. columbi and P. jeffersonii of North America, and the widespread woolly mammoth (Mammonteus primi- genius) of the northern steppes. Great confusion had existed for years in the determination of species of Elephas columbi, E. imperator, and E. primigenius. Superficially these three species are so similar that Cope in 1889 referred the remains of a fine skull of Archidiskodon from Texas to Elephas primigenius columbi, and the present author also saw only resem- blances to Elephas columbi in the fine Indiana skeleton (now the type of Parelephas jeffersonii), whereas Hay referred the same skeleton to Hlephas primigenius. Consequently Professor Osborn made a careful comparison of "Should Mammonteus prove to be invalid, this would leave the subfamily Mammontine without a type genus.—Editor.] ARCHIDISKODON MERIDIONALIS, vat_o'4RNO, ITALY LEST pes Lact ARCHIDISKODON EXI/LI ARCH/IDISKODON SANTA ROSA Harvie IMPERATOR CALIFORNIA TEXAS ARCHIDISKODON MERIDIONALIS NEBRASCENS/S NEBRASKA ARCH/DISKODON MERIDIONALT. DURFORT, FRANCE ae ARCHIDISKODON PLANIFRONS, CHAGNY, FRANCE ms ARCHIDISKODON IMPERATOR SCOTTI NEBRASKA / eo PLANIFRONS, S/IWALIKS, INDIA ra ARCHIDISKODON SUBPLANIFRONS, VAAL RIVER, SOUTH AFRICA ARCH/IDISKODON PROPLANIFRONS, GONG-GONG, SOUTH AFRICA a 7590 105" 20" ales al EQUATOR ve Origin, Migration and Evolution Ae oxlrchidiskodon, «* Aletarchidiskodon PLATE XXI Geologic range: Archidiskodon, Middle Pliocene to Upper Pleistocene; Metarchidiskodon, Lower(?) Pleistocene. ‘ . ix a e _ ce i Ant iA ace! | A SUMMARY 1583 both type and referred specimens of these three species, which he found were united by distinctive cranial charac- ters but separated by various types of dental characters, resulting in their reference by him to the following genera (see his articles in Novitates, 1922.555, 1924.633, 1925.662): Elephas columbi=Parelephas columbi and P. jeffersonii Elephas imperator = Archidiskodon imperator Elephas primigenius= Mammonteus primigenius In the present Memoir the Mammontinz embrace four genera, namely, Archidiskodon Pohlig, 1885, 1888, the most primitive member, with the genotypic species, Hlephas meridionalis Nesti, 1825, of the Val d’Arno, Italy, and FE. planifrons Falconer and Cautley, 1846 [1845] of India; Metarchidiskodon Osborn, 1934, genotype Loxo- donta griqua Haughton, 1922, of the Vaal River, South Africa; Parelephas Osborn, 1924, genotype Hlephas jeffersonii of Indiana, and Mammonteus' Camper-Osborn, 1788-1924, the history and outstanding characters of which are given in the following synopses. Archidiskodon (referring to the archaic molar ridge-plates) of the southern and south temperate zones; also known as the Southern Mammoth, in reference to its remote relationship to the Northern Mammoth. Cranium foreshortened and broadened (brachycephalic), deepened (bathycephalic), heightened (hypsicephalic); through hypsicephaly, orbits and occipital condyles approximated, i.e., brachycranial (A. meridionalis), compressed fore and aft (cyrtocephalic), occipitofrontal apex vertically heightened (acrocephalic); far closer to Mammonteus and Parelephas than to either Elephas or Loxodonta. Nasals pointed. Forehead flattened (A. planifrons), concave (A. meridionalis, A. imperator). Mandible short; prolongation and beaklike depression of the symphysis (A. planifrons); without beak, blunt, obtuse (adult male A. meridionalis—fide Depéret and Mayet, 1923, p. 156); relatively long and shallow (A. hayi); symphysis prominent, ramus relatively slender (primitive stage). Superior tusks large (max. 13+ to 16+ feet in length), incurved, crossing in old males; no inferior tusks. Molars short, subhypsodont; subloxodont in primitive species; ridge-plates extremely broad, widely separated, enamel borders thickened, more or less crenulate; cement usually very thick; A. planifrons without cement outer coating. Ridge-plate formula slowly progressive from M 3° (A. proplanifrons), to M 3 +°+ (A. planifrons), to M 3534, (A. meridionalis), to M 3}235( A. tmperator); succession of premolars, P 4, P 4 (A. planifrons). Dental formula (A. planifrons): 1°°" Dp 34 P $$ M3. This premolar succession does not oceur, so far as known, in any other species of the Elephantide. Ridge-plate formula (A. planifrons): Dp 2* Dp 38% P 3; P 4*** Dp 48} M 1% M 25 M 329+ Horizon.—Middle Pliocene to Upper Pleistocene. Compare this chapter, p. 1540, and figures 1221, 1224, 1239, also Pl. xx1. Dr. Hans Pohlig, to the best of our knowledge, was the first to assign the name Archidiskodonten to elephant molars with archetypal ridge-plates, specifying Elephas planifrons Falconer and Cautley, 1846 [1845], and E. meridionalis Nesti, 1825, as types. This was in 1885. Subsequently (1888, p. 138) he supplemented his descrip- tion, giving EL. meridionalis only as the type, and on page 252 concluded with the introduction of three new generic names— 7 So iio PARELEPHAS f SYEFFERSON// GRANT CO., INDIANA ae PARELEPHAS TROGONTHER// PARELEPHAS COLUMBI/ MOSBACH, GERMANY GEORG/A CIr|? PARELEPHAS TROGONTHERII, rausacn, GERMANY 165" 120" Origin, Migration and Evolution of o Parelephas, * Mammonteus, a Elephas PLATE XXII Geologic range: Parelephas, Upper(?) Pliocene [Lower(?) Pleistocene, see footnote 1 on p. 1049 above] to Upper Pleistocene. Mammonteus, Upper Pliocene to late Pleistocene. Hlephas, Upper Pleistocene and Recent. Fossil representatives of the Elephantine phylum are Hypselephas of India (Lower Pleistocene) and Platelephas also of India (Upper Pliocene or Lower Pleistocene), restorations of which do not appear on the accompanying plate, nor elsewhere in this Memoir, as they were never prepared by Professor Osborn. ew SUMMARY 1585 cephalic), space between condyle and orbit broader than in Mammonteus primigenius; anterior narial opening broad and widely open; divergent maxillo-premaxillary region, tusk sockets less elongate and less parallel than in Mammonteus. Mandible robust, short and deep (bathycephalic); ramus depressed with rounded inferior border; rostrum prominent. Superior tusks long, with remarkable incurvature, crossing in old age (Parelephas jeffersoniz); shorter, more robust (P. floridanus). No inferior tusks. Molars relatively narrow as compared with those of Archidiskodon, with enamel of intermediate thickness, more or less crimped or sinuous. Ridge-plates arcuate, converging toward the summit; consequently the ridge-plate compression depends upon the level at which the count is taken—the count ranges in P. jefferson molars from 7 in 100 mm. (base) to 11% (summit) in the inferior molars, from 7 (base) to 10 (summit) in the superior molars. Relatively few ridge-plates in the Upper Pliocene to Middle Pleis- tocene stages (M 3i{}), progressive Upper Pleistocene stages (P. progressus) with multiple ridge- plates (M 333). Thin cement (P. columbi). Adapted to continental plains or steppe environment, grazing and browsing. Dental formula: I 22° Dp =! M 43 Ridge-plate formule: P. columhi: Dp 2% Dp 3? Dp 4,5; M122 M 23338 M 3332 1234 15-16+ P. jeffersonii: Dp 3°* Dp 422 M 1°** M 22822" M 3 25 19e Comparative Ridge-plate Formule of M 3 in Parelephas Parelephas progressus M 3 22 Parelephas columbi IMS ieee Parelephas jefferson Mes. Parelephas intermedius IMLS). eae Parelephas washingtoni M 3 = Parelephas trogontherit Ma3 Parelephas floridanus M 3425 Parelephas trogontherioides M 3 3*% Horizon.—Upper(?) Pliocene |Lower(?) Pleistocene, see footnote 1 on p. 1049 above] to Upper Pleistocene. Compare p. 1540, also figures 1222, 1224, 1225, and Pl. xxn. Professor Osborn introduces his chapter on Parelephas (Chap. XVII) with the following statement: . it appears that Parelephas is linked with Archidiskodon and Mammonteus in its cranial resemblances and great incurved incisive tusks, while in its grinding teeth and ridge-plate formule it is so nearly intermediate between these two genera as to have been mistaken for an actual connecting link. In the present chapter [X VII] it is shown to be an entirely distinct generic phylum which during the more temperate interglacial periods . . . occupied the same geographic range as that of the true woolly mammoth (Mammonteus) during the glacial periods. [Compare Fig. 1241, below. | While not sufficiently distinctive to preclude the subfamily unity of these three genera, there are certain cranial differences, such as the intensification of characters like hypsicephaly, bathycephaly, and acrocephaly. Their dental diversification, however, definitely separates them generically, for example, the broad, widely sep- arated, thickly enamelled ridge-plates of Archidiskodon, as compared with the relatively narrow, more compressed and less thickly enamelled ridge-plates of Parelephas, and the narrow, extremely compressed, thinly enamelled, hypsodont ridge-plates of Mammonteus; also the multiplication of the ridge-plates of M 3, which in Archidiskodon increase from +°* to +235 and in Mammonteus from {23° to #5, whereas in Parelephas the increment is from wits to 2°. It should also be stressed that, contrary to the normal ridge-plate structure, the superior count generally exceeds that of the inferior in Parelephas. The principal characters by which Parelephas crania may be distinguished from Mammonteus crania are summarized by Professor Osborn (Chap. XVII, p. 1051) as follows: (1) In frontal aspect the crania of Parelephas are relatively broader, more spreading, and more brachycephalic than those of Mammonteus, which are deeper and more bathycephalic; (2) in lateral aspect (a) the orbit is more widely separated from the occipital condyle, (b) the occiput is much more convex, thus throwing the occipitoparietal apex farther forward, (c) the height from the occipital apex to the superior molar crowns is less deep, i.e., less bathyeephalic, (d) the apex formed at the summit of the cranium is less acute, (e) the facial front is shorter and more deeply concave, (f) the maxillo-premaxillary sockets are less 1586 OSBORN: THE PROBOSCIDEA vertical and the tusks emerge in a less vertical plane; (3) in frontal aspect (a) the premaxillary sockets are more expanded at the extremities, whereas in Mammonteus they are more elongate and more closely compressed, (b) the transverse diameter of the frontals is relatively broader than in Mammonteus, (c) the anterior nares are proportionately broader transversely and less deepened vertically; (4) in brief, the proportions of the cranium of Parelephas throughout are harmonious with those of the grinding teeth, i.e., less compressed anteroposteriorly, less bathycephalic and less hypsicephalic than those of Mammonteus. This section would not be complete without some mention of Professor Osborn’s clarification of the phyletic position of the true “Hlephas” columbi of Falconer, 1857, from Georgia, and his own ‘‘Elephas’’ |Parelephas| jeffersonv from Indiana described in 1922. Falconer believed E. columbi to be identical with Leidy’s FE. [Archi- diskodon| imperator, and the present author for several years treated it under the genus Archidiskodon but as a species quite distinct from imperator. Falconer was in error, as the two species are distinct, which will be seen by the following comparison; moreover, according to the views of Professor Osborn, they were not geologically contemporaneous, although the late Doctor Gidley [and many others] believed otherwise. Elephas |=Parelephas| columbi Falconer, 1857-1868. Upper Pleistocene of southern United States and of Mexico. Smaller animal, with narrower grinders (Fig. 887), thin cement outer coating; maximum ridge-plate formula, M 3 ;4°-. Enamel ridge-plates arcuate, converging at summit, giving the appearance in extremely worn grinding teeth of being as far apart in mid-section as in A. imperator. Elephas |= Archidiskodon| imperator Leidy, 1858. Lower {Early and Middle Pleistocene, fide Lugn and Schultz, 1934, pp. 373-876] of southern United States and of Mexico. Larger animal in size, with broader grinders (Fig. 887), very broad enamel plates, and heavy cement outer coating; ridge-plate formula, M 3:=3°. Enamel ridge-plates widely separated. The present author was long misled by the widely separated ridge-plates seen in the types of both Elephas columbi and EF. imperator, but with the opportunity for further study through the acquisition of new and rich materials from Florida and the phosphate beds of South Carolina, he perceived that the type of HE. colwmbi belonged within the phylum Parelephas. Again, the type of the true Elephas |Parelephas| columbi of Georgia, Florida, and South Carolina must not be confused with the Elephas [Parelephas| jeffersonii of the northern states, from which it is separable in its much more primitive ridge formula. The species EL. jeffersonii first described by Professor Osborn in 1922 (Osborn, 1922.555, pp. 11-16) was based on a skeleton from Indiana referred to “‘Elephas columbi”’ by Cope and by Osborn, and to H. primigenius by Hay. In his description he gave the distinctive characters of the grinding teeth of each of the species E. imperator, EF. columbi, and E. primigenius, remarking that these diagnoses left without a name the animal which previously had been described in all the literature (excepting Soergel’s Memoir of 1921) as Hlephas columbi. Unfortunately in his original description of . jeffersonii Professor Osborn chose as paratypes a pair of upper and lower molars of both sides from Zanesville, Ohio, described by Warren in 1855 as Elephas ‘““primigenius,” which he finally made the type of a new subspecies, Parelephas jeffersonii progressus, because of the large number of ridge-plates, namely, M 33°. In the meantime the molars of the aged type specimen from Indiana were cut out of the jaw and sectioned, yielding a ridge-plate formula of M 322, still a much higher ridge-plate count than that of the true ‘“Elephas columbi,”’ namely, M 3,\2;. After the establishment, therefore, of Elephas jeffersoni as a species distinet from 2. columbi and from E. primigenius, Professor Osborn (1924.633, p. 4) reached the con- clusion that this species could be placed neither in the phylum of Archidiskodon nor in the phylum of Mammon- teus; consequently he selected it as the type of a new genus, Parelephas. In summation, Elephas columbi of Falconer, with a ridge-plate formula in M 3 of ,°; and an average specific ridge-plate frequency of 5-6 in 100 mm., as compared with 7-9 (max. 113s) in EH. jeffersonii, 5-7 in LH. imperator, SUMMARY 1587 and 8-13 in EF. primigenius, is to be regarded as a distinct species of Parelephas, namely, P. columbi;! HE. columbi (previously described as tne Columbian Mammoth) is now to be known as Parelephas jeffersonii or the Jeffersonian Mammoth. The skeleton from Brevard County, near Melbourne, Florida, now in the Amherst Museum, be- longs to the true Parelephas columbi. Micration.—A very careful comparison of all the known characters of the Elephas trogontherii phylum of Europe and the Elephas jeffersonii of America, especially the cranial characters, established their close phyletic relationship, justifying the linking of the European and American species in the new and distinct genus Parelephas. The low ridge-plate formula of the true Parelephas columbi of Georgia and South Carolina suggested to Professor Osborn the possibility of the early geologic entrance into America of relatively primitive species of Parelephas, a theory which is supported by the primitive character of the lower jaw of P. washingtoni of Whitman County, state of Washington, the molar ridge-plate formula of which agrees quite closely with that of P. intermedius of southern France. Hence the evidence appears to indicate that the ancestors of P. columbi and of P. washingtonii may have passed across Europe and Asia? and migrated far southward in North America during the 2d and even possibly during the 1st Interglacial period, following the wave of migration into America of Archidiskodon. Profes- sor Osborn was sanguine of the discovery (probably in northern Africa) of an ancestral stage more primitive than the Parelephas trogontherioides of the Upper Pliocene’ of Italy. It will be noted from the accompanying Plate xx that members of this Parelephas phylum are recorded from France, Italy, England, Germany, southern Russia, Asia Minor, Canada (Ontario), United States (Washington, Oregon, Montana, Colorado, Iowa, Kansas, Nebras- ka, Indiana, Illinois, Ohio, Kentucky, South Carolina, Texas), Mexico, and French Guiana—the only representa- tive of the Elephantide that succeeded in reaching the South American continent, as far as we know at the present time. Thus we have an elephant phylogeny and migration second only to that of Archidiskodon. Mammonteus, the Northern or Woolly Mammoth, of the northerly and circumpolar zones. Cranium related to that of Archidiskodon and of Parelephas, with fore-and-aft compression, resulting in bathyceph- aly and hypsicephaly, which exceeds by far that of any other proboscidean or other mammalian skull known. Frontals concave, occipital crest greatly elevated, occiput slightly convex. Mandible with excessively short, deeply depressed ramus, terminating in a deflected and extremely narrow rostrum— in short, harmonic with the extreme hypsicephaly and bathycephaly of the cranium. Elevation of coronoid and of mandibular condyle. Anterior nares small. Superior tusks of large proportions, greatly incurved, crossing in old age; vertically placed tubular maxillo-premaxillary insertions of tusks, which are relatively longer, narrower, and deeper than those of any species of Parelephas. No inferior tusks. Molars with relatively numerous ridge-plates (polydiskodonty), Upper Pliocene stage (Mammonteus primigenius astensis), M 34238; typical Upper Pleistocene stage (M. primigenius), M 324; final progressive stage (M. primigenius compressus), M 37,;; broad, hypsodont, with enamel ridge-plates of minimum thickness, more or less crimped or sinuous, in fact, M 3 is the broadest and deepest probosci- dean molar known (compressus). Ridge-plates compressed in typical superior molars to 10-11-12 in 100 mm., in highly progressive superior molars 13 in 100 mm.; in progressive inferior molars 8-9—10 in 100 mm. As in Parelephas, the molars are arcuate at base, more compressed at summit, consequently the ridge-plate count varies and as a rule should be taken at mid-section, both on the internal and ex- ternal sides. Manus pentadactyl with five digital nails (fide Herz, 1902), manus and pes tetradactyl not pentadactyl (fide Salensky,* 1904, p. 86); four digits (fide Pfizenmayer, 1926, p. 239); total phalanges in ferred to Parelephas columbi with its limited ridge formula, M 37§4, actually belong nearer to P. floridanus stage with its more progressive ridge formula, M 33?}.”’—Editor.] *{Parelephas trogontherit is reported by Tokunaga (Amer. Mus. Novitates, 1933, No. 627, p. 2) as occurring at Honshu (Kazusa, Mikawa, Omi, Shinano), Inland Sea, Japan. See figure 1223 in Chapter XXII above, by Dr. Edwin H. Colbert.—Kditor.] ’[Possibly Lower Pleistocene, see footnote 1 on p. 1049 above.—Kditor. |] 4[See Zalensky, Vladimir Vladimirovich, in Bibliography of Volume I of the present Memoir.—Editor.| 1588 OSBORN: THE PROBOSCIDEA each foot reduced to nine in number, e.g., ¢trinivy pas Dietrich (1912) records five digits in the manus and pes of the large and more primitive Upper(?) Pleistocene M. primigenius fraasi. Tail abbreviate, caudals 21. Dental formula: I =" Dp #} M#8 Ridge-plate formula: Dp 24 Dp 3s Dp 435 M 133 M 2t® M 334 (typical M. primigenius) Horizon.—Upper(?) Pliocene |?Lower Pleistocene] to late Pleistocene. Compare page 1541 of the present volume. The true mammoth is the only extinct proboscidean of which the characters of the soft parts and of the hairy and woolly covering, as well as the nature of the food (with the exception of the Mastodon), are fully known. It is probable that these northerly or woolly mammoths were the first mammalian fossils of northern Eurasia to be discovered and recognized as extinct; the earliest descriptions are naturally lost in obscurity. The typical or true mammoth was found in Siberia long before it was recognized in western Europe. Blumenbach had in mind both Siberian and North German specimens in defining Elephas primigenius in 1799, also Cuvier had the Siberian mammoth in mind in defining Elephas mammonteus (1796 MS., 1799). Out of a host of names applied to the northern mammoth between the years 1696 and 1888, the species Hlephas primigenius of Blumenbach alone sur- vives and is accepted in the scientific literature of the entire world. As to the genus, nearly a century and a half of research since 1799 ‘‘demonstrates that the woolly mammoth belongs not to Elephas, but to a genus of its own, distinct by all the canons of nomenclature from the true modern Elephas.’’ In choosing, therefore, a new generic name for EF. primigenius, Professor Osborn had under consideration the Dicyclotherium of Geoffroy, 1837, the Cheirolites of von Meyer, 1848, and the Synodontherium of Costa, 1850, also the Polydiskodon of Pohlig, 1888, which he regarded as the first really appropriate name. He selected, however, the name Mammonteus (see Osborn, 1924.633, p. 2), a term which he considered as most appropriate and as antedating the generic names just mentioned, but unfortunately this term was used by Camper in 1788 in an adjectival sense (Mammonteum) and applied to the remains of a Mastodon and not toa Mammoth. For a history of the nomenclature of this name and the substitution by Dr. A. Tindell Hopwood of the term Mammuthus Burnett, 1830, type Mammuthus Borealis, see pages 1363 to 1367 of Chapter X XI above. The characters differentiating Mammonteus from Archidiskodon and Parelephas are given on the immediately preceding pages and are also observable by comparing the diagnoses accompanying these three genera, therefore they need not be repeated here. Mammonteus, in a word, is the most acrocephalic, hypsicephalic, bathycephalic, and cyrtocephalic of the Mammontines, with the greatest molar compression and an extremely high ridge-plate formula; also the contour of the body is distinctive, especially in the sloping hind quarters and the sharp notch behind the peaked skull, features brought out even in the drawings of Paleolithic age. Micration.—The geographic distribution of the Northern or Woolly Mammoth has proven to be a most difficult task, owing to the confusion that has existed in the separation of Elephas |[Mammonteus| primigenius from E. |Parelephas| trogontherii and P. jeffersonii in the previous records (see pp. 1041, 1133, 1138, 1140); in fact, all the Pleistocene mammoths of western Europe and North America, including all the species of Parelephas as well as all the primitive and geologically ancient species and subspecies of the true mammoth, have been attributed, or referred, to the typical species Hlephas primigenius. The ancestral relationship to Mammonteus of the Upper Pliocene [?Lower Pleistocene] Mammonteus primigenius astensis of Italy is problematic, as it shares characters of both Mammonteus and Parelephas; its ridge-plate formula, however, is higher (M 31%2°) than that of the contemporary Parelephas species P. trogontherioides (M 3%:\4). “There can be no doubt,” as stated by Pro- fessor Osborn, “that during late Pleistocene (IV Guactau and Postglacial) time there existed all over northern 681 ‘squeydaya UvIPUT SULAT] IO} ZFT ain’y veg ‘seIpuy 4sey oUt puB BISy WEGyNOG :]ISsoy pue yusood ‘(spydajaj0)q ‘spydapasdhiy ‘spydajq) SLNVHAGTG NVIGNT 9 “squeydeye uBolyy BULAT IO} ZPZI GNBY vag “BOIFY JO quaulyu0s 9y} Pue “BIS ‘gdoing, W1eYyNOG :(wopoxoposadsa yy ‘Wopoxojox]0q ‘pyuoporoT) SENVHAATY ANILNOGOXOT SHI, “G “souty UOTyBABIUT (2) 9199T09Y} BTOYMESTS SoorxeyA] PUB ‘SE7eIg PoHU] 94} ‘gdoing useyseq, Puw UIEqyNOS :(spydajai0q) *souUty 9U9904SI9T J 97] UL faqered TOF OF} OF PLEMTTPNOS Suipue}xe woryNqisystp qejodummorg :(snawouun JA) SHLOWINY JA, NUTHLUON CHL SHLOWNVJ) NVIMTHLNODOUT, “FP “OOIXOTAT PUB ‘SEzEIG PHU] 4} ‘aIsy WIN, ‘puvpsug ‘gdoung WeyyNog “Borpy + (wopoysepryr4 V) SHLOWNVA) NUTHLAOS FHL ‘salpuy yseq eyg pue ‘ueder ‘visy U19yseq PU mrayynog : (wopobarg) SLNOGODALY FHL aN FZGT Ut AqaI00g jeorpdersoay) UBL Y 9} Aq paredaad uoryotord rejod YWON SANIT NOILVUDIJ, OILANOUH J, ONIGO TINT VACIOLNOGODALG INV VACIOINVHAGTY AHL JO SAITINVAROS AHL JO NOILOAAINLSIC OILLVWIT() IVUANAL) “TEST PLA EN SIV (06! °F PesiAe4) OEI —“porqoaad guBpS/p/NDa /DYZIUaZ IV/AY [HN 205 «Beep wou gecél ‘P7sd el? uoyday uDipu = sPYy72)9] Pd ‘spydajasdhyy ‘sDyday 7 +4] wopoxojoladsap] uopoxojoenjng b= SF yay pjuopoxoy GUIJUOPOXx0T = SypOUUlDpy ULBYINOG = UOPOYSIPTY Hp. ay] sypourumyy uDitayy uoboay = spydajadn 7 SYJOWIUID)Y ULAYJION = SNAJUOULUD I) 9% ope] St uopobaj9 aul) yadap 1239W 002 | 1590 OSBORN: THE PROBOSCIDEA Eurasia a single species of mammoth to which the name Mammonteus primigenius has been assigned, since it furnished the type to which the earliest as well as the original and later descriptions were applicable.”’ The accompanying plate (Pl. xxir) also figures 1228 and 1241 are of necessity theoretical, but they have been prepared with the utmost care, and only with a view to expressing Professor Osborn’s latest determinations. It will be observed that the woolly mammoth ranged from Lat. 45° (northern France) to 75° (New Siberian Is- lands), according to Tolmachoff and Depéret and Mayet; in the Western Hemisphere from the 40th parallel (New York State on the east) to the western coast, northward to the 70th parallel (Melville Island, fide Sternberg) and Alaska. Superfamily: ELEPHANTOIDEA Osborn, 1921. Family: ELEPHANTID Gray, 1821 Subfamily: LoxopontT1n# Osborn, 1918 Genera: Loxodonta F. Cuvier, 1825, 1827; Palxoloxodon Matsumoto, 1924; Hesperoloxodon Osborn, 1931 Subgenera: Szvalikia Osborn, 1924; Pilgrimia Osborn, 1924, now regarded as synonyms of Palxoloxodon (Cf. Vol. I, Pls. x, x1, also Chap. XIX of the present volume, Figs. 1220-1223, 1228, and Pl. xxim) The subfamily Loxodontine (embracing three genera—Lozodonta, Palxoloxodon, and Hesperoloxodon), like the Elephantine, is distinctive in that certain of its members survive to the present day, namely, the Loxodonta africana of the African continent, the ancestry of which, in the opinion of the present author, is still in doubt, as will be seen by the following quotation from page 1273 above: “‘L. africana cannot be descended from any known typical species of Palxoloxodon but may have sprung from a more primitive ancestral form still to be discovered.”’ While the Loxodonta zulu of Scott, 1907, and the recently described species of Dart (Loxodonta prima, 1929, L. subantiqua, 1929, and L. africana obliqua, 1932) may prove to be ancestral to the living African elephants, Profes- sor Osborn was cautious in expressing his views on the subject (Osborn, 1934.925, pp. 7 and 6): ‘“These occur only on the more recent levels and are clearly related to the existing African elephant, distinguished by the above [following] characters.”’ Loxodonta prima group Crowns relatively narrow, 74 mm. (L. prima) to 92 mm. (L. subantiqua). Enamel relatively thin, coarsely crimped; conelets numerous. Cement thin in middle, thick at edge. Ridge plates per 100 mm. =4 (L. africana obliqua) to 5% (L. subantiqua). Broad typical loxodont sinus expansion, double sinus foldings in contact. Total ridge plates 9 (L. prima) to 12-13 (L. zulu). In Doctor Dart’s description, however, of Loxodonta prima in 1929 (pp. 724-726) he states: “The outstanding character of this tooth (Figs. 25, 26 [Fig. 1149 of the present Memoir]) is its comparability with the living African species. It provides the first indubitable fossil evidence of a very close approximation to the distinctive lozenge- shaped lamellae of the living Loxodont grinding tooth. The absence of such a fossil type up to the present time according to Osborn ‘is a striking circumstance.’ ... There can be no doubt that in this narrow-crowned, loxodont- simulating, few-plated, mountain-inhabiting specimen .. . we have a long-sought ancestral type from which the modern African elephant tooth might reasonably be derived by a progressive widening-out of the loxodont sinus, for which reason I have named it Loxodonta prima. This fossil is of importance not only in demonstrating an advancing, though still simple Loxodont tooth pattern in the Transvaal, but also in revealing Africa, and possibly South Africa as the evolutionary home of the true Loxodonta. Its habitat in the elevated and relatively waterless Pilandsberg suggests the possible reasons for Loxodont persistence, namely, their becoming inured to more arid conditions and more active movement, as compared with their more ponderous Archidiskodont relatives.” LOXODONTA LOXODONTA AFRICANA AFRICANA PUM/L/IO OXYOTIS AFRICA AFRICA PALAEOL DES dt hy LITENSIS, — PA. i, | OXODON PALAEOLOXODON FALCONER], NAMA DICUS PAL AEOLOXODON MNAIDRIENSI/S, INDIA MALTA HESPEROLOXODON = <>) ANTIQUUS GERMANICUS” Zee ’ GERMANY bt HESPEROLOXODON “ANTIQUUS PLAT YRHYNCHUS SPAIN HESPEROLOXODON ANTIQUUS oF upNoR, ENGLAND is oO 15° . iS a 75 90E 105120" 35 150" oa 180" 30 45 ee T LAS AS = Ed ae fe ttt Ee TRAPFLR® 1 Chater | Vf ii Naas My ee) f ie i Re EQUATOR Bae we \ 2 Fe EQUATOR ; 150° 135" 120" Ts is o 15° 30° E 60° 75 90°F 105° 120° 135° 150° 165 180 Origin, Migration and Epelanen Ora +Loxodonta, +Faldeoloxodon, ¢Hesperoloxodon PLATE XXIII Geologic range: Loxodonta, Pleistocene to Recent; Palxolorodon, Upper Pliocene(?) or Lower Pleistocene to Upper Pleistocene; Hesperoloxodon, Upper Pliocene [Lower(?) Pleistocene, see footnote 1 on p. 1049 above] to Upper Pleistocene. SUMMARY 1591 Loxodonta, genotype probably the South African form Elephas africanus Blumenbach, 1797; the extreme South African form was distinguished by Cuvier in 1798 as Elephas capensis. Cranium relatively low, brachycephalic, platycephalic, mesocephalic, bathycephalic, and cyrtocephalic (foreshortened) ; cranial profile and section much more primitive, less hypsicephalic and bathycephalic than in the Mammontine or Elephantine; parietofrontal cranial vertex low, rounded, platycephalic to subacro- cephalic, of persistent primitive form; rostrum short and extremely broad; premaxillaries diverging to the point where the tusks issue from the skull—thus the bases of the tusks are very far apart instead of being close together as in Mammonteus; nasals broad and rounded, narial openings hour-glass shaped; anterior position of suture separating malar from maxillary portion of zygoma. Mandibular ramus elon- gate; symphysis acute, relatively prominent and horizontal. Superior tusks elongate, widely divergent, relatively straight, slightly upeurved and incurved, resulting in a lyre-shaped arrangement; marked sexual disparity in tusks of females. No inferior tusks. Molars low crowned, relatively narrow and with comparatively few ridge-plates, total ridge-plates Dp 4-M 322: enamel borders thick and simple without foldings or plications: cement thin; distinguished by broadly open ‘loxodont sinus’ (lozenge-shaped median expansion) on wear. Ridge-plate formula constant and very conservative, since the M 3 ridge formula of the living African elephant closely corresponds with that of the primitive Upper Pliocene Archidiskodon planifrons. Dental formula: I *** Dp =$ M #3 Ridge-plate formula (typical): Dp 22 Dp 38 Dp 47 M 1! M23, M3 2%; Horizon.— Pleistocene to Recent. Compare page 1541, also figures 1220, 1222, and Pl. xx1m of the present volume. COMPARISON OF LOXODONTA WITH THE GENERA ARCHIDISKODON, PARELEPHAS, MAMMONTEUS, AND HLEPHAS Taking, for example, the middle-aged skull of “Jumbo,” a Sudanese subspecies, we observe that “in the fully adult skull the dome is continuously rounded from the occipital condyles to the broad extremities of the nasals, presenting the widest contrast to the profiles of Archidiskodon, of Parelephas, of Mammonteus, and of Elephas; that while actually brachycephalic, the cranium of Loxodonta is also comparatively mesocephalic or elongate (shown in the relatively long mandibular ramus, { Fig. 1060]), as compared with the deeply depressed mandibular ramus of Hlephas indicus, or with the extremely bathycephalic and abbreviate mandibular ramus of Mammonteus primigentus; that since both superior and inferior molars, M’, Ms, are shorter and less hypsodont, the maxillary and mandibular dental cavities are much less deep, thus accounting for the less hypsicephalic proportions of this part of the skull; the superior aspect of the mandibular rami [ Fig. 1060] also displays the relative prominence and horizontal distinction of the rostrum, again presenting a very wide contrast to the deep, hypsicephalic rostrum of the Mammontinz and of the Elephantine’”’; the superior or frontal aspect of the cranium (Fig. 1061A) shows the short frontal bones, the massive orbital prominences, the broad narial openings, the widely separate maxillo- premaxillary sockets for the enormous incisive tusks, similar to those in Palzoloxodon namadicus and Hesperoloxo- don antiquus. It will be observed in a comparison of the palatal aspect of the Loxodonta cranium with that of the Elephas indicus cranium (Fig. 800) that the former is relatively broader, more brachycephalic in all its dimensions (ef. Chap. XTX, p. 1200 above). Palzxoloxodon.—The origin of the Palzeoloxodonts is obscure. Professor Osborn suggests on page 14 of his article in Novitates (Osborn, 1934.925) that the ‘“‘problematic A. [= Palxoloxodon] andrewsv”’ of Dart, 1929, found at Gong-Gong, Vaal River, may be a “‘primitive or ancestral member of the Palxoloxodon group.” In the present Memoir (cf. p. 1207 above) he also suggests that this generic phylum (which includes Palxoloxodon and its synonyms Sivalikia and Pilgrimia) may have sprung from the giant North African species Elephas atlanticus Pomel, 1879. In any event, there can be no question of its clear generic distinction from Loxodonta. In speaking of Palxoloxodon from the historic standpoint, he remarks (cf. p. 1207) that “It is not impossible that some elephants of the Palxolorodon type survived into early historic times, but it is an open question whether the elephants described from Mesopotamia were of the ancient ‘loxodontine’ or of the modern ‘elephantine’ type, 1592 OSBORN: THE PROBOSCIDEA probably the latter; drawings and inscriptions will probably be found some day which will determine these relations.” Pilgrim (1905) notes that Elephas {[Palxoloxodon| namadicus is entirely absent from the Pliocene Siwalik strata and that there is no ancestral type from which it might arise. Palzxoloxodon, genotype Elephas namadicus naumanni Makiyama, 1924. Cranium of genotype unknown, but compare the Elephas |Palzoloxodon| namadicus of Falconer and Cautley, 1846, 1847, from the Nerbudda, India, which is more hypsicephalic and bathycephalic, and relatively broader and more platycephalic than that of Loxodonta. Parietofrontal cranial vertex somewhat more progres- sive, acrocephalic, than in Loxodonta, with broad, smooth or rugose crest which in Palzoloxodon namad- icus extends like a Phrygian cap down over the frontals almost to the nasals (seen also in the dwarfed Elephas [Palxoloxodon] melitensis and E. [P.| mnaidriensis of Malta). Narial openings very broad and shallow. Premaxillaries extremely broad, with widely divergent, relatively straight, slightly upcurved and incurved tusks toward the extremities. No inferior tusks. Molars moderately hypsodont; relative- ly narrow in primitive stages; superior molars progressively narrow to broad, with numerous ridge- plates composed of thin, plicated enamel foldings. Ridge-plates parallel, closely compressed. Loxodont sinus rudimentary or absent. Dentine areas equal or exceed cement areas by relatively close compression of the ridge-plates. Ridge-plate formula progressive from Palzxoloxodon atlanticus (M 34344), to P. melitensis (M 3,22), to P. namadicus (M 312), to P. namadicus naumanni (M 37%). Dental formula: I *** Dp M #3 Ridge-plate formula (fide Falconer): Dp 4;9, M laqgan M 25 M 3158 Horizon.—Upper Pliocene(?) or Lower Pleistocene to Upper Pleistocene. Compare page 1541 of the present volume, also figures 1220-12238, also Pl. xx111. Sivalikia and Pilgrimia.—In 1924 both Matsumoto in Japan and the present author in America published almost simultaneously the results of their researches on the Loxodontines, neither being aware of the activities of the other. The genus Palzxoloxodon of Matsumoto (described September 20, 1924) anticipated by three months Professor Osborn’s description (December 20, 1924) of Sivalikia and Pilgrimia; hence technically it has priority over the last two genera, which become synonyms of Palxoloxodon. In Chapter XIX of the present Memoir Professor Osborn has explained the morphological reasons for regarding his Sivalikia and Pilgrimia as synonyms of Palxoloxodon Matsumoto. Dwarfed Species of the Mediterranean Islands.—Before taking up the genus Hesperoloxodon, to which Professor Osborn referred members of the ‘Elephas antiquus’ group, the dwarfed species of the Mediterranean Islands deserve some mention. It is interesting to note, that besides the pygmy elephants of Malta, a specimen of nearly normal size is also recorded from that island. Falconer and the older authorities related these insular species to the Hlephas antiquus of the European continent, but subsequent discovery has shown that they are more probably derived from certain of the extinct ancestral African species described in the present Memoir as Palxoloxodon (syn. Pilgrimia); moreover, the cranium and jaw of Elephas |Palxoloxodon] melitensis,’ with broadly overhanging parietofrontal crest, more closely resembles the F. [Palzoloxodon| namadicus of India than the E. |Hesperoloxodon| antiquus of western Europe. A comparison of the type grinding teeth of the dwarfed Mediterranean species with the type grinding teeth of the extinct African species reveals a striking general resemblance in the narrow pro- portions and in the rudiment or absence of the ‘loxodont sinus,’ characters which appear to relate these teeth to the Palzoloxodon (syn. Pilgrimia) of Africa rather than to the typical Loxodonta africana or to the broad-toothed Palzoloxodon namadicus of India. In Europe some of the narrow-toothed varieties of Hlephas [Hesperoloxodon] antiquus may be related to the dwarfed insular elephants. After the discovery and description of the cranium from Pignataro Interamna, Italy, first referred by Professor Osborn to Palzoloxodon antiquus italicus and finally made the type of his genus Hesperoloxodon, he expressed himself as follows (see Chap. XIX, p. 1252): “The entire cranial and dorsal hump silhouette (Fig. 1092) is quite different from that of the African elephant (Fig. 1093)... The cranial profile of Hesperoloxodon italicus is also entirely different from that of Palzoloxodon namadicus (Fig. 1070) and of P. melitensis" (Fig. 1121), both of which are characterized by a very prominent transverse frontal ridge for the '[Subsequently (see p. 1260 above) referred by Professor Osborn to Palzolorodon mnaidriensis.—Editor.] SUMMARY 1593 attachment of the gigantic proboscis. . . . In both cranial and dental characters H. italicus differs profoundly from the African elephant, which is extremely conservative in its structure, in fact, much more conservative than any of the known fossil Pleistocene elephants of Eurasia. ... H. italicus affords additional and positive evidence that the dwarfed elephants of the Mediterranean Islands (P. falconeri, P. melitensis, and P. mnaidriensis) were not derived from the ‘Elephas antiquus’ phylum of Falconer, as hitherto universally believed, but sprang from some undis- covered phylum of elephants of African origin, which gave rise on the one hand to the dwarfed elephants of the Mediterranean Islands and on the other to the gigantic P. namadicus phylum of India and the Far East extending to Japan.”’ Thus, according to the views last expressed by Professor Osborn, we are able to make the following summary: Hlephas namadicus of India (Nerbudda), varieties and subspecies: of which are abundant in Japan =the true Palxoloxodon (syn. Sivalikia of Osborn) Elephas atlanticus, E. jolensis, E. recki, ete. of Africa = Palxoloxodon (syn. Pilgrimia of Osborn) Mediterranean Island species (pigmy and normal size), possibly derived from the Hlephas atlanticus group or the ‘‘Elephas antiquus reckv”’ |= Palzxoloxodon recki| of Dietrich. = Palzxoloxodon (syn. Pilgrimia (Perhaps some of the narrow-toothed varieties of of Osborn) “EH. antiquus” of Europe may be related to these dwarfed species.) Hesperoloxodon.—Elephas antiquus Falconer and Cautley, 1847, described one year after the naming of £. |Palaeoloxodon| namadicus Falconer and Cautley, 1846, is based on a molar, probably from England, although the locality is not recorded. To the best of our knowledge no complete cranium of the true H. antiquus has as yet been discovered, consequently the unearthing of a fine cranial specimen at Pignataro Interamna, Italy, in 1911-1912, and acquired by the American Museum of Natural History in 1929, was of signal importance. Professor Osborn immediately described the specimen, referring it first to Palxoloxodon antiquus italicus, but on comparing it more closely with the cranium of P. namadicus, he observed at once that it was more hypsicephalic and bathycephalic and that it lacked the prominent transverse frontal crest so characteristic of P. namadicus and P. mnaidriensis. On these grounds he provisionally made it the type of a new genus, Hesperoloxodon. Again the discovery in 1926 and 1928 at Steinheim on the Murr, Germany, of two crania referable to Hesperoloxodon antiquus germanicus convinced him more then ever of the distinction of the “Hlephas antiquus”’ group from the Palxoloxodon group. Taken together, these crania furnished the basis for determining the evolutionary history and relationships of the classic Hlephas antiquus, definitely assigned by Professor Osborn in the present Memoir to the genus Hespero- loxodon. Hesperoloxodon, signifying ‘loxodont of the west,’ of the European continent. Cranium domelike with flattened forehead; more hypsicephalic and bathycephalic than that of Palzoloxodon; lacking the prominent parietofrontal crest so distinctive of P. namadicus. Occiput relatively narrow and high (broad and low in P. namadicus). Premaxillaries extremely broad, with widely divergent tusks, slightly upcurved and incurved. Mandibular rostrum abruptly truncated; symphysis wide. Molars hypsodont, ‘loxodont sinus’ vestigial or absent. Ridge formule progressive from M 3 {$;=++ (typical), to M 34522 (germanicus), to M 32; (italicus). Enamel plates relatively thick, more or less crimped or sinuous. Cement present. Dental formula: I °** Dp =* M #3 Ridge-plate formula (Hesperoloxodon antiquus): Dp 23 Dp 3§ Dp 453 M 1*33*° M 2 +3 M 33-4 Horizon.—Upper Pliocene’ to Upper Pleistocene. Compare pages 1542, 1543 of the present volume, also figure 1222, and Pl. xxi. (Possibly Lower Pleistocene (see footnote 1 on p. 1049 above).—Editor.] 1594 OSBORN: THE PROBOSCIDEA In a comparison of the crania of Loxodonta africana, Palxoloxodon namadicus, and Hesperoloxodon antiquus, we observe especially the following characters: Short and extremely broad rostrum; premaxillaries diverging to the point where the tusks issue from the skull—thus the bases of the tusks are very far apart instead of being close together as in Mammonteus primigenius or relatively close as in H. indicus; whereas the premaxillary sockets are relatively of the same length in all three species, the divergence of the sockets in P. namadicus is about the same asin L. africana; the premaxillary sockets are relatively longer and diverge still more widely in H. antiquus; the erania of both P. namadicus and H. antiquus are distinguished from the cranium of L. africana by the greater development of the fronto-occipital crest which in P. namadicus engulfs the frontal bones so that there is a very short space between the lower border of this crest and the extremities of the nasals and the narial openings; the narial openings are extremely broad and shallow, they exhibit approximately the same hour-glass-shaped form in L. africana, P. namadicus, and H. antiquus; the skull of P. namadicus, moreover, is relatively broader and more flattened or platycephalic than the skull of L. africana; this broadening and flattening of the summit of the cran- ium reaches an extreme in the gigantic P. namadicus cranium (cf. Chap. XIX, p. 1209 above). DisTRIBUTION OF LivING ELEPHANTS: LOxODONTA AND ELEPHAS SY S SSS SSS SSS WN . hb E wid Subtspecies f “<) * |' 4 i LEPHAS "| Loxond i —equaror| INDICUS AFRICANA % ' and Sub) Gy 7) aa DF Levett Bradley 1938, betes Living Elephants Fossil Elephants and Mastodonts Fre. 1242. Woripwipe DistrisvuTion OF THE PROBOSCIDEA IN Past AND PRESENT TIME This map on a larger scale constitutes the back end-paper of the present volume Arrican Evppuant: After Blanc, 1897; Bocage, 1890; Buckley, 1876; Chapman, 1868; Chubb, 1909-1919; Claridge, 1915; Cox, 1900; Heilprin, 1887; Hippolyte, 1907; Johnston, 1907; Maydon, 1932; Roosevelt and Heller, 1914; Schultze, 1907; Sclater, 1900. InpIAN EverHant: After Bartholomew, 1911; Blanford, 1888-1891; Blyth, 1872; Bishop, 1921; Champion, 1928; Hornaday, 1885; Hunter, 1868; Jerdon, 1874; Laufer, 1925; Lydekker, 1900-1908; Sanderson, 1907; Sclater, 1899. [Since Professor Osborn’s intensive studies on the subject, recent observations by Pére Teilhard de Chardin, Dr. C. C. Young, Dr. Glover Allen (letter of Dee, 24, 1938, to Dr. Pinkley), and Dr. George Pinkley have led to certain modifications of the map shown in Volume I, figure 6, namely, the area in Borneo, as well as that in southwestern China, has been reduced somewhat. See Chapter XX, p. 1362 above, for the views of Pére Teilhard and Doctor Young regard- ing the existence in historical times of the wild Indian elephant in China.—Editor.] SUMMARY 1595 MIGRATION OF THE LOXODONTINES Loxodonta.—The eighteen existing species and subspecies or varieties of Loxodonta africana (partly distin- guished by size, partly by geographic distribution, and partly by the shape of the ears) are found from the Cape northward to the southern border of the Sahara and the Sudan (see Fig. 1055). Fossil remains are principally of African provenance, that is, from Zululand (L. zulu) and the Transvaal (L. prima, L. subantiqua, and L. obliqua), although two or three localities in Europe have yielded types, e.g., Germany (H. priscus Goldfuss = L. africana) and Italy (L. cornaliae), as shown on the accompanying plate (Pl. xxur). Palzoloxodon.—The Palzoloxodonts, as we have just seen, are divided into two branches, (1) Palxoloxodon (syn. Pilgrimia), embracing such species as atlanticus, jolensis, and recki from the northern and east-central regions of Africa, and transvaalensis, sheppardi, andrewsi, hanekomi, wilmani, kuhni, yorki, and archidiskodontoides of South Africa, as well as the Mediterranean species—melitensis, falconeri, mnaidriensis, lamarmorae, cypriotes, and creticus; and (2) Palxoloxodon (syn. Sivalikia), embracing the namadicus of Falconer and Cautley, type locality the Nerbudda, India, the hysudrindicus of Java, and the various (possibly eight to ten) subspecies of Japan, represented by the genotype of Palzoloxodon, namely, Elephas namadicus naumanni Makiyama. It would appear, therefore, that the phylum Palxoloxodon originated in Africa, migrated northward into Europe, also northeastward, through the Mediterranean Islands, leaving dwarfed descendants resembling either the ‘E. antiquus’ group or the ‘E. namadicus’ group, finally arriving in southern and southeastern Asia developing into the ‘Elephas namadicus’ group, thence eastward to Java and northward to Japan, but as far as we know never reaching America. Hesperoloxodon.—Members of the ‘Elephas antiquus’ group are recorded from Italy (Hesperoloxodon antiquus nanus, H. antiquus ausonius, H. antiquus italicus), from Spain (H. antiquus platyrhynchus), from England (the true H. antiquus especially), from Germany (represented by the H. antiquus germanicus of Steinheim), and from Rumania (by H. antiquus germanicus ref. from Ilfov). Professor Osborn states in Chapter XIX, p. 1258 that some of the narrow-toothed varieties of Elephas [Hesperoloxodon] antiquus in Europe may be related to the dwarfed insular elephants; and again on page 1252 he calls attention to the “lofty cranial dome” of Hesperoloxodon antiquus italicus which resembles that of the Elephas indicus of India and is in close agreement with the “Eléphant tracé en rouge” in the cavern of Pindal, also with the anterior dorsal hump of the Spanish and Algerian elephants (Fig. 1047). Superfamily: ELEPHANTOIDEA Osborn, 1921. Family: ELEPHANTID& Gray, 1821 Subfamily: ELEPHANTIN® Osborn, 1910 Genera: Elephas Linneus, 1735-1758; Hypselephas Osborn, 1934, 1936; Platelephas Osborn, 1934, 1936 (Cf. Chap. XX of present volume, also p. 1543, and figures 1173, 1221, 1228, and PI. xxm) The history of the elephant in art, industry, and science is fascinatingly told by Dr. George F. Kunz in his book on “Ivory and the Elephant.” The early life phases, however, are so beclouded by superstition and myth that we are without positive knowledge of these interesting animals until the time of Homer. Strangely enough, also, the fossil ancestry of the living Asiatic elephant is still unknown, despite the untiring efforts of palzeontolo- gists and zoologists everywhere to solve this problem. Consequently the subfamily phylogeny of most of the elephantoids is better known than that of the Elephantinz, or, as a matter of fact, of the Loxodontine, for, as noted in the preceding section, there is still uncertainty as to the ancestral forms of the living African elephant. Professor Osborn was of the opinion that ‘‘the conclusion is inevitable, that in some as yet unexplored region of 1596 OSBORN: THE PROBOSCIDEA Asia the direct ancestors of 2. indicus were slowly evolving, while in some unexplored area, probably west of central Africa, the very conservative ancestors of L. africana were also slowly evolving.”” As our knowledge stands at present, the Indian elephant suddenly appears fully formed during the Age of Man. The nomenclatural problems in connection with the genotype of Elephas are no less complex. After long consideration Professor Osborn adopted Elephas indicus rather than Elephas maximus in the present Memoir, giving his reasons therefor in Chapter XX, pages 1308-1311 above. Falconer, in his discussion of the unity or plurality of species among the existing Asiatic elephants, namely, types from Ceylon, Nepal, Bengal, and Sumatra, decided in favor of the unity of the species Elephas indicus (Falconer, 1868, Vol. II, p. 269): ‘‘the evidence in every aspect appears to fail in showing that the Elephant of Ceylon and Sumatra is of a species distinct from the Continental Indian form... The result of this range of observation, combined with long osteological study, has been to establish the conviction in my mind that there is but a single species of Asiatic Elephant at present known, modified, doubtless, according to his more northern or southern habitat, but not to an extent exceeding that of a slight geographical variety.’’ He admits that the Ceylon elephants are occasionally imported into Bengal but does not admit, as claimed by Schlegel, that this is an explanation of the variation in the vertebral and rib formule of the Indian elephants. The present author ex- pressed his belief (see p. 1315, Chap. XX above) that the ‘‘wide variations in cranial and vertebral characters as well as in dental and dermal characters and in the shape of the external ear support the subdivision of Elephas indicus into at least four out of the large number (12) of geographic varieties or subspecific forms,’”’ namely: Elephas indicus ceylanicus (cf. Mukna var.) Elephas indicus bengalensis (cf. Dauntela var.) Elephas sumatranus Elephas maximus hirsutus (= E. indicus hirsutus) Professor Osborn left the question of the continental and insular races and subspecies for future investigation. He states on page 1329 that the ‘‘profound cranial differences which divide the Ceylon and Bengal elephants from each other as well as from the Sumatran elephant in all probability will be found to differentiate the Sumatran and other still undiscovered extinct types’; that by ‘‘comparison with the evolution of other Pleistocene un- gulates it appears probable that a very long period of time separated these continental and insular subspecies and species from each other, a period of time equivalent perhaps to nearly half of Pleistocene time or 500,000 years, during which through isolation and segregation the subspecific and specific characters were thoroughly founded. Here again monographic research is essential before we can reach a final conclusion.” As to the extinct forms, it is remarkable that no fossil Pliocene ancestors of the recent Indian elephant have as yet been discovered. Hlephas hysudricus found “below the conglomerates” in the upper Siwaliks of India, according to Dr. Barnum Brown, shows few resemblances in the cranium to the H. indicus (Dauntela var.) and no very marked resemblances in the grinding teeth; it appears to be unique, and in 1936 (see Vol. I, p. 12) it was made one of the types of the genus Hypselephas Osborn. The third genus included within the subfamily Elephantine is the Platelephas of Osborn, genotype Hlephas platycephalus, a very ancient and primitive animal. The inclusion by Professor Osborn of these three genera in the subfamily Elephantine was provisional, because Hlephas hysudricus Falconer and Cautley and E. platycephalus Osborn are at present known chiefly by cranial characters, our knowledge of the dentition of platycephalus being confined to the third superior molar only; they appear, however, to represent generic or subgeneric phyla distinct from the true Elephas. SUMMARY 1597 Characters of these three genera, as far as they are known: ELEPHAS Modernized elephant of India. Highly progressive and distinctive in cranial structure Typified by Elephas indicus Cranium bathycephalic, cyrtocephalic, hypsicephalic, occipitofrontal dome more or less rounded, not acute, with expanding diploé, frontals gently con- cave Premaxillaries relatively narrow, sub- vertical. Orbits large, relatively ele- vated, i.e., directly opposite occipital condyle Superior tusks relatively straight, in- curved, upcurved with bases of tusks relatively close together. No inferior tusks Mandible with extremely abbreviated symphysis. Ramus deeply depressed Molars of intermediate breadth, hypso- dont (less extreme than in Parelephas, Archidiskodon, or Mammonteus); ab- sence of ‘loxodont sinus’ (#. indicus) ; moderately compressed enamel ridges of intermediate thickness, extremely crimped or sinuous in £. indicus. Maximum number of plicated ridge- plates: M 3,24 4-27 Dental formula: I **" Dp $$ Mi Ridge-plate Formule: HyYPsELEPHAS Primitive elephant of India with elevated cranium Typified by Elephas hysudricus Cranium hypsicephalic, brachycephalic; condyles well raised above grinding surface of molars, occiput elevated with broadly transverse rugose frontal crest, relatively flat, sloping backward, frontals deeply concave Premaxillaries relatively narrow or later- ally compressed. Orbits large, depress- ed, near maxillary rostrum, unlike Loxodonta or Elephas Superior tusks relatively straight, in- curved, somewhat divergent at base. No inferior tusks Mandible with elongate, prominent sym- physis; ramus shallow Molars low crowned, long, narrow; ridge- plates with plicated enamel, convexo- concave, reversed above and_ below; widely separated, with cement filled valleys (juvenile); trace of a median ‘loxodont sinus.’ Ridge-plate formula: M 353 17-18-19 1-3 1-3 PLATELEPHAS Elephant of India with flattened cranium, primitive Typified by Elephas platycephalus Cranium relatively elongate, dolichoce- phalic, platycephalic, occipital con- dyles not greatly elevated above level of grinding surface of molars; deeply in- dented supra-occipital border, without rugosity Premaxillaries greatly elongated in front of superior third molars, somewhat divergent. Rostrum somewhat broad. Orbits large, elevated, near frontal profile, set widely apart. Posterior nares deeply indented Tusks imperfectly known Molars imperfectly known: relatively low, ridge-plates directly transverse, as in Elephas. No rudiment of ‘loxodont sinus.’ Ridge-plates broad, depressed, limited in number: M 3184 Elephas (E. indicus, fide Falconer): Dp 2¢ Dp 3% Dp 435 M 133 M 23° M 3,24 12 Hypselephas (EF. hysudricus) : 24-27 (Falconer, 1868): Dp 3°32 Dp 4748+ M 13%42 M 2,48. M 3iv-18 9+-12 (Osborn, 1930): Dp 34g Dp 48% M 134 M 22122 M 328+ Platelephas (as far as known): M 31&# HORIZON.—Elephas, Upper Pleistocene and Recent. Pleistocene. Compare page 1548 of this volume, also figures 1221, 1228, and Pl. xxm. Hypselephas, Lower Pleistocene. Platelephas, Upper Pliocene or Lower 1598 OSBORN: THE PROBOSCIDEA COMPARISON OF HyPSELEPHAS AND PLATELEPHAS WITH ELEPHAS With the exception of the decidedly low position of the orbits, the front view of the Hypselephas hysudricus cranium bears a closer resemblance to the broad-narial variety Elephas indicus bengalensis than to the narrow- narial variety HE. indicus ceylanicus; also the low, narrow grinding teeth of the type of H. hysudricus are entirely different in proportion from those of 2. indicus, they exhibit some resemblance or analogy to the extinct Palzoloxodon of Africa both in their long, narrow proportions and in their ‘loxodont sinus.’ The juvenile cranium of H. hysudricus resembles somewhat closely the juvenile cranium of FE. indicus at the time when the fourth de- ciduous molars are in use. From cranial and juvenile characters alone one would be inclined to regard H. hysudri- cus as an early offshoot in Lower Pleistocene times of the main stem which gave rise to H. indicus of recent times, but the less rugose orbits in a similarly depressed position as that seen in crania in the British and Amherst Museums referable to H. hysudricus, that is, lying directly upon the premaxillary sockets of the tusks instead of being raised above the tusk sockets, demonstrate afresh that H. hysudricus is not to be regarded as ancestral to the collective Hlephas indicus type but that it belongs in a totally distinct phylum or line of descent. In short, H, hysudricus had a modernized cranium with a relatively primitive condition of the molars. The Platelephas platycephalus cranium is of very primitive elephantine affinity—low and flattened—widely different from the elevated Hypselephas hysudricus or the greatly elevated Elephas indicus crania. Both in the cranium and the grinding teeth it is also profoundly distinct from those of Archidiskodon planifrons. Therefore, Platelephas platycephalus, in the opinion of the present author, was not ancestral to either H. hysudricus or E. indicus, nor was it related to Archidiskodon; it would seem to belong to a primitive stage in the evolution of the Elephantine. The specific name has reference to the highly characteristic and primitive lowering of the fronto- occipital profile and the placing of the occipital condyle only slightly above the horizontal level of the maxillary border of the superior grinders. 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Sarre RTPI NVIU3WOND | VWYONNL | Qv¥9 NYNOBAIM ONY pe aes SAILW13SH— 4 | ¢g3q| GYO4SITTIHS 5€S SONI 5 5 NOUVIDNV19 I] Sual4aW o1-06] 2 addais [ | sazovaual_| = uum Sa ee ce ee (eee ASul4 PaaS Cases BST Pee pia 3 Se lg = hee ls 2 pap 0k 12 — ets eee = WS es n (eee a Post SLAGIAL-§ FOREST MEADOW YOUNGEST LOESS is a REGRESSIVE MONASTIRIAN STAGE OF 18-20 METERS.RICH FAUNA) LEVEL6-6m TYRRHENIAN STAGE MARINE DEPOSITS. SHORELINE OF 28-30 METERS MARKED CLIMATIC DIFFERENCES] WARM FAUNA WITH SUBTROPICAL AFFINITIES |COLD NORTH TEMPERATE FAUNA TRANSGRESSIVE PHASE REGRESSIVE PHASE MILAZZIAN STAGE MARINE DEPOSITS. SHORELINE OF 55-60 METERS w on < =x a Ww = nan un py [- 4 Qo no z = in 4 = 7 FOREST BED _OF CROMER CHILLESFORD AND WEYBOURN BEOS MOUTH OF SOMMERIVER ALTITUDE EXISTING BED OF LARGE RIVERS STROMBUS BUBONIUS ZONE LESS WARM THAN TYRRHENIAN STAGE WARM FAUNA REGRESSIVE PHASE SICILIAN STAGE MARINE DEPOSITS. SHORELINE OF 90-100 METERS POLAR AND TEMPERATE INVERTEBRATE FAUNA WITH CYPRINA ISLAND/CA \N THE MEDITERRANEAN ‘RELATIVE — RISS, POLONIAN. WISCONSIN _4 "UPPER DRIFT" 3] ¢ i =e DEPOSITS, ZINTERGLACIAL RISS -WURM DURNTENIAN SANGAMON == OSITION GLACIO- | LOESS DEPOSITS A FLUVIAL GRAVELS 2 INTERGLACIAL MINDEL-RISS TYROLIAN YARMOUTH TINTERGLACIAL SUNZ-MINDEL NORFOLKIAN AFTONIAN REMNANTS OF TERRACES EROSION OF DEPOSITS AND STREAM BED. DEPI OF TERRACES al FA =| =| SNOW LINE OF PRESENT CLIMATE TRANSGRESSIVE PHASE 90-110 METERS |T GLACIATION = [= =~ == ==} PLEISTOCENE ee a] PLIOCENE ——] = a o a JERSEYAN “3 8] ES STEPPE TUNDRA STEPPE TUNDRA STEPPE MEADOW FOREST FOREST MEADOW STEPPE TUNDRA STEPPE MEADOW FOREST FOREST MEADOW TUNDRA STEPPE MEADOW FOREST MEADOW STEPPE TUNDRA STEPPE MEADOW FOREST HISTORICAL PERIOD SST ee AZILIAN MAGDALENIAN GRENELLE CRO-MAGNON SOLUTREAN AURIGNACIAN COLD MOUSTERIAN WARM MOUSTERIAN COLD ACHEULEAN PALEOLITHIC WARM ACHEULEAN LATE CHELLEAN CHELLEAN EARLY CHELLEAN CROMERIAN 1——_ —$ ——-—-—-— — FOXHALLIAN STONE GRIMALDI TUNDRA FAUNA REINDEER MAMMOTH {E. primigenius) WOOLLY RHINOCEROS [R. tichorhinus} LATITUDE 43° LAST WARM AFRICAN-ASIATIC FAUNA E.ANTIQUUS HIPPOPOTAMUS R.MERCKII E.TROGONTHERII THIRD COLD FAUNA LATITUDE 45 ° WARM AFRICAN-ASIATIC FAUNA E.TROGONTHERI E.ANTIQUUS R.MERCKII HIPPOPOTAMUS SECOND COLD FAUNA LATITUDE 48° WARM AFRICAN-ASIATIC FAUNA E.MERIDIONALIS E.ANTIQUUS R.ETRUSCUS HIPPOPOTAMUS MACHAERODUS HEIDELBERG COLD FAUNA LATITUDE 53° E.PRIMIGENIUS QVIBOS MOSCHATUS RANGIFER TARANDUS PILTDOWN 7 | ARCHIDISKODON PLANIFRONS ZYGOLOPHODON BORSONI ANANCUS ARVERNENSIS EQUUS STENONIS WARM AFRICAN-ASIATIC FAUNA [ED 3 RIVER TERRACE | GLACIALAND ES ECHANGES IN SEA LEVEL - Prarie STAGES = |CROSS CORRELATION] INTERGLACIAL 2 CURVE IN PART AFTER GISNOUX [BOULE, LAMOTHE] CURVE IN PARTAFTER PENCK, SOERGEL., DATA FROM DEPERET, GIGNOUX, C.REID DATA FROM PENCK,SOERGEL, BROOKS , LEVERETT, DEPERET CULTURES DATA FROM OSBORN, MAVET, DEPERET, COMMONT, [matte PENCK, OBERMAIER 1929 Tue OsBoRN-REEDS CORRELATION TABLE OF 1922 anp 1929. in dark shading. Alternating glacial advances in light shading. Relative time intervals between the four glacial MAP OF THWESTERN INDIA} ADJACENT TERRITORY. E Fieran 6 70 80 90 ; 100 MILES 21380 023050 | ae aie s eth 423/90 \e ) El \ | ~ A Erwin J. Raisz A.M. by the American Muscum. copie struc- ‘number of nent dental on of them, ults desired al intention phs than as had not yet ‘themselves, only to the inadequate ie Shapes of s that have subject the graphs has Anderson’s e benefit of Anderson. e polarized nining the dut, by the so clearly uich can be in subject, d either its distinctive may be of idually and es radically others are ing animal : MAP OF | ‘NORTHWESTERN INDIA) ADJACENT TERRITORY. SCALE 20 30 40 50 60 7080 _80 109 MILES 20630 \\ 4~ : ‘ Fa c= ( 79° Erwin J. Raisz A.M. t—~— AST OF GREENWICH 7 AMERICA Compiled ron ee OF NATURAL HISTORY 1927 rvey of India | loooooo sheets Nrs, 38,39, 43.44, 52,53. Prare XXV. Map of Northwestern India showing the Siwalik Hills and j i fossils the adjacent Salt Range and the important localities where ve. rtebrate fossils have been recovered by the Ameri rican Museum. APPENDIX TO VOLUME II PROBOSCIDEAN DENTAL HISTOLOGY By GEORGE GAYLORD SIMPSON It was Professor Osborn’s intention to include in this Memoir an appendix devoted to the microscopic struc- ture of proboscidean teeth. For this purpose he had Mr. A. E. Anderson prepare and photograph a number of thin sections of such teeth and write notes on some aspects of their study. It was intended that an eminent dental histologist, the late Dr. J. Leon Williams, should also study these sections and prepare an interpretation of them. Unfortunately no part of this plan was carried far enough for it to be possible now to achieve the results desired without a vast amount of further research impossible to us at this time and going far beyond the editorial intention of publishing the work done under Professor Osborn’s direction without substantial change. Mr. Anderson’s notes were intended and are useful rather as a guide for the study of his photographs than as results of such study. Dr. Williams prepared a few additional photographs of the same sections, but had not yet supplied an accompanying manuscript when he died suddenly on February 23, 1932. The sections themselves, and the photographs of them, pertain to only three genera, Elephas, Phiomia, and Trilophodon, and only to the tusks of these except for two sections of a Trilophodon molar. Even within this limited sphere, they are inadequate to reveal all the important details. For instance, there are no tangential sections and without them the shapes of the enamel rods cannot be determined. Despite this very fragmentary character of the data left, it includes photographs and observations that have not been made obsolete by any publication in the meantime and that do give at least a glimpse at a subject the complete omission of which would be a fault in this Memoir. A selection of the most important photographs has therefore been made for publication here, legends for them have been prepared in part from Mr. Anderson’s notes and in part from independent observation, and this introductory comment written, all without the benefit of Professor Osborn’s direction but making available material that was in his hands. With a single exception (PI. xxviu, Fig. 2, by Dr. Williams) these photographs were made by Mr. Anderson. A few were made with ordinary light, but most by means of a petrographic microscope, some in plane polarized light, and some between crossed nicols. The use of crossed nicols is not only valuable for determining the optical properties and mineralogy of the tissues, a matter of secondary interest here, but also to bring out, by the phenomena of birefringence, details of tissue structure and orientation that are not visible, or not so clearly visible, in ordinary or in plane light. This is a technique of considerable importance, the value of which can be seen, for instance, by comparing figures 3 and 4 of Plate xxvu. Although the two are identical in subject, each reveals important structural features not visible in the other. The histology of a tooth is an important part of its structure. It is impossible really to understand either its gross structure or its functional characters without knowing something of its microstructure and of the distinctive features of the tissues composing it. There is also good reason to believe that histological characters may be of value in the identification of fossils and in the determination of animal affinities. Dental tissues individually and in combination have complex characters, fairly constant within limited taxonomic groups and sometimes radically distinct in different groups. Some of these distinctions seem to be adaptive or habitus characters, but others are probably deep-seated heritage characters and they may prove to be exceptionally reliable helps in tracing animal descent. 1607 1608 OSBORN: THE PROBOSCIDEA The present material gives examples of proboscidean dental tissues that must be more or less typical of that group. This purely descriptive aspect is sufficiently expressed by the photographs and their legends. These data also show that for the three genera in question the microstructure even of a simple transverse section of a tusk is determinative. Elephas, Phiomia, and Trilophodon differ in dental histology and on this basis alone any one of these genera can at once be distinguished from the other two. This fact should stimulate the preparation of the many additional sections necessary to determine to what extent thin sections can be useful in the diagnosis and recognition of taxonomic groups of Proboscidea. The histology of these three genera also shows that they have many characters in common, some of which do not appear to be shared by more distantly related animals. More- over Phiomia and Trilophodon resemble each other more than either resembles Elephas. These facts warrant and urge further study to see how far such resemblances can be used to determine relationships. Although inadequate to establish them definitely, these imperfect data also hint at evolutionary trends of considerable interest functionally and in other ways. This is seen, for instance, in the fine structure of the tusk enamel. In Phiomia (e. g., Pl. xxv, Fig. 2), the rods have a pronounced spiral arrangement which makes for toughness and resistance to injury, suggestive of the very tough enamel of rodent incisors. In T'rilophodon (e.g., Pl. xxrx, Fig. 3), this arrangement persists, but in appreciably weaker form. In EHlephas (Pl. xxv1, Fig. 3), it has practically disappeared and the enamel is weak in histologic structure. Thus these three genera, at least, show progressive simplification and weakening of the enamel in microscopic characters as it becomes more reduced macroscopically and less important functionally. The peculiar enamel stringers in Trilophodon and the relation- ships of cement in Trilophodon and Elephas also suggest potentially fruitful lines of investigation. Another will be found in the evolution of the “‘engine-turning”’ effect of the dentine and another in the probable changes in enamel structure in the molars, with changes from the primitive condition to the heavy cones of some later mastodonts or the lamelle of the elephants. Another of the more interesting of the many points merely glimpsed in these sections is the possible relationship between the dentine cylinders of some mastodonts, such as Platybelodon, and the peculiar dentine modification seen in the most juvenile part of a tusk of Elephas (Pl. xxv1, Fig. 1). CATALOGUE NUMBERS OF SPECIMENS MENTIONED IN THE PROBOSCIDEA MEMOIR, VOLUME II Algiers: School of Sciences Los Angeles: Museum of History, Science and Art Amherst: Amherst College Museum Lyons: Museum of Natural Sciences Amsterdam: Zoological Gardens Berlin: Geological-Paleontological Institute of the University pres M var of Ester 3 exico: Engineering School Museum for Natural History Geological Institute Bologna: University Museum Bonn: Museum Brno: Moravian Government Museum Brussels: Museum Moscow: University Munich: State Zoological Museum iW . 7G Waren : f ; ; Bucharest: Laboratory of Geology of the University New York: American Museum of Natural History Laboratory of Paleontology of the University Ror eee ; ra ; i Zoological Society Buenos Aires: Argentinian Museum of Natural Sciences Norwich: Norwich Castle Museum and Art Gallery Calcutta: Asiatic Society of Bengal Indian Museum Oran: Museum Canterbury: Museum Oxford: Museum Capetown: South African Museum Chicago: Field Museum of Natural History College: University of Alaska (formerly Alaska Agricultural College and School of Mines) Paris: Museum of Natural History Pasadena: California Institute of Technology Philadelphia: Academy of Natural Sciences Columbus: Ohio State University Museum ic PinlosoulieallGoc Savin Collection! American Philosophica Society Seas Wagner Free Institute of Science lerence:” Natural Eistory Moseum Private Collections: Walter W. Holmes, Florida Fukuoka: Kyushu Imperial University ae: HOES, ao ? Percy Madeira, Philadelphia —— Natsume, Minato, Japan Geneva: Museum of Natural History Duce d’Orleans, Wood Norton neta Géttingen: Zoological Institute of the University C. V. A. Peel, Oxford, England Gotha: Museum ————Powell-Cotton, Birchington, England Lord Rothschild, Tring, England? Halle: Museum lowalCity-) University oflowa Quito: Museum of the Central University of Ecuador Ipswich: Natural History Museum Rome: University Museum Jena: Museum Rotterdam: Zoological Gardens Kimberley: McGregor Museum Saint John, New Brunswick: Museum Kobe: Takikawa Middle School Sendai: Tohoku Imperial University Kyoto: Imperial University Sendai (probably): Second High School Stockholm: Museum of King Adolph Frederick La Plata: Museum Stuttgart: Wurttemburg Natural History Collection Leiden: Dubois Collection Natural History Museum Tallahassee: Florida Geological Survey Royal Museum of Geology and Mineralogy Tervueren: Museum of the Belgian Congo Leipzig: Paleontological Collection of the University Tokyo: Geological Institute of the Imperial University State Ethnographical Museum (Grassi Museum) Museum of Education at Ochanomizu Leningrad: Museum of the Institute of Mines, Academy of Sciences of the Uyeno Museum USSR. Turin: Museum Paleozoologic Institute of the Academy of Sciences Zoological Museum of the Academy of Sciences Lincoln: Nebraska State Museum Vienna: Natural History Museum Lisbon: Zoological Gardens uondon: Britis : E His F F 2 Sonoon: 7 Belch) Museum: (Natural History) Washington: United States National Museum Imperial Institute : ‘i India House Zoological Park Museum of the Geological Society Royal College of Surgeons Zoological Society ''The entire collection of vertebrate fossils made by Alfred C. Savin was purchased by the British Museum in 1897.—Editor. | *(Probably at the British Museum.—Editor.] Weimar: Natural History Museum Rebling Collection Schwabe Collection 1609 1610 CAT. NO. XII 711 “XVIT 1384 d4 A 86 A 355 A 426 A 437 OSBORN: THE PROBOSCIDEA ALGIERS: SCHOOL OF SCIENCES PAGE Palxvolocod or 7 Olens7s ty eri aie ae oe he cs ee ere eee eee eee a eee ee eee 1275 AMHERST: AMHERST COLLEGE MUSEUM Rarelennas GOLumurit ele eerie tails eee 1051, 1052, 1079, 1080, 1081, 1106, 1111, 1112, 1603 HAMSeleD hasty SUMGICUS Cl te eerie tien erento arlene eee eee et et ee 1351, 1353 AMSTERDAM: ZOOLOGICAL GARDENS Blephas imnarcus: sumatnanus Celis priser sete tee tete eee eee sede eee 1314 BERLIN: GEOLOGICAL-PALHONTOLOGICAL INSTITUTE OF THE UNIVERSITY Pel eOlOGOM OT TECH TOL swe lovee ove tes WAS Se ee ON eae EE Ene aa 1275 Palzolonodon=reck?: lECtOty PC lscscae hice ee eit ea eel he) IS RTL RO eC Oe ae Toe 1275, 1276 Beri: Museum or Naturau History Loxodontasafricana cyclotistty pes ncise den oo eos ee = Se eee eee SCL nae eee ee eee 1193 ToLnodontanairicamarknochenhaieraty pera. cee c > lise: tee erat tara ee 1193 Botoana: Untversiry Museum VAN ANCUSIOTDETTLCTISES TOR 2! boc osu sascha rh atoere lesfecatio a he sad eh Te LT Eee eee eee 1601 Bonn: Museum Mammonteus (?)vpnimigenvus levth-=ad amsvrty pO =e) iaieto eee eee eee ere eee 1150 Brno: MoraviaAN GOVERNMENT MuseuM WVECUNIMONLCUS: PHUMAG CNS TLebe c0s s Hosea Sr), dee CRT eNO CE ee eR eae 1128, 1139 Brussets: MusktumM IGE PG IRONS | GU CO ALDI WNioa gy Oididia HOE Og awoOEto oo OuEpane sas aaa sno scamyoceobebloogomoD Soc Kod ee 5 1130 Bucuarest: LABORATORY OF GEOLOGY OF THE UNIVERSITY FLESDELOLOTOMONLOMULG UMS) G ENNUANECUSLD YC me a1. cer anei eco elie ciclo: ease nciets eres tees aieiet stones Coker 1235 AnchiGiskOlOTepLanumons mUImMOnNUs hye mamis erira cele ei ete eerie iirc kent ete iene) eee 968, 969 Bucuarest: LABORATORY OF PALHONTOLOGY OF THE UNIVERSITY A CHLAISKOLOTE PLAN NOMS MUMLATUS Lele a Mineo eres) aeiiedete etanete ek a rran iee ee ee 969 Buenos AtrES: ARGENTINIAN Museum or NATURAL SCIENCES GUDTERONUTULS SUPETOUS TEL. eet ies seek ais. o a see Ack. choo Ate skh meee enema eee tne ec gen eR ee a 1546 Catcurra: Astatic Soctery oF BENGAL BP OLLOLOTOAON NAMOAAICUS TEL «oo sic cr Gree cers ugh wo de. Sees SEG RMSE SPCR TG CII carte eee are 1211 CatcutTra: INpIAN Musrum Metralophodon' punjabiensis COby Pe’: «jc ss sacs oat vo eee AS Oe be ae et tales erase ee eee 841 Sieqgolopiiodon Sieg OdONLOTdES ty POP hth. dio a ox ccs Oe eeyeptione tise. + cr enn Stan eae ieee 834, 847 Siegolophodonicartleys Th Fo Oe wiv dete oh caske Os Rawr oops d rks ante wee Ns A en ea ea 842 Lralophodon paleindicus type. owe oc wo ae ko Momusiacy opie e teens ah ln Diep a Oe ena eae nent ee 1546 LEG OLOPNOGON: CAULLENT-COUYCS 2. < ee fees) ace 4 sino si Shans: SNe raf cud dae ollekee reuters or tree eee ene eee 841, 842 CANTERBURY: MusruM EL ESET OLOWOCON: OMG UES TER «5 6. eis cco are nicy Sexe wats Une eee rece ae ee EP Tene ees og 1215 CAT. NO. 201616 197 1240 472 1057 1359 167 213 LIST OF CATALOGUE NUMBERS 1611 Capetown: SoutnH ArricaNn Museum PAGE SOT OCOTLONG | MCOAGLO LOULSEUY DC eens rence Cen eT ee ee 1193 Cuicaco: Firtp Museum or Naturau History CSR ARODROCO. CHOLTOUS GEPLTIUEDS WON 5-010 o bo ook bean enonehoeenwensaneoudnegauoausapceoun 1236, 1237 CoLueGE: UNIVERSITY OF ALASKA MiaimMontedsEprumigenistaLaskens? si COLY DC EME reer eee ee eee ene ern 1159, 1160, 1161 CoutumBus: Onto State Universiry Museum VGSLOd ONRAINETACONU SET Clem me nara Tee Ie AACE act ye an hee ee Re ee 1602, 1605 Cromer: Savin Cotiecrion! Al chidiskodommnentadvonalesicromencnsys Lely pene nena eon ce cin seen 963, 980, 981, 1155 PORCLEDRASH US) MENOGONTREIUUESD we meat coroners ees re eae eos a ee reese 981, 1155 FEES DCL GHILOG OTAMLECTAESHCLLUS OTLULUSH( (2) BLE lige neaete tee eee 981 Denver: CoLtorapo Musrym or Narurau History IRORELEPNLASMVASiNG LONI Cling need: adel yar Nie RE 8 io har sae es toca he ee 1103, 1104 Al Elhid¢SKOdOMNAn Old COOK UY; DO aavar ties APE ee yeh Benen ears oR Dye ee eee 1029 Al ChidishodonmmnenLaLonaliswLeDrascensisity|DC a eerene eae enon a ee nc OeE ene ee 1033, 1035 Fiorence: Natura History Museum EVES PEROVOLOCOMOANILGUUS CLISONULS LY DC aE ear eee E inner tate eee eer eee 1215, 1232, 1233 FACS DET OLO LOC OTN OTULG WLESTCLUSOTUULLS IL ere eee ae aoe 1215 VA CHICUSK OC OTIMEMOLONAISIECLOLYD CHEE ieee eEe ECO rec eee EEE ere eree 938, 970, 972, 975 JNA WOT OTTO TARE DCWMIGS COMM occ nocd oonsunddcovedndoncedssngoabnabooenesduwooosecece 971, 977 Anchiduskodon mendronalisiretin etsahh: sere e carer oe dence eae Te Eo eee 975, 1251 Fuxvuoka: Kyusuu ImperitaL UNIVERSITY Stegodon orzentalis. shodoensis else 2 wh oa a Ra eo Gee SOE OEE 893 GrnrevA: Museum or Natura Hisrory Rhynchothennun tlascaleigenoty pe) (Cast) ser enr neon eee ene eee 1377, 1419, 1561 AOTTINGEN: ZOOLOGICAL INSTITUTE OF THE UNIVERSITY Mammonteusprumigeniisilectoty pes anno Eee ee ee eee ee 1122, 1123, 1141 GotTHa: MusrumM FRESE OLOGOMONKCNELG TUES BRC lie etree 1119, 1122, 1181, 1236, 1365 Haute: Muspum HespenolocodonanizquusygenmanecusmeOlyj Pe ann eh eerie erent ete eee 1235 Iowa Ciry: University or Iowa PRareclepnasyeiensonug Olean as iving COT yo ke let er ta see ee ae 1088 Stegomastodonvaftonvetyperh . ger.ieees 4 ie PT eal eee Oa eae se ene ee oes ome ee Ieee ee 1548 Ipswich: Naturau History Museum Anchidiskodon:2 planifronspreliy 3 <) waka enor neninne eis ae eee eo ree iee 963, 981, 1155 Parelephas(?)itrogontheniuisp horton aco ae) ee eee eRe eee 981, 1155 IMQMTLONLEUSYDIUINIGENILS nOSLENSTagLe Uae ee ee ee eee eee 981, 1155 Jena: Musrum Hesperoloxodonanuquus genmanicus MeOty peserer rer ee eee eee ier eee 1234, 1235 The entire collection of vertebrate fossils made by Alfred C. Savin was purchased by the British Museum in 1897.—Editor.] 1612 CAT. NO. 435 2930 3682 3686 3920 4073 4074 4075 4077 4078 4085 4086 4087 4144 4286 4334 B 85 B 122 E2C B 122 E 3 4402 2280 OSBORN: THE PROBOSCIDEA KimMBerLtey: McGrecor Museum PAGE Paloloxodon andrewst ty Pe. <+ - 6 tec sis 2 on ro a= eleva eee retooled a) =ed isheye d= ei sN ol take 993, 1278 Palolomodon hanekom type). = 2 ose cece ec ako see soe oer elec gene ee teste esol ly lps 993, 1279 Anchidiskodon Onoomaiiype. met sea aac eras hee ee eos ho Ne cet ee ee Raye eee 944, 989 Metarchidiskodon griqua genotype... ...... 06... ee cence en eee ee ete teen ete een ees 994, 995 Archidiskodon subplanifrons type... 0... .0 eee eee ec eee eee ee eee eee 987, 988, 1549 Archidiskodow yorkt type. 1s. 2 sei evry ere sien Fetaree eked eked of -0= alka rol= t=) ve) ole eT -Uaoanae 1015 let lan ECO UO Gai OG Boge cok boemncucet Oa5c0 Unabed onc odounoconocamonduocmbooonmbonad 1082 Lerezic: Srare EranoGcrapuicaL Museum (Grasst MusruM) CELA OA Se WOULD UE So GRIER CEE OIE Or Go Pablo no WOSO OH ASA rpOdbUddeCOOUD .OCCObd oUt 1130, 1228 Lenrncrap: Museum or Tae Institute or Mines, ACADEMY OF ScInNCES OF THE U.S.S.R. Hlephase apin7es HNC Wald ss <5 (or. esse rere eee ee saseren Sieber eke Nod eaten eT ecekedeP ass ek ee eto eee 1393 LENINGRAD: PAL#OZOOLOGIC INSTITUTE OF THE ACADEMY OF SCIENCES Mastodon atavus BOrissiak, ty Pe? << isccic) he cde cesutaye ee tena Se eRe yor es extol eRe cheer ret gene eye re rrr 1418 '\Not determined by the present author.—Editor.] CAT. NO. 1-415 14-26 1-11-8-17E 2-7-17B 2353-38-19 4-12-13 §-9-22 5-11-20 8-7-08 11-3-13 13-24-10-14 16-6-16 18-2-22 1O=OS171 23-6-14 29-25-11-18 BE 595 E 622 M 2004 M 2006 M 2009 M 2010 M 2498 M 2705 M 2817 M 2851 M 2857 M 2882 M 2884 2887 M 2978 M 2979 M 3008 M 3015 M 3039 M 3060 M 3068 M 3070 M 3092 M 3109 M 3110 M 3114 LIST OF CATALOGUE NUMBERS 1613 LENINGRAD: ZOOLOGICAL MusreuM OF THE ACADEMY OF SCIENCES PAGE NVIGITANONLEUSED UINAG CISC Lea eg rs secbn.er oe gee ORR ee Oe OE TN, cic cael ces 1130 MN COUMOLALS (FAT TOUS WE 5 o.o.0 oboe gon bobo boontoe oan déeneceenosensaneboouns 1130, 1148, 1387, 1603 VE CITUMONLEUSEDTAIM GEN ULE Cli ewe ree tos rs see olsen A TOT Re Rt Nee OE ase PT EONS OG sists Si sho Ea ee eee 1388 DUT AIS UGG OS VSCOM Res oer aloes ora ttc opoe Eig oe Oana aaS OM RISA TOGe Stee EN ee 5 1388 Lincotn: Nesraska State Museum LARA OUTS FAVASORU Bina oso50500Gaco beckon bonoec 1012, 1019, 1051, 1052, 1091, 1092, 1093, 1106, 1111 NPRO TOUT. HORTLE ULEIOGE Woe ooo ob ab bh hopes cobanboenvoooooabeconboedeucdcaaseccs 1012, 1027 AIT CHIATSKOCONII DENOQLOTaL Clea eee tele oan te Tee ATCT TCE AGEL RTE ee 1011, 1012 Al chidiskodonninypenaroravelere nye tee ee Tee Ode Oe eR ote eee 1009, 1012 “Al chi disk od ont peralvor stele VA e ONC ye OA a eee eerie ta ee ee 1012 Al chidiskodoniim penacorslelaarere meee corer AOC ee ee ee ee 1013, 1018 Archidiskodon imperator maibeni type... .. . 947, 1009, 1012, 1017, 1020, 1021, 1022, 1027, 1028, 1080, 1111 Airchidiskodonwm pen aloravelsrrpe Pere. et ee RE oe eee 1012, 1013 Archi diseOd ONIN DEROLOTATCUS ATR ee Toca re Sea EE Se ae ee eee 1012 Archidiskodonsim peratornelssery ere en eer Arne ete eT ee Ta te ee 1009, 1012 AChidiSkodOnyim Der alorsLeLse meee ey N AT SN ee Se ee ee fe ae 1013, 1017, 1018 Air chidiskodonwm per ators ener ee Eee Lee 1009, 1012 PAMCHTOLS KOC OTUUNY DENGLOTESCOLLLAUY OC Samet ey tera a Seen 1012, 1026, 1027 Archidiskodonwmperatorsreiarre eek iin ee ee ee Re ec eae ee ee 1012 AT chidisk odor Nay vaty Demy na cracks RoR ee eT Te seit te 1006, 1012, 1023, 1024 Anchidiskodonwmperatonireler nen) ot. ero ee OCS eee 1012 Lisson: ZOOLOGICAL GARDENS Monod ontaxalmcancmmocamorcanvy DC Herre ree oe e EL cer CEE or Eee rn eee 1193, 1409 FOLOCOTUOLATCANaTang Glensisnuy pe =P ee Pre eee eee EEC EEE EEE eee eee errr 1193 Lonpon: British Musrum (Naturat History) Archidiskodon planiironsivelc Wax. cad ads noe ee Oe ee re Ole On eo ee CEE 966 Avchidiskodon :plansfrons tel: 5: okies dee Oa en Ce Oe eee 966 Archidiskodonmendionalis Tel. a... i Nae eo ee eee ee Oe Cee 980 Hesperclocodonianiiquius lec toby De Re n eee tere een 1182, 1217, 1218, 1219 tArchidiskodon=plani{ronsiels., se. dee ae ee ee EE Cerne eee 963 Archidiskodon :planifrons) Coty pean ase eee Oe Eee eee 950, 951 Stegolophodonilydekkervityper(Cast) hee eOeor eee eee eee eee eee 851, 1305 Stegolophodonscaitleyilectotypencc-5 ante ee er OO Eee ee 821, 841 Stegolophodoncautleyrcoty pers. sas ee OE Ce aa eee: 821, 840, 841, 842 Stegolopliodoncautleyz reii(cast)he sac ar ote eee ne ee eee 842 Anancus permmensts teliaea heaton cas Geel eae rae Ce Rr oe Ie Oe Inne ne ener 1548 Anancus perrmensis' paraty peur oc a clscls astute on cee oo ee oe Oe eC eee 1392 Stegolophodon' cautleytcotypes: aera oe ee On Oe eee 841 Steqolophodonicautleymcouypen(Cast) meric rere eae eae 841, 842 Stegodon bombifrons lectoty penn smis ots oe eee a ny ere 830, 864, 865 Stegodon: bombafrons: coty pez. ct cacao ene 830, 858, 865, 891 Stegodon ganesateli.\i52 Bie eRe ak Oe oe Oe Ree eRe ere 858, 871, 882 Stegodon:insignis lectoty pesos nace cen or ae ee Ue en eee 867, 881 Stegodoninsignis:cobype ssca seats os tee ROD me rare Sree ee Ce oe eee eee 867 Archidiskodon:planifrons rete nian: Bom ee ee ee 938, 953, 960, 1352 Archidiskodonmplanuifonsiecto ly pe ae etre eee nanan here nie eerie 950, 951, 952, 953, 967 Archidiskodon:planinonsitel. #ca ina ce Gere eee Bl CO eC eee 953, 959, 960 Palzolorodon ‘namadicus ty pens seta co eee OOOO re OE Oe CC Oe oe 1211 Hypselephasihysudnicus Telistiecrrangeke eaten ee oe CO eee 1341, 1350, 1351 Huypselephasihystadricusirelc sttrerve seve cle tecae Pe SIS Osh eEE e eESOES 1352 Hypselephasihiysudiicus el tars tiers re ote ae ch ee OO ROR oe ee 1354, 1355 1614 CAT. NO. M 3127 M 3146 M 3428 7388 7393 7436 M 8588 M 8591 M 9378 M 9381 M 10520 M 10941 M 12639 M 12641 M 12642 M 14102 14759 16229 17420 18489 21680 27872 27915 29007 32250 32251 32252 32253 32254 32256 33218 33327 33334 36695 39370 39463 39464 40769 41925 41926-7 44140 44304 44306 44312 44895 14.2.16.1 OSBORN: THE PROBOSCIDEA Lonpon: British Museum (continued) PAGE Fiypselephas hysudricus tye ssc. 58 al etccs chet ices, Taules gh ae eet ies Tar oe oor roe ae 1341 Mapselephas Nysudricus PAYAty PCr. ciicesecr ces, ous or antel ser RE om eee ote Mein nee ree 1341, 1342 Metralophodon. punjaviensis| COLY Ped (CASU)h caster ete tee eRe ae ne eee ee 841 Stegodon elephaniordes (—clafir2) icotypel(CASt))- 1-144. ee eee ee 863 Steqodonielephantozdes) (—clifiza)) lectotypel (Gast) siya ee ee eee ee dele k teen etree 862 Hespercloxodon: Antiquis TEL’ §< si. Zckas eeu ot tebe eke ede oe eds eevee helena euaen eee aces eee 989 Palzolocodonicypriotes COLYPC..< 2.64 eos oa 45 ee Soe ace Sone see 1266 Palzolomodon. cyprvotes COUPE «a: 2.¢ fia.c si 21< eve bees nee acne wake eats cxeneneuta setae whe ay ote seen kee 1266 Paloloxodon-Cretveus COLY POs «oy avecog wis Aa oe ele 1 eee us Une TSIe CLIO oe RTL Te EO area oe 1267 Palzolorodon creticus CObYPCls>:« 5 5 «rte wa cls ne ease ge erage ele dee oye eee 1267 Steqodonielephantordes) (—clofizy)) cotypesecis +e tee ee eee 825, 826, 831, 862, 863 “Mammuthus’® pramigenvus Tels. .es,50 o's soe hist se ve Cra needa spaces ler elec era 1366 orodontazulu tele. sey eiys cieihete . s0e. see seen ces cee en ae 835, 848, 849, 850, Pl. XIII SICH OLOPROCONANALROLENSTS LYE ac... <1 dacs) fiers Sai, RASC a ee Eee eee 847, 848, Pl. XIII SEG OMGTH DUNUJONETESTS UY DO's cab lob - Oop aS eC Ec TD OS Ga TT AO Ome ice ceo ne a ce nies SEB be 1077 ORClED ROSH CI eNSOILUCELC Leiner war petite ere eee ie 1086, 1087, 1088, 1089, 1095, 1096, 1097 PAU CRLGIS COG ONEUNU DET ALOTAT CL mee appar rys 1k ha NOG EOSIN TSE eee Oe oe ee Pee O02 FA CHUTES KOA OI RUTU DEN CL OTAL CUA Ia Pee cE NE Poe TE I ere aan .. . 1002 IEC MONLEUSE DUN GENUULSEL Clem enetaey Peerin ka he aeera ee mere cece one eee ae ence 1145 LORE NNOS TON ARSCHOL WO Goo bn a 66 one Alan He OR DO CIGNA ae BESO ae se dps A Ruse carkcies ie Rta Cl Ouse 1097 IROPALV NGS TANASCHOD OF IPS WE CMTOR Ws, sone genaood gegh0 Fananneouseh sabes bu dawon dmc oowbas 1088 TAU CHUGUSOCONNEMUDETOLOTELC Mamet near Roce re rene err ae REE ACR CE RS See ER ee ee 1003 PAU CHUATSKOAOIULTE DEL ULOTARC mttaer oire tata tara, Rae ACR eT eran nee Me I at he 1003 VIQUNINONCUSHDRENLIGCILULSELe lemmeae tara eg Wan er Nee ee Pee ere ee Ree ee aes ...1145, 1146 ORAS NOS FAN ARO. Of JP WOR WAGITIOG BM ooo. 600 sono doer odo UD ep nooo wan ueonoLOmbObbOSOOLHE Ore 1088 IPOTRALIN OS. PANCIMONCG. OR IP, CTS VOAIDUE Wt ogo ont ooo one ncer nbocee bore cobconsodeansboconcoune: 1088 IMiGINODLE NS DRUM GenwUs elena keke ee oe er ene 1091, 1145, 1146 MICIIMONTLEUSYDnUTLGCIUUSELe enter Lene eae ren era er ere 922, 939, 1051, 1091, 1148, 1144 Rarclephasijelf er Soni uerelin werner ery tae ee re ee aera 922, 1052, 1084, 1087, 1088, 1090 Archidiskod on winipenr atone Chace tins ior..o tT ROE OR a Tee I ee 996, 1005, 1078 Archidiskod ontim per atonsreltrrtirys cs Spout ee ee CA OTR ere ate Ee ee 996, 1005 Ranelephasslonmaantuswmelteum an tite: Facies taa tee ee cae ee 996, 1077, 1078, 1079, 1108 Parelephasilondanws rela wer a Se so ho as cee OO eee or 996, 1077, 1078, 1079, 1108 BanelepnassLOonidaniussleleren sincere ee nm Oar en ee eee ee 996, 1077, 1078, 1079, 1108 Alrchidishodonimipenalormeln or ane rrr eee er yea imiercm cieocien ech eee so OsO0S WASHINGTON: ZOOLOGICAL PARK Hlephasandrcusisumainanus mel 020 acest A te ee Oe eee 1314 Wermar: Naturau History Musrtum Parelephas trogonthenw ret.,)<. Ge nose hock sneer ne eee ee ee eee 1056 Parelephasirogontherw els. aclcats ace aoe @ ) ene SPOOR Le eC CREE 1056 Parelephas trogontherit relsrcc.8. 2 ast ees ee ae Re ee eee 1056 Parelephastrogonther tel: tas: +455. nua nee rion ee Oe COD Ona Eco 1056 Hesperolozodoniantiquiusigenmanicus MeOLYy DG Heiner er eee eee ee 1234, 1235 Weimar: REBLING COLLECTION Parelephas trogonthervi tel x. .¢ cn chs eevee eae Oe CI ee ee ee eee 1056 Parélephas trogonthertt-Telics 3.ee so pose ese eS OO IO er PE es se eee 1056 Parelephas trogonthervt welion..c ae os coe ae crcpeacene rece aerate cee heey are ee ae ee eee 1056 Parelephastrogontherw ete). yen eee arco Cea eae eae 2 Oe EOE Gane ee ee 1056 WEIMAR: SCHWABE COLLECTION SUPPLEMENTARY BIBLIOGRAPHY See this Memoir, Volume I, page 761 Adams, Michael 1807.1 Revation D’uN VoyaGcr A LA Mer GLACIALE ET D&COUVERTE DES RESTES D’UN MamMourn. Journ. du Nord, St. Petersburg, No. xxxin, Aotit. Original not seen by the present author. Translated by Sir Joseph Banks (see entry under Michael Adams, this Memoir, Vol. I, p. 762). Agassiz, Jean Louis Rodolphe 1850.1 ON THE Fosstt REMAINS OF AN ELEPHANT FOUND IN VERMONT. Proce. Amer. Assoc. Adv. Sci., 2d Meeting, pp. 100, 101. This reference appears in the Bibliography, Vol. I, p. 762, but without specific determination by the present author, of the fossil elephant remains found on the “slope of Mt. Holly, Vermont.’’ Subsequent research has confirmed the identification of Warren (1855, Pl. xxvi1) as Elephas [Mammonteus] primigentus. Alekseev [Alexejew], A. 1930.1 Die OBERSARMATISCHE SAUGETIERFAUNA VON Expar. I. ACHTIARIA BoRISSIAKII N. sp. Trav. Musée Geol., Acad. Sci. U.R.S.S., VII, pp. 167-204, Pls. 1-v, text figs. 1-6. Allen, Glover Morrill 1936.1 ZooLocicaL ResuLTs OF THE GEORGE VANDERBILT AFRICAN EXPEDITION OF 1934. Parr II,-THr Forest ELEPHANT OF AFRICA. Proc. Acad. Nat. Sci. Phila., LX XXVIII, pp. 15-44, Pls. 1-1v, text figs. 1, 2, folding map. Forest and bush elephants, especially the smaller forest elephant, the so-called “Dbygmy elephant,”’ of the Cameroons Ameghino, Florentino 1880-1881 La Anriqiiepap DEL HOMBRE EN EL Piatra. 8vo, Paris and Buenos Aires: I, 1880, pp. i-xiv, 1-640, Pls. xvi; II, 1881, pp. 1-557, Pls. xvui-xxv, folding table. Separation of the Pampean into three successive horizons: Terreno pampeano lacustre, Terreno pampeano superior, and Terreno pampeano antiquo. Andersson, J. Gunnar 1923.1 Essays on THE Cenozoic oF NoRTHERN Cuina. Mem. Geol. Surv. China, (A), No. 3, March, pp. 1-152 (also 16 pp. in Chinese), Pls. 1-1x, text figs. 1-42, maps 1-1n, 2 folding tables. Describes the Lu Tz Kou beds of Shansi, discovered by Zdansky. Andrews, Charles William 1914.1. On tHE Lower Mrocenr VERTEBRATES FROM BRITISH East AFRICA, COLLECTED BY Dr. Frtrx Oswatp. Quart. Journ. Geol. Soe. London, LXX, pp. 163-186, Pls. xxvii-xxrx, text figs. 1-3. Andrussow [Andrusovy], Nikolai Ivanovich 1905.1 Maxnoriscne Strure. Verh. russisch.-kaiserl. Petersb., (2), XLIII, pp. 289-449, Taf. v, vr. Min. Ges. St. Arabu, N. von 1916.1 ErupES SUR LES FORMATIONS TERTIAIRES DU BASSIN DE LA MER pE MARMARA: CLASSIFICATION ET PARALLELISME DES DERNIERES COUCHES NEOGENES DE LA REGION ET DES REGIONS VOISINES. Compt. Rend. Acad. Sci., CLXII, pp. 332-334. Paris. 1916.2 ExisteNcE DE LA FAUNE A HIPPARION DANS LE SARMATIEN DU BASSIN DE LA MER DE MARMARA ET SES CONSEQUENCES POUR LA CLASSIFICATION DU N&OGENE DANS L’EUROPE SUD-ORIENTALE. Compt. Rend. Acad. Sci., CLXIT, pp. 424-426. Paris. 1919.1 RerMARQUES STRATIGRAPHIQUES SUR LES FORMATIONS TERTIAIRES DU BASSIN DE LA Mer pe Marmara. Bull. Soc. géol. France, (4), XVII, for the year 1917, pp. 390-405, text fig. 1. Arambourg, Camille 1933.1 D&couvERTE D’UN GISEMENT DE MAMMIFERES BURDIGALIENS DANS LE Bassin pu Lac RopOLPHE (AFRIQUE ORIENTALE). Compt. Rend. Soe. géol. France, No. 14, November 20, pp. 221, 222. Mastodon af. angustidens. 1934.1 Lr DrNorHERIUM DES GISEMENTS DE L’Omo. Soe. géol. France, No. 6, March 19, pp. 86, 87. D. Bozasi, p. 87. Compt. Rend. New species: See Arambourg, 1935.1, for type figure. 1935.1 Le DrnorHERIUM DES GISEMENTS DE L’OMO (ABYSSINIE). Bull. Soe. géol. France, (5), 1V, February, pp. 305-310, Pl. xvi. Type figure of Dinotherium Bozast, P|. xvut. Astre, Gaston 1937.1 Sur un ExvepHas antiguus a ForMULE LAMINAIRE ELEVER. Bull. Soc. Hist. Nat. Toulouse, LX XI, Fase. 1-2, June 30, pp. 26-32, leit. New mutation: Elephas antiquus ruthenensis Barbour, Erwin Hinckley, and Harold James Cook 1917.1 Norges on THE Skutt or Meroreopon. Neb. Geol. Surv., VII, Pt. 18, April 15, pp. 165-172, text figs. 1-8. 1917.2 SkuLL or AELURODON PLATYRHINUS, SP. NOV. VII, Pt. 19, April 15, pp. 173-180, text figs. 1-11. The name “‘Valentine”’ given to a series of beds in northern Nebraska. Neb. Geol. Surv., Barbour, Erwin Hinckley, and Charles Bertrand Schultz 1937.1 AN Earty PLeisrocenr Fauna rrom Nepraska. Amer. Mus. Novitates, No. 942, September 10, pp. 1-10, text figs. 1-4. Hay Springs fauna, pp. 3, 4, 6. Beliaeva, E. 1936.1 Eryn Funp von EvepHas In TapscuIikistan. Trudy Paleo- zool. Inst., Akad. Nauk SSSR, V, pp. 103-109, 1 pl., 2 text figs., 1 map. Benedict, Francis G. 1936.1 THe PHystoLocy or THE ExterHanr. Publ. Carnegie Instn. Wash., No. 474, pp. vii+302, Pls. -vmt, text figs. 1-13, tables 1-43. Berkey, Charles P., and Frederick K. Morris 1927.1 GroLtoay or Moncoua. Narurat History or CenTRAL AsIA, II. 4to, American Museum of Natural History, New York, pp. xxxi-+475, 44 pls., 161 text figs. Black, Davidson, Pierre Teilhard de Chardin, Chung-Chien Young, and W. C. Pei 1933.1 Fossm, Man in Cuo1na. Mem. Geol. Surv. China, (A), No. 11, May, pp. i-x, 1-166 (also 5 pp. in Chinese), text figs. 1-82, maps I-vr. Blainville, Henri Marie Ducrotay de 1817.1 Dents. Nouv. Dict. d’Hist. Nat., nouvelle édition, IX, pp. 252-352, synoptic table. Paris. The word Mastodontum used for “Genre Mastodonte”’ on page 276. Blumenbach, Johann Friedrich 1803.3. Das Mammut-OnloricuM, NUN WIRKLICH IN LONDON. Maga- zin f. den neuesten Zustand der Naturkunde, herausgegeben J. H. Voigt, V, January, pp. 1-6. Weimar. 1913.1 Urser DIE FOSSILEN GEBEINE VON ELEPHANTEN UND Mam- MUTSTHIEREN, UND UBER ANDERE PRAADAMITISCHE THIER- UND PFLANZEN-RESTE, BESONDERS AUS DEN HANNOVERSCHEN LANDEN. Annalen der Physik, XLV, pp. 425-436. Leipzig. 1624 OSBORN: Borissiak, Alexei Alexievich 1914.1 Mammirires Fossires pe Sepasropon. I. Mém. Comité Géol., (N. 8.), Livr. 87, pp. i-xii, 1-154, Pls. 1-x, text figs. 1-13. St. Petersburg. 1915.1 Mammirires Fossites pe Stpasroron. II. Mém. Comité Géol., (N. S.), Livr. 137, pp. 1-47, Pls. 1-m1, text figs. 1-3. St. Petersburg. 1936.1 Masropon ATAVUS N. SP., DER PRIMITIVSTE VERTRETER DER Grupre M. anGustipens. Trudy Paleozool. Inst., Akad. Nauk SSSR, V, pp. 171-234, 8 pls., 16 text figs. Bose, B. 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MAMMALS OF CENTRAL AFRICA. Journ. Sei., (5), XVII, pp. 101-118. Amer. 1932.1 THe Ovpuvar Expepirion, 1931. Nat. Hist. Mag., III, pp. 214-225, text figs. 1-11. 1933.1 Miocene Primates FROM Kenya. Journ. Linn. Soc. London, (Zoology), XX XVIII, No. 260, November 7, pp. 437-464, Pl. vi. 1935.2 Fossm ELverpHants AnD Man. Proc. Geol. Assoc., XLVI, Pt. 1, March 28, pp. 46-60. London. Table summarizing ‘‘the distribution of the more important of the larger species of mammals found associated with human remains in Europe" (p. 60). Of the elephants, 2. planifrons, E. antiquus, and E. primigenius are mentioned. 1937.1 Tue Ipentiry or ELEPHAS TROGONTHERIU Ponuia. Bull. Geol. Inst. Univ. Upsala, XX VII, pp. 19-24. 1937.2 Die Fosstten Prerpe von Otpoway. Wiss. Ergebn. Oldoway- Expedition, 1913, herausgegeben von H. Reck, Heft 4, pp. 111-136, Pl. vir. 1940.1 Fossis MamMALs AND PLEISTOCENE CorRELATION. Proc. Geol. Assoc., LI, Pt. 1, March 29, pp. 79-88. London. Hopwood, Arthur Tindell, and J. Reid Moir 1939.1 See Moir, J. Reid, and Arthur Tindell Hopwood. Hopwood, Arthur Tindell, and Guy Ellcock Pilgrim 1939.1 See Pilgrim, Guy Elleock, and Arthur Tindell Hopwood. Hutchinson, G. E., and Hellmut de Terra 1936.1 See Terra, Hellmut de, and G. E. Hutchinson. Issel, A. 1879.1 DerscrIzIONE DI DUE DENTI D’ELEFANTE RACCOLTI NELLA LiguRIA OCCIDENTALE. Ann. Mus. Civ. Storia Nat. Genova, XIV, pp. 153-168, text figs. 1-4. Johnson, F. Walker 1936.1 Tue Srarus or THE NAMb “VALENTINE” IN TERTIARY GEOLOGY AND PaLteontoLtoagy. Amer. Journ. Sei., (5), XXXI, June, pp. 467-475, text figs. 1, 2. Johnston, H. H. [Henry Hamilton) No date Tur OpreninG Up or Arrica. 16mo, Henry Holt and Com- pany, New York, Williams and Norgate, London, 255 pp., 3 maps. 1905.1 A Hisrory or THE COLONIZATION Or AFRICA BY ALIEN Races. 8vo, Cambridge University Press, pp. xiii+349, 8 pls. (maps). Kay, George F. 1931.1 CLassiFICATION AND DURATION OF THE PLEISTOCENE PERIOD. Bull. Geol. Soc. Amer., XLII, March 31, pp. 425-466, text figs. 1-9. Kellogg, Remington 1924.1 Tertiary Peracic MamMats or EasteRN Nortu AMERICA. Bull. Geol. Soc. Amer., XX XV, December 30, pp. 755-766. Age of the Bone Valley formation Khomenko, J. 1913.1 La FAUNE MBOTIQUE DU VILLAGE TARAKLIA DU DISTRICT DE Banpery. I. Les aNcETRES DES Cervinar. IT. GrrarrINAr ET Cavicornia. Annuaire Geol. Min. Russie, XV, Livr. 4-6, pp. 107-143, Pls. vi-rx. Novo-Alexandria. OSBORN: THE PROBOSCIDEA 1914.1 La FrAUNE MBOTIQUE DU VILLAGE TARAKLIA DU DISTRICT DE Benpery. Fissirpepia, RODENTIA, RHINOCERINAE, EQUINAR, SUIDAE, Progposcrpra. Trudy Bessarabskoe obshchestvo estestvoispytatelei, V, pp. 1-55, Pls. tv. Kishinef. Kirk, John 1864.1 List or MAMMALIA MET WITH IN ZAMBESIA, East TROPICAL Arrica. Proe. Zool. Soe. London, pp. 649-660. Koenigswald, G. H. Ralph von 1931.1 Dir BepeuruNe DER EQuIDEN FUR DIE ALTERSSTELLUNG DES RHEINHESSISCHEN DINOTHERIENSANDES. Centralb. Min. Geol. Pal., Abt. B, pp. 42-48, 1 text fig. Stuttgart. 1939.1 Das Pietsrocan Javas. Quirtar (Berlin), II, pp. 28-53, Taf. 1x-x1, Abb. 1-6. Mounted skeleton of Stegodon trigonocephalus in the Geological Museum of Bandoeng. 1939.2 THe RELATIONSHIP BETWEEN THE Fosst. MAMMALIAN FAUNAE oF JAVA AND CHINA, WITH SPECIAL REFERENCE TO EARLY Man. Peking Nat. Hist. Bull., XIII, Pt. 4, June, pp. 293-298, 1 text fig. Kraglievich, Lucas 1934.1 La ANTIQUEDAD PLIOCENA DE LAS FAUNAS DE Monte HermMoso y CHAPADMALAL, DEDUCIDAS DE SU COMPARACION CON LAS QUE LE PRECEDIBRON Y SUCEDIERON. 8vyo, ‘‘El Siglo Ilustrado,’”’ Montevideo, 136 pp., fontispiece and 1 text fig. Designated the Lower Pampean, or HEnsenadense, as ‘‘Upper Pliocene or Pliopleistocene.” Krejci-Graf, Karl 1932.1 PARALLELISIERUNG DES SUDOSTEUROPAISCHEN PLIOZANS. Rundschau, XXIII, pp. 300-339, text fig. 1. Beds of true Sarmatian age, bearing Hipparion. Geol. Lapparent, A. de 1906.1 Trairé pe Gtonoaiz. Fifth edition, 8vo, Masson et Cie., Paris, III, pp. 1289-2015, text figs. 584-883. Remarks concerning the Gningen fossils. Leverett, Frank 1902.1 GuacraL Formations AND DRAINAGE FratTURES OF THE ERIE AND Onto Basins. Monogr. U.S. Geol. Surv., XLI, pp. 1-802, Pls. I-XXVI, text figs. 1-8. Lewis, G. Edward 1937.1 A New Siwauik Correvation. Amer. Journ. Sci., (5), XX XIII, March, pp. 191-204, text figs. 1, 2. Tawi, new name for Boulder Conglomerate. Licent, Emile, and M. Trassaert 1935.1 Tur PLiocenr LACUSTRINE SERIES IN CENTRAL SHANSI. Geol. Soc. China, XTV, pp. 211-219, text fig. 1, 2 maps. Bull. Lugn, Alvin L. 1935.1 Tue PLeistoceNr Grotocy or Nesraska. Neb. Geol. Surv., (2), Bull. 10, pp. 1-223, Pls. 1, u1, text figs. 1-38, tables A-D. 1939.1 CLassiricaTION or THE TERTIARY SysteM IN Nepraska. Bull. Geol. Soe. Amer., L, No. 8, pp. 1245-1275, 1 pl. Use of the long abandoned name Ogallala for a large stratigraphic group of Pliocene age. MacCurdy, George Grant 1924.1 Human Oriains. A Manuva or Prenisrory. S8vo, D. Apple- ton and Company, New York, I, pp. xxxviii+-440, frontispiece, text figs. 1-254; II, pp. xvi+-516, frontispiece, text figs. 255-410. BIBLIOGRAPHY Makiyama, Jiro 1929.1 Chikyi-The Globe, XII, No. 5, pp. 364, 365 (in Japanese). Not available to the present author. New subspecies: Elephas (Palzolorodon) namadicus setoensis. 1938.1 Jaronic Proposcippa. Mem. College Sci. Kyoto Imp. Univ., (B), XIV, No. 1, Art. 1, May, pp. 1-59, text figs. 1-31. New species: Bunolophodon yokotii, pp. 12-14, figs. 5a, 5b. Stegodon shodoensis akashiensis (Takai, 1936) =Parastegodon akashiensis (p. 21). Stegodon insignis sugiyamai (Tokunaga, 1936) = Parastegon sugiyamai (p. 27). Mansuy, H. 1916.1 Sur Que.qurs Mammiréires Fossttes RéicemMMENT Dicouverts EN INpocutNE. Mem. Serv. Geol. Indochine, V, Fasc. II, pp. 1-26, Pls. 1-vur, 1 text fig. Stegodon insignis, S. Cliftii, Elephas sp. ? aff. E. namadicus, E. namadicus, and E. indicus. Mather, Cotton 1717.1. An Exrract or SEVERAL Lerrers FROM Corron Marner, D. D. to JoHN Woopwarp, M. D. anp RicHarp Water, Esq.; S. R. Secr. Phil. Trans., XXIX, No. 339, for April, May, and June, 1714, pp. 62-71. Matsumoto, Hikoshichiro 1939.1 On Some Fosstt ELEPHANTS FROM PROVINCE OF Kazusa, FROM PROVINCE OF SHIMOTSUKE, AND FROM OTHER LocaLities. Dobu- tugaku Zassi (Zool. Mag., Tokyo), LI, No. 10, October, pp. 701-717, text figs. 1-8. New species: Archidiskodon paramammonteus, pp. 704, and 716 (English) Matthew, William Diller 1924.1 Turrp ConrTriBuTION TO THE SNAKE CrEEK Fauna. Bull. Amer. Mus. Nat. Hist., L, Art. II, July 3, pp. 59-210, text figs. 1-63. Matthew, William Diller, and Ruben Arthur Stirton 1930.1 EQuipar FROM THE PLIocENE oF Texas. Bull. Dept. Geol. Univ. Calif., XTX, No. 17, November 29, pp. 349-396, Pls. xiv-Lv1m1. Maxson, John H. 1930.1 A Tertiary Mammauian Fauna From THE Mint Canyon FORMATION OF SOUTHERN CaLirornia. Publ. Carnegie Instn. Wash., No. 404, August, pp. 77-112, text figs. 1-18. Merriam, John Campbell 1919.1 Tertiary Mammanian Faunas or tHE Mouave Desert. Bull. Dept. Geol. Univ. Calif., XI, No. 5, pp. 4387a-e, 488-585, text figs. 1-253. Fauna of the Barstow formation. Merriam, John Campbell, Chester Stock, and C. L. Moody 1925.1 Tur Puiocenr RATTLESNAKE FORMATION AND FAUNA OF Eastern OrgGoN, with Nores ON THE GEOLOGY OF THE RATTLE- SNAKE AND Mascauu Deposits. Publ. Carnegie Instn. Wash., No. 347, October 8, pp. 43-92, text figs. 1-45. Meyer, Hermann von 1841.1 Fossmum KnocHEN von WikEsBADEN. Neues Jahrb. Min., pp. 458-461. New species: Dinotherium minutum, p. 499. Middendorf, Alexander Theodor von 1860.1 Usprsicut per Narur Norp- uNp Ost-Srprriens. Reise in den Aussersten Norden und Osten Sibiriens wahrend der Jahre 1843 und 1844. .., IV, Th. I, Lief. 2, pp. 201-3832. 4to, St. Petersburg. 1627 Moir, J. Reid, and Arthur Tindell Hopwood 1939.1 Excavations ar BruNpDON, SurroLtK (1935-37). Parr I. STRATIGRAPHY AND ARCHAEOLOGY (Moir). Parr II. Fossm Mam- MALS (Hopwood). Proc. Prehist. Soc., (N. 8.), V, No. 1, pp. 1-82, Pls. 1, 0, text figs. 1-18. Moody, C. L., John Campbell Merriam, and Chester Stock 1925.1 See Merriam, John Campbell, Chester Stock, and C. L. Moody. Morris, Frederick K., and Charles P. Berkey 1927.1 See Berkey, Charles P., and Frederick K. Morris. Osborn, Henry Fairfield 1933.901 BroLtoaican INpDUCTIONS FROM THE EVOLUTION OF ‘THE ProgposcipEA. Proc. Nat. Acad. Sci., XIX, No. 1, January, pp. 159-163. 1935.940 THe Recorp ProposcipeaAN Tusk. Nat April, p. 357. (Unsigned article.) Record tusk of Archidiskodon imperator from Texas. Hist.. XXXV, 1936.951 L’AGr GfoLoaiquE DE L’ HOMME DE PitrpowNn (EKoANTHRO- PUS) ET DE L’ HOMME DE TRINIL (PITHECANTHROPUS). Mélanges de Préhistoire et d’Anthropologie offerts au Professeur Henri Bégouen a l’oeeasion de son 70° Anniversaire (20 Novembre, 1863-1933) par ses Hléves, ses Collégues et ses Amis, pp. 23-36, text figs. 1-6. These sheets were received September 28, 1936, with a letter from Comte Bégouen in which he says: “* L'impression fut meme un instant arretée et elle vient de recommencer. Je vous envoie, sous pli separé, en justificatif les premiéres bonnes feuilles. . . .'". No other sheets have been received. 1938.952 EiaHTnEN PRINCIPLES OF ADAPTATION IN ALLOIOMETRONS AND ARISTOGENES. Palaeobiologica, VI, pp. 273-302, text figs. 1-12. Paterson, T. T., Hellmut de Terra, and Pierre Teilhard de Chardin 1936.1 See Terra, Hellmut de, Pierre Teilhard de Chardin, and T. T. Paterson. Peale, Rembrandt 1802.1 AccouNT OF THE SKELETON OF THE Mamoru, A Non-Descripr Carnivorous ANIMAL OF IMMENSE SIZE FOUND IN AMERICA. Sm. 8vo, London, 46 pp. 1803.1 AN Historica Disquisirion ON THE Mammornu, or, GREAT AMERICAN INcoGNITUM, AN Extinct, IMMENSE, CARNIVOROUS ANIMAL, WHOSE Fosstn REMAINS HAVE BEEN FOUND IN Norru America. 12mo, EK. Lawrence, London, pp. viii+91, 1 pl. Pei, W. C., Davidson Black, Pierre Teilhard de Chardin, and Chung- Chien Young 1933.1 See Black, Davidson, Pierre Teilhard de Chardin, Chung-Chien Young, and W. C. Pei. Pfizenmayer, F. W. 1937.1 Mammut-FunpDe IN SIBIRIEN. pp. 279-288, text figs. 6-14. Sangajurach-Mammuts. Natur u. Volk, LXVII, June, Piette, Edouard 1907.1 L’Arr PENDANT L’AGE DU RENNE. pp. iv+12, 100 pls., 128 text figs. 4to, Masson et Cie., Paris, Pilgrim, Guy Ellcock 1931.1 CaraLogur or THE PonriAN Carnivora or Europe IN THY DeparRTMENT or GuoLoey. 4to, British Museum (Natural History), London, pp. vi+174, 2 pls., 30 text figs. 1934.1 CorrELATION OF OsSIFEROUS SECTIONS IN THE Upper Cenozoic or Inpra. Amer. Mus. Novitates, No. 704, March 15, pp. 1-5. 1628 OSBORN: THE Pilgrim, Guy Ellcock—Continued 1939.1 Tue Fosstz Bovinpar or InpiA. Mem. Geol. Surv. India, Palacont. Indica, N.S., XX VI, Mem. 1, pp. iii+356, 8 pls., 35 text figs. APPLICATION OF THE EuROPEAN TIME SCALE TO THE Geol. Mag., LX XVII, pp. 1940.1 THE Upper Tertiary oF NortH AMERICA. 1-27. Pilgrim, Guy Ellcock, and Arthur Tindell Hopwood 1939.1 Are THE Equipar RELIABLE FOR THE CORRELATION OF THE SrwALIKS WITH THE ComNOzoIc STaGEs or NortH America (Pilgrim). APPENDIX ON THE CORRELATION OF CrertraIn TrerTIARY Deposits or INDIA AND Evropge (Hopwood). Ree. Geol. Surv. India, LX XIII, Pt. 4, December, 1938, pp. 437-482. Piveteau, Jean, and Marcellin Boule 1935.1 See Boule, Marcellin, and Jean Piveteau. Pohlig, Hans 1887.3 [Casts or ELEPHANT TEETH, E. ANTIQUUS TYPUS AND E. ANTIQU- US VAR. MINOR.]| Verh. natur. Vereins preuss. Rhein., Jahrg. XLIV, p. 115. Bonn. New subspecies: E. [Elephas) antiquus var. minor. Pontier, G. 1930.1 SCIDIENS. A pROPOS D’ ANOMALIES DENTAIRES OBSERVEES CHEZ LES PROBO- Ann. Soe. géol. du Nord, LV, pp. 2-10, Pls. 1, m1. Roman, F., and J. Viret 1934.1 La Faunr pe Mammirires pU BURDIGALIEN DE LA RomInU (Gers). Mem. Soc. géol. France, (N.S. ),[X, Fase. 2-3, (Mem. No. 21), pp. 1-67, Pls. 1-xrx, text figs. 1-25. Romer, Alfred Sherwood 1933.1 PLrIstoceNE VERTEBRATES AND THEIR BEARING ON THE PrRoBLEM OF Human Antiquiry in NortrH America. In ‘The American Aborigines, Their Origin and Antiquity,” edited by Diamond Jenness. 8vo, University of Toronto Press, pp. 47-83. Roverto, Cayetano 1914.1 Jos Esrraros ArAucanos y Sus Fosines. An. Mus. Nac. Hist. Nat. Buenos Aires, X XV, pp. 1-249, Pls. 1-xxx1, text figs. 1-92. Rusconi, Carlos 1937.1 CoNTRIBUCION AL CONOCIMIENTO DE LA GEOLOGIA DE LA CIUDAD bE BUENOS AIRES Y SUS ALREDEDORES Y REFERENCIA DE SU FAUNA. Actas Acad. Nae. Cien. Cérdoba, X, Nos. 8-4, December 20, pp. 177-384, Pls. 1-x1x, text figs. 1-58. Buenos Aires. Russell, R. Dana, and V. L. VanderHoof 1931.1 A VertreBRaTe Fauna rrom A New PiioceEN® ForMATION IN NortHerRN Catirornia. Bull. Dept. Geol. Univ. Calif., XX, No. 2, February 5, pp. 11-21, text figs. 1-7. Tehama formation named and described. Contains Stegomastodon cf. arizonz. Saheki, Shird 1931.1 ON PareLEPHAS PROTOMAMMONTEUS (MatrsumMoTo) RECENTLY Founp in THE Province or Kazusa. Japanese Journ. Geol. and Geog., VIII, No. 3, February, pp. 125-129, Pl. xv, 1 text fig. New subspecies: Parelephas protomammonteus (Matsumoto) matsumotoi, p 127. Schneider, Carlos Oliver 1929.1 La Disrripuci6n GEOGRAFICA DE LOS MASTODONTES EN CHILE. Actes Soe. Scientifique Chili, XXXVI, pp. 73-83, 1 map. Santiago. PROBOSCIDEA 1930.1 ALGUNOS COMENTARIOS SOBRE MASTODONTES CHILENOS. Revista Universitaria (Univ. Catdélica de Chile), No. 8, Ano XV, pp. 886-893. Santiago. Remarks on the questionable type locality of Mastodon [Cuvieronius] humboldtii. Schroeder, Henry 1928.1 User ELepHAs ANTIQUUS UND TROGONTHERIL AUS DEM Dituvium DER MirretMarK. Jahrb. Preuss. Geol. Landesanstalt, Berlin, XLVITI, for the year 1927, pp. 699-723, Pls. xxxiv-xxxv1, text fig. 1. New subspecies: Elephas primigenius Blumenb. var. n. pachyganalis, p. 718. Schultz, Charles Bertrand, and Erwin Hinckley Barbour 1937.1 See Barbour, Erwin Hinckley, and Charles Bertrand Schultz. Scott, William Berryman 1937.1 A Hisrory or Lanp MAMMALS IN THE WESTERN HEMISPHERE. Revised edition, 8vo, Macmillan Company, New York, pp. xiv-+786, frontispiece, 420 text figs. New family: Stegomastodontide. New subfamilies: Tmlophodontins, Pentalophodonting, Cordillerionine, Stegomastodontine. Serebryakov, A. H. 1938.1 HLerHas MAMMONTEUS Cuvier VERSUS E.. PRIMIGENIUS BLUMEN- BAcH. Bull. Acad. Sci. U. 8. 8. R., Classe des Sci. Math. et Nat., Serie Biol., pp. 1063-1068. He regards the name Hlephas primigenius Blumenbach (1799) as an indis- putable nomen nudum, hence must be rejected, and maintains that 2. mammonteus Cuvier (1799) must be restored, recognizing as the type the skull described by Messerschmidt in 1724, or Adams’ mammoth (Tilesius, 1815). Shikama, Tokio 1936.1 Nore oN PARASTEGODON AKASHIENSIS TAKAI FROM THE AKASI Disrricr. Proc. Imp. Acad., Tokyo, XII, No. 1, January, pp. 22-24, text figs. 1-4. 1937.1 PARASTEGODON INFREQUENS SP. NOY. FROM THE AKAsI District. Japanese Journ. Geol. and Geog., XIV, Nos. 3-4. October, pp. 127— 131 Pljox, 1937.2 Nomenciative Notes ON ParRELEPHAS PROTOMAMMONTEUS (Marsumoro). Japanese Journ. Geol. and Geog., XIV, Nos. 3-4, October, pp. 163-166. New subspecies: Parelephas proximus uehataensis. Shuler, Ellis W. 1934.1 Cotiectinc Fosst, Evepnants ar Dauuas, Texas. Bull. Texas Archaeol. and Palaeont. Soe., VI, September, pp. 75-79, Pl. x11. Soergel, Wolfgang 1921.3 Die URSACHEN DER DILUVIALEN AUFSCHOTTERUNG UND EROSION. 8vo, Borntraeger, Berlin, pp. iv-+-74, 1 text fig. 1925.1 Dre SAvUGPTIBRFAUNA DES ALTDILUVIALEN TONLAGERS VON JOCKGRIM IN DER Prauz. Zeits. Deutsch. Geol. Ges., LX XVII, Heft 3, November 20, pp. 405-438, Taf. xvit, 2 tabellen. Spock, Leslie Erskine 1930.1 New Mesozoic anp Crenozoic FORMATIONS ENCOUNTERED BY THE CenTRAL AstAtTic EXPEDITIONS IN 1928. Amer. Mus. Novitates, No. 407, March 18, pp. 1-8, text figs. 1-6. Stamp, L. Dudley 1922.1 An OvuTLINe or THE TERTIARY GEOLOGY or BurMA. Geol. Mag., LIX, No. 11, November, pp. 481-501, text figs. 1-6. BIBLIOGRAPHY Stehlin, Hans Georg 1908.1 Noricks PALKOMAMMALOGIQUES SUR QUELQUES DEPOTS Mro- CkNES DES BAssINs DE LA LOIRE ET DE L’ALLIER. Bull. Soe. géol. France, (4), VII, for the year 1907, pp. 525-550, text figs. 1-3. Stehlin, Hans Georg, and Auguste Dubois 1932.1 and 1933.1 See Dubois, Auguste, and Hans Georg Stehlin. Sternberg, C. M. 1930.2 MroceNnr GRAVELS IN SOUTHERN SASKATCHEWAN. Trans. Roy. Soe. Canada, (3), X XIV, Sec. 4, pp. 29, 30. The scanty and fragmentary fauna includes an indeterminate Mastodont. Stirton, Ruben Arthur 1933.1 A Critical Revirw or THE Mint Canyon MAmMMaLiaAN Fauna AND ITS CORRELATIVE SIGNIFICANCE. Amer. Journ. Sci., (5), X XVI, December, pp. 569-576. 1936.1 Succession or NorroH AMERICAN CONTINENTAL PLIOCENE MaAmMatian Faunas. Amer. Journ. Sci., (5), XX XII, pp. 161-206. 1939.1 Mrruops anD PROCEDURE LN THE VALENTINE QuESTION. Amer. Journ. Sci., CCX XXVII, pp. 429-483. Stirton, Ruben Arthur, and William Diller Matthew 1930.1 See Matthew, William Diller, and Ruben Arthur Stirton. Stock, Chester 1928.1 Canip AND ProposcIDEAN REMAINS FROM THE Ricarpo DrE- posits, Mowave Desert, Cauirornia. Publ. Carnegie Instn. Wash., No. 393, pp. 39-47, Pls. 1-1v, text fig. 1. Trilophodon sp., consisting of fragmentary skull, incomplete mandibular ramus, humerus, femur, and a caleaneum, found at three distinct localities in the Ricardo deposits. 1936.1 A PLIOoMAsTODON SKULL FROM THE THOUSAND CREEK Beps, NortTHWESTERN Nevapa. Publ. Carnegie Instn. Wash., No. 473, July 10, pp. 35-39, Pl. 1. New species: Pliomastodon nevadanus, p. 37. Stock, Chester, John Campbell Merriam, and C. L. Moody 1925.1 See Merriam, John Campbell, Chester Stock, and C. L. Moody. Stromer, Ernst 1907.1 GroLogiscHeE BrOBACHTUNGEN IM FasUM UND AM UNTEREN Nivrate in Aeypren. Abhand. Senckenb. naturf. Ges., X XTX, Heft 2, pp. 183-148, Taf. xx1. Takai, Fuyuji 1936.1 On a New Fossiu ELepHant rrRoM OKUBO-MURA, AKASHI-GUN, Hyogo Prerecrure, JAPAN. Proce. Imp. Acad., Tokyo, XII, No. 1, January, pp. 19-21, text figs. 1, 2. New species: Parastegodon akashiensis, p. 20. 1936.2 Fossm ELepHants FROM TrpA PREFECTURE, JAPAN. Japanese Journ. Geol. and Geog., XIII, Nos. 3-4, October, pp. 197-203, PI. xxiv, text fig. 1. The author states that from “this district, the following species have thus far been reported: Stegodon orientalis Owen. Parelephas protomammonteus typicus Matsumoto. P. protomammonteus proximus Matsumoto. P. protomammonteus matsumotoi Saheki. Palaeoloxodon naumanni (Makiyama).”’ Takai, Fuyuji, and Shigeyasu Tokunaga 1936.1 See Tokunaga, Shigeyasu, and Fuyuji Takai. Teilhard de Chardin, Pierre 1937.1 THe Post-VitLarRANCHIAN INTERVAL IN NorrH CHINA, Geol. Soe. China, XVII, No. 2, June, pp. 169-176, text fig. 1. Bull. 1629 Teilhard de Chardin, Pierre, Davidson Black, Chung-Chien Young, and W. C. Pei 1933.1 See Black, Davidson, Pierre Teilhard de Chardin, Chung-Chien Young, and W. C. Pei. Teilhard de Chardin, Pierre, and Hellmut de Terra 1936.1 See Terra, Hellmut de, and Pierre Teilhard de Chardin. Teilhard de Chardin, Pierre, Hellmut de Terra, and T. T. Paterson 1936.1 See Terra, Hellmut de, Pierre Teilhard de Chardin, and T. T. Paterson. Teilhard de Chardin, Pierre, and M. Trassaert 1937.1 Tue Proposcipeans or Soura-Easrern SuHansi. (YUSHE BASIN). Pal. Sinica, (C), XIII, Fase. 1, March, pp. 1-84 (also 4 pp. in Chinese), Pls. 1-x11, text figs. 1-6. New species: Pentalophodon cuneatus, p. 11; Stegodon licenti, p. 27. Mastodon intermedius, p. 22; Teilhard de Chardin, Pierre, and Chung-Chien Young 1936.1 ON THE MAMMALIAN REMAINS FROM THE ARCHAOLOGICAL SITE or AnyANG. Pal. Sinica, (C), XII, Fase. 1, pp. iii-61 (also 8 pp. in Chinese), Pls. 1-vu1, text figs. 1-26. Terra, Hellmut de 1936.1 Larr Cenozoic History iy Inptia. 3469, April 25, pp. 686-688. Nature, CXXXVII, No. Terra, Hellmut de, and G. E. Hutchinson 1936.1 Dara on Posr-GuactaL Cumatic CHancrs IN Nortx-Wesr Inpra. Current Science, V, July, pp. 5-10. Bangalore City. Terra, Hellmut de, and Pierre Teilhard de Chardin 1936.1 OBSERVATIONS ON THE UPPER SIwaALik FORMATION AND LATER PLEISTOCENE Deposits IN INprIA. Proce. Amer. Phil. Soe., LX XVI, pp. 791-822, text figs. 1-14. Terra, Hellmut de, Pierre Teilhard de Chardin, and T. T. Paterson 1936.1 CeEnozotc IN InpIaA. 6, pp. 233-236. Jornt GEOLOGICAL AND PREHISTORIC STUDIES OF THE LATE Science, (N. S.), LX XXIII, No. 2149, March Thomas, Oldfield 1895.1 Dr. C. W. L. GuoGgeEr’s ‘‘NATURGESCHICHTE”’ (1841). Nat. Hist., (6), XV, pp. 189-193. An ANALYSIS OF THE MAMMALIAN GENERIC NAMES GIVEN IN Ann. Mag. Tobien, H. 1938.1 MOLASSE SUDWESTDEUTSCHLANDS. Heft 4, May 5, pp. 177-192, Taf, v1. User Hipparion-REsts AUS DER OBERMIOZANEN SiisSWASSER- Zeits. Deutsch. Geol. Ges., XC, Tokunaga, Shigeyasu 1933.2 A List or THE Foss, LAanD MAMMALS OF JAPAN AND KoREA with Descriertions oF New Eocene Forms From Korea. Amer. Mus. Novitates, No. 627, May 27, pp. 1—7, text figs. 1, 2. 1934.1 Fossm ELEPHANT TEETH FOUND AT YOKOHAMA AND Kakio, KanaGawa PREFECTURE. Journ. Geog., XLVI, No. 546, July, pp. 363-371, Pls. vii, rx, 5 text figs. Tokyo. New species: Palaeoloxodon yokohamanus, p. 363, Pl. vii; kwantoensis, p. 365, Pl. rx. Text in Japanese. Parastegodon? 1935.1 A New Fossi. ELEPHANT FOUND IN SHIKOKU, JAPAN. Proc. Imp. Acad., Tokyo, XI, No. 10, December, pp. 432-434, text fig. 1. New species: Parastegodon sugiyamat. 1630 Tokunaga, Shigeyasu— Continued 1936.1 GroLocy or THE DisrRicr OF SHICHINOHE AND Fossin ELrE- PHANT FOUND THERE. Journ. Geog., XLVIII, No. 564, February, pp. 67-70, Pl.1. Tokyo. Original description (in Japanese) of Palaeoloxodon aomoriensis, p. 70. Tokunaga, Shigeyasu, and Fuyuji Takai 1936.1 On a Fossi ELepHANT, PALAEOLOXODON AOMORIENSIS, FROM SHICHINOHE, KaMIKATA-GUN, AOMORI PREFECTURE, JAPAN. Journ. Geol. Soc. Japan, XLIII, No. 511, April 20, pp. 254-258, Pls. x11 (1m), XIV (rv). Type description (in English) of Palaeoloxodon aomoriensis. Trassaert, M., and Emile Licent 1935.1 See Licent, Emile, and M. Trassuert. Trassaert, M., and Pierre Teilhard de Chardin 1937.1 See Teilhard de Chardin, Pierre, and M. Trassaert. Troxell, Edward L. 1916.1 AN Earty PLioceNr ONr-Torp Horse, PLIOHIPPUS LULLIANUS, sp. Noy. Amer. Journ. Sci., (4), XLII, October, pp. 335-348, text figs. 1-7. Oak Creek formation named and described, pp. 345-348. VanderHoof, V. L. 1933.1 AppITIONS TO THE FAUNA OF THE TEHAMA UPPER PLIOCENE OF NorTHERN CaLirorNia. Amer. Journ. Sci., (5), XXV, pp. 382-384. VanderHoof, V. L., and R. Dana Russell 1931.1 See Russell, R. Dana, and V. L. VanderHoof. Van Es, L. J. C. 1931.1 THe Acre or PITHECANTHROPUS. Hague, pp. xii-+142, 4 pls., 11 maps. 8vo, Martinus Nijhoff, The van Riet Lowe, C. 1929.1 Furrner Nores oN THE ARCHAEOLOGY OF SHEPPARD ISLAND. So. Afr. Journ. Sci., XX VI, pp. 665-683, text figs. 1-5. Excellent table of associated fauna during the Pleistocene, including among the Proboscidea Mastodon (Bunolophodon) sp. [=Trilophodon], Archidiskodon subplanifrons, and A. [=Palxolozodon) transvaalensis and A. [ = Palxoloxo- don| sheppardi of the early Pleistocene, to A. broomi of the late Pleistocene. Stellenbosch industry of the Stone Age. OSBORN: THE PROBOSCIDEA Viret, J.,and F. Roman 1934.1 See Roman, F., and J. Viret. Ward, Rowland 1922.1 Row Lanp Warp’s Recorps or Bic GAME witH THEIR DisTRI- BUTION, CHARACTERISTICS, DIMENSIONS, WEIGHTS, AND HORN AND Tusk Measurements. Eighth edition edited by J. G. Dollman and J. B. Burlace, 8vo, Rowland Ward, London, pp. xiii+-527, illustrated. 1928.1 Rowxianp Warp’s Recorps or Bie Game with THEIR DistRI- BUTION, CHARACTERISTICS, DimMeNsIoNs, WEIGHTS, AND HORN AND Tusk MeasurEMents. Ninth edition edited by J. G. Dollman and J.B.Burlace, 8vo, Rowland Ward, London, pp. xiii+-523, illustrated. Weber, Max 1896.1 [1897] VorsrupIEN tiper DAS HIRNGEWICHT DER SAUGETHIERE. Festschrift z. Siebenzigsten Geburtstage von Carl Gegenbaur, ITI, pp. 103-123. Wilmarth, M. Grace 1938.1 Lexicon or GroLoagic NAMES OF THE UNITED STATES (INCLUD- tNG AuasKA). Bull. U. 8. Geol. Surv. 896: Part 1, A-L, pp. 1-1244; Part 2, M-Z, pp. 1245-2396. Young, Chung-Chien, and Pierre Teilhard de Chardin 1936.1 See Teilhard de Chardin, Pierre, and Chung-Chien Young. Young, Chung-Chien, Davidson Black, Pierre Teilhard de Chardin, and W. C. Pei 1933.1 See Black, Davidson, Pierre Teilhard de Chardin, Chung-Chien Young, and W. C. Pei. Zittel, Karl Alfred von 1925.1 TrxrsBook or PataronroLoey. III, MamMatnta. millan and Company, London, pp. viii+316, 374 text figs. Svo, Mae- PEATES: XXAVIESA AX PLATE XXVI PLATE XXVI Fig. 1. Elephas indicus. Complete transverse section near tip of unerupted tusk, probably of an immature female. Ordinary light. Seven times natural size. The dark central area is dentine. In it, to the left, is a triangular area of tubular or cylindrical structure, suggestive of the development in Platybelodon and its allies, but absent in the later dentine of Hlephas. The ‘‘engine-turning”’ effect is not visible in this juvenile dentine. An enamel band encircles about two- thirds of the circumference of the dentine, being absent only to the right and upper right in the photograph. Much of it has been lost in making the section to the left, where it is thickest, but elsewhere it is clearly visible as a relatively transparent and homogeneous tissue. The whole tusk is here encased in a thick layer of cement, irregular in structure but in general with relatively dense, laminated internal and external zones and a thicker, non-laminated intermediate zone, with large canals rather like the Haversian canals of bone. Fig. 2. Hlephas indicus. Area of thin section within the circle on figure 1. Plane light. Nineteen times natural size. The dark, lower region is dentine and the irregular, spotty mass forming the greater part of the photograph is cement. Between these, to the left, is the enamel band, ending naturally near the middle of the photograph. It is sharply defined, the dentine border undulating and the cement border rugose. In the cement, the part to the right clearly shows the three layers, inner and outer dense and vaguely laminated, middle spongy, all with innumerable lacune. Fig. 3. Elephas indicus. Same as figure 2, photographed between crossed nicols. The enamel is characterized by high birefringence and relative simplicity and homogeneity of rod arrangement, with a narrow zone of deflection, dark as photographed, parallel to the dentine border. The cement also shows birefringence but is highly irregular with little consistent orientation except in the inner (lower) layer. Osporn: Tur Prososcipna, II PLatTE XXVI ai an is PLATE XXVII PLATE XXVII Fig. 1. Hlephas indicus. Area within the circle on Pl. xxvi, Fig. 2. Plane light. Three hundred thirty-seven times natural size. In this innermost area of the cement, just above the enamel, lacun# are very numerous and have many branching canaliculi. There are also more sparse, small tubules, one of which appears in the upper right in this figure. Fig. 2. Elephas indicus. Transverse section of about half of a small, mature tusk about 14 inches long. Ordinary light. Four times natural size. The section is taken about 3 inches from the tip of the tusk. Several inches had been worn off, including all the enamel and most of the cement. At this point, there is only a thin band of cement, seen in the figure as a more translucent coating above the dense dentine. The dentine as a whole shows three distinct, superimposed types of structure: (a) radiating tubes, too small to be clearly seen in this figure, directed away from the center of the tusk, (b) well-marked, fine, concentric lamelle, (c) curving, criss-cross markings of coarser character, visible as dark lines in the thinner parts of this section, that give the ‘“‘engine- turning” effect. This last effect appears to be the result of regularly recurring undulations in the courses of the dentine tubes and in the orientation of the surrounding calcified mass. (The prominent white lines are scratches on the section.) Fig. 3. Hlephas indicus. Part of the section within the circle on figure 2. Plane light. Forty times natural size. This shows the whole thickness of the cement on this part of the tusk. In ordinary light the cement here appears vaguely laminated, with fairly numerous lacune but no trabeculz or canals. The boundary against the dentine is smooth and very sharp. Fig. 4. Hlephas indicus. Same as figure 3, with crossed nicols. The effect of strong birefringence in the cement is to bring out a wavy, interwoven, fabric-like structure in this tissue. Fig. 5. Hlephas indicus. A small area near the outer edge of the cement band of the section shown in figures 2-4. Plane light. Three hundred thirty-seven times natural size. This shows a fabric-like texture similar to that seen more plainly and on a larger scale in figure 4. The lacune have very few and very short canaliculi and most of them appear to be empty (or filled with colorless fluid) in this section of a fresh tooth. Contrast with this figure 1, which shows the abundant canaliculi and opaque appearance typical of younger cement. The inner part of the cement in the present section, near the dentine, is about intermediate between the two conditions. Fig. 6. Phiomia wintoni. Transverse thin section about one-half inch from tip of small, worn tusk. Plane light. Thirty-eight times natural size. The lower part of the section is dentine and above this is the whole thickness of the enamel, cut at right angles to its surface. The clear bands in the dentine are caused by infiltration of the embedding medium into the cut ends of tubes and these zones, in fact, have tubes like those seen so clearly in the lower part of the picture. For details of enamel and of enamel-dentine junction see Pl. xxviu, Fig. 2. There is no indication of cement on this tusk. OsporNn: THE Proposcipra, II PLATE X XVII PLATE XXVIII PLATE XXVIII Fig. 1. Phiomia wintoni. Same as Pl. xxvu, Fig. 6, with crossed nicols. & g- 0, The dentine is not birefringent. The enamel is strongly birefringent and variations in extinction angle show the varying orientations of the rods through the greater part of the thickness of the enamel, but near the outer surface (up in the photograph) the illumination is even and shows the rods there to be es- sentially parallel. Fig. 2. Phiomia winlonit. Area within the circle on Pl. xxvu, Fig. 6. Ordinary light. About three hundred times natural size. Photograph by Dr. J. Leon Williams. The small bottom area in the photograph shows part of the granular layer of the dentine. Above this is the sharp enamel boundary, and then the full thickness of the enamel. Natural staining and variable refringence clearly brings out the pattern of the enamel rods, which lie in long, open spirals through most of the thickness of the enamel and become approximately straight and perpendicular to the surface only in the outermost zone, which is on the order of .02 mm. in thickness. The enamel as a whole is here about .5 mm. in thickness. Osporn: THe Proposcipea, II Puate XXVIII PLATE XXIX PLATE XXIX Fig. 1. Trilophodon obscurus. Transverse section of part of tusk. Ordinary light. Four times natural size. The bulk of the section is in the dentine in which the radial and concentric structures are shown, but not the “engine-turning’’ effect, which was, never- theless, present. Above this the whole width of the enamel band is shown. There is no cement on the specimen as preserved, but this probably occurred when the tusk was fresh. Fig. 2. Trilophodon obscurus. Area within circle on figure 1. Crossed nicols. Forty times natural size. In this region the enamel rods are only very gently and obscurely spiral and tend rather to be parallel to each other in simple curves from the dentine to the outer surface. Toward the edge of the enamel band, to the left, the rods are strongly curved and the enamel overhangs a natural pocket. An isolated stringer of enamel occurred beyond this and its edge is barely visible at the left margin of the photograph. The enamel is also marked throughout by fine striations, not very prominent in the photograph, parallel to the outer surface even when this is strongly curved, as at the left of the photograph, and hence approximately at right angles to the enamel rods throughout. The dentine is very feebly birefringent and shows uneven lamination parallel to the surface. Fig. 3. Trilophodon obscurus. Same tusk as figures 1 and 2, thin section across enamel cut in a plane vertical to the surface and parallel to the longi- tudinal axis of the tooth. Crossed nicols. Forty times natural size. Away from the edge of the enamel band, the rods show a slight spiral arrangement much like that of Phiomia (Pl. xxvii) in kind but far less in degree. Striations as seen in figure 2 are somewhat more prominent in this photograph. Fig. 4. Trilophodon (Megabelodon) sp. Transverse thin section of tusk near edge of enamel band. Plane light. Eleven anda half times natural size. The area covered by the lower part of the photograph was occupied by dentine, fragments of which can be seen although most of it has broken away. Above this is the enamel and cement covering. To the left is the edge of the main band of enamel. Beyond this to the right were five long, isolated string- ers of enamel. Transverse sections of two of these are seen in the middle and to the right in the photograph and another is cut by the right margin. Between the main enamel band and the stringer in the middle of the photograph is a mass of cement. From thinner films around the stringers and main band and from the character of the enamel surface it appears that the enamel was all embedded in a coating of cement and was wholly exposed only at the wearing edge. The cement also extends in a thicker coating around the dentine beyond the last enamel stringer, outside the area of the photograph. Fig. 5. Trilophodon (Megabelodon) sp. A different part of the same thin section as figure 4. Crossed nicols. Nineteen times natural size. This photograph ineludes the last two enamel stringers, to the right of the three visible in figure 4. The strong birefringence clearly brings out the peculiar, fan-like divergence of the enamel rods in these stringers, which is equally developed in the other stringers and at the edge of the main enamel band. The stringers are separated and in part surrounded by a mass of cement so feebly birefringent that it can hardly be seen in the photograph. Puate XXIX Tue Proposciwra, II OSBORN: PLATE XXX PLATE XXX Fig. 1. Trilophodon (Megabelodon) sp. Vertical thin section of broken molar tubercle. Ordinary light. Four times natural size. The worn apex of the tubercle is at the top, dentine to the left, and enamel to the right. The enamel here reaches a thickness of about 5 mm. Aside from the mineral staining, largely casual but in part emphasizing natural structure, the most striking thing is the presence and orientation of the numerous enamel striations or laminations. Fig. 2. Trilophodon (Megabelodon) sp. Part of section in circle on figure 1. Crossed nicols. Twenty times natural size. To the left may be seen a small part of the dentine, worn into a pit above. The margin of the enamel is also a wear surface. Within the enamel the striations are emphasized by mineral stain as well as by birefringence and are very prominent, especially in the more internal part of the enamel where they are more wavy than externally. The enamel rods are less clearly distinguishable. In a thin inner zone, very obscure in the photograph (but see Fig. 4), they appear to be spiral. In the bulk of the section they are simple, nearly parallel, arranged in a sweeping curve becoming more horizontal as they pass from the inner (left) to outer (right) side of the enamel coating. Fig. 3. Trilophodon (Megabelodon) sp. Transverse thin section of a molar cusp, from the same tooth as figures 1 and 2 but a different cusp. Ordinary light. Four times natural size. In the center is the circular core of dentine, about 4 mm. in diameter, and surrounding this (except where broken away, lower left) the enamel, 5-7 mm. in thickness. The dentine shows no pronounced structure. The enamel has an inner zone, about 1 mm. in thickness, with vague concentricity but no strong striations, and an outer zone of 4-6 mm. with pronounced concentric striations, which are homologous with those cut in a different plane in figures 1 and 2. Fig. 4. Trilophodon (Megabelodon) sp. Part of section in inner circle on figure 3. Plane light. Forty times natural size. To the left is the dentine with sharply visible tubules and terminating in a granular layer below the enamel. The light band down the middle of the photograph is the thin inner zone of enamel. Mineral staining shows the rods to be oriented in an irregular, highly complex, spiral pattern. Fig. 5. WVrilophodon (Megabelodon) sp. Part of section in outer circle on figure 3. Plane light. Twenty times natural size. This shows the outer part of the enamel, with its well-defined concentric striations and with the rods, not very clear in the photograph, generally radial and much simpler in orientation than in the innermost enamel. PLate XXX II Tue Proposcipea, OSBORN: ARS sii $ Bee eho ee INDEX Page references in small roman figures refer to Volume I only Abel, Othenio, 4, 13, 95, 100-102, 761 abeli (see Trilophodon abelz) Aberdare Mountains, 1193 Abich, Otto Wilhelm Hermann von, 3, 761 Absolon, Karel, 4, 13, 761, 1139, 1168 Abtsdorf, 81 Abyssinia, 117, 1176, 1200, 1417 Academy of Natural Sciences, Philadelphia, 15, 209, 745, 1609, 1619 Academy of Sciences (Zoological Museum, Leningrad), 1128, 1148 Acconci, Luigi, 3, 761, 1187, 1230 Aceratherium, 117, 272, 1429, 1459, 1461, 1479 Acheulean, 1169, 1480, 1433 acrocephaly, 919, 1552 Aculcingo, Cannada de, Mexico, 5387, 558, 740 acutidens (see Mastodon acutidens) Adams County Mammoth, 1009, 1012 Adams, Andrew Leith, 3, 19, 762, 923, 974, 989, 1039, 1040, 1059, 1142, 1198, 1221, 1265. See also Palxoloxodon buski Adams, Michael, 2, 762, 1623. See also ‘Adams skeleton” “Adams skeleton,” 1123, 1130, 1131, 1136, 1147-1149, 1162, 1166, 1603, 1628 adaptive radiation, xiii-xv, 20, 22, 30, 32-35, 46, 333, 684, 933, 936, 983, 1269, 1422, 1423, 1524, Pls. x, x1. See also individual families and sub- families; migrations, origin, phylogeny Addobush, Cape Colony, 1205, 1283 A nocyon, 1082, 1134, 1161 affinis (see Elephas affinis, Zygolophodon borsoni affinis) Afghanistan, 279 Africa, xi, 34-39, 45, 103, 111, 114, 117, 231, 232, 479, 734, 950, 983, 984, 1139, 1190, 1271, 1807, 1422, 1435, 1523, 1553, 1606, Pl. x. See also Faytim, Kalahari Desert Central and East Africa (see Miocene, Quaternary) North Africa (see Eocene, Miocene, Oligocene, Pliocene, Quaternary) South Africa, Archidiskodonts and Loxodonts of, 944, 946. See also Vaal River African elephant, 29, 928, 1334, 1594. See also Loxodonta africana africana (see Loxodonta africana) africanus (see Loxodonta africana) Africanus line, 1178 Afton, Oklahoma, 1002, 1003, 1087 aftoniz (see Stegomastodon aftoniz) Aftonian, 671, 682, 683, 725, 726, 1510, 1511, 1512, 1515, Pl. vit. Interglaciations Agassiz, Jean Louis Rodolphe, 3, 91, 762, 1623 Agricultural and Mechanical College, College Station, Texas, 374, 745 Agricultural College and School of Mines, Alaska (see University of Alaska) Aguilera, José G., 1013 Aichel, Otto, 4, 762 Aichhorn, Sigmund Johann Nepomuk, 91, 762 Ailuropoda bacont, 1451 Ainsworth, Nebraska, 251, 315, 316-319, 324, 433, 443, 444, 446, 601, 602, 606, 726, 728, 738. See also Stegomastodon primitivus Airaghi, Carlo, 4, 762 airawana (see Stegodon airawana) Ajnacsko, Hungary, 114, 159, 160, 210, 638 akashiensis (see Parastegodon akashiensis) Akashi-gun, Japan, 1420 Akeley, Carl Ethan, 13, 15, 1006, 1189, 1190, 1201, 1202, 1239 Akeley, Mary Jobe, 1198, 1239 Aki District, 908 Akimoto-mura, 906 Akira- mura, 818, 906 Akron, Iowa, 682, 683, 741 Alachua clays, 380, 386, 400, 418, 419, 428, 440, 482, 742, 1495, 1496 Alangasi, Ecuador, 537, 567, 568, 571-573, 584, 585, 741, 1521 See also Alaska, 137, 176, 177, 736, 753, 1088, 1091, 1099, 1127, 1134, 1135, 1145, 1156, 1157, 1159, 1169, 1201 Alaska Agricultural College and School of Mines, 1161, 1609, 1611 Alaska College-Frick American Museum Expedition, 1159-1161 Alaska-Yukon (see Yukon) alaskensis (see Mammonteus primigenius alaskensis, and Mastodon americanus alaskensis) Alas-Tuwa, 885, 886 Albany, New York, 136, 1602 Albert Nyanza, 995, 1193 albertensis (see Loxodonta africana albertensis) Alces latifrons, 971, 1155 Alcoi, Spain, 114 Alekseev [Alexejew], A., 1466, 1623 Alexander, Captain, 651 Alexandria Trail (see Faytim) Algeria, 65, 115, 232, 246, 485, 964, 1183, 1184, 1187, 1269, 1274, 1431, 1609, 1610 Algoa Bay, 931 alisphenoid, 916, 917, 920 Allen, Glover Morrill, 1196, 1594, 1623 Allen, Joel Asaph, 13, 15, 1364, 1372 alloiometry, xv, 341, 647, 716, 1545, 1580, 1581, 1606 alluvium, 449, 725, Pl. v1 Alps, 1473, 1475 Altai region, Mongolia, 3938, 461 Altamaha River, 1077 alternation of cones, 627, 650, 1545 Alticamelus, 319, 426, 601, 610 Amajaque, 537, 555, 740, 1082 Amalgamated Phosphate Company, 416 Amebelodon, xv, 11, 30, 226, 231, 249-251, 321, 328, 332-334, 335, 336-338, 445, 459, 461-463, 686, 690, 705, 715, 716, 739, 1381, 1413, 1527, 1550, 1558, 1571, 1600, Pls. v, v1 fricki, xv, 118, 226, 231-246, 251, 260, 288, 321, 329, 330, 331, 332, 334- 336, 337, 338, 443, 444, 445, 460, 705, 706, 711, 714-716, 738, 1411, 1412, 1507, 1604, Pls. v, v1, x grangert, 1413. See Plazybelodon grangeri (?) joraki, 1417. See M egabelodon joraki paladentatus, 226, 248, 251, 260, 288, 289, 298, 309, 310, 311, 329, 331, 460, 706, 711, 739, 755, 1407, 1500, 1501, 1604 sinclairt, 251, 331, 337, 338, 706, 711, 739, 758, 1415, 1507 (Torynobelodon) loomisi (=fricki). See Torynobelodon loomisi (Trilophodon) hicksi, 226, 248, 251, 260, 280, 284, 288, 289, 298, 307, 308, 309, 310, 312, 317, 329, 331, 706, 711, 739, 755, 1407, 1501 Amebelodonti de, 328, 335, 336, 459, 1368 Amebelodontine, 27, 31, 49, 119, 232, 328, 332, 333, 334, 336, 459, 462, 469, 689, 690, 715-719, 734, 739, 1368, 1370, 1526, 1528, 1545, 1571, Pl. x Ameghino, Florentino, 3, 384, 519-521, 539, 550, 579-581, 592, 762, 1517, 1623 America, 1456; S.S. America, 1599 American Cyanimid Company, 160, 161 American elephant, 7, 1382 American Mastodon, 6, 7, 1363, 1374. See also Mastodon americanus American Museum of Natural History, New York, 745-753, 1480, 1609, 1615- 1618 American Philosophical Society, Philadelphia, 753, 1388, 1609, 1619 americanus (see Mammonteus primigenius americanus, Mastodon americanus, Miomastodon tapiroides americanus) Amherst College, 14, 484, 487, 495, 526, 753, 1052, 1079, 1080, 1106, 1353, 1609, 1610 Amilee Creek, Siswan, 1347, 1358, 1359 Amphicyon, 260, 272, 274, 400, 488, 1461 Amsterdam Zoological Gardens, 1314, 1609, 1610 Amthor of Gotha, 1181 1631 1632 Amynodontidx, 824 Anancine, 542, 588, 612, 1370 Anancus, 10, 11, 29, 128, 612-614, 619, 622-625, 627-629, 630, 631, 632, 635, 637, 638, 640, 642, 647, 669, 686, 720, 722, 740, 903, 1375, 1527, 1564, 1601, 1622 arvernensis, 10, 31, 118, 229, 348, 357, 360, 539, 616, 618-622, 624-628, 630, 631, 632, 633-635, 637, 638, 640, 650, 653, 740, 753, 756, 759, 760, 963, 964, 968, 978, 1056, 1177, 13868, 1376, 1393, 1472, 1476, 1564, 1604, 1610, 1620. See also Bunolophodon, Mastodon (Dibuno- don) arvernense arvernensis brevirostris, 613, 614, 617, 620, 625, 627, 634, 636, 643, 758, 1392, 1472, 1548, 1619 arvernensis dissimilis, 125, 618, 625, 632, 1394, 1472 arvernensis macroplus, 10, 618, 625, 628, 630, 631, 632, 1393, 1472 arvernensis progressor, 624, 625, 639, 640, 740, 1403, 1469 bensonensis, 1411. See Cordillerion bensonensis brazosius, 1408. See Trilophodon (?Tetralophodon) brazosius brevirostris (see Anancus arvernensis brevirostris) defloccatus, 1411. See Cordillerion defloccatus elegans (see T'etralophodon elegans) falconeri, 619-621, 624, 625, 628, 630, 632-634, 635, 636, 637, 651, 740 754, 756, 1411, 1472, 1564 gigantarvernensis, 283, 624, 625, 632, 637, 740, 1407, 1472 intermedius, 621, 622, 625, 638, 639, 1387, 1469 macroplus (see Anancus arvernensis macroplus) minutoarvernensis, 283, 625, 632, 637, 740, 1407, 1472 orartus, 1411. See Cordillerion orarius palxindicus, 271 pertmensis, 10, 114, 348, 448, 532, 613, 621, 622, 624-626, 630, 640, 641, 642, 643, 644, 645, 647, 651, 653, 740, 753, 754, 756, 841, 842, 1392, 13938, 1448, 1548, 1604, 1613 properimensis, 360, 624, 625, 630, 642, 643, 645, 647, 653, 740, 749, 1419, 1448, 1604 sinensis, 624, 625, 720, 721, 740, 754, 1418, 1482 Anastasia, Florida, 400 Anca, Baron, 928 ancestrale (see Mearitherium ancestrale) ancestry (see origin) Anchitherium, 400, 467, 902, 1459, 1461, 1478 Ancodon, 53, 1424, 1425 andaranus (see Mastodon andaranus) Andean region, 526, 576, 611, 1516, 1519, 1520. See also Bolivia Anderson, Abram E., xvii, 1607 Anderson, Netta C., 4, 762 Anderson, Robert van Vleck, 4, 762 Andersson, J. Gunnar, 458, 1480, 1481, 1485, 1487, 1623 andicus (see Mastodon andicus) andii (see Mastodon andii) andium (see Cordillerion andium) Andreae, Achilles, 260, 762 Andrews, Charles William, xi, 4, 13, 26, 30, 36, 38, 42-45, 47, 53-60, 69, 71-74, 95, 137, 143, 236, 239-214, 244, 251, 291, 762, 1059, 1162, 1216, 1222, 1226, 1364, 1424, 1623 Andrews, Roy Chapman, 328, 332, 333, 461, 466 andrewst (see Meritherium andrewst, Palxoloxodon andrewst) Andrias scheuzert, 1464 Andrussov or Andrussow, Nikola! Ivanovich, 4, 763, 1466, 1623 Angelhausen, 1050 Angern, Austria (see Mannersdorf) Angola, South Africa, 1193 angolensis (see Loxodonta africana angolensis) anguirivale (see Rhynchotherium anguirivale) anguirivalis (see Serridentinus anguirivalis) Angus, Nuckolls County, Nebraska, 943, 1033 angustidens (see Deinotherium angustidens, Elephas platycephalus angustidens, Trilophodon angustidens) Anjou, Molasse de |’, 115 annectens (see Serridentinus annectens) annular growth rings, 182, 183 ? OSBORN: THE PROBOSCIDEA anteater, 579 antelopes, 444, 498, 508, 579, 1467 Anthony, A. W., 508 Anthony, Harold Elmer, 13, 763 Anthony, Raoul Louis Ferdinand, 4, 13, 715, 763, 1316, 1317 Anthracotheres, 272, 824, 1428, 1441, 1442 Antilospira, 1482 antiqua (see Hesperoloxodon antiquus) antiquissima (see Cordillerion oligobunis antiquissimus) antiquus (see Hesperoloxodon antiquus) antiquus italicus (see Hesperoloxodon antiquus italicus) antium (see Mastodon antium) Antoletherium, 82, 83, 1377 Anyang, China, 1362, 1488 Aomori Prefecture, 1289 aomortensis (see Palxoloxodon aomoriensis) Aphelops, 307, 308, 315, 385, 386, 399, 400 Aphelops Draw, 426, 427 Apolda, 1050 Apterodon, 53, 1424, 1425 Aquino, 1239, 1241 Aquitanian, 116, 117, 659, 903, 1440, 1457 Arabs, 1125 Arabu, N. von, 1466, 1623 Aracan, 927 Aradas, Andrea, 3, 763, 1187, 1204 Aragon, 1468 Arambourg, Camille, 1417, 1436, 1437, 1623 Aransas River, Texas, 562-565, 623 Araucana formation, 551, 1517 Aravalli range, 852 Arbeichan, France, 86 arborense (see Mastodon arborense) Arbutus, 508 Archxohippus, 400, 601 Archer, Florida, 400, 416, 418 Archidiskodon, xii, 10, 11, 22, 25, 32, 33, 113, 515, 602, 891, 904, 905, 913, 914, 932-935, 937-939, 941-944, 946, 947, 950, 953, 981, 983, 986, 996-998, 1000, 1009, 1017, 1039, 1041-1043, 1045, 1046, 1052, 1140, 1146, 1163, 1164, 1178, 1191, 1193, 1209, 1228, 1285, 13804, 1378, 1470, 1476, 1484, 1510-1512, 1527, 1538, 1540, 1547, 1549, 1580, 1582-1584, 1587, 1589, 1602, P. x1. See also Archidiskodon planifrons-meridionalis-imperator phylum, 1058; Hlephas planifrons-meridionalis group, 903 andrewst (see Palxoloxodon andrewst) broomi, 942-946, 983-985, 989, 990, 992, 1412, 1439, 1540, 1612, 1617 extlis, 934, 935, 942, 943, 946, 997, 1030-1032, 1033, 1412, 1514, 1540, 1619 hanekomi (see Palzoloxodon hanekom) haroldcooki, 684, 942, 943, 997, 998, 1029, 1412, 1514, 1540, 1611 hayt, 725, 942, 943, 946, 997, 998, 1003, 1006, 1013, 1014, 1023-1025, 1405, 1514, 1540, 1613, Pl. vin hysudricus (see Hypselephas hysudricus) imperator, 32, 386, 400, 422, 602, 922, 934, 936-939, 941-943, 946, 947, 971, 975, 982, 996, 997, 998-1015, 1017-1019, 1021, 1022, 1025-1027, 1029, 1030, 1041, 1052, 1071, 1072, 1074, 1077, 1078, 1080, 1081, 1084, 1087, 1088, 1094, 1105, 1163-1165, 1394, 1407, 1512, 1514, 1515, 1522, 1540, 1547, 1550, 1582, 1583, 1602, 1605, 1613, 1615-1619, 1621, Pl. vur. See also Elephas columbi var. imperator imperator falconeri, 942-944, 946, 997, 998, 1015, 1016, 1407, 1515, 1540, 1615 imperator maibeni, xiv, 725, 934, 938, 942, 943, 946, 947, 997, 998, 1009, 1012, 1019-1023, 1027-1029, 1080, 1112, 1276, 1277, 1410, 1514, 1540, 1581, 1602, 1605, 1613, Pl. vir. See also Nebraska State Museum imperator scotti, 725, 942, 943, 946, 997, 998, 1012, 1025-1027, 1410, 1514, 1540, 1613, 1616, Pl. vit imperator silvestris, 942, 944, 946, 997, 998, 1015, 1016, 1407, 1515, 1540, 1612 loxodontoides, 942, 943, 984, 985, 991, 992, 1278, 1414, 1439, 1540, 1612, 1618 INDEX Archidiskodon—continued meridionalis, 10, 32, 259, 633, 634, 705, 910, 934, 936, 938, 939, 941-943, 946-948, 961, 963, 964, 968, 969-983, 995, 997, 998, 1002, 1029, 1035, 1037, 1056, 1065, 1095, 1112, 1155, 1164, 1175, 1177, 1182, 1183, 1218, 1220, 1251, 1382, 1394, 1399, 1436, 1476, 1540, 1581, 1583, 1605, 1611, 1613, 1614, 1617-1619. See also British Museum, 972, 1218, 1220; Durfort skeleton; Hlephas meridionalis-E. trogontherti-E. primigenius phylum; Elephas lyrodon meridionalis cromerensis, 905, 942-944, 946, 963, 980, 981, 982, 1014, 1155, 1366, 1407, 1476, 1540, 1611, 1614 meridionalis nebrascensis, 399, 667, 725, 934, 942, 943, 997, 1033-1037, 1416, 1514, 1540, 1581, 1602, 1605, 1611, Pl. vin milletti, 942, 948, 984, 985, 991, 1414, 1439, 1540, 1612, 1618 paramammonteus, 1420, 1540 planifrons, 10, 32, 230, 365, 448, 449, 633, 634, 857, 905, 930, 932, 934, 936, 938, 939, 941-943, 946-949, 950-959, 960-968, 971, 976, 981, 986, 988, 989, 998, 1010, 1024, 1029, 1071, 1155, 1164, 1175, 1177, 1178, 1183, 1291, 1338, 1339, 1352, 1353, 1359, 1378, 1391, 1411, 1430, 1431, 1435, 1448, 1476, 1483, 1540, 1547, 1581, 1583, 1605, 1611, 1613, 1614, 1616, 1617. See also Piltdown planifrons rumanus, 942-944, 968, 969, 1184, 1235, 1408, 1476, 1540, 1610 proplanifrons, 700, 934, 942, 943, 948, 964, 984, 986, 987, 1418, 1540, 1580, 1581, 1605, 1612, 1618 sheppardi, 1411. See Palxoloxodon sheppardi sonoriensis, 942, 943, 997, 998, 1013, 1033, 1414, 1515, 1540, 1618 subplanifrons, 934, 942-945, 983-986, 987, 988, 989, 990, 1412, 1439, 1540, 1549, 1581, 1584, 1605, 1612, 1617 tokunagai, 1409 transvaalensis, 1411. See Palzoloxodon transvaalensis vanalpheni, 942, 943, 984, 985, 990, 991, 992, 1414, 1439, 1540, 1612, 1618 yorki, 942, 943, 984, 985, 992, 993, 1415, 1439, 1540, 1612 Archidiskodonten, 939, 941, 1378, 1583 Archidiskodontine, 937, 1370, 1625 archidiskodontoides (see Palzxoloxodon archidiskodontoides) Arctodus yukonensts, 1135, 1161 Arctotherium, 1515, 1518 arétes, Wulstkanten or spurs, 212 Argentina, 384, 385, 514, 519-521, 524, 527, 530, 531, 536, 537, 541, 543, 544, 550, 579-581, 587-599, 741, 743, 1516, 1518 argentinus (see Notiomastodon argentinus) Arikaree, 426 aristogenes or rectigradations, 225, 276, 277, 341, 613, 627, 647, 781, 1545, 1580, 1581 aristogenesis, origin of new characters, xiv, xv, 1545, 1580, 1581 Aristotle, 1118, 1147, 1164, 1209 Arizona, 535, 537, 543, 565, 678-682, 740, 741 arizonz (see Stegomastodon arizon2) Arizpe, Mexico, 943, 1033 Armenia (see Asia Minor) armeniacus (see Parelephas armeniacus) Armstadt, 1050 Arno, 969, 1236 Arnold, Robert, 713 Arpino, 1239 Arrecifes, Argentina, 536, 537, 570, 579, 596, 598 Arreco, Buenos Aires, 587 Arsinoitherium, 762, 1424 Artaud, Soulange, 2, 763, 1123, 1141 artifacts, 573, 589 Artiodactyla, 272, 399, 678, 680, 887, 1425, 1429 arvernense. See Mastodon (Dibunodon) arvernense arvernensis (see Anancus arvernensis) Ash Hollow formation, 1479 Ashland, Illinois, 1047, 1088, 1097 Ashley, Indiana, 187 Asia, 34, 1307, 1456, 1477-1488. See also Asia Minor, Turgai Asia Minor, 1046, 1047, 1060 Asiatic elephant (see Elephas indicus) Asiatic Society of Bengal Museum, 644, 753, 1609, 1610 1633 Asnot (see Hasnot) Assam, 1315, 1324, 1344 “Astell’”’ (see East India Company) astensis (see Mammonteus primigenius astensis) Astésan, Italy, 634, 961, 964, 1055 Asti d’Auray, 134 Asti (Villanova), Italy, 192, 199, 200, 207, 616, 618, 736, 1470 Astian, 902, 904, 905, 964, 1446, 1447, 1469-1472 Astre, Gaston, 1420, 1623 Asturias, Province, 1184 atavus (see Mastodon atavus) Atbara, Sudan, 1193, 1194 Athanasiu, Sava C., 4, 763, 968, 969, 1400 atlanticus (see Palzxoloxodon atlanticus) Atlas Mountains, 1194, 1195, 1197 atticus (see Turicius atticus) Attock, India, 85, 105, 115, 513, 1395 Augsburg, 114 aurelianensis (see Zygolophodon pyrenaicus aurelianensis) aureliense (see Mastodon aureliense) Aurés, Africa, 232 Aurignacian, 1139, 1168, 1430, 1433 aurore (see Stegodon aurore) ausonius (see Hesperoloxodon antiquus ausonius) australe (see Deinotherium australe) Australia, 1,90. See also Deinotherium australe, Mastodon australis australis (see Deinotherium australe, Diprotodon australis, Mastodon australis, Notelephas australis) Australopithecus africanus, 944, 945 Austria, 639, 729, 961. See also Angern, Levantin, Mannersdorf, Neudorf, Teschen (Schlesien), Wies (Steiermark) austro-germamnica (see Trilophodon angustidens var. austro-germanicus) Autrey, Sadne, 1469 Auvergne, France, 134, 192, 217, 616, 618, 625, 632 Ava, Burma, 268, 824, 825, 843, 844 Avalo y Figueroa, Diego de, 2, 763 Avaray, France, 192, 205, 207 Avinoff, Andrey, 312, 314 Ayacucho, Buenos Aires, 598 Aybelodon, 509, 690, 1382, 1527, 1563, 1601 hondurensis, 118, 479, 480, 503-505, 509, 510, 512, 513, 737, 751, 753, 1382, 1416, 1516, 1604 Aymard, Auguste, 3, 192, 193, 209, 618, 625, 630-632, 763, 942, 1393, 1394 ayore (see Cuvieronius ayore) Azof, 1061 Bach, Franz, 4, 763 Bachkiese, 1045 Bachmann, Isidor, 3, 93, 763 Bies-Bodrog, Hungary, Pl. 1 Baden, Germany, 281, 283, 637, 738, 740 Baigneaux, 1459 Baker County, Oregon, 137, 173, 736 Baldassari, Giuseppe, 2, 763 Balta, Russia, 694 Baltavar, Hungary, 114, 201, 210, 212, 262, 360, 1468 “Baltimore tooth,” 286 Baluchistan, 78, 85, 115, 266, 267, 272, 275-279, 448, 527, 643, 735. Bugti Hills, Churlando, Dera Bugti, Kumbhi Baluchitheriwm, 269, 272, 385, 386, 396, 1441, 1477. See also Paraceratherium Bandoeng, 365. See also Geological Survey of Bandoeng Bangalore, 1322 Banjobora, Japan, 251, 384, 457 Banks, Joseph, 1123, 1147 Barbary, Africa, 232 Barbiani, A., 4, 763 Barbour, Erwin Hinckley, xvi, xvii, 4, 18, 27, 31, 1387, 174, 175, 249, 251, 290, 298, 294, 298, 311, 328, 333, 335-339, 377, 385, 470, 537, 601, 602, 695, 710, 711, 713, 763, 942, 948, 1020, 1021, 1496, 1498, 1623 See also 1634 OSBORN: THE PROBOSCIDEA barbourensis (see Serbelodon barbourensis) barbourt (see Morrillia barbourt) Barcena, 1081 Bardo Museum, 1195 Baretti, Martino, 3, 764 Barikiwa, Tanganyika, 1193 Barkly West, South Africa, 944 Barney, Charles T., 1201 Barnum, P. T., 1199, 1200 barnumbrowni (see Torynobelodon barnumbrownt) Barét, Hungary, 638 Baroth-kopecz, 1468 Barrington, England, 1214 barroisi (see Phiomia barroist) barstonis (see Serridentinus barstonis) Barstovian, 1491-1494 Barstow, California, 319, 447, 742, 1494, 1502 Bartholomew, J. G., 29, 764 Bartolotti, C., 4, 766 Bartonian, 53, 61 Barytherium, 1424 Basel (see Naturhistorisches Museum) basioccipital, 916 basis cranii, 923, 924 basisphenoid, 916, 920 Bassariscops, 315 bat caves, 916, 1462 Batang Serangan, Sumatra, 1330, 1331 Bate, Dorothea M. A., 4, 13, 764, 1183, 1187, 1257, 1266, 1267 Bather, Francis Arthur, 4, 13, 764, 1182, 1222 bathycephaly, 919, 1552 bathymetric maps, 1257, 1304, 1305 Batta-Erd, Hungary, 114, 159, 160, 736, Pl. 1 Battleship Mountain, New Mexico, 251, 324, 325, 446, 738 Baud, J. C., 1330 Bauer, W., quarry, 1253 Baur, George, 3, 764 Bautista Creek, 496 Bavaria, 111, 115, 735. See also Freising, Glonn, Moosburg, Tutzing bavaricum (see Deinotherium bavaricum) Bayerische Ludwig-Maximilians-Universitit, 14 Bayerische Staatssammlung fiir Palaiontologie, 15 Bayle, 91 Baylor University, 373 Beadnell, Hugh, xi, 4, 36, 55, 60, 239, 764 beadnelli (see Palxomastodon beadnelli) “Bear-dogs” (Hemicyon), 1467 Beauce, lac de, 1458 Beaufort County, South Carolina, 137, 171 Beaugency, France, 204, 207 Beauséjour farm, 1275 Bechuanaland, South Africa, 944 Beck, Richard, 4, 764, 944, 994 Beechey, F. W., 3, 764 Beerbower, 1093, 1094 Begouen, Comte, 966 Belaja River, 133, 192 Belgium, 1136 Belgranense, 520, 590, 593 Bellaeva, E., 4, 251, 278, 385, 764, 1411, 1623 Bellecroix, France (see Chagny-Bellecroix) Bellwood, Butler County, Nebraska, 1012 Belomechetskaja, Russia, 461 Belvedereschotter, Austria, 100, 114 Bend (see The Bend) Benedict, Francis G., 1604, 1623 Bengal (see Elephas indicus bengalensis) bengalensis (see Elephas indicus bengalensis) Benghisa, Malta, 1272 Benkleman, Dundy County, Nebraska, 1012 Benson, Arizona, 535, 537, 565, 624, 679, 680, 1504 bensonensis (see Cordillerion bensoneniss) Berckhemer, Fritz, 5, 13, 765, 1236, 1238, 1253 Beresowka River, 1127, 1129, 1130, 1131, 1167 Bering Strait, 1135, 1304 Berkeley, California, 1509. See also University of California Berkey, Charles P., 1477, 1623 Berlin, 14, 1198, 1401, 1609, 1610. See also Museum fiir Naturkunde der Universitat and Geologisches Palaeontologisches Institut und Mu- seum der Universitit, also University of Berlin and Zoological Garden Bermersheim, Rheinhessen, 281, 282, 348, 349, 362, 739 Bernese Jura, 91 Berro, A. C., 587, 615, 765, 1412 Bessarabia, 262, 625, 639, 936, 961, 964, 968. See also Taraklia Beyschlag, Joannes Fridericus, 2, 765 Bhagathoro, Sind, 1442 Bhandar Bone Bed, 279, 448, 451, 657, 658, 737 Bhimbar, India, 622, 625, 643, 659, 740 Bibliography, 2-5, 761-802, 1623-1630 Bibos, 887. See also Stremme Bieber, V., 96, 765 Biedermann, W. G. A., 3, 765 Bien,—, 1451 bifoliatus. See Ocalientinus (Serridentinus) bifoliatus Big-Bone Lick (Strick or Swamp), Kentucky, 131, 134-136, 168, 170, 171, 190, 1088, 1089 Birchington, 1022, 1609, 1619 Birket-el-Qurun (see Faytim) birmanicus (see Stegodon insignis birmanicus) Bishop, Carl W., 29, 765 Bison, 171, 386, 902, 1044, 1049, 1077, 1082, 1107, 1134, 1135, 1161, 1253, 1510, Pl. vii Black, Davidson, 1623 Black Hills, South Dakota, 251, 289, 298, 328, 738, Pl. vir Black Obelisk Shalmeneser, 1207 Black Sea, 1061 Blackburn, Mrs., 1265 Blainville, Henri Marie Ducrotay de, 3, 7, 85, 89, 122, 138, 168, 192, 516, 518, 549, 765, 931, 1123, 1137, 13825, 1623 Blair, William Reid, 14, 1604 Blake, Charles Carter, 3, 765, 997, 1072 Blake, John Frederick, 4, 765 Blake, William Phipps, 3, 765 Blane, Edouard, 29, 765 Blancan, 1503-1505. See also Blanco formation Blanckenhorn, Max, 4, 765. See also Selenka-Blanckenhorn Expedition Blanco formation, 399, 431, 498, 560, 675-677, 1503, 1504 Blanford, William Thomas, 3, 29, 34, 269, 765, 1439 Blésois, Faluns helvetiens du, 109, 251, 254, 261 Blésois, Marnes du, 1458 Blick, John C., 444, 505, 508, 510 blicki (see Blickotherium blickt) Blickotherium, 508, 690, 737, 1382, 1527, 1547, 1562, 1601 blicki, 479, 508, 504, 505, 506, 507, 508, 509, 510, 512, 518, 686, 737, 753, 1416, 1516 euhypodon, 228, 471, 474, 476, 478-480, 482, 483, 484, 489-491, 493, 495, 503, 604, 505, 509, 511-513, 533, 538, 539, 545, 737, 746, 1898, 1500, 1604 Bloemhof, South Africa, 944, 945, 993, 1188, 1279, 1284, 1285 Blora, 894 Blue Nile, 1194 Blumenbach, Johann Friedrich, 2, 6, 7, 135, 136, 167, 168, 765, 1117, 1118, 1122, 1123, 1136, 1141, 1193, 1197, 1325, 13638, 1385, 1623 Blunt, W. G., 1031 Blyth, Edward, 29, 766 Bocage, J. V. Barboza du, 29, 766 Bodard, Jean de, 196 Bohemia, 111, 115. See also Franzensbad INDEX Bohemian Museum of Prague, 91, 753 Bolivia, 514, 530, 537, 550, 551, 568. See also T'eleobunomastodon bolivianus bolivianus (see Cordillerion bolivianus, Teleobunomastodon bolivianus, Tarija, Ulloma) Bolk, Louis, 4, 766 Bologna, 14, 616, 619, 635, 753, 1609, 1610 Bolshaya Baranikha River, Kolyma district, 1128 bombifrons (see Stegodon bombifrons) Bonaerense formation, 520, 551, 1517, 1518 bonaerensis (see Cuvieronius bonaerensis) Bonaparte, Charles Lucien Jules Laurent, 3, 27, 30, 766, 912, 1369 Bondol, Java, 837, 894-896 bondolensis (see Stegodon bondolensis) Bone Valley formation, 160, 161, 285, 386, 400, 428, 430, 440, 482, 742, 1495, 1496 Bongo, 1193 Bonn, 15, 1017, 1609, 1610 Boone County, Kentucky, 136, 736 Bodtherium, 1135, 1161 borealis (see Mammuthus borealis) Borissiak, Alexei Alexievich, 4, 14, 27, 31, 251, 278, 328, 333, 385, 459, 461, 766, 1411, 1413, 1418, 1624 Borna, 1127, 1130, 1167 Borneo, 1304, 1440, 1445. See also Bruni Borntriiger, 1058 Borsod (see Kirdld) Borson, Etienne, 3, 207, 209, 766, 1388 borsoni (see Zygolophodon borsont) Bos, 824, 1449, 1476, 1486. See also Pliocene-Pleistocene boundary Bose, B. K., 1418, 1624 bosei (see Palxoloxodon priscus var. bose?) Boselaphus, 1358 Boskop man, 944, 945, 1285 Bosphorus, 1061 Botti, Ulderico, 3, 766, 1137, 1150, 1399 Boué, Ami, 3, 81, 82, 766 Boulay, France, 109 Boulder, Colorado, 1003 Boulder Conglomerate, 448, 449, 869, 1318, 1338-1340, 1347, 1353, 1358, 1442, 1448, 1445, 1447 Boule, Marcellin, 4, 19, 516, 519, 521, 527-538, 551, 574, 576, 766, 977, 978, 1035, 1037, 1093, 1094, 1268, 1475, 1519, 1624, 1628 Bourdelle, Edouard, 14 Bourg de Bozas, Robert, 4, 766 Bovide, 467 Bowerbank Collection, 1234 Bowman, Isaiah, 14 Boyd County, 251, 298, 738 Boyes, H. D., 308, 310 Bozasi (see Deinotherium Bozasi) brachycephaly, 919, 1552 brachyodonty, 140, 919, 1548, 1552 Brachyodus, 1425, 1426, 1429, 1441, 1442, 1459 brachyopy, 1550, 1552 Brachypotherium, 461, 1479 Bradenton, Florida, 400, 1047, 1105, 1106, 1108 Bradley, D. F. Levett, xvii, 934, Pls. xrv-xvitt. See also charts Brahmaputra River, 853, 927 brains, 47, 1139, 1251, 1335 Branco [Branca], Wilhelm, 4, 551, 766 Brander, Dunbar, 1317 Brandt, Johan Friedrich, 3, 27, 93, 766, 1121, 1123, 1130, 1136, 1369, 1388 brasiliensis (see Cuvieronius brasiliensis) Brassempouy, 1168 Braunkohle von Vordersdorf, 395 Brauns, David August, 3, 766, 818, 1295 Bravard Collection, 631, 633, 634 Brayley, Edward William, 3, 767, 1385, 1624 Brazener, H., 1332 1635 Brazil, 518, 524, 527, 530, 532, 536, 537, 575, 741. See also Minas Geraes Brazos River, Texas, 349, 374, 501, 559, 738, 1047 brazosius. See Trilophodon (?Tetralophodon) brazosius Breislak, Scipione, 3, 767, 1256 Breitenbaum, Hungary, 91 Breslau, 259, 745, 754. See also University of Breslau Breuil, Henri, 4, 767, 968, 1131, 1184 Brevard County, Florida (see Amherst College) brevidens (see Rhynchotherium brevidens) brevirostre (see Anancus arvernensis brevirostris) Brevirostrine, 27, 28, 31, 124-128, 228, 229, 352, 477, 539, 614, 617, 618, 624— 626, 627, 628, 629, 640, 641, 6438, 667, 734, 740, 1528, 1545, 1550, 1563-1565, 1567, Pl. x. See also Anancine, Dibunodontide, Pental- ophodontingw, Anancus, Pentalophodon, Synconolophus Brewster, Florida, 400, 482. See also Ocalientinus (Serridentinus) bifoliatus, Pliomastodon sellardsi, Serridentinus brewsterensis brewsterensis (see Serridentinus brewsterensis) Breyne, John Philip, 2, 767, 1118, 1119, 1165 Bricy, France, 109 Briggs, Caleb, Jr., 3, 767, 1068, 1069 Brihuega, Spain, 134 Brionde, France, 980 Briones, 902 Briscoe County, Texas, 1005, 1017, 1018 Bristow, Boyd County, Nebraska, 251, 288, 289, 298, 299, 301, 315, 738 British Museum, 18, 745, 857, 910, 972, 973, 1193, 1204, 1216, 1218, 1329, 1343, 1348, 1392, 1609, 1611, 1613, 1614, 1619 Brives, Abel, 4, 95, 105, 767 Brno (see Briinn) Broili, Ferdinand, 14, 1314 Bronn, Heinrich Georg, 91, 767 Broom, Robert, 4, 14, 767, 987, 989, 944 broomi (see Archidiskodon broom) Brous, Henry, 424 Brown, Alastair, xvii, 728 Brown, Barnum, 342, 448, 449, 474, 484, 498, 494, 503, 505, 616, 622, 647, 648, 655, 661, 665, 767, 824, 847, 848, 869, 874, 878, 948, 950, 951, 959, 986, 1007, 1008, 1243, 1318, 1339, 13847, 1348, 1354, 1357, 1358 Brown, C. Emerson, 767, 1193 Brown County, Nebraska, 318, 537, 601, 602, 733, 741. Quarry, Devil’s Gulch browni (see Rhynchotherium brownt, Serridentinus browni) Brown’s Park formation, 289, 290, 312, 315, 711, 738, 1491, 1492, Pl. vir Brucksdorf, 1050 Briinn, 1139, 1168, 1609, 1610 Brule formation, 601, 733 Bruni, Borneo, 700, 737, 845, 851, 1305, 1419, 1455 Brunswick Canal, 1047, 1071, 1072, 1077 Brussels, 1130, 1132, 1609, 1610 Brittelen, Switzerland, 115, 204, 1459 Bryan, Kirk, 679, 680 Bryan, William Alanson, 14 Buba (see Flinsch, Margret) Bubalis baini, 984; B. antiquus, 1183; B. palzwindicus, 1449 Bucharest (see University of Bucharest, also Giiceana and Manzati) Buckland, William, 3, 87, 767, 825, 973 Bucklandi (see Tetracaulodon Buckland?) Buckley, Thomas Edward, 29, 767 Budapest, 160, 396, 638, 1251. See also Ungarische geologische Reichsanstalt, and Ungarisches Nationalmuseum Budongo Forest, Uganda, 1190, 1239 Buenos Aires, 517, 519, 526, 527, 580, 531, 537, 548, 544, 575, 579, 581, 586, 587, 590, 591, 593, 597-599, 615, 745, 756, 1609, 1610 Buffalo County, 1012 Buffalo, Kansas, 188, 189, Pl. 1 Buffelus, 887, 1253 buffer-tuskers, 324 Buffon, Georges Louis Leclere de, 2, 133-136, 166, 168, 192, 207, 209, 767 buffonis (see Zygolophodon borsoni buffonis) See also Christmas 1636 OSBORN: THE Bug River, 1136 Bugti Beds (see Bugti Hills) Bugti Hills, Baluchistan, 78, 85, 105, 106, 115, 266, 267, 269 271, 275, 279, 448, 449, 643, 659, 1439-1442. See also Dera Bugti, Gandoi, Kumbhi bugtiense. See Paraceratherium (Baluchitherium) bugtiense Bukit Besar, Nawngchik, 762 bulla, 916, 920 Bullard, Roger Perrin, xvii, 731 Bumiaju, Java, 349, 365, 366, 837, 896, 1453 bumiajuensis (see Tetralophodon bumiajuensis) Bunnell, Charles E., 1134, 1161 Bunolophodon, 10, 11, 194, 195, 266, 269, 271, 272, 765, 944, 945, 984, 994 ayorx, 1377, 1415. See Cuvieronius ayore postremus (see Cuvieronius postremus) yokotw, 1420 Bunolophodont mastodon, 1436, 1439 bunolophodonty, 1545 Bunomastodontide, 25-27, 29, 31, 45, 65, 119, 126, 128, 225, 226, 228-231, 352, 367, 477, 542, 587, 686-688, 690, 705, 730, 737-740, 1367-1369, 1525, 1545, Pl. x Bunomastodontine, 27, 1368 bunomastodonty, 237, 238, 1545 Bunopithecus, 1452 Burdigalian, 107, 115-117, 204, 205, 217, 218, 252-2 735, 742, 902, 903, 1427, 1428, 1441, 1457— Sologne Bureau County, Illinois, 173, 736 Burge, Nebraska, 707, 1496, 1497, 1499 Burg-Graefentonna, 1045 Burgtonna, 1118, 1119, 1122, 1138, 1141, 1165, 1174, 1181, 1217, 1245, 1865 Burleson County, Texas, 479, 501, 537, 559, 737, 740 Burma, 114, 449, 642, 643, 829, 1304, 1440, 1450, 1451, 1487. See also Irrawad- dy, Mandalay, Mingoon, Stegodon insignis birmanicus, Yenangyaung, Burmeister, Hermann, 3, 7, 91, 521, 544-546, 575, 576, 723, 767, 1373, 1374 Burmese elephant, 1332 Burnett, Gilbert Thomas, 767, 1123, 1136, 1363 Burney, Colonel, 863 Burnham, Frederick, 444 burnhami (see Serbelodon burnhamt) Burns, Edward A., xvii Bush elephant, 1196 Bushman, 968; Pre-Bushman (see Boskop man) Busk, George, 3, 767, 907, 1187, 1257, 1263, 1264, 1333, 1334 buski (see Palxoloxodon buskt) Butler County, Nebraska, 1012 Buttikofer, 1314 Buwalda, John P., 4, 14, 486, 487, 767 Buzen, 1489 54, 261, 267, 270, 457, 693, 1460, 1477, 1479. See also Cabinet of Lyceum of Natural History, New York, 1156 Cabrera, Angel, 4, 14, 385, 537, 542, 579, 580, 587-590, 593, 595, 612, 767, 1368, 1370 Cabriéres, 115 Cacciamali, G. B., 768, 1241 Cachihuayco, quebrada von, Ecuador, 537, 567, 568, 585, 1415, 1521 Cenobasileus tremontigerus, 10, 1377, 1396 Caggana, Dominick, xvii Cainello, 1239 Cairo Geological Museum, 745, 755 Calabrian, 818, 902, 904-908, 964, 1178, 1186, 1290, 1293, 1297-1301 Calacoto, Bolivia, 552, 1415 Calcaire de Montabusard (see Montabusard) Calcutta, 266, 745, 756, 844, 1609, 1610 California, 15, 163, 1418, 1609, 1619. See also Bautista Creek, Chanac, Contra Costa, Dry Creek, Etchegoin, Mohave Desert, Mt. Eden Hot Springs, Oak Springs, Ricardo, San Bernardino, San Joaquin, San Timoteo, Santa Rosa Island, Stanislaus County Callaway, Nebraska, 1009, 1012 Callihuayco (see Cachihuayco) PROBOSCIDEA Calomna, Russia, 1136 Caluire, 1063, 1064 Calvert formation, 285 Calvin, Samuel, 4, 172, 682, 683, 768, 1070 Cambay, India, 85, 90, 91, 644, 844, 852, 1390 Cambridge, England, 14. See also University Museum of Zoology Cambridge, Furnas County, Nebraska, 349, 377 Cameloids, 315, 318, 399, 400, 444, 461, 488, 498, 508, 510, 562, 579, 601, 679, 725, 1082, 1135, 1161, 1840, 1518, PL. vii Cameron, G., 1332 Cameroon, South, 1193, 1196 Camp Margetts, 398, 466 Campbell, Franklin County, Nebraska, 1012, 1092 Camper, Peter, 2, 5, 7, 768, 931, 1117, 1126, 1147, 1363, 1383 campester (see T'etralophodon campester) Camporosso, Italy, 1123, 1137 campylotes (see Elephas campylotes) Canada (see Hamilton) canadense (see Harpagmotherium canadense) Cananea, Mexico, 1033 Caneto, |’ Abbé, 3, 768 Cangahua, 1521 Canide, 1518. See also Canis canines, 1547 Canis lupus, 12381; C. nescherensis, 1476 Cantamessa, Fillipo, 3, 768 Canterbury Museum, 1215, 1609, 1610 Canuelas, Province of Buenos Aires, 599 Cap de Bonne Esperance, 1197 Cap Rock Beds, 1496, 1497 Cape Colony, South Africa, 1193, 1197, 1205 Cape Maleka, Crete, 1187, 1267 Cape of Good Hope, 931, 1197 Capellini, Giovanni, 4, 14, 619, 768 Capetown, 1193, 1609, 1611 Capitan, Joseph Louis, 4, 768, 1131 Capitan Sarmiento, Argentina, 594, 598 Capreolus capreolus, 1155 Capromeryx, 1082 Caprovis savinii, 1155 Cardamone cave, Otranto, 1149-1151, 1399 Carette, Eduardo, 4, 518, 519, 523, 569, 570, 768 Carignan, 1086, 1407 Carini, 1182, 1187, 1214, 1257, 1261, 1262 Carlat-le-Comte, 85, 86, 115 Carles, Enrico de, 3, 532, 768, 1519 Carleton, M. M., 1353 Carlos Casares, Buenos Aires, 597, 599 Carman, J. Ernest, 14, 169, 1067 Carnegie Institution of Washington, 14, 1604 Carnegie Museum, 15, 388, 745, 755 Carnivora, 887, 1489 Carolia Beds (Qsar-el-Sagha), 53 carotid canal, 916, 920 Cartennien, 1426, 1427 Caruana, A. A., 1624 Carus, Julius Victor, 93, 768 Casalvieri, 1239 Casanova, 1193 Cass County, 1012, 1047, 1088 Cassegrain, France, 192, 205, 207 Cassino, Tuscany, 114, 660, 1210, 1238, 1239, 1245, 1469, 1470 Castellanos, Alfredo, 4, 589, 590, 595, 768, 1517, 1624 Castelliri, 1239 Castelnau, Francis de, 554, 768 Castelnau-Magnoae, region of Pyrenees, 205, 554 Castillo, 1081 Castor, 48, 161 Castoroides, 1107 INDEX Catalonia, 1468 Catania, 1204 Catawba, Ohio, 169 Caterini, Francesco, 4, 768 Caucasus, North, 251, 385, 460, 461, 729, 743, 964. See also Kuban Cautley, Proby Thomas, 3, 621, 625, 627, 643, 646, 647, 649-652, 768, 940, 1181. See also Falconer cautleyi (see Stegolophodon cautley?) cautleyi progressus (see Stegolophodon cautleyi progressus) cave drawings, 1184, 1194, 1216, 1229 cave fissures and fauna, 1451-1453, 1521 cavendishi (see Loxodonta africana cavendishi) Cayenne, French Guiana, 1046, 1047, 1083 cayennensis (see Parelephas columbi cayennensis) Cedar Mountain Beds, 1509 cement, 1549 Cenozoic, 1480, 1482, 1484, 1490 Central America, 1516 Central Asiatic Expedition, 328, 332, 333, 396, 398, 461, 465, 817, 859 Centralian, 1510 centroversion, 575, 613-615, 1548, 1549 Century Atlas, 1241 Century Magazine, 791. See Christman, Erwin, and Knight, Charles R. Ceprano, 1239 Cerdagne, 1468 Cerro de la Silla, 1082 Cerro de Licto, Ecuador, 567 Cervide, 1518. See also Cervus Cervulus, 887 Cervus, 887, 1155, 1240, 1253, 1261, 1449, 1511 Ceselli, Signor, 1061 Cetacea, 35 Ceylon, 931, 936, 1319, 1328, 1337, 1440, 1450, 1596 Chabel el Ameur, Algeria, 485 Chadron, 318 Chagny-Bellecroix, 633, 634, 934, 961-964, 1024 Chakravarti, D. K., 1188, 1418, 1624 Chakwal, India, 279 Chalampi Madua, Ceylon, 1337 Chalang, quebrada von, Ecuador, 536, 537, 567, 583, 1413, 1521 Chalicomys, 1463 Chalicotherium sinense, 818 Chamberlin, 400 Champion, Frederick Waller, 29, 768 Chanac-Etchegoin, 497, 498, 560, 1509 Chandigarh, 950, 952, 954, 955, 1339, 1340, 1347, 1356, 1367 Chaney, Ralph W., 508, 768 Chang, H. T., 1453, 1624 Changteho, North Honan, 1362 Channel Islands, 1031, 1032 Chantre, Ernest, 3. See also Lortet Chapman, James, 29, 768 Chapman,—, 668 chapmani (see Stegomastodon chapmant) Chappe, l’Abbé, 133 Charleston, South Carolina, 385, 386, 387, 419, 420. See also Phosphate Beds Charnian, 871, 952, 954, 955, 1339, 1340, 1347, 1351 charts, 1422, 1440, 1457, 1477, 1491, 1516, 1523 Cheiroleites, 10, 1123, 1124, 1365, 1375 Chelfor6, arroyo, Ayacucho, 598 Chellean, 1050, 1057, 1065, 1169, 1430, 1433 Cherchel, 1183 Cherichera, Tunisia, 485, 1428 Cherry County, Nebraska, 294, 318, 385, 460, 470 Chevilly, France, 85, 86, 109, 111, 115, 137, 176, 192, 205, 207, 283, 1459 Cheyenne County, 1012 Chi Chia Kou, 1481 Chiba Prefecture, 1300 Chicago (see Field Museum of Natural History) 1637 Chichas, Bolivia, 577 Chiconautla, 1081 Chile, 530, 532, 611. See also Concepcion, Tagua-Tagua chilensis (see Cuvieronius chilensis) Chimborazo, Ecuador (see Cuvieronius ayore) China, 15, 115, 348, 351, 352, 355, 384, 457, 458, 698, 699, 704, 732, 739, 742, 816, 1062, 1440, 1451. See also Fokien, Honan, Kansu, Shanghai, Shansi, Shensi, Szechuan, Yangtze, Yiishe, Yunnan North China, 1477, 1479-1485, 1487, 1488 Chinautla, Guatemala, 385, 432, 742, 1516 Chinglo, North Shansi, 1483 Chinji, 847, 1442-1448 Lower Chinji, 251, 272, 273, 276, 353, 385, 387, 448, 449, 451, 452, 454, 456, 457, 622, 642, 648, 647, 656-658, 731, 737, 738, 741, 742, 835 Upper Chinji, 251, 274, 276, 448, 449, 622, 642, 738 Chinji Bungalow, 272, 273, 279, 385, 452, 454, 456, 479, 625, 642, 643, 645, 647, 657, 737, 738, 740-742 chinjiense (see Rhynchothertum chinjiense) chinjiensis (see Rhynchotherium chinjiense, Serridentinus chinjiensis, Trilo- phodon chinjiensis) Chiquitos, Bolivia, 577 Chishima group, 1305 Chisholm and Leete, bathymetric map, 1304 Chitenay, Sables de, 1458, 1459 Chittagong, 927, 1315 Chleuastochwrus, 704 choerodonty, 629, 1548 Choerolophodon, 10, 249, 261, 344-346, 1379, 1393, 1556. (Choerolophodon) pentelicus Chokrak Beds, 334, 385, 461, 718, 743 “Choonee,”” 1312 Choptank formation, 285, 386, 738 Choukoutien, 1451, 1452, 1454, 1480, 1483-1487 Choulans, 1064 Christiana, South Africa, 984, 1188, 1280, 1281 it Christina (Bone Valley), 400 . Christman, Charles, 324 Christman, Erwin, 38, 48 Christmas Quarry, Nebraska, 317, 318, 443, 742, 1417 Christy, Henry, 3, 784, 1132 chronologic lists of species, 52, 84, 136, 192, 250, 349, 384, 479, 537, 625, 823, 942, 1047, 1136, 1187, 1319 Chuan Tou Kou, China, 732 Chubb, Ernest Charles, 29, 768 Chungkingfoo, 817, 818, 832, 884 Churlando, 266, 272, 275, 277, 279, 448, 449, 1440, 1441 Cierbo, 902, 1446 cimarronis (see Serridentinus serridens cimarronis) Cincinnati, Ohio, 1088, 1090 Cinnamomum, 117, 217 Citellus, 680 Clack County, Oregon, 1088 Clacton (Essex), 1217 Clallam County, Washington, 1047, 1104 Clarendon, 399, 408, 422, 429, 1501. See also Serridentinus productus, S. serridens, S. serridens cimarronis, Tetralophodon fricki Clarendonian, 1495-1501 Claridge, W. Walton, 29, 768 Clark, J. Desmond, 1188, 1420, 1624 Clark, James L., xvii Clarke, John Mason, 4, 14, 15, 132, 769 classification, xii, 10, 12, 17, 19, 20, 23-33, 46, 519, 522, 523, 525-527, 533, 534, 538, 587, 615, 689, 735, 853, 901-932, 1363-1420, 1525, 1526, 1539 Claudius, Matthias, 3, 92, 769 Clay County, Nebraska, 1012 Clery Creek, Alaska, 1135, 1161 Clift, William, 3, 769, 825, 827, 828, 840, 842 cliftii (see Stegodon elephantoides = cliftir) climate, 551, 1373, 1480-1432, 1475, 1476, 1492, 1516, 1518 See Trilophodon 1638 OSBORN: Clovis (valley deposit), 1512 Coalinga, Fresno County, 161, 162, 736. See also North Coalinga Beds Coates, Isaac, 554 Cochise County, Arizona, 535, 565, 623, 678, 740, 741 Cody, Nebraska, 1003 Cohen Collection, 420, 748, 1075, 1076, 1082 Colbert, Edwin H., xvi, 5, 14, 448, 506, 595, 713, 715, 723, 724, 728, 769, 824, 857, 950, 1023, 1049, 1247, 1364, 1367, 1374, 1411, 1416, 1418, 1420— 1521, 1571, 1624 Colintina, Rumania, 969, 1232 Collar, Hubert, 931 Collaud, General, 136 collective names, 10 College Station, Texas, 374, 560, 745, 759 Collinsii (see Tetracaulodon Collinsit) Collinson, Peter, 2, 134, 165, 769 Cologne, 1187 Colombia, 530, 532, 595 Colorado, 729. See also Brown’s Park, Douglas Mountain, Moffat County, Pawnee Buttes, Weld County, Wray, Yuma County Colorado Museum of Natural History, 14, 709, 745, 755, 1103, 1609, 1611 Colorado Springs, 1003 columbi (see Parelephas columbi, also Elephas primigenius columbt) Columbian Mammoth (see Parelephas columbt) Columbus, Ohio, 1609, 1611 Combarelles, 1126, 1131, 1167 Comminge, France, 85, 86, 115 commutatus (see Elephas commutatus) compressus (see Mammonteus primigenius compressus) Compubay (see Cambay) comune (see Elephas primigenius comune) Concepcion, Chile, 5, 122, 123, 516, 537, 541, 549, 575-577, 615, 741 Concordia, Bolivia, 552, 1415 concretionary zone (Brown County), 733 Concud, Spain, 114 cones and conelets, 141, 199, 277, 352, 810, 1545, 1554, Pls. -1v. See also ecto- and entoconelets Congerienschichten, 100 Conglomerate, 482. See also Boulder Conglomerate Congo, 931, 1193, 1196, 1383 connexus (see Trilophodon connexus) conodon (see Gomphotherium conodon) Conohyus, 272 conservativus (see Mastodon arvernensis Croiz. et Job. var. conservativus) Constantine, Province of, 1427 Constantinople, 1466 continental and insular races of Indian elephant, 1315-1328 Contra Costa County, California, 484, 487, 488, 495 Contreras, Francesco, 1007 conules, 25, 139, 144, 145, 225, 226, 277, 341, 686, 1545, 1546. See also trefoils convergence, 911, 930, 933, 1039, 1177, 1350 Cook, Harold James, xvii, 4, 14, 157, 251, 288, 307, 309, 315, 624, 684, 769, 1496, 1499, 1624 Cook, Margaret C., 1624 Cooke, Charles Wythe, 400, 769 Cooke, H. B.S., 1188, 1420, 1624 Cooper, Clive Forster, 4, 14, 95, 103, 105, 251, 267, 269-271, 769, 1214, 1216 1220-1222, 1441. See also Bugti Hills, Churlando Cooper, William, 3, 135, 769. See also Croghan, Longueil cooperi (see Trilophodon coopert) Cope Collection, 298, 342, 394, 401-403, 407, 423, 484, 489, 501, 586, 587, 596, 746, 748 Cope, Edward Drinker, 3, 7, 19, 28, 212, 249, 251, 298, 369, 384, 403, 404, 423, 429, 431, 485, 488, 489, 519, 525, 534, 537-539, 553, 554, 671, 769, 1082, 1100, 1101, 1397, 1399, 1503. See also Cope Collection Copeland farm, 1102 Coralline Crag, 1469 cordillerarum (see Mastodonte des Cordilieres) Cordilleras, 5, 518, 537, 540, 541, 543, 551, 576, 584, 595 THE PROBOSCIDEA Cordillerion, 12, 29, 128, 496, 498, 501, 533, 537, 540-542, 548, 544-549, 552, 553, 560, 571, 576, 584, 587, 588, 595, 685, 690, 722, 730, 731, 740, 1381, 1514, 1515, 1519, 1527, 1549, 1560, 1561, 1566 andium, 10, 12, 29, 118, 228, 229, 418, 477, 501, 514, 516, 617, 518, 519, 521-532, 536, 537, 589-541, 544, 545, 547, 548, 649, 550, 561, 562, 566, 568, 570, 578, 582, 5838, 588, 595, 615, 633, 753, 755-759, 1375, 1386, 1520, 1550, 1604. See Créqui-Montfort Collection, Mastodonte des Cordilitres, Tarija, Weddell Collection andium kraglievichti, 536, 537, 538, 541, 548, 1412 bensonensis, 399, 496, 5385-538, 541, 548, 562, 665, 566, 615, 619, 623-625, 680, 731, 740, 759, 1411, 1505 bolivianus, 522, 524, 5380, 586-538, 540, 541, 548-550, 561, 552, 582, 588, 615, 740, 1400, 1415, 1520. See also T'eleobunomastodon bolivianus defloccatus, 399, 496, 5385-538, 541, 548, 562, 664, 565, 566, 615, 623-625, 740, 759, 1411, 1508, 1514 edensis, 496, 497, 498, 507, 535-538, 541, 548, 559, 560-662, 588, 615, 740, 760, 1406, 1419, 1604 gratum, 399, 496, 536-588, 541, 548, 557, 659, 560, 562, 564, 566, 588, 615, 740, 759, 1405, 1514 oligobunis, 584, 536-538, 541, 548, 549, 664, 655, 564, 588, 615, 740, 757, 1081, 1400, 1515 oligobunis antiquissimus, 534, 536-538, 541, 548, 5538, 656, 556, 559, 562, 564, 566, 615, 740, 757, 1082, 1407, 1515 oligobunis felicis, 535-538, 541, 548, 549, 553, 666, 557, 562, 564, 615, 740, 757, 1082, 1407, 1515 oligobunis intermedius, 534-588, 541, 548, 553, 557, 658, 559, 562, 564, 615, 740, 757, 1082, 1515 oligobunis progressus, 534-538, 541, 548, 553, 658, 562, 564, 615, 740, 757, 1081, 1407, 1515. See Cordillerion tropicus, 1398 orarius, 399, 535-538, 541, 548, 559, 562, 663, 564, 566, 588, 615, 623-625, 740, 757, 1411, 1508, 1514 tarijensis, 536-538, 541, 548, 549, 550, 588, 615, 740, 1401, 1520 tropicus, 431, 525, 582, 588, 587-541, 545, 548, 653, 557, 588, 740, 754, 1081, 1398, 1515 Cordillerionine, 1370, 1371, 1560, 1628 Cérdoba, Province of, 589, 593 cornaliae (see Loxodonta cornaliae) Corocoro, Bolivia, 551, 1415 correlation, new standard of, xiv correspondence in Osborn Library, 9 Corrientes, Argentina, 537, 582, 741 corrugatus (see Synconolophus corrugatus) Corse, John, 2, 770, 1313, 1325, 1336, 1337 Costa, Oronzio Gabriele, 3, 771, 1128, 1241 Cotter, G. de P., 4, 771 cottoni (see Loxodonta africana cottont) cotype, definition of, 9 Couches saumatres 4 congéries, 114 Coues, Elliot, 3, 771 Couper, J. Hamilton, 771, 1071, 1077 Coupet supérieure, 634 Covurlui, 943, 968, 969 Cox, Perey Zachariah, 29, 771 cranial mechanics, 915, 918, 919 cranial sections of elephants, 918, 924 cranium, 933, 1549 Crawford, J., 824, 825, 861 Creodonta, 1425 crepusculi (see Hemimastodon crepuscult) Créqui-Montfort Collection, 527 Crete, island, 1183, 1187, 1257, 1267 Crete, Saline County, Nebraska, 942, 1012, 1023 creticus (see Palzoloxodon creticus) cribriform plate, 920 Crimea, 964 Croatia (see Murinsel) Croghan, George, 135, 136, 166 Croix-Rousse, 1063, 1468 Croizet, |’ Abbé Jean Baptiste, 3, 618, 625, 632, 771 INDEX 1639 Cro-Magnon, 1132, 1168 Cyprus, 1188, 1187, 1257, 1267 Cromer, 963, 981, 1523, 1609, 1611. See also Forest Bed cyptocephaly, 919, 1552 cromerensis, Hlephas meridionalis mut., 1407. See also Archidiskodon meridi- eyrtocephaly, 919, 1552 onalis cromerensis eyrtodonty, 919, 1552 Cromerian, 818, 902, 904-908, 970, 980, 982, 1059, 1186, 1430, 1447. See also Cenozoic, 1453 Archidiskodon meridionalis cromerensis Czechoslovakia, 13 cruziensis (see Megabelodon cruziensis) Cryptomastodon, 1382 Dagshai, 449 martini, 625, 13882, 1417, 1455 Dakota, South (see Black Hills, Dallas) Csakvar, 1468 Dall, William Healey, 3, 400, 418, 772 Cub Creek, Nebraska, 318 Dallas, South Dakota, 251, 304-306, 738 Cucigliana, Tuscany, 1187, 1230 Dallas, Texas, 1004, 1005, 1009, 1399 Cucuron, 114, 1468 Damaliscus, 994 Culbertson, Nebraska, 489 Dana, James Dwight, 3, 772 Culver, Harold E., 4, 771 Dannebrog, Nebraska, 763, 1009, 1012 Cummins Collection, 431, 673, 759 danovi (see Platybelodon danovi) Cummins, W. F., 1501, 1503, 1624 Danube basin, 964 cuneatus (see Pentalophodon cuneatus) Darien, Georgia, 1071, 1072 Cunene, Angola, 1193 Darke County, Ohio, 1097 Curtis Flats, Arizona, 625, 678-680, 741, 1504 Darling Downs, Australia, 85 Curtis, Lincoln County, Nebraska, 948, 1009, 1017, 1019, 1027 Darmstadt Museum, 80, 86, 89, 94, 281, 282, 357, 358, 368, 371, 745, 755 Curtis Ranch, 1504 Dart, Raymond A., 4, 14, 772, 943, 984, 990-992, 1188, 1279-1285, 1287, 1439 Curtognathide, 26, 30, 81-85, 87, 688, 735, 1367, 1368, 1525 darti (see Palxoloxodon darti) Curtognati (see Curtognathidz) Darton, N. H., 1501, 1624 Custer County Mammoth, 1009, 1012 Darwin, Charles, xiv, xv, 12, 1315 Cuttack, India, 1315 Darwinian Period, 3 Cuvier, Frédéric, 3, 6, 771, 1174, 1192, 1325, 1335 Dasypodide, 35, 386, 527 Cuvier, Georges, xiv, 2, 5-7, 84, 86, 119-123, 128, 136, 137, 192, 217, 250, 252, | Daubenton, Louis Jean Marie, 2, 167, 772, 931 340, 516, 537, 549, 576, 618, 771, 915, 931, 970, 1117-1121, 1123, 1136, | Dauntela var. See Elephas indicus Dauntela var. 1148, 1198, 1810, 1364, 1401. See also classification Dauphiny, France (see Vienne) cuviert (see Deinothertum cuviert, and Trilophodon angustidens cuvieri) Davidson,—, 1479 Cuvierian Period, 2, 8, 516 Davies, William, 3, 772 Cuvieroniine, 542, 587, 688, 1370 Davis Ranch, 488 Cuvieronius, 11, 31, 128, 501, 515, 516, 538, 540-542, 546, 567, 569, 574, 575, Dawes County, Nebraska, 1012 576, 584-589, 595, 602, 611, 612, 614, 615, 691, 722, 730, 741, 1380, | Dawkins, William Boyd, 4, 772 1519, 1525, 1527, 1561, 1566, 1567, 1601 Dawson, Charles, 966, 772. See also Hoanthropus dawsoni ayore, 531, 532, 5386-538, 541, 566, 567, 571-575, 683, 584, 585, 598, 611, De Blainville (see Blainville) 615, 741, 757, 1413, 1521, 1604 De Lorenzo, Giuseppe, 4, 14, 772, 1239-1243 bonaerensis, 536-538, 541, 571, 575, 679, 588, 589, 696, 598, 606, 608, 611, decaloph-lophid, 1545 615, 741, 757, 1399, 1519, 1604 Deccan, 267, 268, 1394 bonaerensis (=superbus). See Cuvieronius bonaerensis deciduous premolars, 1270, 1547 brasiliensis, 518, 5386-538, 541, 575, 578, 588, 615, 741, 1521 Deep Creek, 318, 1493 chilensis, 522, 524, 530, 536-538, 541, 575, 576, 681, 682, 588, 615, 741, Deep River Beds, 404, 478, 479, 485, 737, 746, 748, 798, 1498, 1494 1400, 1520 Defay,—, 2, 193, 772 humboldti, 5, 10, 11, 29, 31, 119, 121-123, 228, 501, 516, 517, 519, 521-533, — defloccatus (see Cordillerion defloccatus) 536-542, 544, 547, 549, 569, 575, 676, 577, 578, 580, 587-589, 592, Dehm, Richard, 1624 598, 595, 607, 611, 613, 615, 633, 741, 754, 1380, 1386, 1393, 1520. Deinotheriid#, 26, 83, 85. See also Curtognathidse See also Mastotherium Deinotheriine, 27, 30, 83, 84, 734, 735, 1369, 1526, 1528, 1545, Pl. x (2?) maderianus, 524, 536-538, 541, 575, 576, 579, 681, 589, 593, 595, 615, Deinotherioidea, 22-24, 27, 30, 32, 38, 81-85, 111, 114, 115, 688, 735, 1367, 741, 1400, 1519 1524, 1550, 1553-1555, 1600, Pls. x, x1. See also Curtognathide ptrayuiensis, 5386-538, 541, 575, 682, 589, 615, 741, 1519 Deinotherium, 10, 11, 30, 80, 81, 83-113, 139, 253, 254, 448, 485, 686, 734, 735, platensis, 501, 519-521, 523, 524, 530, 531, 5386-538, 541, 544, 549, 569, 575, 576, 579, 580, 586, 587, 589, 592-599, 611-613, 615, 722-724, 741, 746, 753, 756, 757, 1399, 1519, 1548, 1601, 1604, 1612 postremus, 531, 536-538, 541, 567-569, 571-576, 584, 685, 611, 613, 615, 687, 741, 757, 1415, 1521, 1601, 1604, 1620 rectus, 519, 620, 521, 522, 524, 530, 536-588, 541, 569, 576, 580, 581, 589, 692, 598, 611, 615, 741, 757, 1399, 1519, 1568 superbus, 118, 514, 519, 620, 521, 522, 524, 531, 536-538, 542, 569, 575, 576, 578, 580, 588, 589, 598-595, 597, 599, 687, 722-724, 753, 756, 757, 1899, 1519, 1546, 1601, 1604, 1610, 1618. See also Cuvieronius bonaerensis Cuyama formation, 1509 cyclotis (see Loxodonta africana cyclotis) Cymatotherium, 1374 Cynarctus, 318, 601 Cynocephalus, 448 Cyon, 318 cypriotes (see Palxoloxodon cypriotes) 983, 1177, 1378, 14388, 1485, 1527, 1547, 1550, 1553, 1555, 1600, 1604, Pls. x, x1. See also Antoletherium angustidens, 85, 90, 111, 114, 448, 1890, 1448 australe, 85, 90, 1390 bavaricum, 84, 87, 90-94, 99-102, 106-111, 115, 735, 760, 1387, 1391, 1460, 1464, 1555, 1600, 1604 Bozast, 1417, 1419, 1435, 1437 cuviert, 38, 84, 85, 87, 90, 91, 94, 100, 105, 107-112, 115, 253, 735, 1387, 1391, 1460 (gig.| var. majus, 85, 89, 111, 114, 1391 (gig.| var. medium, 85, 89, 111, 114, 1391 [gig.] var. minus, 85, 89, 111, 114, 1391 giganteum, 80, 84, 86-91, 93, 94, 98, 102, 105-111, 114, 735, 748, 755, 763, 1177, 1373, 1386, 1391, 1469, 1554, 1604 gigantissimum, 84, 85, 95, 96-98, 106, 109-111, 113, 114, 116, 735, 755, 763, 1400, 1470, 1555, 1600, 1604 hobleyi, 38, 85, 103, 104, 111, 112, 115, 735, 983, 1403, 1428, 1429 hopwoodi, 38, 85, 104, 105, 114, 117, 735, 754, 1419, 1435, 1437, 1554, 1555 1640 OSBORN: THE Deinotherium—continued hungaricum, 85, 115, 116, 117, 735, 1381, 1415, 1460 indicum, 85, 91, 98, 103, 105, 109, 111, 114, 272, 274, 448, 454, 642, 735, 1391, 1395 indicum var. gajense, 85, 105, 110-112, 115, 272, 448, 735, 1402, 1403, 1441, 1448 intermedium, 85, 90-92, 94, 111, 115, 735, 1391, 1464 kénigit, 85, 90, 94, 111, 114, 1390, 1391 levius, 85, 100, 107, 109, 111, 115, 1395, 1464 maximum, 84, 111, 114, 1387 medium, 85, 90, 93, 109, 111, 114, 13888, 1469 minutum, 85, 111, 115, 13890, 1627 naricum (see Deinothertum indicum var. gajense) pentapotamix, 82, 85, 93, 103, 105, 111, 115, 272, 448, 454, 756, 1395, 1448 pentapotamicum, 93, 1395 perimense, 85, 105, 111, 114, 448, 1395 podolicum, 85, 87, 137, 735, 1389, 1469 proavum, 85, 87, 90, 92, 94, 111, 115, 1389, 1391 sands, 114, 115 secundarium, 85, 87, 111, 114, 1889, 1464 sindiense, 85, 93, 94, 105, 111, 115, 448, 1397, 1448 styriacum, 1404 uralense (=uralensis), 85, 87, 111, 112, 114, 735, 13889, 1469 DeKay, James Ellsworth, 3, 772, 1137, 1156, 1390, 1410 Delafosse, Wilfrid, 1624 Deli, 1329 Delport’s Hope, Africa, 993, 1188, 1279, 1288 Delsberg, 91, 93 Delsol, Henri, 966, 967 Denny, Charles S., 1492, 1624 Denstedt, 1050 dental histology, 1607, 1608, Pls. xxvi-xxx dental succession, 138, 199, 230-235, 950, 953, 1547, 1548. various species denticules, 828 dentinal lamin (see lamin) dentinal rod-cones (see rod-cones) Denver, 1003. See also Colorado Museum of Natural History Depéret, Charles, 4, 14, 15, 65, 129, 205, 206, 246, 251, 618-620, 693, 772, 773, 943, 980, 1042, 1043, 1049, 1056, 1064, 1137, 1154, 1187, 1221, 1232, 1458, 1459, 1462, 1469-1471, 1473, 1625 depereti (see Miomastodon deperett) Dera Bugti, 115, 251, 267, 269, 275, 279, 738, 1440, 1441 Deraniyagala, P. E. P., 1625 D’Erasmo, Geremia, 4, 14, 773, 1239-1243 Desaguadero (see Rio Desaguadero) Desmarest, Anselme Gaetan, 3, 122—124, 217, 250, 252, 516-518, 537, 576, 577, 773 Desmostylus, 625, 902 Desnoyers, Jules Pierre Frangois Stanislas, 3, 773 Desor, Pierre Jean Edouard, 3, 773 Devil’s Gulch, 294, 317, 318, 601, 733, 1498 Dhok Pathan, 279, 354, 362, 448, 449, 452, 454, 622, 625, 642, 643, 656, 658, 659, 661, 663-665, 737, 739-741, 845, 1442-1448 dhokpathanensis (see Synconolophus dhokpathanensis) diagrams, 426, 601, 728, 903-905 Diard,—, 1312 Dibelodon, 10, 11, 480, 519, 524-527, 537, 539, 540, 543, 575, 1378 cordillerarum, 525 edensis, 1406. See also Cordillerion edensis, and Rhynchotherium shepardi edense humboldtii (see Cuvieronius humboldtit) mirificus (see Stegomastodon mirificus) precursor, 1400. See Serbelodon(?) precursor shepard (see Rhynchotherium shepardi) tropicus, 1398. See Cordillerion tropicus Dibunodon, 905, 1379. See also Mastodon (Dibunodon) arvernense Dibunodontidx, 1369 Dicerorhinus (see Rhinocerotide) See also under PROBOSCIDEA Dicrocerus, 1462 Dicyclotherium, 10, 1123, 1124, 13865, 1373 Didelphis, 400, 579 Dietrich, Wilhelm O., 4, 14, 773, 813, 881, 887, 888, 945, 967, 1122, 1129-1131, 1137, 1140, 1152, 1184, 1187, 1275, 1870, 1411, 1416, 1438, 1466, 1625 Digby, Bassett, 1162, 1164, 1625 Diluvian, 1178 Dinohyus, 400, 1403 Dinotheride (see Dinotheriid) Dinotheriide, 83. See also Curtognathide Dinotheriine (see Deinotheriine) Dinotheriini (see Deinotheriinz) Dinotherina (see Deinotheriine) Dinotherioidea (see Deinotherioidea) dinotherioides (see Trilophodon dinotherioides) Dinotherium (see Deinotherium) Dintesheim, 281 Dipodomys, 680 Dipoides tortus, 298 Diprotodon (see Deinotherium australe, Diprotodon australis, Mastodon australis, Notelephas australis) australis, 1397 dissimilis (see Anancus arvernensis dissimilis) Ditmars, Raymond Lee, 14 divergence (see phylogeny) Dixie Creek, Oregon, 173 Djebel Kouif, 105 Djebel-Chirichira, 232 Djetis Zone, 1454, 1487 Doab Canal, 648, 650 Dobermannsdorf, 961 Dobson, Arthur, 696 dodecaloph-lophid, 1545 Doello-Jurado, Martin, 722, 723, 730, 731 Dolan, Elizabeth, 605 dolichocephaly, 1442 dolichodonty, 237, 1548 dolichopy, 237, 1550, 1552 domestication of the elephant, 1147 D’Orbingy, Alcide, 3, 7, 477, 482, 518, 549, 550, 582, 773 Dorcabune anthracotherioides, 272 Dorcatherium, 272, 274, 1429 Dordogne, 1131, 1132 Dorlhac, J., 3, 618, 680, 773 Dornap, 1137, 1150 Doughty, George D., 1005 Douglas Mountain, Colorado, 312, 385, 738 Douglass, Earl, 1493, 1508, 1625 dredge-tuskers, 335, 338, 471, 717 Driak, Fritz, 1625 Driftwood Creek, Nebraska, 251, 384, 401, 479, 489, 504, 737, 742 Dry Creek, Stanislaus County, California, 479, 487, 497, 737 Dryopithecus, 1426 Du Toit, Alexander Logie, 773, 944 dubius (see Mastodon dubius) Dubois, Eugen, 4, 14, 773, 885, 886, 889, 967, 1187, 1289, 1302, 1403, 1609, 1612 Duboisia (Tetraceros) Kroesentt, 887 Due d’Orleans, 1193, 1609, 1619 Duchére, 1063, 1064 Diirek, Hermann, 1330, 1331 Diisseldorf, 1136 Duggan,—, 1062 Dugongs, 39 Duke University, 14 Duméril, André Marie Constant, 3, 89, 773 Dundy County, Nebraska, 1012 Dunnellon, Florida, 400 Duplin marls, 733 INDEX Dupont, Edouard Frangois, 3, 773, 1130 Durfort, France, 961, 1065 Durfort skeleton, 259, 934, 977-979, 997, 1035, 1037, 1094, 1095, 1226, 1228, 1605 Dusino, Asti, 616, 618 Dutch Creek, Nebraska, 601 Dutch East Indies, 183 Duvaucel,—, 1312 dwarfed elephants, 936, 1271-1273, 1592, 1603. See also pygmy elephants Hales, Nellie B., 4, 14, 773, 931, 1202, 1203 East Anglia, 971. See also Forest Bed, Norwich Crag, Red Crag East India Company, 1312 East Indies, 859, 1304, 1451 Easton, England, 621, 651 Eastview, South Dakota, 251, 304, 328 Echigo, Japan, 818, 906, 1489 Eckingen, 115 Heole des Sciences, Algeria, 1275 ectoconelets, 687 ectotrefoils (see trefoils) Ecuador, 501, 532, 537, 567, 573, 574, 741, 1520, 1521. See also Universidad Central del Ecuador Eden Beds (see Mt. Eden Beds) edense (see Rhynchotherium shepardi edense) edensis (see Cordillerion edensis) Edentata, 35, 399, 400, 680, 887 Edgecombe County (see Tarboro) Edmunds, F. H., 997 Edson formation, 711 Eells, Myron, 1104 eellsi (see Parelephas eellst) Eggenburg, 115, 1459 Egypt, 251, 480. See also Fayim, Moghara, Zoological Gardens Bhik, Julius, 5, 116, 774 Ehringsdorf, 1045, 1181 Eibiswald, 92, 115, 384, 1461 Hichelberger, Grayson, 470 Eichwald, Eduardus, 3, 85, 87, 133, 137, 192, 621, 623, 639, 774, 1123, 1137, 1389, 1393 Eigasima, Japan, 1420 Ejutla, Mexico, 942, 1015 El Tajo, 1082 Elasmodon, 1175, 1177, 1375 Elasmodus, 1175, 1375 Eldar, Russia, 461 Elefante Indiano, 1382 elegans (see Tetralophodon elegans) Elephans etruscus, 972 Italicus, 972 Valdarnensis, 972 elephant, 1, 25, 269, 527, 1147, 1308 Elephant Point, Alaska, 1145 Elephantidx, 26, 32, 34, 126, 230, 400, 808, 809, 888, 911, 912. 913, 914-927, 930, 932, 936, 942, 948, 1202, 1228, 1307, 1364, 1366-1368, 1525, 1547, 1582, 1592 Elephantina, 27, 33, 13820, 1369 Elephantine, 22, 27, 32, 33, 913, 925, 927, 932, 936, 1178, 1227, 1228, 1307, 1320, 1369, 1526, 1538, 1545, 1595, 1596 Elephantini (see Elephantine) Elephantoidea, xii, xv, 22, 23, 25-27, 29, 32, 806, 807, 911, 912, 914, 915, 927, 932, 933, 936, 1307, 1367, 1524, 1545, 1547-1550, 1580, 1582, 1595, 1596, 1602 elephantoides (see Stegodon elephantoides) elephantoides (=cliftii). See Stegodon elephantoides ( =cliftiz) Elephantus indicus, 1310, 1322, 1323, 1383 Elephas, xii, xv, 6, 9, 10, 12, 19, 38, 47, 127, 138, 149, 527, 671, 815, 818, 853, 854, 902, 904, 907, 909, 912-919, 926, 927, 942, 948, 950, 1049, 1164, 1174, 1177, 1178, 1193, 1198, 1209, 1303, 1304, 1308-1311, 1318, 1641 1321, 1322, 1323, 1334, 1335, 1340, 1343, 1366, 13871, 1455, 1472, 1526, 1527, 1548, 1547, 1589, 1595-1597, 1603, 1607, 1608. See also Elephas indicus bengalensis and EF. 7. ceylanicus aff. meridionalis, 945 affinis, 761, 766, 1388, 1393, 1612, 1613 africanus (see Loxodonta africana) africanus albertensis, 1402. See Loxodonta africana albertensis africanus cavendisht, 1402. See Loxodonta africana cavendishi africanus cottoni, 1402. See Loxodonta africana cottoni africanus fossilis, 1183, 1187, 1398 africanus Franssent, 1404. See Loxodonta africana fransseni africanus group, 908 africanus mogambicus, 1409. See Loxodonta africana mogambica africanus orleansi, 1402. See Loxodonta africana orleansi africanus peeli, 1402. See Loxodonta africana peeli africanus priscus (see EHlephas priscus) africanus pumilio, 1402. See Loxodonta africana pumilio africanus rothschildi, 1402. See Loxodonta africana rothschildi africanus selousi, 1402. See Loxodonta africana selousi africanus toxotis, 1402. See Loxodonta africana toxotis africanus Zukowskyi, 1409. See Loxodonta africana zukowskyi americanus (see Mastodon americanus, and Mammonteus primigenius americanus) antiquitatis, 1386. See Hesperoloxodon antiquus germanicus, and Elephas meridionalis antiquitatis antiquus (see Hesperoloxodon antiquus) antiquus ausonius (see Hesperoloxodon antiquus ausonius) antiquus germanicus, 1408. See Hesperoloxodon antiquus germanicus antiquus group, 936, 1180, 1218, 1222 antiquus italicus (see Hesperoloxodon antiquus italicus) antiquus mut. ruthenensis, 1420, 1543 antiquus-namadicus group, 907, 908, 1178, 1185 antiquus Nestii, 1172, 1400. See Parelephas (?) trogontherii nestii antiquus platyrhynchus (see Hesperoloxodon antiquus platyrhynchus) antiquus recki (see Palzxoloxodon reckt) antiquus rumanus, 1408. See Archidiskodon planifrons rumanus antiquus var. insularis, 1182, 1187, 1257, 1269, 1403 antiquus var. minor, 1398 antiquus var. nana, 1397. See Hesperoloxodon antiquus nanus antiquus var. lrogontherioides, 1400, 1404. See Parelephas trogontherioides armeniacus, 1394, 1404. See Parelephas armeniacus asiaticus, 1197, 1319, 1323, 1325, 1383 atlanticus (see Palzoloxodon atlanticus) aurore (see Stegodon aurore) ausonius (see Hesperoloxodon antiquus ausontus) bombifrons (see Stegodon bombifrons) boreus, 1005, 1135 brachyramphus, 1123, 1130, 1136, 1147, 1387 campylotes, 1123, 1136, 1146, 1387 capensis (see Loxodonta africana capensis) (ef. antiquus), 944 cliftii, 1892. See Stegodon elephantoides ( =cliftii) columbi (see Parelephas columbr) columbi-imperator phylum, 1058 columbi var. Falconert, 1407. See Archidiskodon imperator falconeri columbi var. felicis, 1407. See Parelephas columbi felicis columbi var. imperator, 943, 944, 998, 1015, 1017, 1407 columbi var. silvestris, 1407. See Archidiskodon imperator silvestris commutatus, 1123, 1136, 1388 cornaliae, 1396. See Loxodonta cornaliae creticus (see Palxoloxodon creticus) cyclotis (see Loxodonta africana cyclotis) cypriotes (see Palxoloxodon cypriotes) eellsi (see Parelephas eellsi) (Euelephas) antiquus, 1394 (Euelephas) columbi, 1394 exilis (see Archidiskodon exilis) falconeri (see Palxoloxodon falconeri) ganesa (see Stegodon ganesa) 1642 Elephas—continued giganteus, 942, 969, 975, 1121, 1123, 1136, 1388, 1394 giganteus intermedius, 1187, 1397, 1400 gigas, 1319, 1323, 1384 gunnii, 1187, 1397, 1400 haroldcooki (see Archidiskodon haroldcookt) hayt (see Archidiskodon hay?) homotaphrus, 1123, 1136, 1387 hysudricus (see Hypselephas hysudricus) hysudricus hysudricus, 1319 hysudricus-indicus group, 907 hysudricus primitivus, 1319, 1404 hysudrindicus (see Palxoloxodon hysudrindicus) imperator (see Archidiskodon imperator) Indianapolis. See Parelephas(?) mississippiensis(?) indicus, 6, 10, 30, 303, 528, 602, 608, 905, 909, 910, 912, 915-932, 936, 937, 1006, 1020-1022, 1052, 1072, 1095, 1121, 1127-1129, 1131, 1147, 1165, 1173-1175, 1177, 1186, 1200, 1209, 1222, 1227, 1229, 1230, 1249, 1251, 1252, 1259, 1303, 1306-1321, 1822, 1323-1325, 1329, 1335-1337, 1340, 1342, 1344, 1349-1351, 1353, 1359, 1360, 1362, 1371, 1372, 1382, 1383, 1455, 1488, 1526, 1543, 1596-1599, 1605, 1614, 1615, 1618, 1620. See also Elephas maximus, and Messerschmidt indicus bengalensis, 918, 919, 931, 1052, 1092, 1147, 1311-1318, 1315-1319, 1322-1324, 1326, 1827, 1328, 1349, 1350, 1361, 1391, 1405, 1548, 1596, 1605, 1615, 1619 indicus Buski (see Palzoloxodon buskt) indicus ceylanicus, 1310, 1311, 1318, 1315-1319, 1322, 1323, 1327, 1328, 1350, 1361, 1391, 1402, 1522, 1543, 1596, 1615, 1619 indicus (Dauntela var.), 131, 919, 1175, 1813, 1315, 1317, 1819, 1825-1328, 1349, 1350, 1353, 1391, 1392 indicus Heterodactylus, 1313, 1315, 1319, 1389 indicus hirsutus, 1315, 1317-1319, 1331, 1832, 1333, 1404, 1543, 1596, 1614 indicus Isodactylus, 1313, 1315, 1319, 1389 indicus (Mukna var.), 1175, 1311, 1313-1317, 13819, 1825-13828, 1349, 1350, 1353, 1391, 1393 indicus sumatranus, 183, 930, 931, 1314, 1315, 1817-1319, 1322, 1325, 1329, 1330-1332, 1398, 1401, 1548, 1596, 1610, 1612, 1615, 1620, 1621 insignis (see Stegodon insignis) intermedius (see Parelephas intermedius) intermedius-trogontherti phylum, 1065 jacksoni (see Parelephas jacksont) Jjeffersonii (see Parelephas jeffersonit) jolensis (see Palzoloxodon jolensis) jubatus, 1123, 1136, 1385 Kamenskii, 1123, 1136, 1146, 1387, 1391 lamarmorae (see Palzoloxodon lamarmorae) (Lozxod.) priscus, 1187, 1198, 1217, 1219, 1385, 1392, 1394 (Loxodon) zulu. See Loxodonta zulu (Loxodonta) knochenhaueri. See Loxodonta africana knochenhaueri (Loxodonta) oxyotis. See Loxodonta africana oxyotis lyrodon, 942, 943, 969, 975, 976, 977, 1399 macrocephalus, 136, 165, 1383 macrorynchus, 1123, 1136, 1388 maibeni (see Archidiskodon imperator maibent) mammonteus, 5, 771, 1120, 1121, 1123, 1136, 1138, 1146, 1147, 1365, 1383, 1386, 1628 Mammoth, 1120, 1136, 1383 mammouth, 1123, 1384 maximus, 10, 1174, 1308-1311, 1315, 1319, 1323, 1371, 1382, 1543, 1595 maximus hirsutus (see Elephas indicus hirsutus) maximus indicus, 1328 maximus maximus, 1315, 1317, 1319, 1405 maximus sumatranus, 1315, 1317, 1319, 1401 maximus zeylanicus, 1315, 1317, 1319, 1402 melitensis (see Palxoloxodon melitensis) melitensis-atlanticus group, 908, 1178 meridionalis (see Archidiskodon meridionalis) meridionalis antiquitatis, 1400 meridionalis-E. trogontherti-E.. primigenius phylum, 1044, 1058 OSBORN: THE PROBOSCIDEA meridionalis mut. astensis (see Mammonteus primigenius astensis) meridionalis mut. cromerensis (see Archidiskodon meridionalis cromerensis) mindanensis. See Stegodon (Archidiskodon?) mindanensis minimus, 970, 1384, 1392, 1395 minutus, 970, 1395. See Trilophodon angustidens minutus mississippiensis. See Parelephas(?) mississippiensis(?) mnaidrex (see Palxoloxodon mnaidriensis) mnaidriensis (see Palxoloxodon mnaidriensis) namadicus (see Palxoloxodon namadicus) namadicus group, 1180 namadicus namadi (see Palxoloxodon namadicus namadi) namadicus naumanni (see Palxoloxodon namadicus naumanni) Nestii. See Parelephas(?) trogontherti nestii odontotyrannus, 1128, 11387, 1146, 1389 ohioticus (see Mastodon ohioticus) (Palxoloxodon) namadicus setoensis, 1188, 1295, 1408, 1413, 1489 paniscus, 1123, 1136, 1146, 1387 periboletes, 1123, 1136, 1146, 1387 planifrons (see Archidiskodon planifrons) planifrons-meridionalis group, 903 planifrons rumanus (see Archidiskodon planifrons rumanus) platycephalus (see Platelephas platycephalus) platycephalus angustidens, 1319, 1339, 1340, 1348, 1354, 1357, 1382, 1414 platyrhynchus, 1400. See Hesperoloxodon antiquus platyrhynchus platytaphrus, 1123, 1136, 1388 primexvus, 1123, 1136, 1147, 1384, 1387 primigenius (see Mammonteus primigenius) primigenius a lames écartées, 1042, 1064 primigenius alaskensis (see Mammonteus primigenius alaskensis) primigenius americanus, 1399. See Mammonteus primigenius americanus (primigenius bexw. meridionalis) trogontherti, 1058, 1398 primigenius Blum. var. hydruntinus (see Mammonteus primigenius hydruntinus) primigenius Blum. var. trogontherti, 1049, 1057, 1058, 11387, 1138, 1154, 1404, 1407. See Mammonteus primigenius aslensis primigenius Blumenbach var. n. pachyganalis, 1412 primigenius columbi, 1004, 1043, 1399 primigenius comune, 1123, 1137, 1397 primigenius Fraasi (see Mammonteus primigenius fraast) primigenius germanicus, 1123, 1391 (primigenius) Leith-Adamsi. See Mammonteus(?) primigenius leith-adamsi (primigenius) Leith-Adamsi Pohlig var. minor, 1150, 1399 primigenius Matsumoto, 1411, 1416, 1420, 1625 primigenius meridionalis, 1391 primigenius mut. astensis (see Mammonteus primigenius astensis) primigenius Sibiricus, 1123, 1137, 1178, 1391 (primigenius) trogontherii, 1398, 1399 primordialis, 1123, 1136, 1385, 1387, 1624 primus, 1403 priscus, 11, 908, 1181, 1182, 1187, 1191, 1197, 1198, 1217, 1219, 1263, 1271, 1378, 1385, 1392. See Loxodonta priscus-melitensis group pygmaeus, 1123, 1136, 1146, 1387 roosevelti, 997, 1047, 1048, 1065, 1067, 1068, 1083, 1084, 1086, 1087, 1088, 1095-1097, 1099, 1407 rupertianus (see Mastodon americanus rupertianus) scotti (see Archidiskodon imperator scottz) socotrus, 794, 1323, 1384 stenotoechus, 1123, 1136, 1388 sumatranus (see Elephas indicus sumatranus) texianus, 1073, 1394. See Parelephas columbi trogontherii (see Parelephas trogontherit) trogontherii-intermedius phylum, 1049 washingtonti (see Parelephas washingtonit) wiisti (see Parelephas wiistt) Elephasidex, 26, 912, 1368 Eldoran, 1510 Elgg, 115, 134, 192, 193, 199-201, 211-216, 219, 736, 1464, Pl. 11 Elias, Maxim K., 713, 1507 Else, H., 944 INDEX Elsinor Quadrangle, 497 émail epais or pachyganal, 1270-1272 émail mince or endioganal, 1270-1272 Embrithopoda, 1425 Emil Mosonyi Collection (see Mosonyi) Emmenodon, 853, 1375, 1378 Emmons, Ebenezer, 733, 774 emmonst (see Ocalientinus emmonst) enamel, 1608 enamel length (see ganometry) endecaloph-lophid, 1545 Engelhalde, 115 Engelswies, 1461, 1464. See also Trilophodon engelswiesensis engelsuresensis (see Trilophodon engelswiesensis) England. See Easton, English Crag, Nodule Bed, Norwich Crag, Piltdown, Postwick (Norfolk), Ramsey (Essex), Southwold, Suffolk, Whitling- ham English Crag, 620, 1048, 1049 ennealoph-lophid, 1545 Ensenada (see La Ensenada) Ensenadense, 520, 551, 581, 1517, 1518 Entelodon, 386 entoconelets, 141, 199, 277, 687, Pl. 111 entotrefoils (see trefoils) Entre Ridés, Argentina, 593 Eoanthropus dawsoni, 964-966, 968 Eocene, 1421-1426, 1450, 1467, 1555 Eosiren, 39, 42, 53, 1424 Eotherium, 42, 47 Eppelsheim, 84, 85, 88, 89, 114, 281, 735, 739, 1269, 1387, 1388, 1468 Equide, 1, 46, 161, 217, 294, 298, 305, 308, 318, 399, 400, 418, 497, 498, 510, 562, 573, 601, 633, 666, 679, 1518. See also Hquus Equus, 386, 399, 725, 1433, 1444, 1470, 1478, 1480, 1504, 1505, 1510, 1511, 1518, 1519, Pl. vur. See also Plesippus, Pliocene-Pleistocene boundary alaskex, 1134, 1135, 1161 Beds, 372, 373, 386, 725, 998 caballus africanus, 1313 caballus fossilis, 1155 caballus nordicus, 1313 capensis, 984 complicatus, 171, 400 crenidens, 1082 curvidens, 1082 excelsus-E. imperator Zone of Osborn, 998 [fraternus] leidyi, 1077 giganteus, 1515 hemionus, 1486 mosbachensis, 1044 -Mylodon Zone, 1511 namadicus, 1449 onager, 448 robustus, 1476 sanmeniensis, 1486 scotti, 422 sivalensis, 648, 1340 stenonis, 964, 968, 971, 1044, 1155, 1431, 1476 sussenbornensis, 1044, 1065 Erfurt, 1045 Ertemte fauna, 1482 Erzerum, 1060, 1062, 1394 Escanecrabe, 797 Eschscholtz Bay, 1127, 1134, 1135, 1159 Escuela de Ingenieros, Mexico, 1013, 1081 Escuela de Minas, Mexico, 555, 745, 755 Eselsberg, 115 Espafiola Valley, 1492 Espélugues, Lourdes, 1169 Esselborn, 192, 223, 251, 281, 736, 737, Pl. m1 1643 esselebornensis see Trilophodon esselbornensis) Essex, England, 635 Etchegoin formation, 161, 162, 497, 562, 1503 Etchd, 908, 1187, 1298, 1489 Etheridge, Robert, Jr., 774, 781, 1378 Ethiopia, 34, 485, 1423, 1430, 1432 Ethnology Museum, University of the Witwatersrand, 1284 Eubelodon, 11, 128, 515, 531, 538, 540, 541, 601, 611, 615, 685, 686, 688, 691, 1379, 1525, 1527, 1549, 1566, 1568, Pls. x, x1 morrilli, 11, 118, 294, 317, 318, 531, 536-539, 541, 542, 545, 547, 600, 601, 602, 603, 604, 606-611, 615, 733, 741, 751, 752, 758, 1404, 1566, 1568, 1601, 1604 Eucladoceros, 1485 Euelephantine, 27, 33, 935, 936, 1117, 1370 Euelephas, 10, 27, 127, 904, 905, 909, 927, 936, 1072, 1117, 1175, 1177, 1322, 1370, 1375 antiquus, 1394. See Hesperoloxodon antiquus columbi, 1394. See Parelephas columbi indicus (see Elephas indicus) jacksoni (see Parelephas jacksonz) (Parelephas) protomammonteus. See Palxoloxodon protomammonteus primigenius (see Mammonteus primigentus) primigenius astensis (see Mammonteus primigenius astensis) primigenius sibiricus, 905, 907 protomammonteus, 1409. See Palzoloxodon protomammonteus trogontherizi, 1408. See Parelephas trogontheri trogontherti-primigenius group, 906 euhypodon (see Blickotherium euhypodon) Eurasia, 35, 1430, 1466, 1467, 1475 Europe, 822, 1269, 1432, 1456, 1457, 1461, 1465, 1473. See also Vindobonian Eusmilus whitfordi, 601 Evans, George Henry, 4, 774 Everett, A. H., 851, 1455 evolution, 139, 141, 277, 1315, 13873 exilis (see Archidiskodon exilis) exoletus (see Tetralophodon exoletus) expeditions and explorations. See Alaska College-Frick American Museum; Brown, Barnum; Borissiak; Castelnau; Central Asiatic; Cooper, Clive Forster; Fayfm; Field Museum; Frick, Childs; George Vanderbilt African; Gidley, J. W.; Hayden, F. V.; Holmes, Walter W.; Khomenko; Olduvai; Selenka-Blanckenhorn; Sheep Creek; Snake Creek; University of Nebraska extinction, 113, 1162, 1169, 1221, 1476, 1511, 1512 Fabiani, Ramiro, 4, 774, 1183 Fabri,—, 167 Fairbanks, Alaska, 137, 176, 736, 1137, 1150, 1160, 1417, 1603 Fairbanks Exploration Company, 1134 Fairbury, Nebraska, 1003 Fairfield Creek, Nebraska, 317, 318 Fairmount, Indiana, 1047, 1084 Fakenham, 963 Falconer, Hugh, 2, 3, 82, 85, 91, 249, 267, 348, 356, 474, 475, 480, 505, 621, 625, 635, 643, 647, 685, 774, 829, 830, 853, 856-858, 862-864, 866-871, 873, 927, 930, 931, 933, 942, 948-950, 1000, 1060, 1117, 1174, 1177, 1181, 1198, 1219, 1324, 1325, 1839, 1343, 1348, 1625 falconeri, Loxodonta (Pilgrimia), 32. See also Anancus falconeri, Elephas columbi var. falconeri, Palxoloxodon falconeri, Pentalophodon falconert, Rhynchotherium falconeri, Tetralophodon (Lydekkeria) falconert Falkenbach, Otto, 1093, 1094 Falkenham, 981 Faluns de Pontlevoy, 201, 212, 213, 223 families, 19, 26, 83, 689, 1525 Farladani, 961, 968 Farwell, Nebraska, 1012 Fatehjang stage, 449, 1441, 1442 faunal and geological horizons, 278, 385, 386, 451, 1422, 1439; Africa, 34; Alangasi, 573; Alaska, 1134, 1135, 1161; Asti, 637; Bumiaju, 365; “cold fauna,” 1474, 1511, 1513; correlation Africa-Europe, 1430; 1644 OSBORN: THE faunal and geological horizons—continued Djebel Kouif, 105; European succession of faunas, 1513; Freedom, Nebraska, 335; Mediterranean Islands, 1269; Meisen Series, 1420; Melbourne, 400; Minas Geraes, 579; Moghara, 485; North Ameri- ean, 1511, 1513; Norwich Crag, 620, 633, 634, 963; Ulloma, 551; Vaal River gravels, 994; wald, steppe, plains fauna, 1044, 1045; “warm fauna,’”’ 1474, 1475, 1513. See also Ainsworth; Alachua; Anastasia; Araucana; Arikaree; Barstow; Bartonian; Blanco; Bonaerense; Bone Valley; Boulder Conglomerate; Bristow; Brown’s Park; Brule; Brunswick Canal; Bugti Hills; Burdigalian; Chanac- Etchegoin; Chinji; Chokrak Beds; Choptank horizon; Christmas Quarry; Chungkingfoo; Clarendon; Clarendonian; Deep River; Devil’s Gulch; Dhok Pathan; Duplin marls; Ensenadense; Eppel- sheim; Hquus Beds; Etchegoin; Fatehjang; faune pleiocéne récente; Fayim; Fort Niobrara; Fort Pierre; Gaj; Gelben Lehme; Glypto- thertum-Pliohippus-stmplicidens Zone; Godavari; Goodnight horizon; Hawthorn; Hay Springs, Nebraska; Hemphill; Hipparton; Hiramaki; Houldjin gravels; Hsanda Gol; Ione formation; Jilanéik Beds; Jumna; Kali Glagah; Kamlial; Karnul caves; Kendeng-schichten; Khunuk; Kotyhaza; Levantin; Loh formation; Loup Fork; Lujanense; Mas- call; Merychippus; Monterey; Mt. Eden Beds; Murree; Nerbudda; Nodule Bed; Oak Creek Beds; Ocala; Ogallala; Oppeln; Pampean Beds; Pawnee Creek; Permian-Cretaceous; Perrier; Phosphates Beds, South Carolina; Pikermi; Pinjor; Pinole-Tuff-Orinda; Platybelodon grangert Zone; Pliauchenia; Procamelus-Hipparion Zone; Protohippus; Puelche; Quinn limestone; Rattlesnake formation, Oregon; Red Clays; Red Crag; Republican River Beds; Ricardo; R6othliche Sande; Roussillon; San Pedro Valley; Sarmatian; Sheridan or Equus Beds; Siwaliks; Snake Creek; Stegomastodon primitivus; Steinheim; Tapa- suma; Tatrot; Tertiary; Tequixquiac; Thousand Creek; Ticholeptus; Tortonian; Trinil beds; Tung Gur Khara Usu; Val d’Arno; Yen- chingkou; Yorktown faune pleiocéne récente, 633 Fayrer, F. D., 1326 Faytm, xi, xvi, 35-37, 39, 47, 51-60, 64, 65, 72, 137, 238, 465, 685, 735, 739, 1406, 1423-1426, 1523, 1524 felicis (see Cordillerion oligobunis felicis, Parelephas columbi felicis) Felis, 887, 963, 1134, 1135, 1155, 1161, 1476 Felix, Johannes, 4, 475, 477, 534, 556, 557, 774, 931, 944, 994, 1088, 1127, 1131, 1142, 1198 Felixstowe, Suffolk, 963 Fenho stage, North China, 1483 Ferrar, Hartley T., 1524 Yield, Henry, 1236, 1237 Field Museum of Natural History, 15, 546, 568, 755, 1236, 1237 Fig Island, Florida, 1115 Figgins, Jesse Dade, 14, 155, 307, 1037 Filhol, Henri, 3, 473, 774, 1460 filholi (see Serridentinus filholi) Fillmore County, Nebraska, 1012 Finley, Anthony, 136 Fischer de Waldheim, Gotthelf, 5, 7, 134, 192, 250, 252, 517, 537, 775, 1123, 1136, 1146, 1625 fission, 692, 693, 701, 810-812, 1545, 1546, Pls. -1v Fitzinger, Leopold J. F. J., 3, 775 flat-tuskers, 328, 332-334, 459, 715, 717 Flerov, Constantine C., 5, 775, 1128 Fletching, 965 Flinsch, Margret, xiv, xv, xvii, 16, 18, 25, 45, 66, 79, 106, 116, 118, 149, 164, 201, 211, 287, 332, 342, 379, 380, 388, 405, 412, 413, 460, 503, 531, 535, 542, 543, 599, 610, 629, 630, 653, 666, 674, 864, 872, 883, 888, 890, 934, 947, 979, 1037, 1147, 1206, 12388, 1239, 1259, 1523, 1550, Pls. v, 1X, X, XIV-XXIII Flinz, 216, 1466 flora, 217, 508, 717, 719, 927, 1127, 1128 Florence Museum, 616, 619, 756, 961, 969-972, 974-977, 1215, 1233, 1251, 1609, 1611 Florensac, 634 florescens. See Ocalientinus (Serridentinus) florescens PROBOSCIDEA Florida, 400, 482, 966. See Bradenton; Brewster; Lakeland; Pierce; Williston, Levy County Florida Geological Survey, 756, 1609, 1620 floridanus. See Ocalientinus (Serridentinus) floridanus, Parelephas floridanus Flower, Stanley S., 1194 Flower, William Henry, 4, 775, 919, 931, 1131, 1625 Fluvio-marine Beds (see Faytim) Fluvio-marine Crag, England, 635 foetal specimens, 466, 931, 1202, 1310 Foetterle, Franz von, 3, 91, 775 Fokien, China, 817, 885 Font-de-Gaume, 1126, 1131 food, 717-719, 927, 1126. See also under each genus and species foramina, 916, 917, 920 Forest Bed, 963, 969-971, 980, 981, 1042, 1044, 1047-1050, 1056, 1059, 1064, 1065, 1067, 1138, 1140, 1154, 1155, 1187, 1217, 1220, 1222, 1232, 1234, 1290 forest elephant, 1196, 1260 Fort Collins, Colorado, 1003 Fort Hays Kansas State College, 713, 756 Fort Logan, 1493 Fort Niobrara, 318 Fort Pierre, 601 Fort Sibut, 202 (iy Fort Worth, Texas, 564 i Fortis, Giovanni Battista detto Alberto, 2, 775 Foster, J. W., 3, 775, 1069, 1070 Fourtau, René, 4, 95, 108, 251, 260, 479, 485, 775, 1405, 1426 Fourviére, 1063, 1064 Fraas, Eberhard, 4, 103, 775, 994, 1130, 1152, 1237, 1253 Fraas, Oscar Friedrich von, 3, 281, 775 fraasi (see Mammonteus primigenius fraasi) Frade, Fernando, 4, 775, 1192, 1193, 1412 Francis, Mark, 759 francisi (see Rhynchotherium francis?) Frankfort, Germany, 759, Pl. 111 Franklin, Benjamin, 166 Franklin County Mammoth, 1012, 1020, 1091-1093, 1106, 1107, 1111 Fransseni (see Loxodonta africana franssent) Franzensbad, Bohemia, 96, 99-102, 115, 116, 1167 Fraser, Donald McCoy, 1503, 1625 Frederick, Oklahoma (see Archidiskodon haroldcooki, Stegomastodon priestley?) I'reedom, Frontier County, Nebraska (see A mebelodon fricki and A. sinclairi) Iregellae, 1239 Freising, Bavaria, 201, 697, 698, Pl. ur French Congo, 1193 French Guiana (see Parelephas columbi cayennensis) Frenguelli, 520, 590 I’resno County, California (see Pliomastodon vexillarius) Treudenberg, Wilhelm, 4, 14, 19, 587, 553, 555-559, 775, 942, 948, 965, 1014— 1017, 1044, 1046, 1047, 1056, 1080, 1082, 1466, 1506, 1625 Frick, Childs, xvi, 4, 14, 175, 176, 251, 256, 257, 312, 319-327, 335, 374, 375, 384, 385, 387, 407, 419, 483, 435, 440, 441, 443, 446, 447, 473, 479, 496, 500, 501, 505-510, 516, 537, 560, 561, 602, 606-610, 726, 728, 748-753, 776, 1012, 1184, 1135, 1137, 1159-1161, 1406, 1416, 1492, 1493, 1625. See also Mt. Eden Beds fricki (see Amebelodon fricki, Tetralophodon fricki, Trilophodon frickt) Fritsch, Karl Wilhelm Georg von, 3, 620, 776 Frohnstetten, 91 Frontier County, Nebraska (see Amebelodon fricki and A. sinclairt) frozen mammoth of Siberia, 1162, 1165 Fuchs, Theodor, 3, 776 Tiinfkirchen, 91 Firth, Bavaria, 86 Fuhlrott, 1137, 1150 Fukuoka, 1609, 1611 Fulda, Elizabeth Rungius, 305 Fulda, Germany (see Turicius virgatidens) Fullerton formation, 725, Pl. vir INDEX Fulljames, Captain, 642 Fulton, Indiana, 183, 184, 187-189 Furlong, Eustace L., 4, 14, 554, 776, 800, 943, 1082 Furnas County, Nebraska, 377, 739, 1012 Gadari, 844 Gage County, Nebraska, 1012 Giiceana, Rumania, 85, 95, 114, 735 Gaillard, Claude, 14, 257, 259, 273, 1463, 1625 gaillardi (see Trilophodon angustidens gaillard?) Gaj, 78, 266, 267, 270-272, 275, 449, 1440-1442 gajense (see Deinotherium indicum var. gajense) Gamphotherium (see Gomphotherium) Gandoi, Bugti Hills, 269, 275, 279, 1440, 1441 ganesa (see Stegodon ganesa) Ganges, 648, 650, 1312 ganometry, xiv, 967, 968, 1109, 1547, 1580 Gard, France, 977, 978 Garza County, Texas, 1007 Gastaldi, Bartolomeo, 3, 776 Gaudin, Charles, 928 Gaudry, Albert, 3, 19, 91, 192, 193, 212, 218, 232, 250, 255, 263, 776, 844,953, 1042, 1394 gaujaci (see Trilophodon angustidens gaujact) Gavanosy, Russia, 625, 639 Gay, Claudio, 3, 582, 776 gazelles, 1467 Gazin, C. Lewis, 1504, 1505, 1509, 1625 Geierach, Austria, 262 Geikie, Archibald, 1155 Geinitz, Hans Bruno, 3, 90, 776 Gelasmodon, 272 Gelben Lehme, 213, 221, 736, Pl. 111 geneplasmic evolution, 1581, 1582 genera, 1526, 1527. See also genus Geneva (see Muséum d’Histoire Naturelle, Geneva) Gentohyus, 1424 Genoa, 475, 477, 478, 481 Genomastodon, 249, 251, 288-291, 293, 298, 299, 1380, 1557 genotype, 631 genus, 5-11, 20, 29, 33, 252, 525 Geoffroy Saint-Hilaire, Etienne, 2, 89, 776, 1123, 1192, 1325, 1335 Geoffroy Saint-Hilaire, Isidore, 3, 776 geologic horizons (see faunal and geologic horizons) Geological Institute of the City of Mexico (see Museo Instituto Nacional geologico) Institute of the Imperial University (see Tokyo) Institute of the Kyoto Imperial University, 1291, 1300 Institute of Turin, 209 Museum of Bonn, 1017 Society of London, 636, 651, 756, 1353-1355, 1609, 1615 Survey of Bandoeng, Java, 756 Survey of Caleutta, 15 Survey of China, 1480 Survey of Dutch East Indies, 14 Survey of Egypt, xvi, 1524 Survey of India, 731 Survey of Mexico (see Museo Instituto Nacional geologico) Geologisches-Paliontologisches Institut und Museum der Universitat, Berlin, 14, 188, 1275, 1610 Georg August Universitit, 14 George Vanderbilt African Expedition, 1196 Georgensgmiind, 115, 1461 Georgia, 996, 1047 germanicus (see Hesperoloxodon antiquus germanicus) Gerold, Mary M., xvii Gers, 108, 250, 256, 385, 473, 738 Gervais, Frangois Louis Paul, 3, 91, 232, 477, 481, 517, 518, 521, 544, 550, 554, 620, 625, 632, 634, 776 1645 Gez, Juan W., 4, 537, 582, 776, 1405 Gibbes, Robert W., 3, 777 Gidley, James Williams, 4, 14, 15, 400, 408, 410, 418, 422, 424, 565, 619, 623, 624, 667, 668, 673, 675, 677-682, 777, 996, 1005, 1010, 1079, 1134, 1501, 1503, 1504 Giebel, Christof Gottfried Andreas, 91, 777, 1625 Giels, Colonel, 1061 Gift, Dora E., 1084, 1095 gigantarvernensis (see Anancus gigantarverensis) giganteum (see Deinotherium giganteum, Mastodon giganteum) giganteus (see Elephas giganteus, Leptodon giganteus, Mastodon giganteus, Trilophodon giganteus) gigantissimum (see Deinotherium gigantissimum) gigantorostris (see Tetralophodon gigantorostris) Gila formation, 679 Gilchrist County, Florida, 400 Gill, Theodore Nicholas, 3, 777 Gilmore, Charles Whitney, 4, 14, 777, 1134, 1145 giraffids, 1467 Girard, Charles, 3, 30, 91, 777, 912 Glacial Age (see Glaciations, Interglaciations, also Quaternary) Glaciations, 725, 971, 1044, 1057, 1065, 1083, 1084, 1094, 1097, 1101, 1131, 1133, 1135, 1138, 1140, 1149, 1155-1157, 1169, 1271, 1472-1476, 1510, 1512, 1516, 1588, Pls. vir, xxtv Gleiwitz, 1044 Glendive, Montana, 1088 Glenwood Springs, Colorado, 1003 Glocker, Ernst Friedrich von, 3, 91, 777 Gloger, Constantin Wilhelm Lambert, 3, 7, 10, 777, 1374 Glonn, Bavaria, 698 Glorieux, sabliére Cassegrain, 693 Glyptodonts, 386, 399, 400, 579, 666, 679, 680, 1082 Glyptotherium-Pliohippus simplicidens Zone, 673 Gmelin, Johann Fridrich, 2, 777 Gmiind, Bavaria, 84, 109, 115, 735, 1464 Gnathabelodon, 11, 27, 686, 690, 705, 706, 710, 711, 739, 13882, 1527, 1557 thorpei, 686, 706, 707, 711, 712, 718, 714, 739, 756, 1382, 1418, 1507 Gnathabelodontine, 27, 31, 119, 225, 689, 690, 710, 1371, 1525, 1557 Gnathalodontine (see Gnathabelodontine) Gobi, 376, 444, 461, 464, 471, 666, 685 gobiensis (see Serridentinus gobiensis) Godavari, 448, 449, 874, 1211, 1213, 1249, 1447, 1449 Godbey, Allen H., 14, 1207 Godman, John D., 3, 777 Godmani (see Tetracaulodon Godmant) Goémor, Hungary, 638 Goppert, Heinrich Robert, 3, 777 Goriach, 115, 1461, 1462 Gottingen (see Georg August Universitat and University of Gottingen) Gokayama, Japan, 1188, 1298 Gola Forest, Sierra Leone, 761 Gold Stream, 1135, 1161 Goldfuss, Georg August, 3, 777, 1187, 1198, 1385, 1392, 1394 Gomphotheriide, 542, 587, 601, 768, 1368, 1369, 1525, 1526 Gomphotheriine, 590, 731, 738, 1370 Gomphotherium, 10, 11, 481, 560, 1373, 1374, 1525 conodon, 1403 elegans (see Tetralophodon elegans) emmonst (see Ocalientinus emmonst) floridanus. See Ocalientinus (Serridentinus) floridanus gratum (see Cordillerion gratum) priestleyi (see Stegomastodon priestleyt) serpentirivale, 251, 1415 Gong-Gong, South Africa, 943, 986, 993, 1188, 1278, 1415, 1418 Gonnesa, Sardinia, 1187 Goodnight horizon, 399, 422, 1501 Goodwin, George G., 777, 1599, 1625 Gordon Creek, Nebraska, 318 Goshen, Wyoming, 1012 1646 OSBORN: Gotha, 1, 1118, 1122, 1181, 1236,.1365, 1609, 1611 Gozo Island, 1149 gracile (see Meeritherium gracile) Gracios, Honduras, 479, 508, 509 Graells, D. Mariano de la Paz, 4, 777, 1187, 1231, 1400, 1401 Grand Ducal Museum, Florence, 969 Grand Island, 725, 946, Pl. v1 Grande Miamis, 136 Grandian, 1510 grandincisivum (see Tetralophodon grandincisivus) grandis (see Mastodon grandis) Granger, Walter, 4, 13, 14, 39, 328, 332, 333, 355, 385, 397, 458, 468, 695, 777, 817, 859, 876, 1452, 1481, 1524 grangeri (see Amebelodon grangeri, Mastodon grangeri, Platybelodon grangeri, Stegodon orientalis grangert) Grant County, Indiana, 1047, 1084 Grant, Madison, 14 Grant, [R.] E., 3, 777, 1390 Grass Roots (Snake Creek), 427 Grassi-Museum, Leipzig, 1130 gratum (see Cordillerion gratum) graviportal, 1581 Gravitelli, 1468 Gray, Asa, 3, 778 Gray, John Edward, 3, 32, 778, 912, 1174, 1191 Gray’s Thurrock, 1050, 1182, 1187, 1217, 1219, 1226 Grayson, Nebraska, 1003 Great Bone Lick (see Big Bone Lick) Greater St. Louis Museum, 14 Grebeniki, 1468 Greece, 1147. See also Pikermi, Samos Green Collection, 1220, 1234 Greensburgh, Maryland, 250, 285, 286, 738 gregorii. See Trilophudon (Tatabelodon) gregorii Gregory County, South Dakota, 304, 738 Gregory, William King, xvi, 4, 14, 20, 39, 40, 102, 778, 857, 911, 915-926, 1100, 1200, 1247 Grenoble, France, 86 Grewingk, Constantin Caspar Andreas, 3, 778 griqua (see Metarchidiskodon griqua) Griqualand West, 943, 994 Griquatherium cingulatum, 944, 994 Grive-Saint-Alban, Isére, 85, 115, 1395, 1443, 1462, 1463 Guadalajara, 999, 1001, 1005 Guatemala (see Chinautla) guatemalensis (see Serridentinus quatemalensis) Guayaquil, Ecuador, 573 Guebar-Rechim, Algeria, 1184 Giimbel, K. W. von, 1466, 1625 Giinz, 725, 1056, 1430, 1457, 1473, Pl. vin Guettard, Jean Etienne, 2, 165, 217, 778 Guiana, French, 527, 1521 Gumbotill, 725 “Gunda,” 1093, 1094, 1337, 1598 Gunduck River, 852 Gunn, John, 3, 778, 1187, 1396 gunnii (see Elephas gunnir) Gunter, Herman, 14, 415, 1107 Guntersdorf, Austria, 262, 395 Gunzberg, 115, 1464 Gur Tung Khara Usu, 466, 1477 Hackettstown, New Jersey, 185, 186 Haddon, Albion A., xvii Hadsu District, 907, 908 Hagenbeck, 1193 Hagerman, Idaho, 1503-1505 Halbinsel Krim, 262 Halicore, 42 THE PROBOSCIDEA Halifax County, North Carolina, 733 Halitherium, 42, 91 Hall, James, 3, 778 Hall, William, 1019 Halle, 1050 Halle Museum, 1235, 1609, 1611 Halter, C. R., 14, 285 Hamilton, Canada, 1069 Hanareyama, 902 Hanchow, 898 Hanekom, C. J., 1279 hanekomi (see Palzoloxodon hanekomi) Hangman’s Creek, Oregon, 180 Hannibal, 1197 Hannosura, 908 Happisborough, 1059, 1222, 1226 Harlan County, Nebraska, 251, 338, 349, 351, 372, 385, 1012 Harlan, Richard, 286, 778, 1071, 1077 haroldcooki (see Archidiskodon haroldcookt) Harpagmothertum, 517, 775, 1364, 1372, 1380 canadense, 136, 165, 775, 1372, 1384, 1408 Harpagonotherium (see Harpagmotherium) Harrison County, Iowa, 137, 172, 736, 1012 Hartilyangar, 449 Hartman, E. J., 1026 Hartnagel, Chris Andrew, 4, 778 Harz, Germany, 11238, 1136, 1141 Haslemere, 1337 Hasnot, India, 251, 279, 385, 387, 449, 451, 658, 737, 740, 741, 844, 1419 hasnotensis (see Serridentinus hasnotensis, Trilophodon hasnotensis) hasnoti (see Synconolophus hasnot?) Hastings, Nebraska, 1009 Hatcher, John Bell, 419 Hatt, Robert Torrens, 14, 1228 Haug, Emile, 3, 103, 117, 485, 778, 1049, 1436, 1437, 1473 Haughton, Sidney Henry, 4, 778, 943, 984, 1184, 1187, 1188, 1282, 1288, 1439 Hauser, Franz, 3, 91, 778 Haute-Garonne, 93, 108, 115, 192, 199, 201, 206, 736 Haute-Loire, 208 Hawthorn formation, 400 Hay, Oliver Perry, 4, 137, 349, 372-374, 479, 501, 537, 562, 564, 622-624, 671, 684, 733, 778, 942, 1002-1005, 1039, 1077, 1083, 1084, 1095, 1138, 1134, 1142, 1364, 1371, 1374, 1511, 1512, 1625 Hay Springs, Nebraska, 1010, 1012, 1093 Hayasaka, Ichiro, 4, 779 Hayasi-zaki, Japan, 1420 Hayden, Ferdinand Vandeveer, 3, 403, 669-671, 779, 998 Hayes, Seth, 4, 779 hayt (see Archidiskodon hayi) Hays, Isaac, 3, 142, 192, 193, 207, 669, 780, 1388 Hays, Kansas, 713 Haysti (see Tetracaulodon Haysit) Hazard, Frank 489, 490 Hazen, Guy E., 507 Hebron, Nebraska, 1003 Heck, Ludwig, 4, 780, 1193 Hedin, Sven, 718 Heian-nando, Korea, 1489 Heidelberg, 1050 heights of Proboscideans, 1604, 1605 Heilprin, Angelo, 29, 780 Heinthurm, 1050 Helarctos namadicus, 1449 Helena, Montana, 1003 Helladotherium, 1467 Heller, Edumund, 4, 29, 780, 1175, 1192, 1193 Helvetian, 107, 115, 201, 261, 384, 395, 742, 902, 903, 1441, 1443, 1460 Hemicyon, 380, 1461, 1467 INDEX Hemimastodon, 10, 269, 272, 477, 902, 1379, 1403, 1556 annectens (see Serridentinus annectens) crepuscult, 271, 272, 457, 756, 903, 1402, 1441 Hemphill Beds, 399, 1509 Hemphillian, 1502, 1503 Hensel, Reinhold Friedrich, 92, 780 heptaloph-lophid, 1545 Herault, France, 93, 625, 634 Herbolzheim, 283, 624, 625, 740 Hermann, Adam, 1093 Hernando, Florida, 400 Herndon County, Kansas, 1012 Heroditus, 1209 “Herr B.,” 761, 1388 Herz, Otto, 4, 780, 1127, 1131 Hesiod, 1164, 1209, 1308 Hesperoloxodon, 12, 13, 32, 33, 913, 914, 1173, 1178, 1180, 1186, 1191, 1212, 1217, 1221, 1228, 1247, 1303, 1381, 1411, 1503, 1527, 1549, 1589, 1598, 15955 Plex antiquus, 32, 633, 634, 905, 910, 928, 932, 936, 963, 969, 971, 972, 975, 981, 1023, 1042, 1044, 1045, 1057, 1058, 1095, 1118, 1119, 1121, 1122, 1131, 1138, 1141, 1149, 1153, 1155, 1165, 1174, 1175, 1177, 1178, 1180-1184, 1186-1188, 1191, 1192, 1202, 1206, 1209, 1210, 1214-1230, 1232-1235, 1243-1245, 1247, 1249-1252, 1258, 1261, 1262, 1268-1273, 1275-1277, 1290, 1295, 1365-1367, 1377, 1378, 1392, 1394, 1397, 1400, 1416, 1420, 1475, 1476, 1538, 1542, 1592-1595, 1603, 1605, 1610, 1611, 1613, 1614, 1618, Pls. xxim, xxtv. See also Hlephas (Loxod.) priscus, Elephas priscus, and Palxoloxodon antiquus (andrews??) antiquus ausonius, 634, 905, 932, 1059, 1060, 1172, 1178, 1184, 1187, 1208, 1210, 1214, 1215, 1221, 1222, 1232, 1233, 1271, 1290, 1476, 1543, 1611 antiquus germanicus, 1180, 1184, 1187, 1188, 1206, 1210, 1217, 1221, 1222, 1233-1238, 1244, 1251, 1253-1256, 1271, 1276, 1277, 1294, 1408, 1476, 1541, 1603, 1605, 1610, 1611, 1620, 1621 antiquus italicus, 12, 969, 1180, 1188, 1206, 1208, 1210, 1211, 1217, 1221, 1234, 1236, 1238-1252, 1255, 1271, 1381, 1416, 1476, 1542, 1603, 1617, 1620 antiquus nanus, 1187, 1230, 1397, 1476, 1543 antiquus platyrhynchus, 11838, 1187, 1206, 1208, 1230, 1231, 1232, 1476, 1541, 1605 Hesse, Curtis J., 780, 1509, 1625 Hesse Darmstadt, 86 Heterodactylus (see Elephas indicus Heterodactylus) Heurn, F. C. von, 5, 780 hexabunodont, 78, 132, 139, 145, 1544, 1545 hexaloph-lophid, 1545 hickst. See Amebelodon (Trilophodon) hicksv Hidalgo, Valley of Amajaque, Mexico, 537, 555, 740, 1082. See also Vera Cruz Hidvég, Hungary, 210 Higashi-Higasa Bed, 1301 Higashi-Kanamachi, 906 Higashi-Tonami District, 908, 1188, 1298 Hilber, V., 780, 1404, 1625 Hill, Russell S., 369 Hills, Richard C., 155 Hillsboro, New Brunswick, 1575 Hilzheimer, Max, 4, 780 Himalayas, 642, 648, 653, 852, 940 Hingenau, Otto von, 3, 91, 780 Hipparion, 274, 294, 298, 318, 324, 363, 364, 380, 386, 399, 400, 418, 444, 451, 458, 497, 498, 508, 510, 622, 633, 643, 661, 663-665, 671, 678, 704, 740, 1268, 1431, 1433, 1444, 1445, 1447, 1457, 1465, 1466, 1478, 1480, 1481, 1483, 1484, 1492, 1497, 1499, 1502, 1506 punjabiense Quarry (see Hipparion) ?Hipparion Zone, 1503 Hippidion, 551, 1518, 1521 Hippolyte-Boussac, P., 29, 780 Hippopotamus, 36, 47, 48, 86, 166, 167, 887, 944, 964, 1056, 1240, 1431, 1433, 1434, 1436, 1449 Hiramaki formation, 457, 742, 902 1400, 1647 Hira-mura, 908, 1188, 1298 hirsutus (see Elephas indicus hirsutus) Hishiike, 907, 908 Historie Bluff (see Eschscholtz Bay) Hitachi Province, 905, 907, 908 Hitchcock County, Nebraska, 251, 384, 401, 489, 737, 742 Ho Shun Hsien, China, 699 Hobley, C. W., 103, 104 hobleyi (see Deinotheriwm hobley?) Hodgson, Bryan Houghton, 3, 780, 1313, 1398 Hodgson, William Brown, 3, 780, 1077 Hodkinson,—, 944, 1285 Hoernes, Moriz, 3, 92, 780 Hoffmann, R. W., 1122 Hofmann, Adolf, 4, 780 Hofmuseum in Wien, 1400 Hog Creek, McLennan County, Texas, 373 Hokkaido, 907, 908, 1299, 1305, 1333, 1489 Holdrege formation, 725, Pl. vit Hollaender, Ludwig, 93, 780 Holloman, A. H., 684, 1029 Holmes, Walter W., 14, 400, 1076, 1106, 1107, 1609, 1618, 1619 Holmesville, Nebraska, 1003 holotype, 9 Homer, 1164, 1209, 1307, 1308 Homo heidelbergensis, 1044, 1056 Homo kanamensis, 85, 105 homceotype, 218 homonym, 1416 homotaphrus (see Elephas homotaphrus) Honan, China, 384, 458, 853 Honduras, 480, 482, 484, 508, 527, 532, 535, 1516. See also Aybelodon hondur- ensis and Blickotherium blicki hondurensis (see Aybelodon hondurensis) Honshv, 1305, 1489, 1490 Hooker County, Nebraska, 998, 999, 1003 Hoopston, Illinois, 180 Hopkins, Barbara, xvi Hopwood, Arthur Tindell, xvi, 4, 14, 117, 262, 267, 272, 349, 353, 356, 513, 624, 625, 650, 686, 698, 699, 702, 704, 720, 731, 732, 781, 807, 816, 853, 894, 898, 899, 965, 966, 995, 1049, 1059, 1062, 1117, 1119, 1164, 1363, 1365, 1366, 1374, 1418, 1427, 1429, 1432-1434, 1436, 1473, 1476, 1626, 1627 hopwoodi (see Deinotherium hopwoodt) Hornaday, William Temple, 4, 14, 29, 781, 1093, 1094, 1193, 1259 Horner, William E., 3, 781, 1389 horse, association with man, 1 Horsethief Canyon, Nebraska, 607 Hoshangtun Cave, Yunnan, 1453, 1487 Hosono, 906 Hot Spring, 1418 Hot Springs, New Mexico, 137, 175, 736 Houldjin gravels, 398, 466 Hovey, Edmund Otis, 781 Howard County Mammoth, 1009, 1012 Howorth, Henry Hoyle, 3, 781, 1125, 1169 Hsanda Gol, Mongolia, 396, 1477 Hsien Shui Ho, 732 Hsien T’ien P’o, 732 Hsuan Hu Hsien, 699 Huallago River, Peru, 554 Huang Tui Kou, 699 Hubbard, T. R., 1332 Hudson River, 135 human association with Proboscideans, xiv, 1. See also artifacts, Aurignacian, Cuvieronius postremus, Cuvieronius superbus, Paleolithie art and the mammoth humboldianus (see Mastodon humboldtianus) Humboldt, Alexander von, 549, 568, 576, 577 Humboldt County, Nevada, 154, 692, 736 1648 Humboldtide, 26, 30, 119, 128, 225, 515, 575, 601, 602, 667, 687, 688, 691, 722, 741, 1368, 1369, 1525, 1526, 1566, 1568. See also Humboldtine humboldtii (see Cuvieronius humboldtii, Mastotheriwm humboldtit) Humboldtine, 31, 119, 225, 228, 515, 536, 537, 540, 542, 575, 593, 601, 602, 605, 611-615, 617, 622, 624, 625, 628, 667, 689, 691, 722, 734, 741, 1371, 1526, 1528, 1545, 1566, 1568, Pl. x humboldtius (see Mastodon humboldtius) humerus, 1251, 1276 Hunan, 853 Hung Kureh Beds, 397, 718, 1481 Hungarian National Museum (see Ungarisches Nationalmuseum) hungaricum (see Deinotherium hungaricum) Hungary, 91, 638, Pl. m. See also Ajnicské, Baltavdr, Batta-Erd, Budapest, Gomoér, Kotyhiza, Levantin, Pestszentlorincz, Rakoskeresztur, Szabadka, Tasniad, Ungarische geologische Reichsanstalt, Ungarisches Nationalmuseum Hunter, William Wilson, 29, 781 Hupeh, 853 Hurlbert sand pit, 1023 Hutchinson, G. E., 1626 Hutchinson, Henry Neville, 4, 781 Huxley, Thomas Henry, 919 Hyena, 274, 579, 704, 818, 887, 971, 1155, 1476 Hyznarctos, 508, 963, 1503 Hyznodon, 1424, 1425 Hyzxnognathus, 508, 510, 1515 hydruntinus (see Mammonteus primigenius hydruntinus) hyodon (see Mastotheriwm hyodon) Hyogo Prefecture, 1420 Hyotherium sindiense, 272 hyperlongirostrines (see Trilophodon) Hypohippus, 294, 318, 400, 488 Hypolagus edensis, 508 Hypselephas, 12, 913, 914, 1318, 1321, 1339, 1340, 1349, 1382, 1527, 1543, 1589, 1597 hysudricus, 10, 12, 448, 874, 905, 910, 936, 952, 982, 1065, 1175, 1177, 1307, 1318-1322, 1327, 1338-1357, 1359-1361, 1382, 1391, 1414, 1448, 1451, 1543, 1549, 1596-1598, 1603, 1610, 1613-1617 hypsicephaly, 919, 1552 hypsodonty, 919, 1548, 1552 Hyracoidea, 35, 36, 41, 42, 1425, 1428, 1429, 1467. See also Hyrax Hyraz, 48 Hystrizx, 887, 1425, 1451 hysudricus (see Hypselephas hysudricus) hysudrindicus (see Palzoloxodon hysudrindicus) Ichikawa, Shoichi, 893 Ictitherium, 704 “Idaho” formation, 1505 ideotype, 9, 1068, 1084 Ides, Eberhard Ysbrant, 2, 781, 1124, 1147, 1162, 1165 Ikadachi-mura, Japan, 818, 906 Tle-en-Dodon, 108, 115, 198-200, 206, 736, 1464 Ilford, 1050, 1132 Ilfov, 969, 1184, 1188, 1232, 1233, 1235 Illiger, Carolus, 2, 30, 781 Illinoian, 170, 725, 1099, 1510, Pl. vin Illinois (see Ashland, Walnut) Ilm gravels, 1045, 1050, 1245 Imantag, 583 Imbaburra, 122, 123, 516, 518, 549, 740 Imba-numa, 908 ; immigrant types as diagnostic time markers, 1431 imperator (see Archidiskodon imperator, Elephas columbi var. imperator) Imperial Institute, London, 1193, 1609, 1614 Imperial Mammoth (see Archidiskodon imperator) Imperial Museum of Uéno, 1291, 1298 incisors, 1547, 1550. See also tusks Indarctos, 704, 1467 OSBORN: THE PROBOSCIDEA Indersdorf, Bavaria, 698 indeterminate names, 10, 11. See also classification and nomenclature India, 111, 114, 115, 268, 448, 449, 480, 502, 829, 831, 1482, 1440, 1442-1447 India House, 1324, 1609, 1614 Indian elephant, 29, 927, 1311, 1834-1336, 1594, 1604, 1605. See also Hlephas indicus Indian Museum, Calcutta, 513, 745, 756, 855, 856, 1395, 1608, 1610 Indiana (see Ashley, Fulton, Jonesboro, Rochester) Indianapolis. See Parelephas (?) mississippiensis (?) indicum (see Deinotheriwm indicum) indicus (see Elephas indicus) Indigirka River, 1119, 1120, 1136 Indo-China, 1440, 1453 Indricotherium, 461, 1478 Indus River, 85, 279, 527 infrequens (see Parastegodon infrequens) Ingram, Bruce 8., 1322 Inland, Clay County, Nebraska, 1012 Inland Sea, Japan, 1188, 1299, 1489 inopinatus (see Trilophodon inopinatus) insectivores, 1425 insignis (see Stegodon insignis) Institut des Mines, St. Petersburg (Leningrad), 133, 1393, 1609, 1612 Institute of Geology, Florence, 1215, 1233 Institute of Geology and Paleontology, Japan, 1299, 1333 Instituto Geologico (see Museo Instituto Nacional geologico, Mexico) insularis (see Elephas antiquus var. insularis) Interglaciations, 170, 373, 671, 682, 725, 726, 934, 970, 971, 980, 998, 1003, 1033, 1042-1050, 1056, 1057, 1062, 1065, 1067, 1071, 1135, 1138, 1140, 1149, 1150, 1155, 1161, 1210, 1217, 1221, 1234, 1244, 1245, 12538, 1256, 1271, 1473, 1476, 1510, Pls. vim, xxiv intermedia (see Cordillerion oligobunis intermedius) intermediate molars, 812, 1547, 1548 intermedium (see Deinotherium intermedium) intermedius (see Anancus intermedius, Cordillerion oligobunis intermedius, Elephas giganteus intermedius, Mastodon intermedius, Palzomastodon intermedius, Parelephas intermedius) International Commission on Zoological Nomenclature, 5, 6, 81, 1173, 1364, 1525 International Congress of Zoology (Fifth), 13809, 1310, 1371, 1543 Tone formation, 487 Iowa, 725. See Akron; Harrison County; Missouri Valley; Mastodon pro- genius; Stegomastodon aflonix Iowa City, 1609, 1611 Tren Dabasu, 385, 387, 398, 466, 742, 1416 Tren Gobi, 718 Irondiquoit River, 1156 Irrawaddy, 114, 449, 642, 643, 737, 824, 825, 842, 844, 853, 861, 874, 1450, 1451, 1455, 1487 Trushi, 899, 900 Ise Province, Japan, 818, 906, 1489 Isére, France, 85, 115 Ishikari, 908, 1299, 1333 Ishim, Russia, 461 Ismail, 461, 625, 639 Tsodactylus (see Elephas indicus Isodactylus) Isoletta, 1239 Isone, 907, 1188, 1298 Issel, A., 1123, 1137, 1626 Isserville, Algeria, 246, 485, 739, 1426, 1428 Issoire, France, 634 italicus (see Hesperoloxodon antiquus italicus) ° Italy. See Astésan, Asti, Dusino, Olivola, Piedmont, Tossano, Val d’Arno, Villafranea (San Paolo), Villanova Itchatucknee River, Florida, 1115 Itsukaichi, 908 ivory industry, 1162-1164, 1308 Iwaki, 1489 INDEX Jabi, 844 Jack, Robert Logan, 781, 1378 Jackson County, Ohio, 1047, 1068, 1069, 1389 Jackson, J. B.S., 3, 781 jacksoni (see Parelephas jacksoni) Jacquemin, Emile, 3, 89, 781 Jager, Georg Friedrich, 3, 91, 781 Jalisco, 999, 1001 Jammu, 274, 448, 449, 659, 845, 1188 Janensch, Wilhelm (or Werner), 4, 781, 833, 885, 886 “Jap,” 1604 Japan, 115, 193, 215, 280, 457, 729, 734, 818, 853, 907, 1062, 1185, 1289, 1292 1293, 1304, 1305, 1477, 1489, 1490, 1528 Jardin de Zoologique de Lisbonne, 1193 Jardin des Plantes, 341, 548, 1200, 1312 Java, 833, 1185, 1289, 1304, 1305, 1440, 1451, 1453-1455 javanicus (see Stegodon ganesa var. javanicus) javanoganesa (see Stegodon javanoganesa) Jebel el Qatrani beds, 1423 Jeddo (see Yedo) Jefferson County, Nebraska, 1012, 1017, 1018 Jeffersoni (see Mastodon jeffersont) Jeffersonian Mammoth (see Parelephas jeffersoniz) Jjeffersonii (see Parelephas jeffersonii) Jena Museum, 1234, 1609, 1611 Jentink, F. A., 3, 781, 1329 Jentzsch, Carl Alfred, 3, 192, 208, 209, 781 Jerdon, Thomas Claverhill, 29, 782 Jesup, Morris K., 1095, 1523 Jesup, Mrs. Morris K., 420 Jhelum River, 853 Jilanéik Beds, 278, 461, 738, 1478, 1479 Jobert, Antoine C. G., 3, 618, 625, 632, 782 Jockgrim, 1044, 1056 Johannesburg, South Africa, 14 John Day region, 155 Johnson, Frank Walker, 707, 1496, 1626 Johnson, Martin, xvii, 1170, 1189 Johnston, Henry Hamilton, 14, 29, 103, 782, 1195, 1626 Johnstown, Nebraska, 318 Jol ou Julia easaroea, 1183 Joleaud, Léonce, 4, 117, 261, 782 jolensis (see Palxoloxodon jolensis) Jonesboro, Indiana, 1047, 1084 Jong, J. J. de, 1282 Jongh, H. Munniks de, 183 jorakt (see Megabelodon jorakz) Joubert,— de, 2, 86, 167, 782 Jourdan, Claude, 3, 85, 192, 618, 632, 782, 1040, 1041, 1062, 1394, 1397 jubatus (see Elephas jubatus) Juliette, Florida, 400, 418 “Jumbo,” 179, 931, 1018, 1021, 1022, 1095, 1131, 1176, 1193, 1194, 1199, 1200, 1225, 1227, 1230, 1249, 1402, 1591, 1603, 1605 Jumna, India, 448, 648, 650 junior. See Palxoloxodon (Archidiskodon?) tokunagat mut. junior Jura, Bernese, 91 Jussieu, Antoine de, 84, 86, 262 K. K. Geologische Reichsanstalt, Vienna, 760 K. K. Naturhistorisches Museum, Vienna, 760, 1620 Kabylie, 65, 246, 251, 739, 1400 Kadayan, 700, 1455 Kadonosawa, Japan, 902 Kapfnach, 115, 201, 216, 219, 1464 Kaga, 834, 892, 893, 1489 Kage District, Japan, 818, 906 Kairouan, 1428 Kaisen, Peter, 1134, 1161 Kaiso Bone-beds, 945, 983, 995, 1273, 1287, 1432, 1435, 1436 1649 Kakio, 897, 1418 Kalachilta Hills, India, 449 Kalahari Desert, 968 Kalgan-Urga Trail, 466, 1477 Kali Glagah, Java, 365, 896, 1454 Kalka, Siwaliks, 950, 952, 954, 955, 1339, 1340, 1345, 1347 Kama River, 133 Kamchatka, 1304, 1305 Kamenez-Podolsk, 694 Kamenskii (see Elephas Kamenskit) Kaminog6-mura, Japan, 384, 457 Kamlial, 266-270, 275, 279, 448, 449, 642, 1441-1445, 1447, 1448 Kanagawa Prefecture, 897, 907, 1303, 1333, 1334, 1418 Kanam, Africa, 105 kanamensis (see Homo kanamensis) Kandy, 1306 Kangra, 274, 448, 449, 845 Kani District, Japan, 384, 457 Kankyo-d6, Japan, 1420 Kansan, 725, 726, 946, 1510, Pl. vir Kansas, 251. See also Buffalo, Edson formation, Hays, Long Island (Phillips County), Loup Fork horizon, McPherson, Ness County, Ogallah, Pendennis, Republican River Beds, Sappa Creek, University of Kansas Kansu, North China, 458, 702, 732, 738, 742, 1418 Kaoko District, 1193, 1409 Karaturgaj, 461 Karnul caves, 448, 449 Karriger, H. S., 1019 Karungu, 85, 104, 105, 115, 735, 1428, 1429, 1432 Kasauli, 449 Kashiwazaki, Japan, 818, 906 Kasumiga-ura, 907, 908 Kato, T., 4, 782, 818, 1298, 1609, 1619 Kaup, Johann Jacob, 3, 30, 80-89, 349, 357, 782, 1388 Kavirondo Gulf, Africa, 85, 105 Kay, George Frederick, 14, 683, 725, 1510, 1626 Kazusa, Japan, 818, 906-908, 1047, 1188, 1297, 1298, 1300, 1420, 1489 Keikido, Korea, 1489 Kellogg, Remington, 1626 Keltschau, 91 Kendeng-schichten, 366, 833, 874, 885-888, 890, 1186, 1187, 1303, 1403 Kennedy, Ildephons, 2, 86, 782 Kent, England, 1215, 1222 Kentucky, 131, 185. See also Big Bone Lick Kenya Colony, 1189, 1193, 1429, 1432 Kerr, Robert, 2, 7, 185-137, 168, 170, 782, 1363 Kertch (Bessarabia), 262 Kerynia Hills, Cyprus, 1187, 1257, 1266 Kessingland, Suffolk, 943, 980, 1407 Kettenheim, Germany, 281 Kettleman Hills, California, 161, 162 Keyapaha, 305, 318, 1498 Khajuri, Bugti Hills, 78, 79, 1441 Khanoos, Armenia, 1060, 13894 “Khartoum,” 1194, 1239, 1250, 1336, 1604 Khartum, Sudan, 983, 1285 Khenchela, Tunisia, 485, 1428 Kherson, 1065 Kholobolehi Nor, Mongolia, 385, 387, 397, 742, 1481 Khomenko, J., 4, 461, 625, 639, 782, 1403, 1466, 1626 Khunuk formation, 397, 398, 742, 1481 Kilimanjaro, 1204 Kilpatrick Pasture, 426, 427 Kimberley, South Africa, 948, 944, 1278, 1282, 1609, 1612. See also McGregor Museum Kimitsu District, 906-908, 1188, 1297, 1298, 1300 Kimmswick, Missouri, 1602 King Adolph Frederick (see Stockholm) 1650 Kingfisher, Oklahoma, 1003 Kinlock, Oklahoma, 1003 Kirdld, Hungary, 116 Kirk, John, 1626 Kislakovsky,—, 694 Kitayama, Japan, 251, 280, 738, 905 Kittl, Ernst Anton Leopold, 3, 95, 99, 782 Kiyohawa-mura, 908 Klahn, Hans, 4, 14, 192, 195, 223, 251, 281, 283, 348, 349, 362, 625, 783, 1406, 1407, 1461 Klassen, Stephen, xvii Klipdam, South Africa, 994 Klipstein, August von, 3, 88, 783 Knight, Charles R., xvi, 39, 40, 190, 224, 259, 546, 672, 679, 1040, 1116, 1126, 1522 Knight, Wilbur Clinton, 783 knochenhaueri (see Loxodonta africana knochenhauert) Kobe, 1609, 1612 Koch, Albert C., 3, 85, 90, 165, 783, 1374, 1389 kochii (see Missourium kochii, Tetracaulodon kochit) Kollner, Karl, 93, 783 kénigii (see Deinotherium kénigit) Koenigswald, G. H. Ralph von, 5, 783, 816, 837, 894, 896, 1403, 1417, 1451, 1465, 1626 Koerner, H. A., 1494 Kohat District, Sind, 1442 Koito River, 1301 Koken, Ernst, 3, 349, 355, 783, 817, 884, 1398 Kokubo, 906, 907, 1188, 1298, 1609, 1619 Kolyma-Beresowka River, 1128, 1130 Kondratiev, Mrs., 1128 Koninklijke Akademie van Wetenschappen, Amsterdam, 14 “Koomareah,” 1314 Korea, 1489 Koru, Kenya Colony, 1429 Koshiba Bed, 1301 Kotalkund, India, 449 Kotyhiza, Hungary, 85, 116, 735 Kotzebue Sound, 1135 Kowak Clays, 1135 Kowalewsky, Wladimir, 113, 783 Kraglievich, Lucas, 590, 1517, 1626 kraglievichii (see Cordillerion andium kraglievichit) Krantz, 667 Kranzkloof farm, 1188 Krasnoie, Podolia, 135 Krejci-Graf, Karl, 1466, 1626 Krems, 961 Krober-Roubia, Algeria, 1187, 1275, 1400 Krohn, Ernst C., 14 Krottling, Germany, 281 Kriiger, Johann Friedrich, 3, 783, 1187, 1256 Kuala Pila District, Malay, 1332 Kuan Tao Kou, 721 Kuban region, North Caucasus, 385, 460, 461, 743 Kuchu (Kutsha), 718, 719 Kuft, 1524 kuhni (see Palxoloxodon kuhnt) Kuji District, 905 Kullu, Punjab, 1353 Kumbhi, Sind, 251, 266, 267, 270-272, 275, 279, 1440, 1441 Kunz, George F’., 4, 14, 783, 1204, 1308, 1595 Kushalgarh, India, 82 Kusthardt, Gustav, 14, 1314, 1330, 1331 Kuwung, 894 Kwangsi, 1453, 1487 Kwang-Tung, 853 kwantoensis (see Parastegodon kwantoensis) Kyoto Imperial University, 893, 907, 1291, 1300, 1609, 1612 OSBORN: THE PROBOSCIDEA Kyushu, 1305, 1489 Kyushu Imperial University, 893, 1609, 1611 La Chaux-de-Fonds, 91 La Croix, Frangois Antoine Alfred, 14, 1123 La Ensenada, Argentina, 520, 537, 580, 581, 592, 741 La Grive-Saint-Alban (see Grive-Saint-Alban) La Mancha, 1468 La Monte, Francesca, xvii, 1240 La Plata, Argentina, 520, 530, 595, 757, 1609, 1612 La Prietas, 1033 La Romieu, 1459 Laaerberg, Austria, 221, 961 Labuan Deli, 1329 Laeépéde, Bernard Germain Etienne de la Ville, 2, 783 Lake Albert, 945, 1190, 1193, 1239, 1402, 1435 Chad, 11938 Leopold IT, 1198 Nyasa, 983, 1432, 1486 Paradise, 1170 Qurun, 1406 Rudolf, 983, 1198, 1402, 1429, 1432, 1436 Tagua-Tagua, 1400 Victoria, 1419, 1428, 1433, 1436 Lakeland, Florida, 285 Lamarck, xiv lamarmorae (see Palxoloxodon lamarmorae) Lambe, Lawrence Morris, 4, 784, 1134 lamin, dental, 445, 715, 1571 laminar frequency, 1077, 1088 Lancaster County, Nebraska, 696 land connections, 1257, 1268, 1269, 1304, 1305, 1328, 1490 Lane County, Kansas, 1088, 1089 Lang, Charles, 1111, 1243 Lang, Herbert, 4, 14, 784, 1162, 1164, 1165, 1192 Lang, William Dixon, xvi, 14 Langley, M. A., 1103 Languedoc, 340 Lankester, Edwin Ray, 784, 919, 1401 Lapparent, A. de, 1464, 1626 Larkana, Sind, 115, 250, 266-268, 738, 1394 Larroque, José, 587, 596 Lartet, Edouard, 3, 7, 87, 108, 133, 192, 193, 206, 219, 250, 256, 263, 631, 784, 1046, 1132, 1187, 1394, 1461 Laskerew, 1065 latidens (see Mastodon angustidens var. latidens, Stegolophodon latidens) Laufer, Berthold, 29, 784 Laurent, Louis, 14, 1046 Laurillard, Charles Léopold, 3, 7, 256, 475, 477, 481, 784, 931 Lausanne, 115 Laveta, Colorado, 1003 Lay Creek, Moffat County, Colorado, 1103 Le Blane, M., 980 Le Grand Mastodonte, 5, 135, 1386, 517, 13864, 1372, 1384, 1385 Le Hon, Henri, 3, 784 Le Mar, John, 338 Le Petit Rosey (see Petit Rosey) Leakey, L. 8. B., 85, 105, 784, 1433 Léberon, 1269 lectotype, 9 Leghorn, 1061 Lehri, Punjab, 737, 834, 844, 846, 847 Leiden, 15, 891, 1330, 1609, 1612 leidii. See Ocalientinus (Serridentinus) floridanus leidii Leidy, Joseph, 3, 7, 187, 165, 168, 171, 285, 400, 416, 784, 942, 998, 1364, 1382 Leipzig, University of, 556, 557, 760, 1082 Leitha Chalk, 91 leith-adamsi. See Mammonteus(?) primigenius leith-adamst INDEX Leith- Adamsia, 960, 1381 Siwalikiensis, 942, 943, 947, 949, 953, 959, 960, 1291, 1381, 1412 Lena River, 1130, 1136, 1162 Leney, Frank, 4, 785 Leningrad, 14, 15, 745, 756, 1167, 1248. See also Institut des Mines, Palseozo- ologic Institute of the Academy of Sciences, Zoological Museum of the Academy of Sciences Lenk, Hans, 4, 475, 477, 785 Leoben, Steiermark, 92 Leporide, 680 Leptarctus, 400 Leptobos, 887, 1449 Leptodon, 35, 1377 giganteus, 1187, 1377, 1397, 1400 minor, 1187, 1377, 1397 leptodon (see Mastotherium leptodon) Leptomeryx, 400 Leptothertum, 579 Lepus, 680 Lesson, René Primiveére, 3, 518, 537, 578, 785, 1390 Levantin, 114, 154, 159, 160, 205, 396, 638, 643, 1374 Leverett, Frank, 1083, 1084, 1099, 1626 Leviathan missouriensis, 1390 missourti, 165, 783, 1389, 1390 levius (see Deinotherium levius) Levy County, Florida, 384, 418, 742 Lewis, Bailey, xvii Lewis, G. Edward, 1442, 1445, 1626 libyca (see Eosiren, Trilophodon angustidens libycus) Libytherium, 1431 Licent, Emile, 1482, 1626 licenti (see Stegodon licentt) Lichtenstein, Martin Heinrich Karl, 3, 785 Liddell, Henry George, 785, 1308 Lierre, Province d’Anvers, 1130 ligoniferus (see Trilophodon ligoniferus) Limpopo River, 1188, 1287 Lincoln County Mammoth, 943, 1009, 1012, 1017, 1019, 1027, 1028 Lincoln, Nebraska, 1609, 1613 Lindgren, Waldemar, 173, 487, 785 Lingle, Wyoming, 1012 Link, Heinrich Friedrich, 2, 785, 1128, 1136 Linnean ‘Special Creation’ system, xiii, 2, 6, 12, 20 zoological classification, 19, 20, 516 Linneus, Carolus, 2, 6, 9, 516, 785, 1308, 1309 Linz, 1466 Liquidambar, 217 Liri, Valle del, 1210, 1239, 1241, 1242 Lisbon, 217, 1193, 1609, 1613 Lista, Ignacio, 590 Listriodon, 272, 274, 732, 1429, 1459, 1461 Intopterna, 35 living elephants, 1192-1196, 1199-1202, 1205, 1307-1318, 13825-1333, 1392, 1398, 1401, 1402, 1409, 1412, 1590, 1594, 1595 Llano Estacado (Staked Plains), 384, 399, 422, 429, 431, 479, 494, 673, 732, 741, 742 Lockhart, Charles Frangois, 3, 91, 786 Lockhart, W., 1479 Loénnberg, Einar, 14, 1310, 1311, 1323 loess, 1487, 1488, 1516, 1517 Logan, William E., 3, 786, 1088 Loh formation, Mongolia, 115, 384, 387, 396, 742, 1318, 1477 Loire, 112, 115 Loiret Department, France, 91 Loir-et-Cher, France, 108 Lombardy, 1219 Lombez, France, 250, 738, 1464 Lomnicki, Aloys Maryan, 4, 786 Long Island, Phillips County, Kansas, 414, 415, 488 1651 longirostre (see Tetracaulodon longirostre) Longirostrine, 24, 27-29, 31, 119, 121, 124-126, 128, 228, 231, 232, 234, 235, 250, 254, 327, 338, 477, 539, 545, 546, 689, 690, 705, 734, 737, 1370, 1871, 1525, 1528, 1545, 1555, Pl. x longirostris (see Tetralophodon longirostris) Longueil, 131, 135, 136 Loochoo Island, 1489 Loomis, Frederick A., 4, 14, 400, 403, 478, 487, 488, 786 loomisi (see Torynobelodon loomis?) Lophiodontide, 824 lophodonty, 159 Lortet, Louis, and Ernest Chantre, 3, 124, 192, 194, 618, 786, 1064 Los Angeles Museum, California, 14, 975, 1007, 1009, 1010, 1609, 1615 Loup Fork horizon, 490, 555, 623, 669, 671, 999, 1002, 1003, 1501 Loup Fork of Madison Valley (see Madison Valley Beds, Montana) Loup River, Nebraska, 498, 560 Lourdes, 1169 Louverné, 1064 Loveland formation, 725, Pl. vit Lowe, C. Van Riet, 946, 1630 Lox—(disko)don, 941, 1378, 1583 Loxo-(disko)donten, 1378 Loxodon, 927, 1072, 1174, 1175, 1191, 1198, 1375 loxodont sinus, 1191, 1192, 1219, 1273, 1277, 1287, 1288, 1549 Lozodonta, 3, 6, 10, 11, 18, 853, 904, 907, 913, $14, 916, 924, 926, 927, 929, 930, 937, 942, 1042, 1173-1175, 1178-1180, 1185, 1186, 1189, 1191, 1192, 1195, 1199, 1203-1205, 1277, 1334, 1335, 1340, 1343, 1372, 1373, 1527, 1541, 1549, 1589-1591, 1595, 1603 africana, 10, 13, 32, 202, 230, 905, 908, 910, 915, 921-925, 928, 930-932, 937, 1005, 1029, 1052, 1072, 1095, 1120, 1121, 1128, 1129, 1131, 1147, 1170, 1172-1175, 1178, 1182, 1183, 1186, 1187, 1191, 1193-1203, 1205, 1209, 1216, 1219, 1227, 1249, 1251, 1252, 1271, 1273, 1283, 1286, 1307, 1311, 1312, 1321, 1325, 13829, 1334-1336, 1378, 1383, 1385, 1391, 1430, 1522, 1541, 1549, 1590, 1594, 1596, 1605, 1618. See also Messer- schmidt, and pygmy elephants africana africana, 797, 1175, 1193 africana albertensis, 1190, 1193, 1195, 1201, 1202, 1239, 1402, 1603, 1614, 1618 africana angolensis, 1193, 1195, 1412, 1613 africana capensis, 1121, 1187, 1192, 1193, 1195, 1197, 1198, 1383 africana cavendisht, 1193, 1195, 1614 africana cottont, 1193, 1195, 1196, 1202, 1203, 1402, 1619 africana cyclotis, 1193, 1195, 1196, 1401, 1522, 1610 africana cyclotis pumilio, 590 africana fransseni, 1193, 1195, 1196, 1404, 1620 africana knochenhauert, 1193, 1195, 1401, 1610 africana mogambica, 1193, 1195, 1409, 1613 africana orleansi, 1193, 1195, 1402, 1619 africana oxyotis, 179, 1147, 1176, 1193-1195, 1199-1201, 1227, 1229, 1230, 1239, 1249, 1336, 1401, 1402, 1615, 1618 africana peeli, 1190, 1193, 1195, 1198, 1201, 1202, 1239, 1248, 1402, 1603, 1617-1619 africana pumilio, 15, 1184, 1198, 1195, 1252, 1259, 1260, 1402, 1605, 1618 africana rothschildi, 1193, 1195, 1402 africana seloust, 1193, 1195, 1402, 1614 africana toxotis, 1193, 1195, 1402, 1611 africana var. obliqua, 984, 1188, 1273, 1287, 1288, 1321, 1415, 1439, 1541, 1612, 1618 africana zukowsky?, 1193, 1195, 1409 antiqua (see Hesperoloxodon antiquus) (antiqua) zulu. See Loxodonta zulu cornaliae, 1187, 1204, 1205, 1396, 1476, 1541 griqua (see Metarchidiskodon griqua) hysudrindica (see Palxoloxodon hysudrindicus) namadica (see Palxoloxodon namadicus) naumanni (see Palzoloxodon namadicus naumannt) (Palxoloxodon) namadicus. See Palxoloxodon namadicus (Palzoloxodon) namadicus (Yabet). See Palxoloxodon namadicus yaber OSBORN: THE Loxodonta—continued (Palxoloxodon) Tokunagai junior mut. See Palzoloxodon (Archidiskodon?) tokunagai mut. junior (Pilgrimia) melitensis. See Palxoloxodon melitensis prima, 984, 985, 1188, 1273, 1287, 1307, 1321, 1415, 1439, 1541, 1590, 1612, 1618 priscus-melitensis group, 908 subantiqua, 984, 985, 1188, 1288, 1416, 1439, 1541, 1612 zulu, 798, 983-985, 1184, 1187, 1273, 1277, 1281, 1285-1287, 1402, 1436, 1437, 1541 Loxodonte, 6, 1191, 1197, 1373. See also Loxodonta Loxodonten, 1378 Loxodontine, 22, 27, 32, 913, 925, 926, 928, 932, 936, 1171, 1178, 1174, 1178- 1180, 1186, 1191, 1210, 1227, 1228, 1271, 1277, 12891304, 1305, 1320, 1369, 1378, 1526, 15388, 1545, 1590, 1595, 1603 loxodontoides (see Archidiskodon loxodontoides) Lozano, Diaz, 1081 Lu Tzii Kou Beds, Shansi, 1480, 1623 Lueas, Frederic Augustus, 4, 14, 15, 400, 416, 418, 786 Lueas, Jannette May, xvii Ludolf, Heinrich August [Hiob?], 2, 5, 7, 786, 1118, 1120, 1125, 1147, 1164, 1165 Lugn, Alvin L., 786, 1497, 1500, 1626 Lujanense, 520, 590, 593, 1517, 1518 Lull, Richard Swann, 4, 7, 9, 234, 297, 488, 526, 527, 539, 575, 786, 1046 lulli (see Megabelodon lull) Lumbres, 1251 Lund, Peter Wilhelm, 3, 518, 537, 578, 579, 736, 1390 Luparello, Sicily, 1268, 1270-1272 Lutra, 887 Lycée de Riazan, 134 Lyceum of Natural History, New York, 1156, 1390 Lydekker, Richard, xii, 29, 82, 85, 105, 250, 267-269, 349, 353, 354, 362, 636, 685, 786, 818, 832, 837, 840, 853, 865, 871, 885, 907, 974, 1039, 1040, 1142, 1221, 1332 lydekkeri (see Serridentinus lydekkeri, and Stegolophodon lydekkert) Lydekkeria, 343, 344, 349, 351-363, 379, 739, 1381, 1559, 1560 Lyell, Charles, 3, 787, 1071, 1072, 1077 Lyons, France, 84, 86, 1609, 1615. See also Muséum des Sciences Naturelles, Université de Lyon Lyons, Henry, xvi, 1524 lyonsi (see Meeritherium lyonsi) lyrodon (see Elephas lyrodon) Maarel, Franciscus Hendricus van der, 5, 14, 27, 31, 349, 356, 365, 787, 816, 837, 887, 889, 1416, 1454 Macacus, 887, 1155, 1489 MacCurdy, George Grant, 1131, 1167, 1626 Machexrodus, 971, 1155, 1431, 1461 MacIntosh, William, 1575 Macrauchenia, 1519 macrocephalus, 1383. See Mastodon americanus macrodon (see Mastodon macrodon) macrognathus (see Trilophodon macrognathus) macroplus (see Anancus arvernensis macroplus) macrorynchus (see Elephas macrorynchus) Macrotherium, 1459, 1461 Maddren, Alfred Geddes, 4, 787, 1134 Madeira, Percy, 1202, 1609, 1619 Madeleine, La, 1132, 1167 maderianus. See Cuvieronius(?) maderianus Madison Valley Beds, Montana, 1491, 1508 Maeotic, 1468 Magdalenian, 1126, 1132, 1167 Magello, 975 Magliabechi, A., 1118 Maiben, Hector, 302, 304, 312, 1019 maibent (see Archidiskodon imperator maibent) Mainz Museum, 282, 283, 362, 745, 756 PROBOSCIDEA Majer, Istvan, 4, 787 Major, Charles Immanuel Forsyth, 3, 787, 11838, 1187, 1232 majus. See Deinotherium [gig.| var. majus Makiyama, Jir6, 4, 787, 893, 908, 1185, 1187, 1291, 1292, 1295, 1296, 1301, 1408, 1420, 1627 Malafrasca, 1215 Malan loess, 1483, 1485-1488 malar, 916 Malay Peninsula, 927, 1332 Malbattu, 1065 Malta, 1182, 1257, 1262, 1263, 1265 Mammaliferous Crag, near Norwich, 973 mammille, 828 Mammont, 5, 7, 775, 1164 mammonteo, 1147 Mammonteum (see Mammonteus) Mammonteus, 5, 7, 10, 11, 32, 909, 913, 914, 916, 923, 924, 926-930, 932, 933, 935, 938, 942, 981, 1039, 1040-1043, 1045, 1089, 1090, 1117, 1118, 1124, 1126-1128, 1133, 1185-1138, 1140, 1146, 1147, 1149, 1157, 1158, 1163, 1167, 1178, 1193, 1209, 1228, 1304, 13438, 1363, 13865-1367, 1370, 1372, 1381, 1512, 1514, 1527, 1541, 1548, 1550, 1582, 1587-1589, 1603. See also Mammoth, Mammut primigenius, 5, 7, 10, 32, 33, 170, 171, 534, 725, 905, 907-910, 915, 922, 924, 927-931, 936-971, 976, 982, 1041, 1043-1045, 1050, 1051, 1053, 1056, 1058, 1065, 1067, 1068, 1070, 1073, 1074, 1077, 1088, 1091, 1095, 1097, 1099, 1112, 1116, 1118-1124, 1127-1147, 1149, 1155, 1156, 1162-1169, 1173, 1175, 1177, 1178, 1181, 1199, 1201, 1209, 1221, 1227-1229, 1248, 1344, 1362, 1365, 1366, 1373, 1388, 1387, 1389, 1390, 1397, 1475, 1476, 1488, 1518, 1514, 1522, 1541, 1582, 1588-1590, 1605, 1610-1613, 1616, 1618, 1621, 1628, Pl. vir. See also “Adams skeleton,’ Burgtonna, Messersehmidt primigenius alaskensis, 1134, 1137, 1138, 1159-1161, 1417, 1541, 1611, 1618 primigenius americanus, 999, 1000, 1137, 1138, 1140, 1156, 1157, 1166, 1390, 1410, 1541 primigenius astensis, 905, 932, 938, 981, 1127, 1137, 11388, 1140, 1142, 1149, 1150, 1164, 1155, 1158, 1178, 13866, 1407, 1476, 1541, 1611, 1620. See also Hlephas primigenius Blum. var. trogontherit primigenius compressus, 922, 925, 931, 932, 938, 939, 1045, 1088, 1099, 1127, 1131, 1184, 1135, 11387, 1138, 1140, 1142-1145, 1155, 1156, 1157-1159, 1164, 1166, 1167, 1169, 1366, 1409, 1514, 1541, 1616 primigenius fraast, 1130, 1137, 1138, 1140, 1149, 1162, 1153, 1253, 14038, 1476, 1541, 1603, 1605, 1620 primigenius hydruntinus, 1137, 1149, 1150-1152, 1153, 1399, 1476, 1541 (?) primigenius leith-adamsi, 1137, 1138, 1140, 1149, 1150, 1399, 1476, 1541, 1610 primigenius leith-adamsi var. Minor, 1150, 1399 primigenius primigenius (see Mammonteus primigentus) primigenius sibiricus, 905, 1128, 1137, 1178, 1391 mammonteus (see Elephas mammonteus) Mammonth, 517 Mammontheum, 1365 Mammontinx, 27, 32, 127, 913, 925, 932, 935, 936, 937, 939, 942, 1116, 1117, 1136, 1159, 1163, 1227, 1228, 1370, 1526, 1538, 1545, 1547, 1582-1584, 1587, 1588 Mammontova Kost (see Mammotovoi kost) , Mammoth, 1, 5-7, 182, 183, 928, 1116, 1119-1121, 1125, 1127-1129, 11381, 1132, 1135, 1139, 1140, 1162, 1164, 1165, 1167, 1169, 1363, 1487, 1490. See also Archidiskodon imperator, Columbian Mammoth, frozen mammoth of Siberia, Mammonteus, Mammontine, Maydell Mam- moth, Moravia, Parelephas jeffersonii, Sceleto Elephantino Tonne Mammoth (see Elephas Mammoth) Mammotovoi kost, 5, 7, 786, 1120, 1125, 1164, 1165, 1197 Mammouth, 5,7, 1164 ohioticum (see Mammut ohioticum) mammouth (see Elephas mammouth) Mammut, 6-8, 10, 135, 169, 526, 527, 1124, 1141, 1364, 1865, 1372 cautleyt, 527 francisi (see Rhynchotherium francis?) INDEX 1653 Mammut—continued Maseall zone, 155, 485, 1494 ohioticum, 6, 7, 136, 165, 168, 169, 1166, 1364, 1372, 1383, 1387 Mascara, 1274 oregonense (see Mastodon oregonensis) Mashonaland, 1193, 1402 progenium (see Mastodon progenius) Masritherium, 1426 stbiricum, 1123, 1136, 1387 Mastelephas, 290, 1380 Mammuthus, 1117, 1363, 1865-1367, 1372, 1373, 1588, 1614 Mastodon, xii, xiv, 5-8, 10-12, 28, 29, 33, 43-45, 127, 128, 131, 132, 136, 138— borealis, 1117, 1123, 1136, 1366, 1373, 1387, 1588 primigenius, 1366, 1367 Mammutide, 1368, 1526, 1572, 1574, 1575, 1578 Mammutine, 1364, 1371 Mamonteum, 1364 man, 1490, 1512, 1514, 1521, 1547, 1606 Manatee, 39 Manatee County, Florida, 400, 1047, 1108 Manchhar, 266-269, 448, 449, 643, 738, 853 Mandalay, 874, 875 Manis palxojavanica, 887 Manitoba, 736 Mannersdorf, Austria, 360, 361 Mantell Collection, 121, 252, 254, 340 Manzat!, Rumania, 96-98, 114 Mapleton, Iowa, 1003 maps: Afghanistan, 642; Africa, 964, 983; Alaska, 1134; Amebelodon, 321, Pl. xv; Amebelodontine, 734, 1528, Pls. vir, xv; Anancus, 642, 1528, Pl. xvi; Archidiskodon, 934, 942, 964, Pl. xx1; Asia, 461, 853, 1304, 1305; Big Bone Lick, 136; Brevirostrine, 624, 734, 1528, Pl. xv1; Bugti Hills, 275; Burma, 824; Channel Islands, 1032, circumpolar, 914, 1135, 1589; Devil’s Gulch, 601, Pl. vir; Deinotheres, 84, 734, 1528, Pl. x1v; Ecuador, 567; Eden (Mt.) deposits, 498, 560; Ele- phantide, 914; Elephantine, 29, 1318, 1538, 1594, Pl. xxm; Elephan- toidea, 914, 941, 1589; Faytim, 37, 52; Gnathabelodontine, 711, Pls. vu, xv; Humboldtine, 536, 624, 734, 1528, Pl. xvir; India, 279, 642, 852, 1528, 1538; Java, 366; Kendeng, 366, 887; Liri Valley, 1239, 1241; Longirostrine, 250, 734, 1528, Pls. vir, xv; Loxodonta (living elephants), 1195; Loxodontine, 1186, 1538, Pl. xx1u; Mammontine, 942, 952, 1047, 1078, 1133, 1135, 11386, 1164, 1538, Pls. xx1, xx; Mastodon americanus in North America, 177; Mastodontine, 137, 176, 734, 1528, Pl. xrx; Mediterranean (bathymetric), 1257; Metarchi- diskodon, 942, P|. xx1; Moeritheres, 734, 1528, Pl. x1v; Miocene, 201, 204, 254; Mio-Pliocene, 400; Notiomastodon, 536, Pl. xvi11; Notoro- strine, 536, 734, 1528, Pl. xv1 (Cordillerion); Paleomastodontine, Pl. xrx; Parelephas, 536, 1047, 1133, 1135, Pl. xxm; Phiomia, 321, Pl. xv; Platybelodontine, 384, 414, 1528, Pls. vir, xv1i1; Pleistocene of Europe, 941, 1042; Pliocene, 263, 318; Rhynchorostrine, 480, 513, 734, 1528, Pl. xv1; San Timoteo, 498, 560; Serridentine, 384, 386, 414, 734, 1528, Pl. xvu1; Sheep Creek to Snake Creek, 426; Siberia, 1162, 1164; Siwaliks, 940, 952, 1339; Staked Plains, 399, 422; Stegolopho- dontine-Stegodontine, 823, 914, 1538, 1589, Pls. xix, xx; Tetra- lophodontine, 349, 734, 1528, Pl. xv; Trilophodon, 321, 1528, Pl. xv; Tung Gur, 398, 466; United States, 386 (east coast), 1003 (west of Mississippi River); worldwide distribution, past and present, 29, 1594; zodgeographic realms, 35; Zygolophodontine, 176, 193, 215, 734, 1528, Pl. xrx. Maragha, Persia, 114, 261, 262, 349, 739, 853, 1062, 1400, 1443, 1468 Margetts, Camp, Mongolia, 398, 466 Margniole, 1063 Maria-Theresiopel, Hungary, 159, 160 Marietta College, 1067 Marion County, Florida, 418 Marnes de 1’Orléanais (see Orléanais) Marnes du Blésois (see Blésois) Marschall, Augusto de, 3, 787 Marseilles, (see Muséum d’Histoire Naturelle, Marseille) Marsh, Othniel Charles, 3, 787 Martaban, Burma, 824 Martin, Johann Karl Ludwig, 3, 788, 818, 832, 885, 890 martini (see Cryptomastodon martini) Marty, 256, 259 Maryland (see Greensburgh) 142, 165, 169, 170, 175, 180-190, 200, 211, 269, 502, 517, 575, 685, 688-690, 736, 983, 1141, 1147, 1166, 1365, 1372, 1374, 1377, 1489, 1490, 1512, 1514, 1527, 1545, 1546, 1574, 1602, Pls. 1, 1. See also Mastodon americanus, Mastodonte, Warren Mastodon, Whitfield Mastodon acutidens, 137, 165, 169, 185, 186, 189, 191, 211, 692, 694, 696, 697, 736, 748, 880, 1419, 1514, Pl. 1 affinis, 1397. See Zygolophodon borsoni affinis americanus, 5, 7, 10, 30, 120, 122, 127, 131-1387, 189-142, 150, 153, 156, 160, 161, 164-171, 176-178, 180-191, 205, 208, 211, 230, 232, 399, 502, 610, 681, 687, 696, 698, 699, 735, 746, 748, 751, 758-760, 844, 1121, 1134, 1135, 1161, 1177, 1363, 1364, 1372, 1374, 1382, 1383, 1385, 1388, 1395, 1482, 1503, 1514, 1522, 1546, 1550, 1575, 1602, 1605, 1611, 1614, 1616, 1619, Pl. 1. See also Brunswick Canal, China, Hillsboro, Le Grand Mastodonte, Mammut, Mastodon giganteum, M. ohioticum, Mastodonte de VOhio, Mastotheritwum megalodon, Melbourne, Ohio- tncognitum, Otisville, Otoe County, Phosphate Beds, Shawangunk, Warren Mastodon, Whitfield Mastodon americanus alaskensis, 165, 176, 177, 736, 753, 1417, 1514 americanus plicatus, 137, 165, 169, 173, 736, 746, 1410 americanus rugosidens, 137, 165, 171, 736, 745, 748, 1399, 1514, 1619 americanus rupertianus, 137, 736, 1393, 1514 andaranus, 267, 1395 andicus, 588 andi, 588 andium (see Cordillerion andium) andium Kraglievichti (see Cordillerion andium kraglievichii) angustidens (see Trilophodon angustidens) angustidens Cuy. mut. ase. pygmexus (see Phiomia pygmexus) angustidens Cuvier var. libyca (see Trilophodon angustidens libycus) angustidens minus (see Trilophodon angustidens minutus) angustidens mut. pontileviensis (see Trilophodon pontileviensis) angustidens sivalensis, 650 angustidens steinhermensis (see Trilophodon steinheimensis) angustidens var. austro-germanica (see Trilophodon angustidens var. austro-germanicus) angustidens var. latidens, 1401 antium, 544, 545, 549 arborense, 1390 argentinus (see Notiomastodon argentinus) arvernensis (see Anancus arvernensis) arvernensis Croiz. et Job. var. conservativus, 625, 640, 1403 arvernensis var. progressor (see Anancus arvernensis progressor) atavus, 1418, 1479, 1612 atticus (see Turicius atticus) aureliense, 1401 aus Alangasi (see Cuvieronius postremus) australis, 1390 bolivianus (see Cordillerion bolivianus) bonaerensis (see Cuvieronius bonaerensis) borsoni (see Zygolophodon borsont) Brasiliensis (see Cuvieronius brasiliensis) brevidens (see Rhynchotherium brevidens) brevirostre (see Anancus arvernensis brevirostris) buffonis (see Zygolophodon borsoni buffonis) (Bunolophodon) angustidens Cuv. f. subtapiroidea (see Serridentinus subtapiroideus) ‘ (Bunolophodon) angustidens Cuy. forma typica (see Trilophodon angusti- dens) (Bunolophodon) grandincisivum (see Tetralophodon grandincisivus) (Bunolophodon) longirostre forma sublatidens (see Stegolophodon sublatidens) cautleyi (see Stegolophodon cautley?) chapmani (see Stegomastodon chapmant) 1654 Maslodon—continued chilensis (see Cuvieronius chilensis) cordillerarum (see Mastodonte des Cordiliéres) cuviert, 165, 1388, 1395 (Dibunodon) arvernense, 195, 630, 638, 1379 dissimilis (see Anancus arvernensis dissimilis) dubius, 114, 283, 349, 357, 358, 755, 1388, 1389 elephantoides (see Stegodon elephantoides) engelswiesensis (see Trilophodon engelswiesensis) esselbornensis (see Trilophodon esselbornensis) euhypodon (see Blickotherium euhypodon) falconeri. See Tetralophodon (Lydekkeria) falconert, and Pentalophodon falconert floridanus. See Ocalientinus (Serridentinus) floridanus gaujaci (see Trilophodon angustidens gaujact) gigantarvernensis (see Anancus gigantarvernensis) giganteum, 7, 119, 122, 135-137, 165, 167, 169, 170, 209, 1364, 1372, 1385 giganteus, 1395 gigantorostris (see Tetralophodon gigantorostris) grandis, 114, 283, 349, 357, 358, 359, 746, 755, 1389 grangeri, 137, 165, 169, 692, 695, 696, 725, 736, 758, 1417, 1514 hasnoli (see Synconolophus hasnoti) humboldii (see Cuvieronius humboldti) humboldtianus |humboldianus), 537, 796, 1385 humboldtii (see Cuvieronius humboldtiz) humboldtius, 1393 intermedius, 1420, 1482, 1483. Jjeffersoni, 165, 1388 latidens (see Stegolophodon latidens) ligoniferus (see Trilophodon ligoniferus) longirostris (see Tetralophodon longirostris) lydekkeri (see Serridentinus lydekkert) macrodon, 136, 1385 maderianus. See Cuvieronius(?) maderianus (Mammut) americanus forma praetypica, 1406, 1514. americanus praetypica matthewt (see Pliomastodon matthew?) maximus, 137, 1385, 1386 merriamt (see Miomastodon merriamt) microdon, 192, 1385. See Mastotheriwm microdon minor, 122, 250, 252, 1385, 1386 minus, 124, 1386 minutoarvernensis (see Anancus minutoarvernensis) minutus (see Trilophodon angustidens minutus) mirificus (see Slegomastodon mirificus) moodiet, 137, 165, 169, 174, 725, 736, 758, 1416, Pl. vii obscurus (see Trilophodon obscurus) ohioticum, 1385, 1387 ohioticus, 6, 7, 126, 132-135, 137, 165, 168, 169, 1374, 1375, 1391, 1395, Pl. 1. See Mammut ohioticum, Mastodon pavlowr oligobunis (see Cordillerion oligobunis) oligobunis var. antiquissima (see Cordillerion oligobunis antiquissimus) oligobunis var. Felicis (see Cordillerion oligobunis felicis) oligobunis var. intermedia (see Cordillerion oligobunis intermedius) oligobunis var. progressa (see Cordillerion oligobunis progressus) oregonensis, 137, 173, 736, 759, 1410, 1514 pandionis (see Trilophodon pandionis) parvus, 1386, 1388 pavlowi, 126, 131-135, 137, 165, 169, 692, 694, 736, 760, 1419, 1476, Pl. 1 pentelicus. See Trilophodon (Choerolophodon) pentelicus perimensis (see Anancus perimensis) perimensis var. sinensis. See Tetralophodon (Lydekkeria) sinensis pirayuiensis (see Cuvieronius pirayuiensis) platensis (see Cuvieronius platensis) podolicum (see Deinotherium podolicum) proavus (see Serridentinus proavus) productus (see Serridentinus productus) progenius, 137, 165, 169, 172, 175, 190, 399, 736, 760, 1514 punjabiensis (see Tetralophodon punjahiensis) See Anancus intermedius See Pliomastodon OSBORN: THE PROBOSCIDEA pyrenaicus (see Zygolophodon pyrenaicus) raki, 137, 165, 169, 175, 736, 751, 1417, 1514 rectus (see Cuvieronius rectus) rhomboides, 537, 1385 rugatum, 165, 1390 Sahendi, 1400 Senodon, 250, 252, 1385 serridens (see Serridentinus serridens) shepardi (see Rhynchotheriwm shepardt) Simorrense, 121, 250, 252, 254, 1393 sivalensis (see Pentalophodon sivalensis) spencert (see Rhynchotheriwm spencer?) stegodontoides (see Stegolophodon stegodontoides) sleinhermensis, 251, 281, 282, 284, 738, 1406 successor (see Stegomastodon successor) superbus (see Cuvieronius superbus) laptiroides (see Turicius tapiroides) taptroides americanus (see Miomastodon tapiroides americanus) tapiroides-minus (see Turicius tapiroides-minus) larijensis (see Cordillerion tartjensis) (Tetralophodon) latidens, 1404 (Tetralophodon) perimensis, 1398 (Tetralophodon) sivalensis, 1404 (Trilophodon) angustidens Cuy. var. paleindicus. indicus (Trilophodon) Falconert. See Tetralophodon (Lydekkeria) falconeri (Trilophodon) floridanus. See Ocalientinus (Serridentinus) floridanus (Trilophodon) pandionis, 1404 turicense, 1386 turicensis (see T'uricius turicensis) vellavus (see Zygolophodon borsoni vellavus) vialetti (see Zygolophodon borsoni vialetir) virgatidens (see Turicius virgatidens) von Chimborazo (see Cuvieronius ayore) wahlheimensis (see Turicius walheimensis) zaddachi (see Zygolophodon borsoni zaddacht) Mastodonadae, 778, 912, 1368 Mastodonta, 6 Mastodonte, 5, 6, 119, 122, 127, 167, 517, 1864, 13865, 1372, 1527 Mastodonte a dents étroites, 121, 122, 250, 252-254, 518, 633, 1384 Mastodonte de Alangasi, 1412. See Cuvieronius postremus Mastodonte de VOhio, 120, 122, 135, 136, 165, 517, 633, 1174, 1384, 13885 Mastodonte des Andes, 550 Mastodonte des Cordiliéres, 121-123, 516, 517, 525, 637, 543, 649, 568, 588, 722, 1384, 1386 Mastodonte humboldien, 121-123, 616, 617, 537, 576, 577, 633, 722, 1384 Mastodonte Petit (see Petit Mastodonte) Mastodonte tapiroide, 633, 1385, 1386. clus tapiroides Mastodontide, 18, 25, 26, 30, 45, 119, 126-128, 148, 225, 532, 686-689, 691, 700, 735-737, 912, 13867, 1368, 1524, 1525, 1547, Pl. x Mastodontina (see Mastodontine) Mastodontine, 24, 27, 28, 30, 119, 120, 126, 128, 131-133, 135, 137-140, 156, 170-177, 191, 689, 690, 692-697, 718, 734, 736, 816, 1369, 1526, 1528, 1545, 1551, 1574, 1575, Pls, 1, x : Mastodontoidea, 22-24, 26, 27, 30, 32, 33, 114, 115, 118, 119, 127, 128, 686, 688, 689, 700, 734, 735, 1367, 13871, 1547-1551, 1555-1579, 1600, Pls. 3552.9) Mastodontoideum (see Tetracaulodon Mastodontoideum) Mastodontum, 1372 Mastotherium, 10, 11, 137, 192, 250, 617, 537, 1364, 1372 hyodon, 10, 617, 537, 548, 549, 587, 588, 1372, 1385, 1386 humboldtii, 618, 537, 575, 1872, 1385 leptodon, 10, 250, 252, 618, 1372, 1384 megalodon, 10, 137, 165, 517, 13873, 1384, 1385 microdon, 10, 192, 618, 1373, 1385, 1386 ohioticum, 1384 Matanzas River, Argentina, 597, 599 Mather, Cotton, 1627 See Trilophodon palx- See also Petit Mastodonte, and Turi- INDEX Mather, William W., 3, 788, 997, 1067 Matschie, Paul, 4, 14, 15, 788, 1192, 1193, 1197, 1383 Matsumoto, Hikoshichiré, 4, 41-45, 48, 64, 77, 143, 251, 280, 384, 457, 788, 816, 818, 821, 822, 834, 835, 839, 890, 901-909, 943, 964, 1185, 1186, 1188, 1290-1292, 1627 matsumotoi. See Parelephas protomammonteus (Matsumoto) matswmotot Matsuoka-mura, 906 Matthew, Christina D., 86 Matthew, William Diller, 4, 15, 19, 20, 102, 187, 157, 161, 208, 218, 251, 256, 298, 305, 355, 360, 400, 488, 546, 548, 578, 601, 637, 640, 789, 858, 1422, 1444, 1499, 1504, 1518, 1627 matthewi (see Pliomastodon matthew?) Mauer, 1044, 1045, 1050, 1065 Mauretania, 1195 Mauri River, Bolivia, 552 maximum (see Deinothertum maximum) maximus (see Elephas maximus, and Mastodon maximus) Maxson, John H., 1446, 1509, 1627 Maxwell, Marius, 1239 Maydell Mammoth, 1131 Maydon, H. C., 29, 789 Mayence, 217 Mayet, Lucien, 4, 15, 95, 107, 124, 194, 195, 204, 205, 212, 217, 631, 789, 943, 961, 1042, 1043, 1056, 1137, 1154, 1187, 1232 McGregor Museum, Kimberley, 15, 987, 1278-1281, 1287, 1458, 1609, 1612 McGrew, Paul O., 470, 800, 1496 McKee, James R., 171 McLennan County, Texas, 373 McPherson, Kansas, 349, 373, 739 Mead, H. L., 430 Meagher County, Montana, 479, 485, 737 Mececyon trinilensis, 887 Mecklenburgian, 1473 medilongirostrines (see T'etralophodon, Serridentinus) Medio, arroyo del, 599 Mediterranean area, 1456, 1461, 1465 Mediterranean Islands, 936, 1182, 1252, 1257-1272, 1473, 1592, 1593, 1603 medium. See Deinotherium medium, and Deinotherium |gig.| var. medium Medway, 1222 Meerut, 1312 Mefferd, Ralph LeRoy, 728 Megabelodon, 44, 45, 249, 251, 289-291, 294, 295, 297, 686, 690, 705-707, 737, 738, 1379, 1527, 1556, 1600 cruziensis, 251, 257, 319, 323, 324, 326, 327, 329, 436, 706, 707, 711, 738, 751, 1412, 1417, 1498, 1600, Pl. vi joraki, 251, 257, 297, 319, 326, 327, 329, 706, 707, 735, 738, 751, 1412, 1417, 1493 lulli, 226, 229, 248, 250, 251, 260, 287-291, 294-298, 317, 318, 321, 329, 330, 388, 501, 679, 686, 706-709, 711, 714, 738, 758, 1379, 1380, 1404, 1412, 1498, 1499, 1600, 1604 phippsi, 251, 257, 287, 289, 290, 315-319, 323, 326, 329, 388, 705, 706, 709, 711, 733, 738, 751, 755, 1412, 1498, 1600, 1604, Pl. vii Megaceros, 1253 megalodon (see Mastotherium megalodon) Megalohyrax, 1424 Megalonyz, 171, 399, 400, 508, 579, 725, 1503, 1512, Pl. vim Megalonyx-Mylodon fauna, 1515 Megalonyx Zone, 1511 Megatherium, 399, 418, 551, 725, 1077, 1521, Pl. vin Meisen Series, Japan, 1420 Melbourne, Florida, 400, 1005, 1079 Melchingen, Germany, 91, 1468 Meles taxus, 1231 Melitz (see Palxoloxodon melitensis) melitensis (see Palzoloxodon melitensis) Melville Island, 1590 Menelek, 1201 Menges, 1193 Mercedes, Argentina, 594, 596, 597 1655 Merced-Etchegoin, 902 Merck, Johann Heinrich, 2, 789, 1120 _Merian, Peter, 91, 789 meridionalis (see Archidiskodon meridionalis) Merriam, John C., 4, 15, 152, 154, 155, 487, 497, 561, 790, 1494, 1509, 1627 merriamt (see Miomastodon merriamt) Merrill, Elmer Drew, 14, 717 Merrill, George Perkins, 4, 15, 286, 287, 418, 420, 790, 1395 Merychippus, 315, 318, 380, 386, 400, 403, 426, 488, 510, 601, 1446, 1467, 1497, 1499 sejunctus Zone, 315, 403 Merycodus, 315, 318, 444, 680 Merycops, 272, 1428 Mesocyon, 386 Mesohippus, 601 Mesohippus-Oreodon Zone, 601 Mesopotamia, 1207, 1209, 1591 Messerschmidt, Daniel Gottlieb, 767, 1118, 1119, 1121, 1186, 1149, 1165, 1628 Messinien, 114 metachinjiensis (see Serridentinus metachinjiensis) metaconules (see conules) metaloph-lophid, 139, 141, 812, 1545 Metaphiomys, 1425 Metarchidiskodon, 942, 983, 994, 1382, 1527, 1549, 1584, Pl. xr griqua, 942-946, 983-985, 994, 995, 1184, 1187, 1278, 1285, 1406, 1436, 1439, 1540, 1584, 1612 Metasinopa, 1425 metatype, 9 Metoldobotes, 1425 :. Metoreodon, 601 Metoreodon Zone, 318 Mexico, 15, 4938, 533, 534, 548, 552, 1013, 1080, 1082, 1506, 1609, 1615. See also Aculeingo, Amajaque, Arizpa, Ejutla, Escuela de Minas, Hidalgo, Michoacan, Museo Instituto Nacional geologico, San José de Pimas, Tequixquiac, Tlascala, Tlatlaya, Uhde Collection, Valley of Mexico, Vera Cruz, Zumpango Meyer, Hermann von, 3, 84, 85, 87, 192, 194, 212, 213, 221, 553, 790, 1123, 1136, 1395, 1627 Michoacan, Mexico, 532, 537, 553, 740, 1013 Microbunodon, 272 microdon (see Mastotherium microdon) microscopy, 1607, 1608, Pls. xxvi-xxx Microteinz, 680 Middendorf, Alexander Theodor von, 1162, 1165, 1627 Midway, Florida, 400 migrations: Africa, 22, 34, 36; Amebelodontine, 1572; Archidiskodonts, 934, 964, 982, 983, 996, 998; Asia, 22, 1304, 1490; Brevirostrine, 643, 720; Deinotheres, 485, 734, 1528; Ethiopian region, 34, 35; Gnathabelodon- tine, 1557; Hesperoloxodon antiquus, 1295; Humboldtine, 1568; Longirostrine, 1557; Mammontine, 1584, 1587, 1588; maps, 35, 176, 177, 215, 321, 414, 734, 1528; Mastodon, 176; Mastodontinw, 1575; Mastodontoidea, 734, 1528; Mceritheres, 734, 1528, 1553; North and South America, 527, 536, 562, 611, 685; Notiomastodontine, 1572; Notorostrine, 1561; Paleoloxodonts, 1180, 1210, 1268, 1292; Parele- phas, 1044, 1046, 1049, 1067, 1071; Platybelodontine, 414, 1572; reversal of migration route, 982, 983; Rhynchorostrine, 485, 513; Serridentine, 396, 414; Stegodontine, 822, 1579; Stegolophodon- tine, 822, 1578; Stegomastodon, 680, 685; Tetralophodontinx, 379, 1560; Trilophodon, 284, 321, 822; Zygolophodontine, 215, 822, 1577 Mikawa, 907, 908, 1489 Milazzian, 818, 893, 902, 904-906, 908, 1186, 1473, 1474 Milford, Nebraska, 137, 174, 736 Miller, Gerrit Smith, 15, 1134 Miller, Hugh, 92, 790 Millett, W., 987, 989, 991 milletti (see Archidiskodon millettt) Milligan, Florence 8., xvii Milne-Edwards, 1460 1173, 1656 Milwaukee, Wisconsin, 1097 Minami-Akita District, 908 Minas Geraes, Brazil, 537, 578, 579, 741 Minato Town, Japan, 818, 906, 1297, 1420 Mindanao, 833, 892 mindanensis. See Stegodon (Archidiskodon?) mindanensis Mindel, 725, 1430, 1457, 1473, 1474, Pl. vim Mingoon, 874, 875 Minnechaduza, 1497 Mino, Province of, 384, 457, 742, 907, 1333, 1488 minor. See Elephas antiquus var. minor, Elephas (primigenius) Leith-Adamsi Pohlig var. minor, Leptodon minor, Phiomia minor, Trilophodon angustidens minutus Mint Canyon Beds, 1445, 1446, 1509 minus (see Palzomastodon minus, and Trilophodon angustidens minutus) minutoarvernensis (see Anancus minutoarvernensis) minutum (see Deinotherium minutum) minutus (see T'rilophodon angustidens minutus) Miocene, 20, 32, 33, 201, 204, 254, 279, 400, 426, 448, 642, 711, 1426-1429, 1439-1447, 1456, 1457, 1464-1466, 1477-1480, 1489-1491, 1493, 1499, Bishvat x cr Miohyrax oswaldi, 1428 Miomastodon, 11, 28, 132, 137-139, 151-157, 211, 688, 690, 698, 694, 736, 1509, 1527, 1545-1547, 1575, 1602 depereti, 131, 137, 192, 196, 284, 693-695, 736, 758, 1419, 1460, Pl. 1 matthewt (see Pliomastodon matthew?) merriami, 11, 132, 137, 140, 150-156, 159, 164, 170, 690, 692, 693, 697, 736, 748, 751, 755, 1380, 1381, 1406, 1492, 1494, 1502, 1605 tapiroides americanus, 133, 137, 140, 150-154, 156, 692, 736, 1380, 1406, 1469 Miosiren, 42 “Mirghi,” 1314 mirificus (see Rhabdobunus mirificus, Stegomastodon mirificus) Mirzapur, 950, 952, 954, 955, 964 Mishima, Kazusa, 1188, 1289, 1300 Mississippi River, 498, 560, 1003 mississippiensis. See Parelephas (?) mississippiensis (?) Missouri Leviathan, 1389 Missouri Valley, 172, 736, 844 Missourian, 1374, 1389 missouriensis (see Leviathan missouriensis) missourii (see Leviathan missourit) Missourium, 1374, 1602 kochii, 165, 783, 1389 theristocaulodon, 165, 783, 1389, 1390 Mitsugo-shima, 818, 893, 906 Miura Peninsula, 908, 1301 Miura series, 1300 Mixson’s Bone Bed, 384, 386, 416, 419 Miyagi District, 845, Pl. 1v Miyata, 908 Mijousse River, Russia, 1393 Mnaidra Gap, Malta, 1187, 1257 mnaidre (see Palxoloxodon mnaidriensis) mnaidriensis (see Palzoloxodon mnaidriensis) mocambicus (see Loxodonta africana mogambica) models (see Abel, Christman, Flinsch, Knight) Meeris, Lake (see Fayfim) Meeritheriide, 26, 30, 45, 65, 69, 71, 688, 735, 1867, 1368, 1525, 1552 Merritheriine, 30, 71, 734, 735, 1369, 1370, 1526, 1528, 1545, 1552, Pl. x Meeritheriini (see Mceritheriine) Meeritherioidea, 20, 23-27, 30, 32, 39, 69, 83, 688, 734, 735, 1367, 1524, 1528, 1550, 1552, 1553, 1600, Pls. x, x1 Maritherium, 10-12, 20, 22, 24, 32, 33, 36, 38-43, 46-52, 54, 56, 57, 61, 65, 71-15, 77-79, 83, 139, 202, 688, 734, 735, 1379, 1424, 1524, 1527, 1545-1547, 1550, 1553, 1554, 1572, 1600, Pls. x, x1 ancestrale, 51, 52, 65, 76, 735, 754, 1408, 1426, 1604 andrewsi, 51, 52, 55, 61, 62, 65, 66, 68, 73-76, 77-79, 145, 341, 688, 735, 746, 754, 755, 1408, 1426, 1544, 1604, 1616 OSBORN: THE PROBOSCIDEA gracile, 39, 51-53, 65-67, 65, 68, 73, 74, 77, 688, 735, 746, 755, 1401, 1426 lyonsi, 10, 39, 46, 51-65, 65, 66, 70-73, 77, 78, 145, 735, 746, 755, 1401, 1426, 1544, 1616 trigodon, 51-53, 67, 58, 65, 68, 73-75, 77, 78, 145, 688, 735, 746, 747, 754, 1401, 1402, 1426, 1544, 1604, 1616 trigonodon (see Meritherium trigodon) Mosskirch, 85, 91, 115, 281, 1390, 1463 Moffat County, Colorado, 251, 312, 385, 738, 1103 Moghara, 35, 103, 115, 251, 260, 479, 485, 737, 738, 1426, 1427 Moginand, India, 815, 1318, 1338 Mogok fissures, 1451, 1452, 1487 Mohave Desert, 251, 385, 387, 447, 508, 718, 742, 1494, 1502 Moir, J. Reid, xvii, 1627 Molasse de l’Anjou, Molasse du Royans, Grauen Susswasser Molasse, Obere Susswasser Molasse, 115; Miocene Molasse, 91 Monastirian, 902, 905, 907, 1186, 1290, 1333, 1473, 1474 Moncamp, 205 Mongolia, 251, 334, 384, 385, 397, 398, 460, 461, 742, 743, 1477, 1488. See also Altai region, Camp Margetts, Gobi, Houldjin gravels, Hung Kureh Beds, Iren Dabasu, Kalgan Urga Trail, Kholobolehi Nor, Khunuk formation, Loh formation, Serridentinus gobiensis, Serridentinus mongoliensis, Tairum Nor, Tsagan Nor, Tung Gur Khara Usu mongoliensis (see Serridentinus mongoliensis) monophyly (see phylogeny) Monroe County, New York, 1137, 1156 Monroe Mastodon, 175 Montabusard, Caleaire de, 5, 115, 122, 123, 192, 193, 199, 200, 204, 217, 223, 518, 736, 1458, 1459, Pl. m1 Montana (see Deep River Beds, Glendive, Smith River, Spokam Bar) Mont-Ceindre, 1462 Monte Bamboli, 115, 201, 1464 Monte Hermoso, Argentina, 385, 537, 590, 743 Monte Verde, 908 Monterey, 902, 1082 Monti Pisani, Tuscany, 1187, 1230 Montpellier, 617, 624, 625, 632-634, 745, 756, 1443, 1446, 1470, 1471 Montredon, 93, 1468 Moodie, Roy Lee, 15, 174, 780 moodiet (see Mastodon moodiet) Moody, C. L., 1509, 1627 Mook, Charles Craig, xvi, 15, 237 Mooknah. See Elephas indicus (Mukna var.) Moore, J. E., 1106, 1107, 1111 Moosburg, Bavaria, 698, Pl. 111 Moosh, 1060 Moravia, 1, 1139, 1169 Moravian Government Museum (see Morayské Zemské Museum) Moravské Zemské Museum, 13, 1139, 1610 Moreno, Francisco J. P., 3, 518, 537, 579, 741, 790, 1399 Morgan, John Pierpont, x Morimentu, 1257, 1266 Morocco, 232 Moropus, 315 Morren, Charles F’. A., 3, 790, 1123, 1136 Morrill, Charles H., 302, 304, 335, 337, 338, 470, 602, 695, 1019, 1025 morrilli (see Eubelodon morrillt) Morrillia, 231, 344, 345, 351, 352, 372, 377, 379, 391, 690, 739, 1381, 1514, 1548, 1549, 1601 barbouri, 226, 307, 345, 349-361, 356, 368, 369, 372, 377-379, 390, 391, 725, 739, 758, 1406, 1408, 1527, 1559, 1604, Pl. vir Morris, Frederick K., 1477, 1623, 1627 Moshach, 1088, 1042, 1044, 1045, 1048, 1050, 1052, 1056, 1065 Moscow (see University of Moscow) Moselle, 1624 Mosonyi, Emil, 432 Mossel Bay, Cape Colony, 1193 Mossom, Stuart, 400, 790 Mostert, John, 1287 INDEX Mt. Angel, Oregon, 1088 Blanco, Texas, 384, 387, 431, 445, 494, 671, 673, 675, 677, 737, 742 Coupet, 625, 630, 632, 634 Eden Beds, 496-498, 508, 561, 1502, 1503 Eden Hot Springs, 479, 496-498, 507, 537, 560, 740, 1406 Hanare, Kuji District, 905 Holly, Vermont, 762, 1623 Kenya, 1202 Léberon (Vaucluse), 114, 1468 Pisgah, Iowa, 1003 Tomuro, 25, 834, 892, 893 Mousterian, 1430, 1432, 1433 Mozambique, 1193 M’Paa, 1193 Mukna. See Hlephas indicus (Mukna var.) Mulberry, 400 Mulberry Canyon, Texas, 677 Mundesley, 1397 Munich, 14, 15, 216, 360, 697, 731, 846, 1314, 1330, 1331, 1609, 1615 Murchison, Charles, 3, 643, 644, 646, 790, 951, 1220 Muride, 680 Murinsel, Croatia, 196, 221 Murphy Canyon, Nebraska, 318 Murray, James Augustus Henry, 790, 1209, 1372 Murree, 448, 449, 1441 sashi Province, Japan, 908, 1489 Musashino (Upper), 907 Musée Académique, Geneva, 475. Geneva Crozatier, Puy, 630, 745, 756 de Bruxelles (see Brussels) d’Orléans, Avaray, 207, 745, 756 du Palais Carignan, 1056, 1154 Museo Argentino de Ciencias Naturales, Buenos Aires, 526, 544, 546, 590, 591, 745, 756, 1609, 1610 de Historia Natural, Mexico, 15 Instituto Nacional geologico, Mexico, 544, 745, 757, 1014, 1081, 1165, 1609, 1615 Nacional Bernardino Rivadavia (see Museo Argentino de Ciencias Naturales) Nacional de La Plata, 745, 757, 1609, 1612 Museos de la Universidad Central del Ecuador, Quito, 15, 745, 757, 1609, 1620 Museum Adolphi Friderici Regis, 1808-1310, 1609, 1620 Muséum d’Histoire Naturelle, Geneva, 474, 475, 477, 505, 551, 552, 745, 757, 1377, 1419, 1609, 1611 d’Histoire Naturelle, Marseille, 14, 15, 1046, 1083, 1609, 1615 des Sciences Naturelles, Lyon, 14, 273, 745, 758, 1063, 1064, 1396, 1609, 1615 National d’Histoire Naturelle, Paris, 13, 14, 138, 134, 166, 196, 213, 252, 254-259, 271, 330, 341, 388, 473, 517, 529, 547, 549, 550, 577, 613, 627, 693, 745, 757, 758, 934, 977, 978, 1035, 1037, 1132, 1133, 1316, 1609, 1619 Museum fiir Naturkunde der Universitit, Berlin, 14, 548, 745, 758, 1609, 1610 Museum of Education, Ochanomizu, Japan, 1298, 1609, 1620. See also Tokyo Imperial University of History, Science and Art, Los Angeles, 1609, 1615 of Natural History, Berlin (see Museum fiir Naturkunde) of the Belgian Congo (see Tervueren) of the Geological Society of London (see Geological Society of London) Sebae, 1310 Museum, St. John, New Brunswick (see St. John, New Brunswick) Muskingum County, 1097 Mutsu Province, 907, 1188, 1333 Muybridge, Eadweard, xvii, 790 Mwelle District, 1193 Mylagaulus, 601 Mylodon, 171, 399, 400, 573, 725, 1520, Pl. var Mylohyus gidleyz, 400 Mysore, 1311, 1319, 1322 mythology (see origin) See also Muséum d’Histoire Naturelle, 1007, 1008, 1013, 1657 Nachtigall Falls, 1193 Nagahama, 818, 906, 1188, 1297, 1420 Naganuma, 902, 1301 Nagasaki, 906 Nagri, 449, 518, 1442-1447 Nahun, India, 624, 625, 648, 650, 740 Nakao, 908 Nakatadotsu-gun, 893 namadi (see Palxoloxodon namadicus namadi) namadicus (see Palxoloxodon namadicus) Namadus River, 1175, 1181, 1211 nana (see Hesperoloxodon antiquus nanus) Nandtr Madméshwar, India, 12138 Nannippus, 1497 Narahara, Ushinosuke, 898, 897, 1301 Narbada (see Nerbudda) Narbonne, France, 93 nares, 916 Nari, 275, 1440, 1442 naricum (see Deinothertum indicum var. gajense) Narita, 902, 906-908, 1298, 1300 Nasmyth, Alexander, 3, 790 Natal, 1402 Nathot, India, 274, 735, 737, 747, 749 nathotensis (see Stegolophodon nathotensis) National Geological Survey of China, 15 Natsume, Mr., 1609, 1619 Natural History Museum, Leiden, 1609, 1612 History Museum, Vienna (see K. K. Naturhistorisches Museum, Vienna) Naturhistorische Staatsmuseum, 360, 361. See also K. K. Naturhistorisches Museum, Vienna Naturhistorisches Museum, Basel, 15 Naturhistoriska Riksmuseum, Stockholm, 14 Naumann, Edmund, 3, 790, 818, 833, 892 naumanni (see Palzoloxodon namadicus nawmannt) Nawngchik (see Bukit Besar) Neals, Florida, 400 Neanderthal hunters, 1168 Nebenhiigeln, 218, 219 nebrascensis. See Archidiskodon (Ocalientinus) nebrascensis Nebraska, 623, 725. See also Ainsworth; Angus, Nuckolls County; Boyd County; Bristow; Brown County; Burge; Cambridge, Furnas County; Christmas Quarry; Devil’s Gulch; Driftwood Creek; Furnas County; Harlan County; Horsethief Canyon; Lancaster County; Loup River; Milford; Niobrara River; North Loup River; Otoe County; Pender; Red Willow County; Sand Canyon; Snake Creek; Snake River, Cherry County; Springview; Talmadge, Otoe County; Thurston County; Trail Canon; Valentine “Nebraska” of Scott, 318 Nebraska State Museum, 745, 758, 1012, 1018, 1025, 1165, 1609, 1613 Nebraskan age, 418, 725, Pl. vu Nebraskan Glacial, 996, 1510 Necrodasypus, 35 Negri Sembilan, Malay, 1332 Nelson, Nels C., xvii, 589 Nelumbium, 717 Neofiber, 400, 680 Neohippus, 573, 1521 Neolithic, 907, 1186 neomorph, 225 Neotomeniz, 680, 1082 neotropical Proboscideans, 1516, 1518 neotype, 9 Nepal, 13138, 1315 Nerbudda, 91, 448, 449, 835, 862, 874, 908, 1175, 1181, 1187, 1211, 1214, 1447, 1449, 1451 Ness City, 1003 Ness County, Kansas, 1001, 1005, 1006 meridionalis nebrascensis, Serridentinus 1658 Nester, Alfred T., 15 Nesti, Filippo, 2, 790, 942, 970, 1392 nestii (see Parelephas trogontherii nestit) Neudorf, 134, 210, 260, Pl. 1 Neusiedler See, 91 Neuville, Henri, 4, 790 Nevada (see Humboldt County, Thousand Creek) nevadanus (see Pliomastodon nevadanus) New Brunswick, 1575, 1609, 1620 Jersey (see Hackettstown) Mexico (see Battleship Mountain, Hot Springs, Ojo Caliente, Santa Cruz, Santa Fe) Siberian Islands, 1590 York Botanical Garden, 15, 717 York State Museum, 14, 15 York Zoological Park, 14, 1194, 1201, 1239, 1598, 1604 York Zoological Society (see New York Zoological Park) new families (see Humboldtide, Serridentide) species (see Anancus properimensis, Deinotherium hopwoodi, Mastodon acutidens, Miomastodon depereti, Pentalophodon falconeri, Rhyncho- therium browni, Stegolophodon lydekkeri, Stegomastodon primitivus, Tetralophodon fricki, Trilophodon hasnotensis) subfamilies (see Humboldtine, Notiomastodontine, Paleomastodontine) subspecies (see Mammonteus primigenius alaskensis) superfamilies (see Stegodontoidea) Newberry, Florida, 400 Newburgh, New York, 179, 190, 1387 Newton, Edwin Tulley, 4, 636, 790 Newton, Richard Bullen, 4, 791 Ngandong, 1454, 1455 Ni Ho Chang, China, 721 Nicol:is de los Arroyas, 520, 579 Nicolueci, Giustiniano, 3, 791, 1241 Niederésterreichische Landessammlungen, Vienna, 15 Nieman River, 1137 Niezabitowski, Edward Lubicz, 4, 791 Nihowan Beds, 1482-1484, 1486, 1487 Niki, 844 Nikolsburg, 91 Nikolsdorf, 210 Nile River, 45, 66, 717, 1426, 1574 Nine Mile Bottom, South Carolina, 175 Ninohe District, 907, 1188, 1333 Niobrara River, 294, 317, 318, 601, 670, 1496, 1497 nipponicus (see Parastegodon nipponicus) Nishiyagi, Japan, 1420 Nizza della Paglia, 1055, 1060, 1061 Noack, Th., 4, 791, 1193 Noback, C. V., 1604 Nodule Bed (Ramsey, England), 635 Nogrsid, Hungary, 85, 116, 735 nomenclature: Cabrera on, 587-589; Cope on, 525; definition of, 21; history of, 135, 1363-1420; of Elephantidw, 1308; of families, 1368; of genera, 1371-1382; of Hesperoloxodon, 1247; of Japanese and Javanese species, 1289; of Loxodontinw, 1173, 1174; of Mammoth (Mam- monteus), 1117, 1863-1367, 1372; of Mastodon, 1117, 1363-1367; of Notorostrin#, 516, 518; of species, subspecies, and varieties, 1382- 1420; of subfamilies, 1869; of superfamilies, 1367; Osborn on, 12, 13, 19, 20; principles and rules of, 5-11, 17, 18, 1363, 1372; Weinsheimer on, 94. See also classification, phylogeny nomina nuda, 480, 493, 625 Nomura, 8., 1301 Nooitgedacht farm, 1287 Noorpoor, 1396 Nopachtus coagmentatus, 551 Nordenskidéld, Erland, 4, 518, 522, 539, 546, 547, 561, 562, 570, 759, 791, 1162, 1165 Nordlingen, 115 Norfolk Bone Bed, 964 OSBORN: THE PROBOSCIDEA Norfolk, England, 964. See also Forest Bed, Norwich Crag Norfolkian, 980 North America, 1490-95, 1507, 1510-1514, 1518 North Carolina (see Halifax County, Tarboro) North Coalinga Beds, 1509 North Loup River, Nebraska, 318 Northern Mammoth (see Mammonteus) Norwich Castle Museum and Art Gallery, 973, 1897, 1609, 1618 Norwich Crag, Norfolk, East Anglia, 620-622, 632-634, 636, 740, 961, 963, 971, 973, 1055, 1155 Notelephas, 1378 australis, 1378, 13897 Nothrotherium, 1082, 1503 Notiomastodon, 12, 33, 381, 515, 536-588, 541, 590, 592, 614, 615, 687, 691, 729, 730, 743, 971, 1381, 1519, 1527, 1572 argentinus, 384, 515, 619-621, 524, 530, 531, 5386-5388, 541, 650, 569, 587, 588, 591, 592, 615, 730, 737, 743, 1399, 1520, 1605 ornatus, 384, 385, 515, 531, 536-5388, 541, 542, 588, 690-592, 595, 614, 615, 700, 731, 743, 756, 757, 1381, 1414, 1519, 1605, Pls. 1x, x1 Notiomastodontine, 381, 384, 515, 581, 541, 542, 688, 689, 691, 730, 743, 1370, 1371, 1526, 1545, 1572 Noto Province, 908, 1489 Notorostrine, 24, 27, 31, 119, 121, 128, 228, 515, 516, 518-574, 588, 612, 615, 689, 690, 722, 734, 740, 1525, 1526, 1528, 1545, 1550, 1560, 1561, Pl. x. See also Cordillerion Notus, 539 Nouel Collection, 207 Nouvelle-Grenade, 530 Nova Scotia, 1575 Novo Elisavetovka, 1468 Novogrodek, Russia, 1137 Nuckolls County, Nebraska, 943, 1033 Nuphar, 717, 719 Nyasa (see Lake Nyasa) Nyasaland, 1432 Nymphaea, 717, 719 ’Nzoi River, Kenya Colony, 1189, 1190 Oak Creek Beds, 305, 1498, 1630 Oak Springs, Contra Costa County, 495 Oaxaca, 942, 1015 Oberdorf bei Weiz, Styria, 1404 Obleoukhow, Mr., 694 obliqua (Loxodonta africana var. obliqua) oblique tuskers, 329, 331, 705, 706, 738 obliquidens. See Ocalientinus (Serridentinus) obliquidens obscurus (see T'rilophodon obscurus) Ocala, 400, 1077, 1495 Ocalientinus, 384, 385, 407, 414-421, 433, 435, 686, 691, 729, 742, 1382, 1527, 1547, 1569, 1570, 1601 emmonsi, 251, 385, 483, 733, 742, 1415, 1508 ojocaliensis, 232-234, 312, 319-323, 326, 327, 384, 385, 387, 388, 391, 406, 433-440, 444, 456, 729, 742, 750-752, 1382, 1417, 1493, 1605, 1617 (Serridentinus) bifoliatus, 385-387, 393, 414, 415, 416, 431, 433, 742, 745, 756, 1413, 1496 (Serridentinus) florescens, 384, 385, 387, 393, 397, 433, 730, 732, 742, 751, 1413, 1481 (Serridentinus) floridanus, 118, 228, 373, 380, 383-388, 391-394, 400, 411, 413, 416-419, 421, 432, 433, 435-437, 440, 455, 469, 501, 539, 742, 748, 759, 1398, 1496, 1605 (Serridentinus) floridanus leidii, 234, 235, 383, 384, 400, 414, 419, 433, 742, 751, 759, 1411, 1496 (Serridentinus) obliquidens, 286, 287, 384-387, 393, 400, 419-421, 432, 433, 742, 746, 748, 759, 1395, 1410, 1508 (Serridentinus) republicanus, 328, 369, 384, 385, 387, 393, 414, 415, 421, 433, 436, 436, 437, 742, 746, 748, 1410, 1500, Pl. vir Ochanomizu, 1298, 1609, 1620 Ochsenvater der Louisianer, 86 octaloph-lophid, 1545 INDEX Odessa, 1466 Odocoileus, 171, 386, 400, 680 odontotyrannus (see Elephas odontotyrannus) (Eningen, 115, 134, 201, 1464-1466 officinalis (see Stegodon officinalis) Ogallah, Kansas, 713, 739, 1418 Ogallala, 1497, 1501, 1507 Ohio (see Jackson County, Zanesville) Ohio, 7, 10, 166, 1872. See Mastodon, Mastodonte de V Ohio, Ohio-Incognitum Ohio River, 135-137 Ohio State University, 14, 15, 164, 169, 758, 1067, 1605, 1609, 1611 Ohio-Incognitum, 6, 7, 135, 136, 165, 168, 171, 1166, 1363-1365, 1372, 1383 ohioticum (see Mastodon ohioticum) ohioticus (see Mastodon ohioticus, Trilophodon ohioticus) Ojo Caliente, New Mexico, 324, 385, 387, 435, 440, 447, 742 ojocaliensis (see Ocalientinus ojocaliensis) okapi, 1467 Okeene, Oklahoma, 1003 Oken, Lorenz, 3, 5, 8, 791, 1364 Okhotsk Sea, 1305 Okimisone, Japan, 818, 906 Okine, 1296 Oklahoma (see Archidiskodon haroldcooki and Stegomastodon priestleyt) Okubo-mura, 1420 Oleott Hill, 426, 427, 492 Old Stone Age, 1163, 1167, 1168 Oldham, 88, 642 Oldoway Tuffe, 1187, 1275, 1432, 1433 Olduvai, Tanganyika, 85, 104, 105, 114, 135, 1419, 1432-1435 oligobunis (see Cordillerion oligobunis) Oligocene, 37, 51, 53, 116, 117, 1421-1426, 1450, 1457, 1493, 1555, Pls. 1, 1 Oliphants River, 1287 Olivola, Italy, 634 Olsen, George, 39, 1524 Omi, Japan, 818, 906, 1489 Omo, 782, 1417, 1436, 1437 Onohippidion, 1518, 1519 ontogenetic cranial changes, 919 Onuki-mura, 906, 907, 1188, 1298 Oort, E. D. van, 15, 1329, 1330 Oppeln, 251, 260, 738, 1463, 1464 Oppenoorth, W. F. F., 15 Optima formation, 1509 Oran, 1195, 1609, 1619 Orange County, New York, 1387, 1602 Orange River, South Africa, 1193, 1198 Orange sand, Dallas, 1399 orarius (see Cordillerion orarius) orbitosphenoid, 917, 920 Oreamnos, 1135 Oregon, 137, 497, 498, 736, 1088 oregonensis (see Mastodon oregonensis) Oreodon culbertsoni, 733 Orient, 1439-1456, 1466, 1487 orientalis (see Stegodon orientalis) origin: African center of, xi, 34-39, 45, 734, 983, 1273, 1422, 1423, 1523, 1524, 1528, 1538, 1580; ancestral types, 18, 1183, 1423; ancestry or adaptive ascent, xii, xv; Archidiskodonts, 948, 950, 996, 1273; Asia, 34; Biblical interpretation, 1; Cretaceous phase, 22, 39, 1553, Pl. x; Cordillerion, 685; Darwinian divergent-evolution principle of descent, 12; diphy- letic theory, 22; dwarfed Mediterranean species, 1271-1273, 1592; elephants, 2, 21, 25, 27, 527, 1801; Elephas indicus, 1307, 1595; Eocene phase, 22, 685, 1553; Eubelodon, 685; Loxodonts, 1287, 1307, 1590; Mastodontine, ancestor of, 692; Meritherium, 12, 24; monophyletic theory, 12, 33; mythological, 1; Palzeoloxodonts, 1180, 1210, 1252, 1258, 1273, 1293; polyphyletic theory, 18, 22; primary stocks, 21, 22; special creation system, 2, 6, 12, 19, 20; Stegodonts, 12, 25, 27, 821, 822, 936; Stegomastodon, 685. See also adaptive radiation, Africa, and various families and subfamilies 1659 Orignac, 1468 Orindan formation, 1509 Orléanais, Marnes del’, 1458 Orléanais, Sables de, 109, 112, 115, 131, 134, 192, 201, 203-207, 217, 223, 250, 252-254, 267, 270, 283, 692, 736, 738. See also Burdigalian, Musée d’Orléans orleanst (see Loxodonta africana orleans?) ornatus (see Notiomastodon ornatus) orthocephaly, 919, 1552 Orycteropus, 715, 716, 1467 osagit (see Tetracaulodon osagit) Osborn, Henry Fairfield: Africa, homeland of Proboscidea (see Africa); Asia, secondary homeland, 34; cranial mechanics (wire sectional method), 915-926; expedition (see Fayfim); families described by, 31, 689, 722, 729, 1368, 1369; genera described by, 131, 157, 212, 353, 377, 393, 543, 575, 654, 994, 1048, 1126, 1179, 1217, 1340, 1358, 1380-1382, 1405-1414, 1416-1419; invasions of African fuana into Europe, 34, 35, 983, 1271; nomenclature, principles of, 6, 1363; origin of Probo- seidea (see Africa); phylogenetic classification (see classification, phylogeny); polyphyletic theory (see polyphyly); publications, 4, 791, 1627; species described by, 85, 137, 251, 259, 279, 285, 304, 349, 372, 375, 377, 384, 385, 396-398, 401, 414, 415, 419, 425, 429, 430, 432, 452-457, 463, 473, 479, 488, 491, 493, 494, 500, 502, 560, 623, 625, 636, 647, 657, 661, 665, 673, 682, 693, 694, 696, 700, 835, 847, 848, 851, 874, 875, 948, 984, 986, 987, 989, 1083, 1101, 1108, 1137, 1157, 1159, 1188, 1245, 13819, 1359; species named for, 64, 244, 298; subfamilies describ- ed by, 27, 30-33, 71, 137, 191-197, 389, 543, 627, 689, 691, 730, 837, 1369-1371; superfamilies described by, 22, 24-27, 30-33, 69, 83, 912, 1367 osbornt. See Phiomia osborni, Trilophodon (Genomastodon) osborni Osborn-Tullberg-Stehlin theory of African origin (see Africa) Osburn, Raymond C., 15, 1067 Ostend, 1220, 1234 Osterode (Harz), Germany, 1122, 1123, 1136, 1141 ostrich (see Struthiolithus) Oswald, Felix, 4, 794 Otisville, New York, 183-185 Otoe County, Nebraska, 696 Otomi, 906 Otranto, Italy, 1137, 1149-1151 Otsuka, Y., 1301 Ottumwan, 1510 Oubangui Chari, Africa, 202 Ouganda, 485 Ouse River, 968 Ovibos, 190, 971, 1134, 1135, 1155, 1161 Ovibos-Rangifer Zone, 1511 Ovts, 1135 Owen, Richard, 3, 88, 90, 95, 620, 794, 831, 860, 997, 1072, 1390, 1397, 1452 Oxford, 931, 973, 1609, 1619. See Peel, C. V. A. Oxydactylus, 315, 318, 386, 400 oxyotis (see Loxodonta africana oxyotis) Oyster beds (Baluchistan), 1441 Ozaki, Tomekichi, 893 Pa P’an Shan, 732, 1418 Pabbi Hills, 845 Pachyderms, 89 pachyganalis (see Elephas primigenius Blumenbach var. n. pachyganalis) paladentatus (see Amebelodon paladentatus) Palearctic Realm, 1477 palxindicus (see Anancus palxindicus, Trilophodon palxindicus) paleobotanical evolution and migration, 717 Palzxocherus, 1459 Paleolithic art and the mammoth, 1, 1168, 1238, 1252, 1285 Palzxoloxodon, 11-18, 22, 25, 894, 901, 904, 907, 909, 913, 914, 984, 986, 1042, 1062, 1173, 1178-1180, 1186, 1191, 1207, 1209, 1210, 1212, 1217, 1221, 1257, 1258, 1273, 1277, 1278, 1289, 1290, 1291, 1292-1305, 1380, 1527, 1541, 1549, 1589, 1591-1593, 1595, 1603 1660 Palxoloxodon—continued (?)andrewst, 943, 984, 985, 993, 1188, 1278, 1305, 1415, 1439, 1542, 1591, 1612, 1618 antiquus, 936, 1247 antiquus (andrewsi?), 1222, 1249, 1250, 1416. antiquus antiquus antiquus, 1247 antiquus germanacus, 1247, 1254 antiquus italicus, 1247, 1416. See Hesperoloxodon antiquus italicus antiquus namadicus (see Palzoloxodon namadicus) antiquus typicus, 1247 antiquus var. insularis (see Elephas antiquus var. insularis) aomoriensis, 1289, 1419, 1542 (Archidiskodon?) tokunagai mut. junior, 943, 1185, 1188, 1289, 1293, 1299, 1300, 1414, 1542, 1612 archidiskodontoides, 984, 985, 1188, 1277, 1282, 1283, 1416, 1439, 1542, 1612 atlanticus, 905, 1180, 1183, 1184, 1187, 1207, 1210, 1263, 1271, 1273, 1274, 1305, 1391, 1397, 1398, 1542, 1593, 1619 ausonius (see Hesperoloxodon antiquus ausonius) buski, 907, 909, 1188, 1315, 1318, 1319, 1333, 1334, 1408, 1412, 1489, 1542, 1620. See also Kanagawa Prefecture creticus, 32, 1183, 1187, 1257, 1259, 1263, 1267, 1269, 1402, 1476, 1542, 1614 cypriotes, 32, 1188, 1187, 1257, 1259, 1263, 1264, 1266, 1267, 1269-1271, 1401, 1476, 1542, 1614 darti, 1188, 1420, 1542 falconeri, 32, 1179, 1182, 1187, 1206, 1252, 1257-1259, 1263-1265, 1267— 1273, 1476, 1542, 1603, 1605 hanekomi, 943, 984, 985, 993, 1188, 1277, 1279, 1415, 1439, 1542, 1612 hysudrindicus, 366, 887, 902, 905, 967, 1186, 1187, 1289, 1302, 1303, 1318, 1403, 1542, 1612, 1620 jolensis, 1188, 1187, 1263, 1271, 1273-1275, 1305, 1400, 1480, 1542, 1593, 1610 kuhni, 984, 985, 1188, 1277, 1281, 1415, 1439, 1542, 1612, 1618 lamarmorae, 1183, 1187, 1257, 1259, 1266, 1267, 1269, 1398, 1476, 1542, 1603 melitensis, 32, 1172, 1173, 1180, 1182, 1183, 1187, 1206, 1214, 1252, 1267— 1263, 1265, 1267-1273, 1395, 1476, 1541, 1542, 1592, 1603, 1605, 1614, 1615 mnaidriensis, 1180, 1182, 1183, 1206, 1252, 1257-1260, 1262-1265, 1267, 1269-1274, 1395, 1476, 1541, 1542, 1603, 1605, 1614, 1619 namadicus, 10, 12, 13, 448, 449, 874, 877, 902, 905, 906, 908, 909, 932, 936, 1172, 1175-1178, 1180, 1181, 1183, 1185-1187, 1191, 1192, 1206-1215, 1217, 1218, 1247, 1249, 1250, 1252, 1268, 1272, 1273, 1290, 1303, 1305, 1340, 1344, 1362, 1392, 1449, 1451-1453, 1455, 1484, 1485, 1487, 1488, 1541, 1593, 1564, 1603, 1605, 1610, 1613, 1615, 1629 namadicus namadi, 1185, 1187, 1289, 1293, 1295, 1296, 1408, 1409, 1413, 1542 namadicus naumanni, 11, 902, 905, 907-909, 1178-1180, 1185-1187, 1289— 1291, 1293-1296, 1305, 1380, 1408, 1489, 1542, 1612 namadicus yabet, 1185, 1188, 1289, 1293, 1299, 1408, 1414, 1489, 1542, 1620 priscus, 905 priscus var. boset, 1188, 1418, 1624 protomammonteus, 306, 909, 1047, 1185, 1186, 1188, 1221, 1289, 1293, 1297, 1299, 1301, 1409, 1411, 1489, 1542, 1619, 1629 protomammonteus proximus, 907, 909, 1047, 1185, 1188, 1289, 1293, 1298, 1299, 1411, 1542, 1619, 1629 recki, 945, 983, 1095, 1184, 1186, 1187, 1271, 1273, 1276-1277, 1283, 1305, 1405, 1433, 1435-1437, 1542, 1593, 1603, 1605, 1610 sheppardi, 943-946, 983-985, 993, 1188, 1277, 1284, 1285, 1411, 1412, 1439, 1542 tokunagaz, 905, 908, 909, 1185, 1186, 1188, 1289, 1291, 1292, 1298, 1305, 1409, 1484, 1489, 1542 transvaalensis, 943-946, 983-985, 993, 1188, 1277-1279, 1284, 1285, 1411, 1412, 1439, 1542, 1618 wilmani, 984, 985, 1188, 1277, 1280, 1281, 1415, 1439, 1542, 1612, 1618 yokohamanus, 1188, 1289, 1301, 1408, 1417, 1542, 1629 yorki, 984, 985, 1188, 1280, 1415, 1439, 1542, 1612, 1618 See also Hesperoloxodon OSBORN: THE PROBOSCIDEA Palzomastodon, 10-12, 28, 29, 36-39, 41-48, 50-52, 54, 59, 63, 65, 77, 128, 131, 132, 137-139, 141, 143, 144, 146, 149, 238, 532, 688, 689, 691, 692, 701, 735, 812, 1879, 1424, 1524, 1527, 1545-1547, 1572-1574, 1602 barroist, 1402. See Phiomia barroisi beadnelli, 10, 36, 41, 44, 50-54, 58, 60, 61, 65, 66, 137, 138, 142, 148, 146—- 149, 164, 238, 244, 246, 247, 735, 746, 747, 754, 755, 1401, 1426, 1600, 1602, 1605, Pl. 1 intermedius, 51-54, 63-65, 77, 137-141, 143-147, 341, 735, 747, 748, 1406, 1426, 1544, 1545, 1616, Pl. 1 minor, 1401. See Phiomia minor minus, 63, 1401 parvus, 41, 51-54, 69, 60, 65, 137, 138, 148, 146, 147, 149, 735, 746, 748, 754, 1401, 1426 wintont, 1401. See Phiomia wintoni Paleomastodontide, 26, 30, 143, 689, 691, 1368, 1545 Paleomastodontine, 27, 30, 36, 65, 119, 128, 131, 137, 143, 689, 691, 735, 1371, 1526, 1572 Palzomeryx, 380, 1461 Paleontological Institute, Vienna (see Paliontologisches Institut der Uni- versitiit, Wien) Museum of the University, Breslau, 745, 754 Society of America, 935 Paliontologisches Institut der Universitit, Wien, 396, 745, 760 Palxoryx, 274 Palzxotragus, 704, 1467 Paleozoologic Institute of the Academy of Sciences, Leningrad, 1418, 1609, 1612 Palembang, Sumatra, 1329, 1330 Palermo Museum, 1260 Pallas, Peter Simon, 2, 87, 138, 192, 794, 1117, 13866, 1389 Palmer, Rupert William, 4, 95, 103, 794 Palmer, Theodore Sherman, 4, 7, 15, 680, 631, 794 Pampas, 527, 576 Pampean Beds, 1516-1519 Pampeano, 520, 537, 579-581, 589, 590, 593-595, 598, 599, 611, 741 Panama, Isthmus of, 515 pandionis (see Trilophodon pandionis) paniscus (see Elephas paniscus) Pao Té Hsien, Shansi, 699, 704, 1480, 1481 Paracamelus, 461 Paraceratherium (Baluchitherium) bugtiensis, 448, 449, 1441 Paraguay, 524, 530, 575 Parahippus, 318, 386, 400, 488 parallelism, 545, 911, 930, 933, 1039, 1177 paramammonteus (see Archidiskodon paramammonteus) Parana River, 527, 595 Paraphiomys pigolti, 1428 Parapithecus, 1425 Paraplatybelodon, 459, 1382 Parastegodon, 816, 818, 823, 835, 837, 838, 840, 853, 893, 897, 901-905, 909, 947, 1380, 1405, 1418 akashiensis, 893, 1420, 1539 aurore (see Stegodon aurore) infrequens, 1420, 1612 insignis (see Stegodon insignis) kwantoensis, 816, 823, 837, 897, 1539, 1629 latidens (see Stegolophodon latidens) mindanensis. See Stegodon (Archidiskodon?) mindanensis nipponicus, 1420 sugiyamai, 816, 823, 837, 899, 900, 1418, 1539, 1629 paratype, 9 paredensis (see Rhynchotherium shepardi edense) Parelephantine, 937, 1370, 1625 Parelephas, 11, 32, 33, 335, 905, 909, 913-915, 923-926, 932, 933, 935, 937, 938, 942, 946, 981, 996, 997, 1039-1048, 1050-1052, 1055-1108, 1133, 1 135, 1146, 1163, 1193, 1228, 1297, 1304, 1322, 1381, 1514, 1527, 1540, 1548, 1582, 1584, 1585, 1587, 1589, 1603 armeniacus, 10, 1042, 1043, 1045-1049, 1055, 1060-1062, 1080, 1233, 1476, 1540, 1614 INDEX Parelephas—continued columbi, 32, 127, 173, 400, 420, 534, 671, 725, 748, 936, 941-944, 946, 996, $97, 1000-1002, 1005, 1013, 1014, 1020, 1041, 1046-1048, 1051, 1052, 1058, 1059, 1067, 1070-1082, 1084, 1085, 1087, 1088, 1090, 1091, 1094, 1105, 1106, 1108, 1140, 1394, 1514, 1541, 1582, 1583, 1586, 1587, 1603, 1605, 1610, 1614-1616, 1619, 1621, Pl. vi1r. See also Amherst Museum columbi cayennensis, 527, 536, 997, 1046-1048, 1067, 1083, 1105, 1414, 1521, 1541, 1615, 1617 columbi felicis, 942, 944, 946, 997, 1015, 1047, 1067, 1080, 1082, 1083, 1407, 1515, 1541, 1612 eellsi, 1047, 1067, 1104, 1411, 1514, 1541 floridanus, 996, 997, 1046-1048, 1051, 1052, 1067, 1070, 1078, 1105-1115, 1414, 1541, 1603, 1605, 1618, 1620, 1621 intermedius, 1040, 1041, 1043, 1045-1049, 1051, 1052, 1062-1065, 1067, 1071, 1135, 1140, 1395, 1476, 1540, 1603, 1605, 1615 jacksoni, 127, 997, 1046-1049, 1067-1069, 1071, 1072, 1083, 1084, 1089, 1389, 1541 Jjeffersonii, 12, 32, 33, 40, 170, 171, 439, 725, 921, 922, 924-926, 931, 932, 938, 939, 996, 997, 1004-1006, 1012, 1020, 1040, 1041, 1043-1048, 1051, 1052, 1053, 1067, 1068, 1070, 1077, 1078, 1083-1097, 1099, 1103, 1109-1114, 1131, 1133, 1135, 1140, 1144, 1163-1165, 1227, 1228, 1277, 1407, 1514, 1541, 1582, 1603, 1605, 1611, 1613, 1615-1617, 1621, PI. vul jeffersonii progressus (see Parelephas progressus) (?) mississtppiensis (?), 1047, 1067, 1068, 1070 progressus, 938, 1040, 1045, 1047, 1048, 1067, 1070, 1084, 1085, 1097-1099, 1140, 1828, 1409, 1541, 1615 protomammonteus (Matsumoto) matswmotoz, 1188, 1289, 1300, 1416, 1542, 1629 protomammonteus (Matsumoto) typicus, 1411, 1420. protomammonteus protomammonteus proxtimus, 1411. proximus protomammonteus proximus proximus, 1411 proximus uehataensis, 1420, 1541 textanus, 127, 942, 943, 1047, 1067, 1068, 1073 trogontherii, 32, 902, 9095, 907, 909, 910, 932, 936, 938, 976, 981, 1038, 1040— 1045, 1047-1052, 1056-1059, 1062-1065, 1067, 1095, 1133, 1135, 1140, 1149, 1155, 1164, 1178, 1186, 1210, 1221, 1251, 1299, 1398, 1408, 1450, 1476, 1489, 1540, 1603, 1605, 1611, 1621 (2) trogontherit nestiz, 981, 1047, 1048, 1059, 1060, 1155, 1209, 1222, 1232, 1233, 1400, 1476, 1540, 1614 trogontheriordes, 1042, 1043, 1045-1049, 1054-1056, 1061, 1067, 1108, 1140, 1400, 1476, 1540, 1620 washingtonii, 939, 1006, 1047-1049, 1051, 1053, 1067, 1068, 1070, 1071, 1084-1086, 1090, 1091, 1100-1104, 1111, 1144, 1408, 1514, 1541, 1587, 1611, 1615, 1621 wiisti, 1047, 1048, 1065, 1066, 1403, 1476, 1540, 1615 Paris Museum (see Muséum National d’Histoire Naturelle) Parkalta, India, 1418 Parker, Rushton, 1222 Parschlung, 92 Parsons, L. E., 1222 Parthenon, 1257 parvus (see Palxomastodon parvus) Pas de Calais, 1251 Patagonia, 1516 Paterson, T. T., 1627, 1629 Pati-Ajam, 625 Pavlodar, Russia, 461 Pavlow, Marie, 4, 15, 126, 132-134, 137, 694, 760, 794, 961, 968, 1058, 1065, 1400, 1403 pavlowi (see Mastodon pavlowi, also Mastodon ohioticus) Pawnee Buttes, Colorado, 403, 479, 488, 737, 742 Pawnee Creek, Colorado, 151, 155, 156, 312, 315, 384, 385, 408, 440, 478, 485, 486, 488, 737, 742. See also Pawnee Creek formation Pawnee Creek formation, 1491, 1492 Pawnee Loup branch (see Platte River) Payette, 1509 See Palzxoloxodon See Palzoloxodon protomammonteus 1661 Peabody Museum (see Yale University) Peace Creek or Peace River, 386, 400, 1495 Peale, Charles Willson, 1602 Peale, Rembrandt, 1602, 1627 Pearson, Helga Sharpe, 732, 794, 1480 peccaries, 308, 400, 497, 498, 562, 579 Peel, C. V. A., 1193, 1609, 1619 peeli (see Loxodonta africana peelt) Pegu, 824, 927, 1450 Pei Hou Kou, China, 699 Pei, W. C., 1451, 1623, 1627 Pekalongan, 366 Pekarna Cave, 1139, 1168 Peking, 876, 869 Pelorvis oldowayensis, 1433 Penck, A., 1473, 1474, Pl. xxiv Pendennis, Kansas, 1088, 1089 Pender, Thurston County, Nebraska, 137, 695, 736 Pennant, Thomas, 2, 7, 168, 794, 1382 pentaloph-lophid, 141, 1545 Pentalophodon, 10, 11, 31, 128, 612, 621, 622, 625-629, 641, 642, 647, 649, 650, 653, 667, 669, 690, 722, 740, 1177, 1376, 1527, 1545, 1549, 1564, 1565, 1601 cuneatus, 1420, 1482 falconert, 448, 449, 622, 624-626, 628, 640-643, 646, 648, 649, 651-653, 720, 740, 746, 754, 1419, 1448, 1604 sinensis (see Anancus sinensis) stvalensis, 10, 31, 228, 348, 355, 448, 449, 532, 539, 621, 622, 625-630, 635, 640, 643, 645, 647-654, 720-722, 740, 754, 756, 1177, 1376, 1389, 1448, 1604 Pentalophodontine, 1370, 1371, 1563, 1628 pentapotamiz (see Deinotherium pentapotamix) pentapotamicum (see Deinotherium pentapotamicum) pentelicus. See Trilophodon (Choerolophodon) pentelicus Peorian, 725, 1510, Pl. vit Pepper, Miss, 644 Peraceras Zone, 305, 369, 1495 Perey, Mabel Rice, xvii Pergamino, Argentina, 520, 537, 580, 593, 597, 741 periboletes (see Elephas pertboletes) pericones, 675 Périgord, 1132 Perim Island, 85, 91, 105, 114, 448, 621, 624, 625, 642, 643, 735, 737, 740, 853, 1445, 1447, Pl. 1v perimense (see Deinotheriwm pertmense) perimensis. See Anancus perimensis, and Mastodon (Tetralophodon) perimensis Perissodactyla, 19, 399, 680, 1429, 1489 Permian-Cretaceous reptiles, 461 Permo-Triassic, 1482 Pérols, 980 Perrier, Auvergne, 632-634, 740, 961 Perry, George, 794, 1323, 1384 Persia, 527. See also Maragha Peru, 523, 530, 554 Pestchana, Russia, 131, 135, 137, 694, 736, Pl. 1 Pestszentlérinez, Hungary, 361, 362 Peters, Carl F., 3, 93, 794 Petersdorf, 1044, 1045 Peterson, Olof August, 4, 15, 251, 290, 312, 385, 794, 1412 Petit Mastodonte, 122, 123, 1384, 1386, 1388. See Trilophodon angustidens minutus, and Turictus tapiroides Petit Rosey, 208 Petite Tartarie, 133, 166 Petékovice, 1139 Petronievics, Branislav, 4, 76, 794, 1408 Peuelchense, 590 Peyrony, D., 4, 795, 1131 Pfalz, 1044 Pfizenmayer, E. W., 4, 795, 1127, 1128, 1131, 1162, 1164, 1627 1662 Philadelphia (see Academy of Natural Sciences and Wagner Free Institute of Sciences) Philippi, Rudolph Amandus, 4, 518, 522, 537, 551, 576, 581, 795 Philippine Islands, 891, 892, 1304, 1456. See also Mindanao Philipps County, Kansas (see Long Island) Phillips, Philip Lee, 4, 136, 795 Phiomia, 10, 11, 18, 28, 29, 31, 33, 39, 41-47, 49-53, 65, 77, 140, 148, 147, 149, 225, 226, 228, 231, 232, 236-238, 247, 277, 334, 465, 686, 690, 715, 739, 903, 1379, 1524, 1527, 1547, 1550, 1555, 1558, 1571, 1600, 1607, 1608, Pl. 1x barroisi, 52, 61, 65, 1402 latidens (see Stegolophodon latidens) minor, 51—54, 58-60, 61, 63, 65, 146, 149, 225, 227, 229, 236-239, 241-245, 739, 746, 747, 754, 1401, 1402, 1426, 1604 osborni, xv, 40, 51-53, 64-66, 142, 146, 225-227, 231, 236-239, 241, 242, 244-247, 276, 278, 305, 321, 330, 334, 336, 341, 352, 359, 465, 706, 715, 716, 739, 746, 747, 755, 1406, 1555, 1604, Pls. v, vr pygmeus, 65, 225, 226, 245-247, 250, 251, 739, 748, 1400, 1426, 1604 serridens, 20, 52, 55, 56, 61, 65, 146, 226, 236, 237, 239-241, 245, 253, 739, 746, 755, 1401, 1426, 1604. See Phiomia wintoni (cf. serridens) stegodontoides (see Stegolophodon stegodontoides) wintont, 38, 40, 43, 48, 51-53, 59-61, 65, 146, 147, 149, 225, 226, 230, 236— 245, 706, 715, 716, 739, 747, 748, 754, 755, 1402, 1426, 1604, Pl. v wintondi (cf. serridens), 240, 241, 244, 246, 255, 346, 739, 747, 1402 Phiomys, 1425, 1428 phippsi (see Megabelodon phippst) Phlaocyon, 315 Phoenicians, 1308 Phosphate Beds, 386, 400, 419, 420, 482, 742, 996, 1072, 1075-1077, 1087, 1106, 1495, 1508 Phyllotillon, 272 phylogeny: Ameghino on, 519-521; Boule on, 527, 532, 533; Cabrera on, 588; Carette on, 519, 523; Cope on, 19, 28, 525, 538; divergence, xii, 12, 686, 911, 919, 933; Fischer de Waldheim on, 517; Lull on, 526, 527; Nordenskidld on, 522; of Amebelodontine, Gnathabelodontine, and Longirostrine, 231, 360, 706; of Brevirostrine, 360, 629, 630; of Bunomastodontide, 228; of Deinotheres, 112; of Stegodonts, 820, 821; of Tetralophodontinz, 360; Osborn on, 119, 538, 539, 689, Pls. X, X1; principles of, xiii, 12, 18, 20, 22, 27, 32, 33, 36, 38, 39, 41, 44; 46, 527, 593, 935; Spillman on, 573. See also adaptive radiation, classi- fication, origin phylum, 19 Physeter, 969 Picard, Emile, 1123 Pichincha, Ecuador, 537, 567, 585, 741 Pictet, Francois Jules, 3, 7,91, 795 Piedmont, Italy, 192, 207, 209, 616, 736, 1047, 1055 Piedmont, Moravia, 1139 Pierce, Florida, 251, 285, 384, 386, 400, 482, 738, 1408 Pierre Shale, 308 Piette, Edouard, 1168, 1169, 1627 Pigafetta, 1118 pigmy (see pygmy) Pignataro Interamna, 1188, 1216, 1234, 1238, 1239, 1245 Pikermi, 91, 92, 114, 192, 199, 200, 212, 214, 218, 220, 250, 262, 638, 659, 736, 737, 1443, 1444, 1468 Pilandsberg, 1188, 1287 Pilgrim, Guy Ellcock, 4, 15, 19, 78, 79, 85, 251, 271, 448, 449, 625, 658, 659, 795, 819, 824, 835, 846, 1212, 1338, 1403, 1404, 1440-1442, 1444, 1446, 1468-1471, 1626-1628 Pilgrimia, 32, 905, 984, 986, 1178, 1179, 1188, 1257, 1273, 1280, 1381, 1592 archidiskodontoides (see Palzxoloxodon archidiskodontoides) kuhni (see Palxoloxodon kuhni) subantiqua (see Lorodonta subantiqua) wilmani (see Palzoloxodon wilmani) yorkt (see Palxoloxodon yorkt) Pilloy, J., 1168, 1169 Piltdown, 964-968, 1163 Pindal, 1131, 1184, 1252, 1595 OSBORN: THE PROBOSCIDEA Pine Creek, Washington, 1047, 1100-1103 Pinellas County, Florida, 400 Ping Fan Hsien, 732 Pinjor, 448, 449, 643, 815, 866, 869-871, 948, 950, 964, 1318, 1442, 1443, 1445, 1447, 1448, 1454 pinjorensis (see Stegodon pinjorensis) Pinkley, George, 1049, 1594 Pinole-Tuff-Orinda, 498, 560, 1509 Pirayui arroyo, Paraguay, 575, 582 ptrayuiensis (see Cuvieronius pirayutensis) Pithecanthropus erectus, 366, 833, 885-888, 966, 968, 1186, 1302, 1303, 1451, 1453 Pits (Snake Creek), 427 Pittbridge, Texas, 479, 501, 537, 541, 559, 737, 740 Pittsburgh, 15. See also Carnegie Museum Piveteau, Jean, 1475, 1628 Pjatovsky, K. J., 1128 plains fauna, 1044, 1045 Plaisancian, 114, 643, 818, 902, 904-906, 1446, 1457, 1469, 1470 planifrons (see Archidiskodon planifrons) Plata, Rio de la, 595 Plate Ltd., 1306 Plate, Ludwig, 4, 795 Platelephas, 12, 913, 914, 1321, 1868, 1882, 1527, 1543, 1589, 1597, 1603 platycephalus, 952, 1307, 1318-1321, 1339, 1340, 1347, 1348, 1352, 1353, 1358-1360, 1361, 1382, 1414, 1448, 1548, 1596-1598, 1616 platensis (see Cuvieronius platensis) Platte County, 1012 Platte River, 947, 998, 999, 1003 Platybelodon, 12, 251, 328, 333, 334, 385, 414, 444, 445, 459-461, 463, 465-469, 687, 691, 714, 715, 729, 730, 743, 1381, 1413, 1478, 1527, 1547, 1550, 1571, 1601, 1608 barnumbrowni (see Torynobelodon barnumbrownt) danovi, 12, 31, 334, 339, 384, 385, 459-463, 471, 472, 718, 748, 756, 1381, 1413, 1469, 1605 grangeri, 31, 118, 251, 328, 332-334, 376, 384, 385, 444, 445, 459-472, 666, 715, 743, 752, 758, 1418, 1478, 1571, 1601, 1605 Platybelodon grangert Zone, 334, 398 Platybelodontinw, 27, 31, 119, 225, 228, 328, 334, 338, 384, 385, 459, 462, 469, 689, 691, 715, 717, 718, 734, 743, 13870, 1526, 1528, 1545, 1570, 1571, LEAL >.< platycephalus (see Platelephas platycephalus) Platygonus, 399, 498, 508, 725, 1082, 1508, Pl. vii platyrhynchus (see Hesperoloxodon antiquus platyrhynchus) platytaphrus (see Elephas platytaphrus) Playa del Barco, Argentina, 590 Pleistocene (see Quaternary) Plesippus, 161, 399, 666, 667, 678-680, 1431, 1504, 1505, 1510, 1518 Pliauchenia, 318, 369, 380, 386, 399, 431, 490, 498, 508, 562, 679 plicatus (see Mastodon americanus plicatus) Pliny, 1118 Pliocene, 263, 318, 386, 399, 1431, 1444, 1446, 1454, 1457, 1465-1472, 1480— 1483, 1489, 1497-1499, 1501-1506, 1516, 1596. See also Pliocéne ancien, Pliocéne récente, Pliocene-Pleistocene boundary Pliocéne ancien, 964, 1469 Pliocéne récente, 633, 634, 964, 1469 Pliocene-Pleistocene boundary, 1049, 1055, 1301, 1431, 1444, 1472, 1473, 1483, 1484, 1504, 1510 Pliohippus, 294, 305, 306, 318, 369, 399, 426, 427, 444, 498, 507, 508, 510, 562, 610, 678, 680, 1498, 1502, 1518 coalingensis Zone, 1503 proversus Zone, 1503 Pliohyrax, 35, 1429, 1467 Pliomastodon, 11, 28, 30, 182, 187, 138, 141, 152, 167, 159, 161, 162, 210, 211, 386, 440, 462, 492, 688, 690, 736, 1881, 1527, 1545-1547, 1575, 1602 americanus praetypica, 114, 133, 137, 138, 140, 141, 151, 154, 159, 160, 210, 396, 692-694, 736, 755, 1406, 1469, Pl. 1 matthewi, 133, 137, 140, 141, 150-154, 166-159, 162, 385, 425-429, 473, 697, 736, 749, 1406, 1500 INDEX Pliomastodon—continued nevadanus, 1418, 1502, 1629 sellardsi, 137, 160-162, 400, 692, 736, 756, 1415, 1496, 1619 vexillarius, 137, 157, 161-164, 692, 736, 760, 1415, 1503, 1602, 1605 Plionarctus edensis, 508 Pliopithecus, 1459, 1461 Plum Creek, 317, 318 Plymouth County, Lowa, 682, 683, 741 Pniel Estate, Vaal River, 1188, 1281 Podolia, Russia, 85, 87, 115, 132, 135, 137, 692, 694, 735, 736, 1136, Pl. 1 podolicum (see Deinotherium podolicum) Pohlig, Hans, 4, 10, 11, 15, 19, 622, 667, 795, 839, 853, 939, 1042, 1056, 1123 1137, 1172, 1248, 1251, 1628 Pojoaque, New Mexico, 320, 435, 440 pojoaquensis (see Trilophodon pojoaquensis) Polgardi, 1468 Polk County, Florida, 285, 415, 480, 482, 692, 738, 742 Polonian, 1473 Polydiskodon, 10, 11, 941, 1123, 1124, 1322, 1365, 1583 Polydiskodonten, 1123, 1378 polydiskodonty, 1159 polyphyly, 12, 138, 22, 33, 41, 527, 686, 820, 821, 933, 935, 936 Pomel, Auguste, 3, 91, 192, 193, 209, 232, 250, 252, 795, 1183, 1184, 1187, 1194, 1195, 1274, 13938, 1397 Pomeroy, Daniel E., 1170 Pompeckj, Joseph Felix, 4, 524, 549, 551, 796, 881 Pondaung, 449, 824, 1450 Pontale von Carini, 1214 Pontecorvo, 1239 Pontian, 107, 114, 159, 160, 360, 361, 698, 699, 704, 739, 899, 902-905, 1268, 1269, 1448, 1445, 1447, 1457, 1465-1468, 1470, 1480, 1482-1484. See also Violet sands Pontier, G., 4, 61, 796, 1628 pontileviensis (see Trilophodon pontileviensis) Pontlevoy, 108, 196, 201, 213, 254, 738, PI. 11 Pontlevoy-Thenay, 253 Ponzi, Professor, 1061 Poppelack, F., 3, 796 Populus alexanderi, 508 Port Elizabeth, 931 Port Williams, Washington, 1047, 1104 Porta San Lorenzo, 1236 Portis, Alessandro, 4, 796, 1400 Post, Texas, 947, 1007 post-metaloph-lophid, 812, 1545 postremus (see Cuvieronius postremus) Postwick, Norfolk, 620 Potwiar District, Punjab, 349, 354, 739, 844, 1447, 1449 Powell, Jefferson County, Nebraska, 1012 Powell-Cotton, P. H. G., 1022, 1193, 1609, 1619 Power River, Wyoming, 1003 Poysdorf, Austria, 395 precursor. See Serbelodon (?) precursor, Stegodon trigonocephalus praecursor praetypica (see Pliomastodon americanus praety pica) Prague-Briinn railway (see Bohemian Museum of Prague) Pray, L. L., 1237 Prebelgranense, 520 precampester (see Morrillia barbourt) Piedmosti, 1139, 1168 Preez, J. du, 1282 premolars, 138-140, 142, 144, 170, 1221, 1547, 1554 Prestwich, 973, 1155 Pretoria, South Africa, 14 Price, Annette L., xvii priestleyt (see Stegomastodon priestleyt) prima (see Loxodonta prima) primzovus (see Elephas primexvus) primary stocks, 21, 22 Primates, 887, 1424, 1426, 1471, 1489 ’ 1663 primigenia, 972 primigenius (see Mammonteus primigenius) primitivus (see Blephas hysudricus primitivus, Stegomastodon primitivus) primordialis (see Elephas primordialis) primus (see Hlephas primus) Prinsep, James, 648 priscus. See Hlephas (Loxod.) priscus, Elephas priscus, Paleoloxodon priscus, Palxoloxodon priscus var. boset private collections, 759, 1609, 1619 privately printed publications, 1371 Proano, Doctor, 532, 574 proavum (see Deinotherium proavum) proavus (see Serridentinus proavus) Proboscidea, xiv, 1, 2, 22, 23, 29, 39, 47, 642, 1147, 1425, 1456, 1490, 1523, 1524, 1527, 1545, 1546, 1594, 1606. See also adaptive radiation, classi- fication, extinction, families, migration, nomenclature, origin, phy- logeny, subfamilies, superfamilies Procamelus, 318, 380, 399, 400, 418, 510, 601 Procamelus-Hipparion Zone, 158, 318, 429, 742 Prochenia edensis, 508 prochinjiensis (see Serridentinus prochinjiensis) procyonids, 400 Prodinotherium, 1381. See Deinotheriwm hungaricum hungaricum (see Deinotherium hungaricum) prod-tuskers, 249, 288, 313, 326, 328-331, 334, 444, 705, 706, 738 productus (see Serridentinus productus) progenium (see Mastodon progenius) progenius (see Mastodon progenius) progressa (see Cordillerion oligobunis progressus) progressor (see Anancus arvernensis progressor ) progressus (see Cordillerion oligobunis progressus, Parelephas Serridentinus progressus, Stegolophodon cautleyt progressus) Prohylobates, 1426 Promastodon, 10, 1379 Prome, 825 Pronothrotherium, 1503 Propalxomeryx sivalensis, 272 propathanensis (see Synconolophus propathanensis) properimensis (see Anancus properimensis) proplanifrons (see Archidiskodon proplanifrons) Propliopithecus, 1424 Propotamocherus, 272 pro-protoloph-lophid, 812, 1545 Prorastomus, 42 Prostren, 42 Prostegodon, 815, 820, 823, 839, 840, 901, 902, 904, 909, 1380 latidens (see Stegolophodon latidens) Prosthenops edensis, 497, 508, 562 Protauchenia, 1521 protoconules (see conules) Protohippus, 204, 298, 306, 318, 369, 399, 403, 424, 429, 510, 1506 Protolabis, 488 protoloph-lophid, 139, 141, 812, 1545 protomammonteus (see Palxoloxodon protomammonteus) Protopithecus, 579 Protragocerus, 274 Prouteaux, M., 5, 796 proversion, 575, 586, 594, 612-615, 621, 627, 649, 667, 668, 670, 1548, 1549 proximus (see Parelephas protomammonteus proximus) Prozeuglodon, 1424 Prussia, 736 Pseudelurus, 1428, 1459, 1461 Pterodon, 1424, 1426 pterygoid, 916 Ptolemaia, 1425 ptychodonty, 1548 ptychodus (see Synconolophus plychodus) P’u Fang Yen, China, 704 Puebla, Mexico, 537, 556, 740, 1013, 1082 progressus, 1664 Puelche, 551 Puelchense stage, 1517 Puerto Madero in Buenos Aires, 537, 581, 741 pullung daunt, 1325 pumilio (see Loxodonta africana pumilio) Punin, Valley of Chalang, Ecuador, 532, 537, 567, 583, 741, 1413 Punjab, 115, 250, 266-269, 279, 349, 354, 362, 642, 650, 737-739, 853, 940, PI. Iv. See also Gadari, Hasnot, Jabi, Lehri, Niki, Potwdr District, Siwaliks punjabiensis (see Tetralophodon punjabiensis) Puntali, 1270 Puschkin, Mussin, 1388 Puy Courny, 114 Puy, France, 618, 630, 632, 633. See also Musée Crozatier Puy-de-Dome, 134, 618 Puy-en-Velay, 625, 631, 632 pygmeus (see Elephas pygmaeus, Phiomia pygmeus) pygmy elephants, 1182, 1184, 1193, 1196, 1215, 1252, 1264, 1271. See also dwarfed elephants pyrenaicus (see Zygolophodon pyrenaicus) Pyrenees, 108 Qsar-el-Sagha (see Fayam) Quackenbush, Lee Schuyler, 4, 796, 1127, 1134, 1135 Quarantaine, La, 1064 quarries. See Ainsworth, Bauer, Christmas Quarry, Devil’s Gulch, Faytim, Snake Creek, Tung Gur Khara Usu Quatal Canyon (see Cuyama formation) Quaternary, or Pleistocene, 190, 725, 1140, 1169, 1183, 1210, 1321, 1339, 1422, 1423, 1429-1439, 1442, 1447-1457, 1470, 1472-1476, 1483-1490, 1495, 1506, 1510-1521, 1630, Pl. vu. See “Tdaho” formation, Pliocene-Pleistocene boundary, Villafranchien Quebrada von Cachihuayco (see Cachihuayco) Quebrada von Chalang (see Chalang) Quenstedt, Friedrich August von, 3, 91, 93, 796 (uercus hannibali, 508 Quercy, 1425 Quetepee, 557 Quex Museum, 1022 Quincy, Florida, 400 Quinn, James, 316 Quinn limestone, 733 Quito, Ecuador, 122, 123, 516, 527, 537, 549, 567-569, 585, 740. Museos de la Universidad Central del Ecuador Qurun, Lake, 51, 63, 1406 also See also Rachoy, Joseph, 92, 796 Rademan, Luke, 1278 radiograph, 1336 Raffles Museum, 1332 Rafinesque-Schmaltz, Constantine Samuel, 2, 136, 192, 250, 252, 537, 796 Railroad Ridge, Humboldt County, 1418 Rainey, Paul, 1202 Raisz, Edwin, J., 279, 650, back end paper Rak, Joseph, 324, 500, 507, 535, 560 raki (see Mastodon raki) Rikoskeresztur, Hungary, 114, 159, 160, 210, 396, 638 rameau de |’ Elephas primigenius, 1049, 1064 rameau de |’ Elephas trogontherii (see Parelephas trogontherii) Ramsey, England, 635 Rancho La Brea, 1007, 1009, 1010 Randan, 964, 980 Randublatung, 894 Rangifer, 171, 190, 1135, 1476 Rapid City, South Dakota, 709 Rappenfluh, 115 Rattlesnake Canyon, Nebraska, 317, 318, 325 Rattlesnake formation, Oregon, 497, 498, 560, 562, 1509 Raven, Henry C., 1205 OSBORN: THE PROBOSCIDEA Rawlins County, Kansas, 342, 369, 739 Ray, John, 2, 796,1118, 1308, 1309, 1371 Raymond Collection, 416, 431 Reading, England, 14 Reartes, Valley of, 589 Réaumur, René Antoine Ferchault de, 2, 84, 86, 796 Rebling Museum, 1056, 1609, 1620 Recent, 1510 Reck, Hans, 4, 796, 1276, 1286, 1433 recki (see Palxoloxodon reckt) rectidens (see Rhynchotheriwm rectidens) rectigradations or aristogenes (see aristogenes) rectus (see Cuvierontus rectus) Red Clays, China, 458, 742 Red Crag, 632-636, 963, 964, 1055, 1056, 1155, 1472 Red River, Texas, 429 Red Willow County, Nebraska, 372 Reeds, Chester Albert, 4, 15, 796, Pl. xxiv Rees, H., 1278 Refugio, Texas, 563, 565 Reichenberg, Bavaria, 86 Reid, Clement, 1155 Reiss, Wilhelm, 3, 796 Rembang, 894 Rennes, 91 Renshaw, Graham, 15, 1314, 1331 Repelin, Jules Joseph, 15, 1046 Republican City, Harlan County, 338, 1012 Republican River Beds, Kansas, 328, 349, 369, 372, 400, 414, 471, 479, 489, 490, 498, 560, 737, 739, 742, 1500, Pl. vi republicanus. See Ocalientinus (Serridentinus) republacanus restorations. See Abel; Andrews, Charles William; Barbour; Breuil; Capitan; Christman, Erwin; Dolan; Flinsch; Gregory; Knight; Madeleine; Peyrony; Sterling retroversion, 575, 613, 614, 1548, 1549 Reuss, August Emmanuel, 3, 91, 796 Révil, Charles, 1064 Revilliod, Pierre, 4, 15, 474, 537, 551, 552, 796 Reyes, Alicia E., 4, 15, 796, 1013, 1080, 1081 Reynolds, Nebraska, 1003, 1017, 1018 Rhabdobunus, 622, 667, 668, 1879, 1394 mirificus, 1404, 1409. See Stegomastodon mirificus Rhagatherium, 1424 Rheinheimer, Philip, 307 Rheinhessen, Germany, 223, 281, 282, 362, 736, 737, 739, Pl. 111 Rheinsande, 1044, 1045 Rhenoster spruit, 1287 Rhinoceros antiquitatis, 1162, 1164 etruscus, 633, 964, 1210, 1475. See also Rhinocerotidie elruscus var. astensis, 637 kendengindicus, 887 leptorhinus, 633 megarhinus, 1476 merckit, 1210, 1237, 1240, 1253 palxindicus, 616 sinensis, 818 swwasondaicus, 887 sondaicus, 887 tichorhinus, 1475, 1476 unicornis, 1449 Rhinocerotidx, 19, 46, 298, 385, 386, 400, 448, 467, 508, 510, 616, 633, 704, 1044, 1056, 1065, 1155, 1210, 1340, 1476. See also Rhinoceros Rhodesia, 1188, 1193, 1420, 1432 rhomboides (see Mastodon rhomboides) Rhone (Montpellier), 634 Rhone Valley, 1062 Rhynchorostrine, 24, 27, 28, 31, 128, 228, 230, 400, 475-483, 485-513, 539, 689, 690, 731, 734, 737, 1362, 1370, 1526, 1528, 1551, 1561, 1562, Pl. x. See also Muséum d’ Histoire Naturelle, Geneva INDEX Rhynchotheriine, 731, 1370, 1561. See also Rhynchorostrine Rhynchotherium, 10, 11, 83, 260, 399, 426-428, 440, 447, 474-489, 491-503, 505, 507, 511, 513, 5338, 537-539, 544, 545, 690, 737, 807, 1377, 1378, 1527, 1561, 1562, 1601, Pls. x, x1. See Muséum d’Histoire Naturelle, Geneva angutrivale, 157, 385, 425-429, 473, 479, 480, 483, 491-493, 502, 511, 737, 749, 751, 1411, 1500 brevidens, 31, 228, 404, 447, 476-480, 482, 483, 485-489, 493, 502, 511, 539, 737, 746, 748, 1399 browni, 474, 476, 478-484, 489, 493-495, 503-505, 507, 508, 611, 512, 533, 545, 558, 737, 748, 1405, 1419, 1506, 1604 chinjiense, 448, 476, 479, 480, 483, 493, 495, 502, 503, 611, 513, 737, 749, 1414, 1448 chinjiensis (see Rhynchotherium chinjiense) (Dibelodon) edensis, 1411, 1413. See Rhynchotherium shepardi edense dinotherioides (see Trilophodon dinotherioides) euhypodon (see Blickotherium euhypodon) falconert, 230, 232, 399, 479, 480, 482-484, 488, 489, 494, 495, 500, 503, 504, 509, 611-513, 545, 595, 737, 746, 1408, 1494, 1504, 1604 francisi, 479, 480, 483, 501, 502, 737, 759, 1410, 1514 paredensis, 479, 496, 498-6501, 507, 511, 748, 1413 proavus (see Serridentinus proavus) rectidens, 479, 480, 483, 488-489, 493, 737, 1408, 1492 serpentirivale, 479 shepardi, 127, 476-480, 482-484, 487-489, 493, 495, 501, 511, 525, 532, 534, 539, 737, 751, 753, 760, 1507 shepardi edense, 228, 230, 232-234, 479, 480, 483, 493, 496-508, 510, 512, 562, 737, 748, 749, 759, 760, 1406, 1411, 1503, 1507, 1601, 1604. See also Cordillerion edensis, Rhynchotherium paredensis spenceri, 115, 261, 476, 479, 480, 483-485, 493, 503, 504-512, 513, 737, 1405, 1427, 1604 tlascalx, 10, 31, 118, 228, 229, 474-480, 482-483, 493, 503-505, 507, 512, 539, 737, 753, 757, 1082, 1405, 1419, 1506, 1604, 1611 Riazan, 134 Ricardo, 385, 387, 444, 498, 560, 718, 742, 1446, 1502, 1629 Richardson County, 1012 Richardson, John, 3, 137, 796, 1393 Richthofen, von, 355, 1487 ridge-crests or plates, evolution of, 139, 352, 675, 809, 812, 1138, 1140, 1141, 1199, 1221, 1270, 1271, 1325, 1348, 1547 Ries (Noérdlingen), 115, 201 Riggs, Elmer Samuel, 15, 568 Rignano, 1232 Rijks-Museum van Natuurlijke Historie, Leiden (see Leiden) Rikuzen, Province of, 280, 738, 845, Pl. 1v Rio Desaguadero, valley of, Bolivia, 537, 552, 1415 Riobamba, Ecuador, 532, 572, 583 riograndensis. See Trilophodon (Tatabelodon) riograndensis Riss, 725, 1271, 1430, 1457, 1473, 1474, Pl. vir Riuge, Japan, 818, 906 Riukiu, 1305 Rixdorf, 1045, 1050 Roccasecea, 1239 Rochecardon, valley of, 208 Rochester, Indiana, 137, 185, 186, 692, 696, 736, 1137, 1157, 1158, Pl. 1 Rochester, New York, 1137, 1156 rod-cones or tubules, dentinal, 334, 338, 459, 465, 715, 716, 1570, 1571 Rodentia, 23, 26, 34, 298, 467, 579, 601, 678, 680, 887, 1425, 1428, 1489 Rothliche Sande, 384, 731, 817, 885 Roigneau, Marcelle, xvii Roman, Frédéric, 4, 797, 1459, 1628 Rome, 1351, 1609, 1620. See also University of Rome Romer, Alfred Sherwood, 4, 797, 1431, 1511, 1512, 1628 Roosevelt, Kermit, 1189, 1190 Roosevelt, Theodore, 4, 29, 797, 1175, 1190, 1192, 1193 roosevelti (see Elephas roosevelti) Rosebud, 400 Rostock (see University of Rostock) Rothe Tone, 817, 885 1665 Rothschild Museum, 1193, 1609, 1619 rothschildi (see Loxodonta africana rothschildi) Rotterdam (see Zoological Garden, Rotterdam) Rouault, Marie, 3, 91, 797 Roussillon, France, 618, 1446, 1470, 1471 Roverto, Cayetano, 1517, 1628 Royal College of Surgeons, London, 1609, 1615 Museum of Geology and Mineralogy, Leiden (see Leiden) Society, London, 1118, 1119, 1181 University of Bologna (see University of Bologna) Royans, Molasse du, 115 Rozier, Francois, 2, 84, 86, 797 Roziéres, 1065 Rudolf, or Rudolph, Lake (see Lake Rudolf) Riitimeyer, Ludwig, 3, 34, 797 rugatum (see Mastodon rugatum) rugosidens (see Mastodon americanus rugosidens) Rumania, 111, 1466. See Colintina, Deinotherium gigantissimum, Giiceana, Ilfov, Tulucesti (Covurlui), University of Bucharest rumanus (see Archidiskodon planifrons rumanus) Rusconi, Carlos, 1517, 1628 Russell, Kansas, 1003 Russell, R. Dana, 1505, 1628 Russia. See Altai region, Belomechetskaja, Bessarabia, Bug River, Calomna, Caucasus, Eldar, Gavanosy, Gobi, Ishim, Ismail, Kamenez-Podolsk, Karaturgaj, Kuban region, Leningrad, Loh formation, Pavlodar, Pestchana, Petite Tartarie, Podolia, Semipalatinsk, Stavropol, Tung Gur Khara Usu, Ural Mountains, Volga, Volhynia, Zwenigorod ruthenensis (see Elephas antiquus mut. ruthenensis) Rye Valley, Oregon, 173, 736 Saalekiese, 1045 Sables de Chitenay (see Chitenay) Sables de la Sologne (see Sologne) Sables de l’Orléanais (see Burdigalian, Orléanais) Saboyama (see Sawoyama) sabre-tooth tiger, 708, 709 Saffron Walden Museum, 931, 1215, 1234 Sagami, 908, 1489 Sagawa Town, 908 Saghatherium, 1424 Sahamma, 908, 1187, 1295 Sahara Desert, 1194, 1195 Saheki, Shiré, 1188, 1289, 1300, 1411, 1416, 1628 Sahendi (see Mastodon Sahendt) Saida village, 899, 900 St. Andrea, Italy, 1123 St. Chinian, 93 St. Claire, 1063 St. Cyr, 1063 St. David, Arizona, 679 St. Foy, 1064 St. Frajou, France, 205 St. Gaudens, 115, 201, 219, 257, 1464-1466 St. Gerand-le-Puy, 1458 St. Hilaire (see Geoffroy) St. Jean de Bournay, 115, 263 St. John, Mr., 1149 St. John, New Brunswick, 1334, 1609, 1620 St. Just, 1064 St. Lary, France, 86 St. Louis, 14 St. Louis and Brownsville Railroad, 563 St. Martial, 961 St. Prest, 969 St. Prestien, 634, 961, 982, 1430 St. Rambert I’Ile Barbe, 1064 St. Romain au Mont d’Or, 1063 St. Simon’s Island, 1077 1666 Sainzelles, Puy, 634 Sal forests, 927 Salem, Massachusetts, 1599 Salensky (see Zalensky, Vladimir) Saline County, Nebraska (see Crete) Salix coalingensis, 508 Salles-la-Source, 1420 Salmendingen, 91, 1468 Salonica, 1468 Salt Creek (see Jackson County) Salt Range, 85, 105, 266, 268, 269, 275, 279, 363, 616, 642, 845 Salta, Argentina, 510, 550, 596 Saltillo, 1082 Salto, Argentina, 599 Sammlung Mexicanischer Alterthiimer (see Uhde Collection) Samos, 35, 114, 261-266, 737, 1448, 1468 Samotherium, 1467 Sampoeng Zone, 1454, 1455 “Samson,” 1337 Samwel Cave, Shasta County, 1082 San Bernardino County, California, 444, 479, 496, 537, 560, 737, 740, 742 San Chuang, China, 721 San Felipe de Austen, Texas, 250, 251, 349, 374, 738, 1047 San Francisco area, 1509 San Francisco River, Brazil, 527, 578 San Ildefonso, New Mexico, 408, 440 San Isidro, Spain, 201, 219, 1187, 1231, 1232, 1400, 1464 San Joaquin Valley, California, 497 San José de Pimas, Sonora, 474, 479, 493, 494, 737, 1033, 1419 San Miguel Island, 1032 San Nicolas de los Arroyos, Province of Buenos Aires, 520, 537, 579, 592, 741 San Pablo, 902 San Pablo Bay, 1509 San Paolo de Villafranea, 634, 1042, 1055, 1056, 1060, 1061, 1137, 1154, 1232 San Patricio County, Texas (see Sinton) San Pedro Valley, Arizona, 565, 623, 624, 678-680, 682, 683, 1504. See also Benson, Curtis Flats, Curtis Ranch San Romano, 1187, 1214, 1215, 1232 San Teodoro, 928, 1270 San Timoteo, 496-498, 560, 1502 Sanaga River, South Africa, 1193 Sanborn, Elwin R., xvii, 1194, 1259 Sand Canyon, Nebraska, 338, 743 Sanderson, George P., 4, 29, 797 Sandford, Kenneth Stuart, 4, 797 Sang Chia Liang Kai, Shansi, 704 Sanga-jurach mammoth, 1131 Sangamon, 170, 725, 1510, Pl. vin Sangre de Cristo Mountains, 1492 Sankawa valley, 906 Sanmenian, 1452, 1482, 1484-1486 Sanna-Solaro, J. M., 93, 797 Sannoisian, 903 Sansan, 87, 115, 213, 233, 246, 252, 256, 257, 259, 264, 271, 1459-1461 Santa Barbara channel, 1031 Santa Chiara monastery, 1204 Santa Cruz, California, 1032 Santa Cruz de la Sierra, Bolivia, 577 Santa Cruz, New Mexico, 251, 322, 323, 326, 385, 440, 738 Santa Fé de Bogota, 530, 544, 593 Santa Fé, New Mexico, 251, 289, 319, 320, 323, 324, 326, 380, 384, 387, 404, 408, 433, 435, 436, 440, 447, 711, 738, 742, 743, 1435, 1491-1493, 1501, Pl. vir Santa Margarita, 902 Santa Rosa Island, 943, 1031, 1032 Santee Beds, 171 Sanuki Province, 893, 906, 1187, 1296, 1300 Sadne, France, 425, 634 Sappa Creek, Kansas, 342, 349, 369, 739 OSBORN: THE PROBOSCIDEA Sapporo, 907, 1333 Sarasota County, Florida, 400, 1106 Sardinia, 1182, 1183, 1187, 1257, 1266 Sarmatian, 257, 458, 461, 732, 902, 903, 1443, 1447, 1457, 1464-1466, 1480 Sarmato-Pontian fauna, 1468 Sasage bed, 1301 Saskatchewan (see University of Saskatchewan, Wood Mountain gravels) Sato, Denzo, 4, 797, 818 Saugus-Tulare, 902 Saussure, H., 475 Savin, Alfred C., 1155, 1611 Savin Museum, 963, 980, 981, 1609, 1611 Sawoyama, 902, 905 Saxe-Weimar, 1245 Saxonian-Mindel, 1057, 1473 Saxony, 5, 122, 123, 193, 250, 252, 518, 738, 1464 Scandinavian glaciation, 971, 1155 Seanian, 1473 scaptobelodonts (see prod-tuskers) Sceleto Elephantino Tonne, 1118, 1138, 1181, 1217 Scelidotherium, 551 Schinz, Heinrich Rudolf, 3, 124, 192, 193, 213, 219, 736, 797 Schlagel Creek, Nebraska, 318 Schlegel, Hermann, 3, 797, 930, 931, 1312 Schlesien, 114, 195, 737, 835, 846, 1044, PI. 1v Schlesinger, Giinther, 4, 15, 19, 95, 101, 120, 121, 195, 198, 234, 261, 262, 349, 360, 384, 394, 621, 629, 630, 638, 797, 835, 846, 941, 961, 1403 Schlesische Friedrich-Wilhelms-Universitit, 15 Schlosser, Max, 4, 15, 61, 74, 197, 212, 384, 457, 735, 742, 797, 817, 819, 822, 885, 1401, 1424 Schlotheim, Ernst Friedrich von, 3, 795, 1123, 1136, 1387 Schmidtgen, O., 4, 798, 1042, 1044, 1095 Schneider, Anton, 160, 161 Schneider, Carlos Oliver, 1628 Schonnbrunn, Oberbayern, 360 Scholl, Johann Baptist, 3, 349, 357, 358, 798, 1388 School of Mines (see Escuela de Minas, Mexico; South Dakota) Schouteden, Henri, 4, 798, 1193 Schroeder, Henry, 1411, 1628 Schuchert, Charles, 4, 9, 798 Schultz, C. Bertrand, xvii, 348, 669, 682, 725, 726, 786, 798, 942, 946, 1027, 1507, 1623, 1628, Pl. vii Schultze, Leonhard, 29, 798 Schwabe Collection, 1057, 1609, 1620 Sciuride, 680 Sciuropterus gibberosus, 260 Sclater, Philip Lutley, 3, 29, 798, 930 Sclater, William Lutley, 4, 29, 798, 1175 Sclerocalyptus cordubensis, 551 Scott, Robert, 785, 1038 Scott, William Berryman, 4, 798, 984, 1184, 1187, 1286, 1367-1369, 1371, 1402, 1437, 1493, 1628 scolti (see Archidiskodon imperator scottz) Seattle (see University of Washington) Seba, Albert, 2, 798, 1310, 1323 Sebastopol, 1445, 1465, 1468 Sebu, 1195 Second High School, Sendai, 1299, 1609, 1620 section names, 10, 249 sections, 937, 945, 985 secundarium (see Deinotherium secundarium) Seguenza, Luigi, 4, 798 Seki-mura, 906 Selenka-Blanckenhorn Expedition, 886 Sella turcica, 920 Sellards, Elias Howard, 4, 15, 160, 400, 798, 996, 1005, 1078, 1079, 1108 sellardsi (see Pliomastodon sellards?) selousi (see Loxodonta africana seloust) Seminole, Pinellas County, Florida, 400 INDEX Semipalatinsk, Russia, 461 Semur, 980 Senckenberg Museum, Frankfort, 180, 181, 745, 759 Sendai, Japan, 14, 15, 1609, 1620 sendaicus (see Trilophodon sendaicus) Seneca, Thomas County, Nebraska, 699, 741, 942, 998, 999, 1003 Senegal, 1197 Senéze, 961, 964, 980, 1024 Senkaibo, 1420 Senodon (see Mastodon Senodon) Serbelodon, 326, 384, 385, 414, 441, 443, 444, 445, 686, 691, 717, 729, 730, 742, 1382, 1527, 1570, 1601 barbourensis, 317, 385, 387, 391, 436, 440, 441-445, 447, 733, 742, 752, 1382, 1417, 1498 burnhami, 118, 385, 387, 441, 444, 445, 718, 719, 742, 749, 1417, 1502, 1605 (2?) precursor, 384, 387, 393, 399, 431, 432, 445, 742, 748, 759, 1400, 1501, 1504 Serebryakoy, A. H., 1628 serpentirivale (see Gomphotherium serpentirivale) serrate crests or spurs, 156, 383, 393, 687, 689, 729, 730 Serrengetisteppe, 1187, 1275-1277 Serres, Pierre Marcel Toussaint de, 3, 617, 634, 798 serridens (see Phiomia serridens, Serridentinus serridens) Serridentide, 26, 30, 119, 128, 225, 381, 384, 385, 389, 432, 515, 531, 541, 590, 614, 687-689, 691, 729, 741-743, 1368, 1369, 1525, 1568, 1569 Serridentine, 27, 31, 33, 126, 128, 156, 225, 228, 326, 333, 381-391, 393, 414, 433, 446, 451, 469, 515, 615, 689, 691, 731, 734, 741, 1370, 1525, 1528, 1545, 1568-1570, Pl. x Serridentinus, 11, 198, 223, 229, 252, 256, 278, 280, 328, 341, 367, 381-389, 391, 392, 393, 394, 401, 414, 423-482, 440, 451-458, 483, 489, 492, 513, 544, 687, 688, 691, 729, 730, 741, 742, 1380, 1527, 1547, 1569, 1601 angutrivalis, 157, 158, 384, 385, 387, 393, 414, 425-429, 473, 492, 742, 748, 749, 1410, 1500 annectens, 115, 251, 280, 384, 393, 457, 458, 742, 901, 903, 909, 1409, 1489 barstonis, 251, 319, 384, 385, 387, 447, 742, 750, 1417, 1495 bifoliatus. See Ocalientinus (Serridentinus) bifoliatus brewsterensis, 383-387, 393, 400, 430, 431, 440, 742, 745, 1410, 1496 browni, 31, 272, 280, 384, 385, 387, 393, 448, 450-455, 457, 732, 733, 742, 749, 750, 1410, 1448 chinjiensis, 384, 385, 387, 393, 448-452, 456, 457, 458, 742, 749, 1413, 1448 filholi, 256, 258, 384, 385, 393, 473, 742, 757, 1417, 1460, 1619 florescens, 1413. See Ocalientinus (Serridentinus) florescens floridanus. See Ocalientinus (Serridentinus) floridanus fricki, 1412. See Trilophodon frickt gobiensis, 333, 334, 384, 385, 387, 388, 393, 398, 399, 742, 752, 1416, 1478, 1605 guatemalensis, 384, 385, 387, 393, 432, 527, 742, 748, 1411, 1516 hasnotensis, 384, 385, 387, 393, 448, 450-452, 456, 457, 741, 749, 750, 1413, 1448 lydekkeri, 31, 384, 393, 451, 456-458, 731, 742, 817, 1401, 1482, 1615 metachinjiensis, 384, 385, 387, 393, 421, 448, 450, 451, 452, 454-456, 741, 749, 1413, 1448 mongoliensis, 31, 115, 228, 384, 385, 387, 392, 393, 396, 421, 461, 742, 749, 1408, 1477 obliquidens. See Ocalientinus (Serridentinus) obliquidens (Ocalientinus) nebrascensis, 157, 317, 384, 385, 387, 393, 426-429, 473, 492, 623, 625, 733, 742, 749, 751, 752, 1500 (?) precursor. See Serbelodon(?) precursor proavus, 127, 153, 156, 270, 384, 385, 387, 393, 394, 401, 403, 404, 413, 456, 472, 478, 488, 489, 526, 532, 742, 746, 1492, 1605 prochinjiensis, 384, 385, 387, 393, 413, 448, 450, 451, 452, 456, 457, 742, 749, 1413, 1448, 1605 productus, 11, 31, 127, 227-230, 232-235, 244, 246, 305, 326-328, 359, 380, 382, 384, 385, 387, 388, 390-394, 399, 402-413, 435, 436, 440, 441, 451, 455, 456, 473, 483, 486, 489, 508, 526, 527, 532, 538, 539, 687, 742, 746, 748, 750, 751, 759, 1493, 1501, 1546, 1601, 1605, 1615 progressus, 227, 248, 251, 298, 328, 381, 382, 384, 391, 393, 401-403, 407, 729, 742, 746, 1408, 1500 republicanus, 1410. See Ocalientinus (Serridentinus) republicanus 1667 serridens, 11, 31, 227, 228, 245, 246, 367, 382, 384, 387-389, 393, 399, 404, 405, 410, 411, 422-425, 429, 430, 440, 441, 451, 456, 483, 526, 532, 742, 746, 748, 1398, 1501, 1506, 1605 serridens cimarronis, 384, 387, 429, 430, 439, 456, 742, 759, 1400, 1501 serridens life zone, 422 simplicidens, 1403. See Trilophodon simplicidens sublapiroideus, 382, 384, 385, 387, 393-396, 413, 451, 455, 732, 733, 742, 745, 760, 903, 1405, 1605 wimani, 250, 384, 458, 731, 732, 742, 1418, 1480, 1482 Setit River, Abyssinia, 1200 seloensis. See Elephas (Palxoloxodon) namadicus setoensis Seward County, Nebraska, 736, 743, 1012, 1025, 1026 Shackelford, J. B., 396 Shan Plateau, 1451 Shanafelt, Marjorie, 1023 Shang Dynasty (see Anyang) Shanghai, 817, 860, 1479 Shansi, 458, 625, 699, 704, 721, 739, 740, 817, 853, 885, 964, 1164, 1418, 1420, 1451, 1482, 1487 Shantiiin, 1271 Shawangunk, 175, 183 Sheep Creek, 400, 426, 427, 1499 Shensi, 458, 699, 853 Shepard, C. N., 487, 488 shepardi (see Rhynchotherium shepardi) shepardvi. See Rhynchotherium falconert, Tetracaulodon (Tetrabelodon) shepardii sheppardi (see Palxoloxodon sheppard) Sherborn, Charles Davies, 4, 15, 798, 1122, 1308, 1383, 1408 Sheridan County Mammoth, 1011, 1012, 1093 Sheridan or Equus Beds, 372, 373, 1011, 1161 Shichinohe-gawa, Japan, 1419 Shikama, Tokio, 1411, 1420, 1628 Shikoku, Japan, 837, 899, 900, 1305 Shimek, Bohumil, 4, 682, 798 Shimo6sa, 908 Shimoosa, 1489 Shimosuyeyoshi beds, 1301 Shimotsuke, 1489 Shinano, 1489 Shirado, 902 Shiwogama, 845, 905, Pl. 1v Shoédo Island, 818, 835, 893, 906, 1187, 1296 shodoénsis. See Stegodon orientalis shodoénsis short-jawed Bunomastodonts (see Brevirostrine) shovel-tuskers, 328, 330-332, 334, 445, 470, 706, 715, 717. dontine, Serbelodon, and Trobelodon Sh6zu-shima, 908 Shuler, Ellis W., 1628 Shui Chiian P’u, China, 702, 1418 Siam, 1322 Sian, China, 702, 1418 Siberia, 198, 215, 1091, 1117-1120, 1122, 1136 sibiricum (see Mammut sibiricum) sibiricus (see Mammonteus primigenius stbiricus) Sibut, Fort, 202 Sicilian, 906-908, 1050, 1065, 1182, 1430, 1469, 1470, 1473, 1474 Sicily, 1182, 1183, 1209, 1257, 1261, 1269 Sierra de Cérdoba, Argentina, 589 Sierra Leone (see Gola forest) Sierra, Lorenzo, 4 Siestan formation, 1509 Sihaf valley, 1440 Silesia, 134. See also Schlesien Silhet, 1315 silvestris (see Archidiskodon imperator silvestris) Simba, 1170 Simionescu, Ioan, 5, 798 Simla, 449, 648, 650, 940, 1338, 1339, 1358, 1359 See also Amebelo- 1668 OSBORN: THE Simorre, 5, 254, 259, 340, 518, 736, 738, 1463, 1466 simorrense (see Turicius turicensis simorrensts) simorrensis (see Turicius turicensis stmorrensis) simplicidens (see Trilophodon simplicidens) Simpson, George Gaylord, 5, 15, 39, 400, 440, 482, 590, 595, 725, 798, 1107, 1159, 1222, 1368, 1369, 1374, 1411, 1416, 1420, 1495, 1500, 1525, 1607 Sinanthropus, 1451, 1485 Sinclair Draw, 426, 427 Sinclair, William John, 4, 798 sinclairi (see Amebelodon sinclairi) Sind, 85, 93, 105, 115, 250, 251, 266-269, 449, 738, 844, 853, 1440, 1442 sindiense (see Deinotherium sindiense) sinensis. See Anancus sinensis, Stegodon sinensis, Tetralophodon (Lydekkeria) sinensis Sining Fu, 702, 1418 Sining Hsien, 732 Sinker Creek, 1505 Sinkiang Province, 718 Sino-Malayan fauna, 1451 Sinopa, 1424 Sinton Collection, 565 Sinton, San Patricio County, Texas, 535, 537, 562-565, 623, 740 Sinzow, J. F., 1065 Sioux County, Nebraska, 137, 157, 384, 385, 425, 427, 429, 473, 479, 491, 492, 736, 737, 742 Sirenia, 35, 39, 40, 47, 89,91. See also Cryptomastodon martini, Eotherium Sismonda, Eugenio, 3, 618, 620, 637, 798 Siswan, 868-870, 952, 954, 955, 986, 988, 1339, 1340, 1347, 1358, 1359 Sivacherus, 272 Sivalikia, 901, 905, 1178, 1179, 1207, 1209, 1214, 1215, 1380, 1592 Siva-Malayan fauna, 1451 Sivapithecus indicus, 274 Sivatherium olduvaiensis, 1435 Siwalikiensis (see Leith-Adamsia Siwalikiensis) Siwaliks, 268, 273, 274, 278, 279, 348, 363, 385, 448, 449, 451, 454, 456, 502, 616, 621, 622, 625, 642, 643, 645-648, 650, 653, 656, 657, 660, 661, 664, 665, 737, 740-742, 808, 852, 853, 934, 936, 940, 943, 1339, 1340, 1347, 1349, 1442-1444, 1446, 1447, 1449, 1451, 1487 Six-mile Swamp, 1077 Sjara-osso-gol, 1487 skeletal material, 48, 69, 70, 88, 91, 97, 99-101, 112, 113, 116, 130, 147-149, 163, 164, 178-181, 439, 678, 682, 1079, 1600-1603 Skinner, Morris, xvii, 316, 726, 728 Skull Springs Beds, Oregon, 1509 Sloane, Hans, 2, 799, 1118, 1119, 1165, 1181 Smendou, Algeria, 232, 485, 1427, 1428 Smilodon, 400, 725, 1503, 1518, 1519, Pl. vit Smilodontopsis-hyenoides, 1515 Smith, James, 1149 Smith River, Montana, 479, 737, 1493 Smith, William, 3, 620, 799 Snake Creek, 152, 154, 156-158, 162, 251, 304, 384, 385, 389, 400, 425-429, 473, 479, 491, 492, 497, 498, 560, 562, 711, 736, 737, 1492, 1494, 1499 Snake River, Cherry County, Nebraska, 251, 288, 289, 295-297, 317, 318, 460, 470, 707, 738, 743 Soan, 1449 Soblay, lignites de, 134 Sémmerring, Samuel Thomas von, 3, 86, 799 Soergel, Wolfgang, 4, 15, 19, 799, 961, 1014, 1043, 1045, 1057, 1142, 1182, 1187, 1221, 1403, 1628 Sokolow, 1065 Solilhae, 1065 Solo River, Java, 366 Sologne, Sables de, 1458 Solutré, 1168 Somaliland, 1193, 1402 Sonora, Mexico, 474, 479, 493, 737, 1033 108, 115, 122, 123, 134, 192, 207, 213, 214, 219, 223, 246, 250, 252, PROBOSCIDEA sonoriensis (see Archidiskodon sonoriensis) Sorachi, 908, 1299 Sorensen, Carl, 316, 1107 Soret, France, 634 South Africa (see Africa) South African Museum, 11938, 1609, 1611 South America, 515, 516, 518-533, 567, 593, 1516, 1518. See also Argentina, Bolivia, Chile, Ecuador, Parana River, San Francisco River South Carolina, 729, 736, 739, 996. See also Charleston, Phosphate Beds South Dakota, 298, 304, 305, 708, 709. See also Black Hills, Eastview, Rapid City Southwold, England, 621, 651, 1059 Soyama, Japan, 1188, 1298 Spain, 134, 1131, 1184 species, subspecies, varieties, list of, 1527 spectabilis (see Trilophodon spectabilis) Speegleville, Texas, 373 Spencer, Robert, 485 spencert (see Rhynchotherium spencer?) Spillmann, Franz, 4, 15, 5382, 537, 567-574, 588, 585, 799, 1413, 1520 Spock, Leslie Erskine, 466, 718, 799, 1478, 1628 Spokam Bar, Montana, 943, 1017, 1407 spoonbills, 707, 1412. See also Megabelodon Springview, Nebraska, 305, 317, 318, 328, 414 spurs, 1545, 1546 Squalodon bariensis, 217 squamosal, 916, 917 Sredne-Kolymsk, 1128 Staats-Sammlung fur Paliontologie und Historische Geologie, Munich, 697, 748, 759 Staked Plains (see Llano Estacado) Stamp, L. Dudley, 824, 1628 Stampian, 903, 1441 Standard Oil Company of California, 161 Stanislaus County, California, 478, 479, 487, 497, 737 Staplehurst, Seward County, 943, 1012, 1025, 1026 State Zoological Museum, Munich, 1609, 1615 Stauffer, Clinton R., 4, 799 Stavropol, Russia, 461 Stearns, Robert Edward Carter, 799, 1031 Steelport River, South Africa, 1188, 1287 Steer Creek, Nebraska, 318 Stefanescu, Gregorit, 3, 15, 85, 96-98, 799 Stefanescti, Sabba, 4, 799, 943, 968, 969, 1184, 1188, 1234, 1408 Stefani, 634 Stegodon, 10-12, 25, 32, 33, 197, 806, 807, 810, 811, 814, 815, 818, 821, 837, 838, 853, 854, 858, 859, 874, 888, 891, 898, 904, 906, 909, 913, 914, 927, 942, 964, 965, 1177, 1304, 1375, 1435, 1436, 1455, 1484, 1485, 1524, 1527, 1539, 1547, 1579, 1589, 1602 aff. bombifrons, 817, 898 airawana, 366, 809-814, 818, 821, 823, 833, 834, 836, 838, 854, 855, 857, 859, 874, 881, 885-889, 891, 892, 904, 967, 1302, 1399, 1403, 1539, 1546, 1605, 1615, 1618 airawana (=javanicus), 874 (Archidiskodon?) mindanensis, 823, 833, 836, 838, 854, 892, 905, 942, 1380, 1399, 1456, 1539 aurorx, 25, 812-816, 818, 823, 834, 835, 836, 838, 854, 891, 892, 893, 905, 1380, 1405, 1489, 1539, 1620 bombifrons, 10, 114, 448, 449, 527, 642, 809, 811-815, 817-820, 8238, 830, 836, 838, 853-860, 863-866, 888, 891, 904, 906, 909, 953, 1392, 1448, 1489, 1539, 1605, 1613, 1615, 1617 bondolensis, 816, 823, 837, 894-896, 1416, 1539 cliftii. See Stegodon elephantoides (= cliftir) elephantoides, 114, 650, 809-812, 814, 823-825, 826-829, 836, 838, 854, 861-863, 1305, 1386, 1448, 1451, 1453, 1539 elephantoides (=cliftii), 10, 114, 448, 527, 701, 808, 809, 811, 812, 814, 815, 818-821, 823-828, 831, 836, 838, 853, 854, 860-863, 888, 904, 906, 909, 1305, 1392, 1448, 1449, 1539, 1614, 1615 INDEX Stegodon—continued ganesa, 10, 32, 83, 448, 449, 809, 811, 812, 814-816, 818-820, 823, 829, 830, 836, 838, 853-859, 869-875, 882, 883, 904, 939, 1340, 1392, 1448, 1449, 1539, 1550, 1605, 1613, 1614, 1616. See also Stegodon insignis ganesa ganesa var. javanicus, 814, 823, 833, 836, 888, 855, 856, 859, 887, 889, 1403, 1539 insignis, 10, 32, 448, 449, 527, 809, 811-820, 823, 828, 829, 836, 838, 853— 859, 866-873, 877, 878, 881, 885, 890, 904, 905, 939, 1177, 1391, 1448, 1449, 1453, 1539, 1605, 1613, 1616, 1617 See also Stegodon insignis ganesa insignis birmanicus, 814, 823, 824, 835, 836, 838, 854, 874-877, 1305, 1414, 1451, 1539, 1617 insignis-ganesa, 811, 823, 873, 874. 877, 1391, 1392 javanoganesa, 1403 licentt, 1420, 1482, 15389 mindanensis-aurore group, 903 officinalis, 816, 823, 837, 898, 1418, 1539, 1617 orientalis, 812-814, 817, 818, 823, 832, 836, 838, 854, 855, 884, 885, 902, 904, 906, 909, 1301, 1395, 1451-1458, 1483, 1484, 1489, 1539, 1614, 1629 orientalis grangeri, 808, 809, 811-815, 818, 823, 835, 836, 838, 854, 859, 875-881, 884, 891, 1305, 1414, 1539, 1547, 1605, 1616 orientalis shodoénsis, 814, 823, 835, 836, 838, 855, 893, 894, 904, 906, 909, 1409, 1539, 1611, 1612, 1620 pinjorensis, 448, 814, 823, 835, 836, 838, 854, 855, 881-883, 891, 938, 1360, 1414, 1448, 1539, 1605, 1616 shodoénsis akashiensis, 893, 1420, 1539 sinensis, 115, 812-814, 817, 818, 823, 831, 838, 854, 855, 860, 861, 898, 902, 904, 909, 1395, 1489, 1539, 1614 trigonocephalus, 818, 823, 833, 836, 838, 854, 855, 857, 859, 860, 874, 886, 889-891, 904, 1398, 1403, 1455, 1539, 1605, 1612 trigonocephalus precursor, 816, 823, 837, 896, 1417, 1455, 1539 ytishensis, 816, 823, 837, 897, 898, 1418, 1539 zdanskyi, 816, 823, 837, 899, 1418, 1482, 1483, 1539, 1617 Stegodonten, 1378 Stegodontide, 27, 806, 807, 894, 1867-1369, 1525, 1526, 1579, 1548, 1551 Stegodontineg, 25, 27, 32, 114, 115, 126, 195, 197, 527, 807, 808, 809, 812-816, 819-824, 837, 888, 913, 936, 1304, 1305, 1369, 1524, 1526, 1538, 1545, 1579, 1580. See also Stegolophodontine Stegodontoidea, 22, 25, 191, 806, 807, 837, 1367, 1524, 1526, 1539, 1548, 1550, 1552, 1555, 1579, 1602, Pl. x1 stegodontoides (see Stegolophodon stegodontoides) Stegolophodon, 11, 27, 33, 191, 195, 211, 690, 700, 701, 737, 806, 807, 809, 810, 812, 814, 815, 819-824, 827, 837-840, 853, 896, 898, 904, 1304, 1378, 1379, 1527, 1578, 1579, 1602, Pls. 1v, x1 cautleyi, 32, 114, 197, 365, 448, 527, 622, 643, 644, 700, 701, 737, 753, 754, 809, 810, 812, 814, 815, 819-821, 823, 832, 836, 838, 840-842, 845, 848, 854, 904, 1398, 1448, 1610, 1613, 1618, Pl. 1v cautleyi progressus, 448, 642, 737, 814, 815, 823, 835, 836, 838, 848-861, 854, 1414, 1448, 1605, 1616, Pl. 1v latidens, 10, 11, 114, 195, 197, 348, 448, 458, 527, 532, 700, 701, 737, 809, 810, 812, 814, 815, 817, 819-821, 823-828, 836, 838, 839, 842-847, 854, 874, 885, 903-905, 909, 1380, 1386, 1448, 1451, 1455, 1489, 1617, Pl. 1v lydekkeri, 100, 701, 737, 754, 814, 823, 836-838, 844, 847, 851, 854, 885, 1305, 1419, 1455, 1456, 1613, 1615, Pl. 1v nathotensis, 448, 737, 814, 815, 823, 835, 836, 838, 847, 848, 1414, 1448, 1616, Pl. 1v stegodontoides, 448, 701, 737, 810, 812, 814, 820, 821, 823, 834, 836, 838, 846, 847, 854, 903, 904, 1404, 1448, 1610, Pl. 1v sublatidens, 114, 195, 197, 737, 809, 814, 822, 823, 835, 836, 838, 846, 854, 1405, 1470, 1620, Pl. tv Stegolophodontine, 27, 30, 191, 197, 689, 700, 737, 805, 806, 819-823, 1371, 1526, 1538, 1545, 1546, 1578 Stegomastodon, 11, 31, 228, 229, 367, 377, 482, 531, 541, 575, 586, 588, 589, 612— 615, 622, 624, 628, 667-669, 676, 685, 686, 691, 722, 727, 741, 1379, 1380, 1513, 1514, 1525, 1527, 1545, 1548, 1549, 1564, 1567, 1601 aftoniz, 228, 593, 613, 615, 623-625, 667, 671, 674, 682, 683, 725-727, 741, 745, 760, 1409, 1514, 1548, 1605, 1611, 1615, Pl. vir arizone, 118, 228, 229, 377, 399, 575, 612, 615, 623-625, 666, 672, 674, 678— 683, 727, 741, 751, 759, 1410, 1505, 1506, 1514, 1601, 1605 bonaerensis (see Cuvieronius bonaerensis) chapmani, 127, 615, 622, 623, 625, 669, 683, 727, 741, 753 1669 mirificus, 11, 31, 127, 228, 399, 477, 526, 531, 532, 538, 539, 586, 612, 615, 622-625, 628, 666-671, 673-676, 682, 683, 725-727, 741, 745, 746, 759, 998, 1379, 13894, 1504, 1505, 1514, 1566, 1605, Pl. vin nebrascensis, 1410. See Serridentinus (Ocalientinus) nebrascensis platensis (see Cuvieronius platensis) priestley, 251, 615, 624, 625, 684, 741, 755, 1415, 1508, 1514 primitivus, 614, 617, 625, 724-728, 741, 746, 751, 1419, 1514, Pl. vim successor, 399, 539, 547, 554, 586, 615, 623-625, 667, 668, 671, 673-677, 683, 726, 727, 741, 748, 759, 1400, 1504 texanus, 399, 575, 586, 612, 615, 623-626, 667, 668, 672-677, 683, 726, 727, 741, 746, 750, 1409, 1504, 1605 Stegomastodontide, 1369, 1525, 1526, 1566, 1628 Stegomastodontine, 1371, 1566, 1628 Stehlin, Hans Georg, 4, 15, 475, 799, 1458, 1474, 1625, 1629 Steiermark, 91, 115, 384 Steinheim, Germany, 115, 251, 281, 738, 1045, 1058, 1130, 1137, 1152, 1233, 1236-1238, 1258, 1403, 1461, 1463, 1464 steinheimensis (see Trilophodon steinheimensis) Stellenbosch, South Africa, 14 Steneofiber, 260, 1459, 1461 Steno, 1118 Stenomylus, 315 stenotoechus (see Elephas stenotoechus) steppe fauna, 1044, 1045 Sterling, Lindsey Morris, xvi, 25, 110, 305, 1100 Sternberg, Charles Hazelius, 4, 291, 711, 713, 739, 800, 1100-1102 Sternberg, Charles M., 800, 1508, 1629 Stewart, Alban, 1017 Stirton, Ruben Arthur, 15, 161, 718, 800, 1446, 1496, 1627, 1629 Stock, Chester, 4, 15, 487, 800, 943, 1418, 1509, 1627, 1629 Stockholm, 14, 539, 546, 547, 561, 562, 745, 759, 1609, 1620 Stone Age, 1131 Stone, Witmer, 15 Strand, 1193, 1409 Strauss, F. C. J. von, 3, 89, 800 Stremme, Hermann, 4, 800, 886, 887, 889, 1186, 1303 Stromer, Ernst, 4, 37, 697, 698, 800, 1424, 1629 Strong, William, 312 Strozzi, Carlo, 1061 Struthiolithus, 1487 Strzlecki, Count P. E. de, 1390, 1397 Stur, Dionys, 92, 800 Stuttgart, 13, 745, 759, 1130, 1152, 1153, 1236, 1253, 1255, 1609, 1620 Stylohipparion, 1433, 1434 stylomastoid foramen, 916 Styria, 223, 742. See also Oberdorf styriacum (see Deinotherium styriacum) subantiqua (see Loxodonta subantiqua) subfamilies, 22, 27, 30, 31, 689, 735-748, 1526 sublatidens (see Stegolophodon sublatidens) suborders (see superfamilies) subplanifrons (see Archidiskodon subplanifrons) sub-shovel-tuskers (see Serbelodon and Trobelodon) subtapiroidea (see Serridentinus subtapiroideus) successor (see Stegomastodon successor) Sud Oranais, Algeria, 1184 Sudan, 1176, 1193, 1194 Suess, Eduard, 3, 92, 800 Siissenborn, 1042, 1044, 1045, 1047, 1050, 1055-1058, 1065, 1067, 1398 Suez, 1269 Suffolk, England, 625, 634, 635, 651, 740, 943, 963, 964, 1059, 1401 Sugiyama, Tsurukichi, 900 sugiyamat (see Parastegodon sugiyamai) Suina, 251, 266, 271, 272 suleus, 139, 140, 211, 689, 692, 1546, Pls. 1-1v Sulza, 1050 Sumatran elephant (see Hlephas indicus sumatranus) 1670 sumatranus (see Elephas indicus sumatranus) Sundevall, 1377 Sunkareah, 1314 superbus (see Cuvieronius superbus) superfamilies or suborders, 19, 23, 28, 140, 735, 1367, 1524 Surakarta, Java, 833, 890, 1398 Sus, 274, 887 namadicus, 1449, 1489 scrofa, 1155 Sussex, 965 Sutton, Nebraska, 1003 Suwo, Japan, 818, 906 Swabey, E. C., 696 Swan River, Canada, 137 Sweden (see Stockholm) Swinhoe Collection, 884, 885 Swinton, William E., xvi, 15 Swisher, George, 1084 Switzerland, 115, 134, 760, Pl. m1. See also Elgg, Zurich Sydney-on-Vaal, South Africa, 943, 987, 990, 991, 1188, 1282 Sykes, Colonel, 268 Sylhet, 927 Sylvilagus (?), 680 Symbos, 171, 725, 1135, 1161, Pl. vit Synconolophus, 262, 268, 274, 347, 613, 622, 625, 627-630, 649, 654-657, 720, 740, 1381, 1527, 1548, 1549, 1565, 1601 corrugatus, 114, 539, 613, 622, 624, 625, 627, 628, 630, 643, 654, 655, 658- 661, 740, 756, 1404, 1448, 1604 dhokpathanensis, 12, 31, 118, 347, 364, 448, 616, 622, 624, 625, 628-630, 642, 643, 653-656, 658, 661-664, 740, 749, 750, 1381, 1414, 1448, 1604 hasnoti, 114, 448, 538, 613, 622, 624, 625, 630, 648, 653, 654, 655, 658-661, 740, 749, 756, 1404, 1448, 1604 propathanensis, 448, 450, 624, 625, 643, 654, 657-659, 665, 740, 749, 1414, 1448 plychodus, 448, 622, 624, 625, 630, 643, 656-658, 740, 749, 1414, 1448 Synodontherium, 10, 1123, 1124, 1365, 13876 Syria, 1269 “Systema Nature,”’ 6, 135, 1308-1310 Szabadka, Hungary, 159, 160, 1406, Pl. 1 Szalay, Tibor, 5, 116, 800 Szechuan (or Szechwan), 815, 837, 853, 875, 884, 898, 1452, 1487 Szentloriez, Hungary, 114, 159, 160 Tabata, 908, 1296 Tables, 111, 114, 115, 448, 1388; 1, 111; m, 114-115; 111, 814; 1v, 836; v, 854; v1, 949; vir, 954; viir, 984; 1x, 985; x, $86; x1, 1037; x1, 1109; xin, 1113; xiv, 1114; xv, 1161; xvi, 1343; xvir, 1348 Tabriz, 1061 Taga, 902 Taganrogh, Russia, 1393 Tagua-Tagua, Chile, 523, 537, 581, 582, 741, 1400, 1520 Tairum Basin, 718, 719 Tairum Nor, Mongolia, 251, 332, 385, 463, 743 Taiwan, 1305, 1489 Takai, Fuyuji, 1420, 1629 Takamori, 902 Takikawa Middle School, 1609, 1612 Tallahassee, 1609, 1620 Talmadge, Otoe County, Nebraska, 696 Tamaki-mura, 908 Tambla tooth, 482, 484, 535 Tamise, Belgium, 1136, 1388 Tanana River, Alaska, 1159 Tanganu, Rumania, 969, 1184, 1188, 1233, 1235 Tanganyika, 85, 945, 1184, 1187, 1193, 1275-1277, 1419, 1432 taoensis (see Trobelodon taoensis) Taos, New Mexico, 441 Tapalqué, Argentina, 592 Tapasuma, Honduras, 479, 508-510, 737, 1516 OSBORN: THE PROBOSCIDEA tapiroides (see Turicius tapiroides) tapiroides-minus (see Turicius tapiroides-minus) tapirs gigantesques, 84, 86, 96, 98, 99, 109, 1554 Tapirus, 48, 83, 98, 170, 400, 579, 818, 887, 1077, 1515 proavus, 1389 Tapyroides (see Tetracaulodon Tapyroides) Tarai, 927, 1313 Tarakla, 1466 Taraklia Kischinev, 1468 Tarboro, North Carolina, 286, 287, 386, 419, 420, 1395 Targioni-Tozzetti, Giovanni, 2, 800, 969, 1382 Tarija, 514, 517-519, 527, 587, 544, 546, 548, 550, 568, 570, 740, 743, 763, 1519, 1520 tarijensis (see Cordillerion tarijensis) Tarrapota, Peru, 554 Tartars, 1164 Tartarie (see Petite Tartarie) Tasnid, Hungary, 152, 156, 736 Tatabelodon, 1382, 1557 gregoru. See Trilophodon (Tatabelodon) gregorvi riograndensis, 1417. See T'rilophodon (Tatabelodon) riograndensis Tatrot, 279, 448, 449, 642, 950, 1442, 1445-1448, 1454 Tatsunokuchi formation, 280, 902 Tatu bellus, 400 Taubach, 558, 969, 1042, 1045, 1050, 1052, 1181, 1184, 1188, 1217, 1233-1235, 1245, 1251, 1253, 1294 Taungs district, Bechuanaland, 944 Tawi, 1442, 1445, 1447, 1626 Tébessa, Egypt, 105 Tecamachaleo, Puebla, 1047, 1082 Tecucit, Rumania, 85, 95 Tehama formation, 1505, 1506 Tehuichla, Mexico, 555, 1506 Teilhard de Chardin, Pierre, 5, 15, 800, 950, 1362, 1409, 1420, 1446, 1449, 1451, 1478, 1480, 1488, 1485, 1594, 1623, 1627, 1629 Teleobunomastodon, 543, 552, 1381 bolivianus, 536, 537, 548, 551, 652, 1381, 1415 Teleoceras, 261, 273, 307, 308, 335, 380, 385, 386, 399, 400, 418, 461, 510, 1082, 1426, 1441, 1442, 1459 Temblor, 902 Temminck, Coenraad Jacob, 3, 800, 930, 1314, 1325, 1329, 1393 Temnocyon, 318 Tenasserim, 927, 1594 Tenjinbayashi, 13889, 1419 Tentzelius, Wilhelmus Ernestus, 1, 2, 800, 1118, 1119, 1165, 1181 Tepexpan, Mexico, 947, 1013, 1081 Tephrocyon, 318, 680 Tequixquiac, 537, 554, 740, 948, 1013, 1016, 1080-1082, 1515 Ternifine, 1183, 1187, 1274 Terra, Hellmut de, 950, 1446, 1447, 1449, 1451, 1626, 1629 Terraces, 1050, 1065, 1278, 1284, 1285, 1289, 1430, 1473, 1474, 1488 Terreno pampeano antiquo, lacustre, superior, 1517, 1623 Tertiary, 279, 314, 461, 1422, 1450, 1456, 1457, 1470, 1488, 1484, 1492, 1497, 1508, 1509 Tervueren Museum, 11938, 1609, 1620 Teschen (Schlesien), 737, 809, 835, Pl. 1v Testudo, 53, 117 tetartoloph-lophid, 141, 812, 1545 Tetrabelodon, 11, 231, 249, 251, 290, 625, 526, 532, 1378, 1495, 1502, 1509 abeli, 1410. See Trilophodon abeli andium (see Cordillerion andium) angustidens (see Trilophodon angustidens) angustidens proavus (see Serridentinus proavus) angustidens var. chinjiensis (see T'rilophodon chinjiensis) brevidens (see Rhynchotheriwm brevidens) (Bunolophodon) ayorx. See Cuvieronius ayore corrugatus (see Synconolophus corr ugatus) crepusculi (see Hemimastodon crepuscult) dinotherioides (see Trilophodon dinotherioides) INDEX Tetrabelodon—continued euhypodon (see Blickotherium euhypodon) longirostris (see Tetralophodon longirostris) lulli (see Megabelodon lullz) macrognathus (see Trilophodon macrognathus) osborni. See Trilophodon (Genomastodon) osborni serridens (see Serridentinus serridens) serridens cimarronis (see Serridentinus serridens cimarronis) shepardii. See Rhynchotherium falconeri, also Tetracaulodon (Tetrabel- odon) shepardii willistoni. See Trilophodon (Genomastodon) willistont tetrabunodonty, 77, 78, 1382, 139, 145, 1545 Tetracaulodon, 165, 777, 783 Bucklandi, 165, 1390 Collinsvi, 165, 1388 Godmani, 165, 1388 Haysii, 165, 1390 kochii, 165, 783, 1390 longirostre, 349, 357, 782, 1373, 1387, 1389 Mastodontoideum, 165, 777, 1373, 1387 osagit, 165, 783, 13889 Tapyroides, 165, 783, 1373, 1389 (Tetrabelodon) shepardii, 488 Tetraconodon, 272 Tetralophodon, 10, 11, 28, 29, 44, 45, 127, 225, 226, 228, 231, 307, 343-353, 356-376, 379, 389, 391, 538, 546, 629, 661, 690, 703, 903, 1375, 1376, 1400, 1527, 1559, 1560, 1600 arvernensis (see Anancus arvernensis) (?) brazosius. See Trilophodon (Tetralophodon?) brazosius bumiajuensis, 31, 349, 351, 365, 366, 379, 739, 753, 756, 1416, 1455, 1600, 1604 campester, 226, 227, 229, 306, 342, 348-352, 356, 359, 363, 365, 369-372, 376-379, 385, 390, 391, 490, 526, 532, 538, 679, 739, 746, 758, 1396, 1397, 1500, 1604 elegans, 226, 348, 349, 351, 352, 356, 368, 369, 372-374, 379, 739, 745, 759, 1405, 1604 exoletus, 704, 705, 739, 1418, 1482 fricki, 348-351, 356, 369, 374-376, 379, 399, 739, 751, 1419, 1501, 1604 gigantorostris, 282, 283, 348, 349, 362, 739, 756, 1406, 1469 grandincisivus, 114, 118, 226, 229, 342, 347-349, 351, 356, 357, 359-362, 739, 760, 1405, 1469, 1604 longirostris, 10, 31, 114, 125, 226, 231, 232, 255, 305, 342, 344-346, 348, 349, 350-353, 355-361, 367-371, 378, 389, 527, 532, 660, 739, 746, 1387-1389, 1469, 1604 (Lydekkeria) falconeri, 114, 226, 343, 348, 349, 351, 353-357, 362, 370, 379, 448-451, 532, 703, 720, 739, 750, 754, 756, 903, 1396, 1448, 1560, 1604 (Lydekkeria) sinensis, 115, 226, 348, 349, 351, 355, 379, 703, 739, 817, 1398, 1482, 1559, 1604 (Morrillia) barbouri, 1406. See Morrillia barbourt perimensis (see Anancus perimensis) precampester, 349-351, 368, 372, 378, 758, 1408 punjabiensis, 114, 226, 342, 347-363, 356, 357, 362-365, 368, 370, 378, 448, 647, 661, 739, 748, 749, 754, 756, 841, 1398, 1448, 1560, 1604, 1610, 1614 Tetralophodontine, 27, 31, 119, 228, 231, 343, 347, 351, 356, 377, 390, 689, 690, 734, 739, 1370, 1371, 1526, 1528, 1545, 1548, 1555, 1559, 1560, Pl. x texanus (see Stegomastodon texanus) Texas, 384, 431, 532, 543, 559, 680, 729, 737, 740, 741. See also Aransas River, Blanco formation, Brazos River, Clarendon, College Station, Fort Worth, Hog Creek, McLellan County, Llano Estacado, Pittbridge, Post, Red River, Refugio, San Felipe de Austen, Sinton (San Patricio County), University of Texas, Waco Texas Geological Survey, 429, 431, 673, 675, 745, 759 texianus (see Parelephas columbi) Thames valley, 1050, 1059 Thayer County, 1012 The Bend, Vaal River, 944, 945, 989 The Pits, 427 Theobald, William, 3, 644, 658-660, 800, 1332 1671 Theophrastus, 1147 Therisiopel, 210 theristocaulodon (see Missourium theristocaulodon) Thevenin, Armand, 4, 766, 800, 1519 Thiede, Thuringia, 1187, 1256 Third Asiatic Expedition (see Central Asiatic Expedition) Thomas County, Nebraska (see Seneca) Thomas, Michael Rogers Oldfield, 4, 15, 800, 1374, 1629 Thomas, Philippe, 3, 800, 1187, 1398 Thomson, Albert, 429, 1012 Thone, Rothe (see Rothe Thone) Thorn, West Prussia, 192, 208, 736 Thorpe, Malcolm R., 710 Thorpe, Norwich Crag, 963 thorpet (see Gnathabelodon thor per) Thousand Creek, Humboldt County, Nevada, 137, 152, 145-156, 497, 498, 560, 562, 736, 1418, 1502 Thousand Isle Creek, 155 Thrace, 1466 Thuringia, 1045, 1118, 1137, 1150, 1187 Thurston County, Nebraska, 692, 695, 736 Tiao Kou, Kansu, 702, 1418 Tibet, 794, 852, 940 Ticholeptus, 315, 485 Tieh Chang Kou, 719 Tientsin, China, 384, 855 Tilesius von Tilnau, Wilhelm Gottlieb, 3, 800, 931, 1128, 1130, 1131, 1147, 1148, 1162 Tillman County (see Archidiskodon haroldcooki and Stegomastodon priestleyt) Tilloux, 1270 time-scale, 278 Tipperah, 1315 Tipton County, Indiana, 1012, 1088 Tiraspol, 961, 1047, 1065, 1066, 1403 Tiseo, Saverio, 1239, 1243 Titanotheriide, 824, 918 Tji Djoelang, 1454 Tji Sande zone, 1454 Tkatchenko, M. J., 1128 Tlaseala, Mexico, 474, 475, 482, 493, 534, 737 tlascale (see Rhynchotherium tlascalz) Tlatlaya, Mexico, 557 Tobien, H., 1465, 1466, 1629 Togane Town, Japan, 818, 906 Togari, 902 Tohoku Imperial University, Sendai, Japan, 14, 15, 1333, 1609, 1620 Tokunaga, [Yoshiwara] Shigeyasu, 4, 800, 816, 818, 894, 897, 899, 1188, 1289, 1301, 1417-1419, 1490, 1629 tokunagai (see Palxoloxodon tokunagat) tokunagai junior mut. See Paleoloxodon (Archidiskodon?) tokunagat mut. junior Tokyo, 902, 907, 908, 1298, 1609, 1620 Tokyo Imperial University, 15, 893 Tolmachoff, Innokenty Pavlovich, 4, 15, 801, 1162, 1164, 1169 Tonami district, 1188 Tonna (see Burgtonna) Tonnellier, 167 topotypes, 9, 254, 266 Tortonian, 115, 201, 219, 315, 902, 903, 1443, 1447, 1460, 1462-1464, 1466 Torynobelodon, 290, 333, 338, 385, 470, 471, 686, 691, 716, 729, 730, 743, 1381, 1527, 1550, 1571, 1602 barnumbrownt, 31, 317, 318, 384, 385, 445, 460, 470-472, 711, 715, 719, 743, 758, 1416, 1498, 1605, Pl. vir loomisi, 250, 251, 290, 333, 338, 339, 384, 385, 445, 465, 471, 472, 711, 715, 716, 743, 758, 1881, 1413, 1507, 1571, Pl. vu Tosa, 908 Tossano, Italy, 634 Totomi, Japan, 908, 1187, 1295, 1489 Touraine, Faluns de la, 134, 205, 221 Tournan, 246, 252, 256, 259, 271 1672 Toxodontia, 35, 1519 toxotis (see Loxodonta africana toxotis) Trail Cation, Hitchcock County, 479, 489 trans-Caucasian region, 1466 transitional mastodons, 828, 885 Transvaal, 943, 983, 984, 1287 transvaalensis (see Palxoloxodon transvaalensis) Trassaert, M., 1409, 1420, 1482, 1483, 1620 Travertines (see Interglaciations, Thuringia) trefoils, 29, 140, 141, 225, 226, 277, 519, 554, 686, 1545, 1546 Trego County, Kansas, 713, 739, 1418 tremontigerus (see Cenobasileus tremontigerus) Tricaulodon, 781 Trichechus americanus (see Manatee) Trichecodon, 963 trigodon (see Meritherium trigodon) trigonocephalus (see Stegodon trigonocephalus) trigonodon (see Meritherium trigodon) Trilophodon, 9-11, 28, 31, 42-45, 112, 113, 127, 225, 226, 228, 231, 232, 238, 248-252, 254, 257, 258, 263, 264, 266, 275, 277, 284, 288, 289, 293, 298, 312, 315, 316, 319, 321, 326-331, 333, 341, 343-346, 351, 367, 369, 378, 383, 386, 388, 389, 391, 400, 412, 413, 436, 513, 538, 546, 686, 690, 737, 822, 901, 909, 945, 1200, 1374, 13875, 1378, 1461, 1463, 1502, 1509, 1525, 1527, 1547, 1555-1557, 1600, 1607, 1608, 1629, Pl. vir abeli, 226, 248, 250, 251, 260, 288-290, 298, 311, 312, 317, 329, 331, 601, 706, 711, 738, 758, 1410, 1498 angustidens, 5, 10, 35, 37, 38, 112, 115, 117, 121-123, 214, 224-226, 231, 232, 234, 235, 244, 246, 247, 249-260, 270, 277, 281, 284, 287, 330, 340, 341, 344-346, 357, 360, 379, 388, 395, 418, 486, 519, 525-527, 532, 633, 637, 648, 686, 738, 751, 754, 758, 812, 1374, 1428, 1441, 1464, 1479, 1557, 1604 angustidens cuvieri, 121, 209, 226, 250, 252, 254, 738, 1398, 1460 angustidens gaillardi, 250, 251, 257, 259, 273, 287, 388, 738, 758, 1413, 1464, 1604 angustidens gaujact, 250, 252 angustidens libycus, 112, 115 1427 angustidens minutus, 121, 122, 123, 124, 225, 250, 252, 254, 633, 738, 1386, 1389, 1464 angustidens palxindicus, 106, 115, 1448. See also Trilophodon palzindicus angustidens var. austro-germanicus, 226, 250, 251, 254, 259, 260, 738, 754, 903, 1403, 1464 angustidens var. chinjiensis. See Trilophodon chinjiensis atavus, 1479 (?) barstonis, 1417. See also Serridentinus barstonis chinjiensis, 250, 251, 272, 273, 275, 276, 278, 284, 287, 330, 334, 341, 388, 448, 449, 454, 738, 749, 1404, 1448, 1604, 1616 (Choerolophodon) pentelicus, 10, 114, 193, 214, 250, 255, 261-266, 284, 345, 346, 737, 750, 1393, 1469 connexus, T02, 732, 738, 1418, 1480 cooperi, 106, 112, 115, 250, 251, 266, 269-271, 276-278, 284, 287, 341, 448, 449, 702, 703, 738, 746, 753, 754, 756, 1416, 1441, 1480, 1604 cruziensis (see Megabelodon cruziensis) dinotherioides, 248, 250, 251, 288, 289, 291, 293, 329, 331, 478, 484, 711, 738, 754, 1403, 1500, 1506, Pl. vin engelswiesensis, 250, 251, 260, 281, 738, 1406, 1464 esselbornensis, 250, 251, 260, 281, 737, 755, 1407, 1469 falconeri. See Tetralophodon (Lydekkeria) falconert fricki, 250, 251, 257-290, 312-315, 329, 331, 385, 388, 706, 711, 738, 755, 1412, 1492, 1604, Pl. vir gaillardi (see Trilophodon angustidens gaillardz) (Genomastodon) osborni, 226, 248, 250, 251, 260, 284, 287-290, 298-304, 309, 312, 314, 315, 317, 319, 321, 324-326, 329, 331, 388, 436, 444, 706, 733, 738, 751, 752, 758, 1380, 1405, 1498, 1604, Pl. vir (Genomastodon) willistoni, 226, 229, 248, 250, 251, 284, 288-290, 292-294, 317, 329, 331, 601, 711, 733, 738, 758, 1380, 1404, 1498, Pl. vir giganteus, 226, 227, 248, 250, 251, 284, 287-289, 298, 304-307, 309, 312, 321, 328, 329, 331, 369, 360, 382, 388, 403, 412, 413, 711, 738, 748, 1406, 1498, 1600, 1604, Pl. vir 54, 738, 1393, 1464 25: 226, 250, 251, 260, 261, 394, 686, 738, 1405, , OSBORN: THE PROBOSCIDEA Trilophodon—continued hasnotensis, 250, 251, 279, 448, 449, 454, 737, 749, 1419, 1448 hicksi, 1407. See Amebelodon (Trilophodon) hicksi inopinatus, 250, 251, 278, 279, 385, 461, 738, 1412, 1479 joraki (see Megabelodon jorakz) ligoniferus, 248, 250, 251, 288, 289, 298, 312, 328, 329, 331, 403, 711, 738, 746, 1405, 1507 macrognathus, 118, 226, 248, 250, 251, 272, 274, 275, 276, 284, 341, 448, 449, 642, 738, 756, 1404, 1448, 1604 (Megabelodon) lulli. See Megabelodon lulli obscurus, 11, 127, 250, 251, 285-287, 321, 381, 383, 386, 392, 394, 420, 448, 487, 738, 745, 1380, 1395, 1508 ohioticus, 1177, 1395. See also Mastodon americanus paladentatus. See Amebelodon paladentatus palzindicus, 106, 115, 226, 250, 266-271, 274-278, 284, 341, 448, 449, 527, 642, 687, 703, 738, 746, 749, 756, 1398, 1407, 1441, 1489, 1546, 1610, 1615 pandionis, 79, 115, 226, 250, 262, 266, 267, 268, 270, 448, 622, 645, 654, 655, 738, 754, 817, 13894, 1441, 1448 phippsi (see Megabelodon phippsi) pojoaquensis, 226, 232-234, 251, 284, 289, 290, 319-323, 329, 331, 385, 436, 438, 706, 711, 738, 750, 1411, 1493, 1617, Pl. vir pontileviensis, 115, 196, 250-252, 260, 261, 283, 284, 287, 686, 693, 738, 819, 1405, 1408, 1460, 1604 progressus (see Serridentinus progressus) pyrenaicus (see Zygolophodon pyrenaicus) sendaicus, 226, 250, 251, 280, 321, 738, 901, 903, 909, 1409, 1489 (Serridentinus) leidii, 1411. See Ocalientinus (Serridentinus) floridanus leidit (Serridentinus) pojoaquensis. See Trilophodon pojoaquensis simplicidens, 11, 250, 251, 285, 321, 381, 383, 386, 392, 394, 400, 440, 738, 745, 1408, 1496 spectabilis, 702, 703, 738, 1418, 1482 steinheimensis, 250, 251, 281, 282, 738, 759, 1406, 1464 (Tatabelodon) gregorti, 250, 251, 287, 317, 324, 325, 444, 706, 733, 738, 752, 1417, 1498, 1604 (Tatabelodon) riograndensis, 250, 251, 287, 324, 325, 447, 706, 711, 738, 750, 1382, 1417, 1493, 1604, Pl. vir (Tetrabelodon) shepardi edensis, 1406. Rhynchotherium shepardi edense (?Tetralophodon) brazosius, 349, 374, 399, 564, 623, 738, 745, 1408 wimani (see Serridentinus wimant) Trilophodontide, 1368, 1369, 1525, 1526, 1555 Trilophodontine, 1371, 1555, 1628 trilophodonty, 39, 189-141 Trimingham, 981 Tring, 1198, 1609, 1619 Trinil beds, 365, 366, 833, 874, 885-887, 967, 1302, 1403, 1453-1455 Trinity College, Hartford, 15 Trionyx, 117 Tripoli, 232, 1246 tritoloph-lophid, 139, 141, 812, 1545 Trobelodon, 441, 446, 729, 743, 1382, 1527, 1570, 1601 tavensis, 326, 327, 384, 385, 441, 442, 444, 446, 447, 557, 748, 750, 752, 1382, 1417, 1493 Trochtelfingen, 1468 Trogontherian Mammoth, 914 trogontherii (see Elephas primigenius Blum. var. trogontherit, and Parelephas trogontheri) trogontherioides (see Parelephas trogontherioides) Trogontherium, 963, 1056, 1155 tropicus (see Cordillerion tropicus) Trouessart, Edouard Louis, 4, 7, 801, 1141, 13864, 1399 Troxell, Edward Leffingwell, 15, 305, 1498, 1630 Truckee Beds, 1509 Tsagan Nor, Mongolia, 397, 719 Tschobrotschi, 1468 Tsinling, 1484 Tsukiyoshi, 902, 908 See also Cordillerion edensis and INDEX Tsurumi-gawa, 1301, 1417 tubules (see rod-cones) Tuckey, H. P., 1282 Tubingen, 91 Tufts College, 1199 Tule Canon, Brisco County, Texas, 1005, 1006, 1010, 1017, 1018 Tullberg, Tycho Fredrik Hugo, 4, 801 Tultschin, Podolia, 1389 Tulucesti (Covurlui), 943, 968, 969, 1184 tundra, 1161, 1167 Tung Gur Khara Usu, 332, 398, 399, 444, 460, 461, 463, 466-469, 718, 742, 743, 1478 Tunisia, 232, 485, 1269, 1428 Turgai, 251, 278, 385, 738, 1418 turicensis (see Turicius turicensis) Turicius, 10-12, 30, 191, 192, 198-202, 205, 211, 212, 215-217, 223, 254, 382, 383, 686, 690, 698, 699, 718, 736, 819, 820, 822, 1381, 1461, 1463, 1527, 1545, 1546, 1576, 1602, Pl. 11 alticus, 114, 192, 193, 199, 200, 213, 214, 218, 220, 223, 263, 736, 1377, 1394, 1469, Pl. m1 tapiroides, 5, 10, 30, 115, 119, 120, 122, 123, 133, 134, 191-196, 198-201, 203-205, 209-218, 221, 232, 395, 633, 736, 758, 819, 822, 1375, 1377, 1427, 1428, 1460, 1605, 1619, Pl. nt tapiroides-minus, 192, 1389 turicensis, 10, 30, 115, 124, 134, 164, 191-202, 204, 205, 209, 210, 212-216, 218, 219, 221, 253, 697, 698, 736, 760, 819, 822, 1460, 1464, 1605, PI. m1 turicensis simorrensis, 115, 192, 193, 197, 207, 213-215, 219, 220, 736, 821, 822, 1410, 1464 virgatidens, 30, 134, 191-193, 199-201, 211-214, 221-223, 736, 822, 1395, 1470, 1605, Pl. m1 wahlheimensis, 192, 193, 282, 736, 1407, 1469, PI. 11 Turicum, 212 Turin, 209, 745, 759, 1056, 1061, 1154, 1609, 1620 Turnau, Steiermark, 92 Turtle River, 1077 Tuscany, 114, 969, 1187, 12380 tusks, 202, 253, 1007, 1167, 1547, 1550, 1551, 1608 Tusuque, New Mexico, 322, 323 Tutova, district of, 96 Tutzing, Bavaria, 201, 697, 698 Twin Creek, Kentucky, 1088, 1089, 1096 Tyler, Ruth, xvi tympanic bulla (see bulla) type, definition of, 9, 631 type revision, principles of, 816 typha, 719 Typotheria, 35, 590, 595, 1518, 1519 Tyrrhenian, 818, 902, 904-906, 908, 1050, 1065, 1290, 1299, 1474 Uasin Gishu plateau, 1189, 1190 Ube coal-field, Japan, 818, 906 Udden, Johan August, 4, 801 Udiapur, 1326 Ueda, F., 1300, 1301 Uéhata, 906 uehataensis (see Parelephas proximus uehataensis) Uganda, 485, 1190, 1239, 1432 Ugo, 908, 1489 Uhde Collection, 553, 554 Uhle, Max, 4, 567, 569, 573, 574, 801 Uichteritz, 1045 Ulloma, Bolivia, 537, 551, 740, 1400, 1520 Ulm, 115 Umegase bed, 1300, 1301 Underwood and Underwood, 1599 Ungarische geologische Reichsanstalt, Budapest, 745, 755 Ungarisches Nationalmuseum, Budapest, 159, 160, 745, 755 Ungulata, 887 unicorn, 1118 1673 Union City moraine, 1084, 1085 Unionide, 1240, 1442 United States Biological Survey, 15 Geological Survey, 497 National Museum, 14, 15, 745, 759, 1609, 1621 Smelting and Mining Company, Alaska, 1135, 1159 unity, or plurality, of species, 1072, 1315, 1596 Universidad Central del Ecuador, 15, 501, 572, 583, 745, 757 Nacional de Mexico, 1007 Universita di Napoli, 14, 1239 Université de Lyon, 14, 15 University Museum of Pisa, 1061 Museum of Zoology, 14, 731 of Alaska, 1161, 1609, 1610 of Arizona, 565 of Berlin, 14, 881 of Bologna, 14, 1610 of Bonn, 15, 1610 of Breslau, 15, 553, 554, 754 of Bucharest, 15, 745, 755, 968, 969, 1235, 1609, 1610 of California, 14, 15, 163, 745, 759 of Gottingen, 1122, 1141, 1609, 1611 of Iowa, 14, 745, 760, 1609, 1611 of Kansas, 745, 760 of Leipzig, 745, 760, 1609, 1612 of London, 15 of Moscow, 15, 126, 745, 760, 1065, 1066, 1609, 1615 of Nebraska, 14, 15, 294, 338, 708. See also Nebraska State Museum of Pittsburgh, 15 of Prague, 1139 of Reading, 14 of Rome, 1236, 1247, 1609, 1620 of Rostock, 14 of Saskatchewan, 997 of Southern California, 15 of Texas, 15 of Washington, 745, 760 of Witwatersrand, 14, 1284 Unterlangkat, 1330, 1331 Unyoro, Uganda, 1190, 1239 Upland, 1033 Upnor (see Hesperoloxodon antiquus) uproot-tuskers (see prod-tuskers) Upsala University, 1479 Uquiana or Araucana, 1517 Uraha Hill, Africa, 983 Ural Mountains, 85, 87, 735, 13889, 1555 uralense (see Deinotherium uralense) Ureuqui, 583 Ur-Nile, 66 Ursavus, 1467 Ursus arctos, 1253 arvernensis, 963 speleus, 1155, 1231, 1476 Uruguay, 524, 530, 537, 593, 595 Ury@ District, Japan, 1299 Uyeno Museum, 893, 1298, 1609, 1620 Vaal River, 934, 943-945, 983, 984, 1188, 1278, 1281, 1282, 1285, 1288, 1438, 1439 Vacek, Michael, 3, 93, 125, 139, 194, 195, 198, 203, 209, 222, 525, 801 Vainsot, Colonel, 634 Vaise, 1064 Val d’Arno, 134, 616, 619, 634, 942, 961, 964, 969, 1187, 1215, 1443, 1444 Val di Comino, 1241 Valentine, 317, 318, 385, 711, 738, 1445, 1496, 1497, PI. vir Valier, Montana, 1003 Valley Company, 1012 Valley of Mexico, 474, 479, 482, 493, 537, 553, 1081 1674 OSBORN: THE Van Alphen, 991 van der Maarel (see Maarel) Van Es, L. J. C., 1453, 1630 van Riet Lowe (see Lowe) vanalpheni (see Archidiskodon vanalphent) Vanasswegenshoek-Bloemheuvel, 992, 1280 Vanderbilt (see George Vanderbilt African Expedition) VanderHoof, V. L., 1505, 1628, 1630 Vaqueros, 902 Varian, W. H., 1337 variegated beds, 1339, 1345, 1347, 1358 Vaucluse, 114 Vaufrey, Raymond, 5, 801, 1183, 1260 Vaughan, Thomas Wayland, 4, 801 Velay, France, 192 Veles, 1468 Velhas, Valley of, Brazil, 537, 578, 579, 741 vellavus (see Zygolophodon borsoni vellavus) Venice, Florida, 400, 1005, 1077 Ventimiglia, Italy, 1123, 1137 Vera Cruz, 555 Vera Cruz-Hidalgo, 1506 Vergennes, Comte de, 134 Vermont (see Mt. Holly) Vernay, Arthur S., 1249, 1311, 1337 Vernay, le, 1063, 1064 Vero, Florida, 400, 1005, 1079 Verri, Antonio, 3, 801, 1187, 1232 vertebral distinctions, 930, 931, 1131, 1147, 1193, 1227, 1311-1313 Vestonice, 1139 Vevers, F’. M., 15, 1336 vexillarius (see Pliomastodon vexillarius) vialetii (see Zygolophodon borsoni vialetii) Vialette, France, 192, 208, 634 Vicary,—, 1439 Victoria College, Stellenbosch, 14 Victoria Falls, Rhodesia, 1188, 1420 Victoria Nyanza, 103, 104, 115, 735 Victoria, Texas, 1001, 1005, 1009 Vienna, 13, 15, 99-101, 360, 621, 745, 760, 964, 1609, 1620 Vienne, Dauphiny, 84, 86 Vieselbach, 1045 Villa de Guadalupe, 1081 Villada, Manuel Maria, 4, 554, 555, 801, 1016, 1082 Villafranea (San Paolo), Italy, 634 Villafranchien, 634, 740, 964, 1043, 1047-1049, 1055, 1233, 1290, 1431, 1444, 1447, 1454, 1457, 1472, 1473, 1480, 1483-1486 Villanova, Asti, Italy, 199, 207, 209, 736, 1470 Villefranche d’Asterac, 202, 250, 251, 257, 259, 273, 738, 1413, 1463, 1464 Vilna, Russia, 1137 Vindhya range, 852 Vindobonian, 205, 257, 732, 1444, 1457, 1460, 1463, 1464, 1466 Violet sands, 897 Viret, J., 1459, 1628, 1630 virgatidens (see Turicius virgatidens) Virgin Valley formation, Nevada, 150-152, 154-156, 693, 736, 1494 Viverra, 1463 Voitsberg, 115 Volga, 1136, 1388 Volhynia, Russia, 621, 625, 638, 639, 740 Volney, 136 vomer, 916 von Koenigswald (see Koenigswald) Waco, Texas, 373, 374 Wadi Rayan Series (see Faytim) Wadia, D. N., 4, 801, 825 Wagner, Andreas, 3, 91, 192, 193, 213, 220, 263, 736, 801, 1394 Wagner Free Institute of Science, Philadelphia, 1609, 1619 PROBOSCIDEA Wagner, George, 4, 448, 801 Wahlheim, 192, 223, 282, 736 wahlheimensis (see Turicius wahlheimensis) Wakayama, 907, 1333 Walcott, Charles D., 1314 Wald fauna, 1044, 1045 Waldeck’s Plant, Vaal River, 764, 944, 945 Waldelefant, 1253 Walker, J. W., 1022 Walla Walla, Washington, 1102 Wallace, Alfred Russel, 3, 527, 578, 801 Wallihan, A. G., 1103 Walnut, Illinois, 137, 173, 736 Walsh, Jeremiah, 1243 Walton, 1059 Wanhsien, 815, 817, 859, 875 Ward, Rowland, 1022, 1201, 1239, 1332, 1337, 1630 warm fauna (see faunal and geological horizons) Warren, John C., 3, 142, 286, 307, 746, 801, 1097, 1375, 1393, 1602 Warren Mastodon, 130, 164, 174, 175, 179-188, 185, 190, 211, 489, 610, 680, : 681, 1602, 1605 Warren Museum, 130, 487 Warther, George A., xvii Washington, D. C., 14, 15, 745, 759, 1088, 1314, 1609, 1621 washingtonii (see Parelephas washingtoni) water-holes, 666, 679 Watson, David Meredith Seares, xii, 15, 801 Wauneta, Nebraska, 1003 Wayland, E. J., 4, 801 Wayne Township, Drake County, 1097 Weber, Max, 15, 1252, 1329, 1630 Weddell Collection, 550 Wegner, Richard Nickolaus, 4, 251, 259, 801 weights of elephants, 1604 Weimar, 558, 969, 1042, 1045, 1047, 1050, 1056, 1057, 1181, 1184, 1188, 1217, 1233-1235, 1245, 1253, 1609, 1620, 1621 Weinsheimer, Otto, 3, 81, 85-95, 801 Weissenfels, 1045 Weithofer, K. Anton, 3, 19, 640, 801, 915, 933, 971, 974, 989, 1172, 1233, 1241 Weiz (see Oberdorf) Weld County, Colorado, 403, 488, 737, 742 Weller horizon, 312 Weller ranch, 385 Wellington, Kansas, 1002, 1003 Wellington Valley, Australia, 1390 Wentworth County, Ontario, 1069 West, Andrew F., xvi Westhofen, Germany, 251, 281, 737 Wetmasut, 825 whale, 616, 619 Whipple, Ralph W., 1067 White Beds, 677 white rhinoceros, 1194 Whitfield Mastodon, 180, 181, 759 Whitfield, Robert Parr, 4, 801 Whitlingham tooth, 620, 621 Whitman County, Washington, 1047, 1088, 1100-1103 Wiehle, Mr., 1322 Wies, Steiermark, 91, 92, 115, 384, 394, 742, 1464 Wiesbaden, 1050, 1056 Wiesberg, 281 Williams, Coleman Shaler, 15 Williams, J. Leon, 1607 Willis, Bailey, 1483 Williston, Levy County, Florida, 384, 386, 400, 416, 419, 742 willistoni. See Trilophodon (Genomastodon) willistont Wilman, M., 15, 987, 993, 994, 1281, 1282 wilmani (see Palxoloxodon wilmant) Wilmarth, M. Grace, 1509, 1630 INDEX Wiman, Carl, 732 ; : eae : a F wimani (see Serridentinus wimant) Windsorton, South Africa, 944, 945 Winge, Herluf, 4, 27, 30, 33, 71, 801 Winnipeg, 137, 736 Winterthur, 134, 216, 218, 745, 760 Winton, W. E. de, 60 wintoni. See Phiomia wintoni, and Phiomia wintoni cf. serridens wire sectional method, 911, 919-927 Wisconsin (IV Glacial), 170, 173, 190, 725, 996, 1040, 1083, 1094, 1097, 1099, 1101, 1133, 1135, 1510, 1512, Pl. via Wiseton, Saskatchewan, 997 Witsen, Nicolaes, 2, 801, 1125 Wolfsheim, 281 Wolockowoi rugzei, 1120 Wood Mountain gravels, 1508 Wood Norton, 1193, 1609, 1619 Woodring, 1446 Woodward, Arthur Smith, 4, 15, 802 Woodward, G. M., 952, 966, 967, 1223 Woodward, H., 884, 1132 Woolly Mammoth (see Mammonteus primigenius) Woolly Rhinoceros (see Rhinoceros antiquitatis) Wortman, J. L., 4, 802 Wray, Yuma County, Colorado, 251, 288, 289, 307-310, 460, 739, 1500 Wu Hsiang Hsien, 699 Wu Lan Kou, 699 Wiirm, 725, 1271, 1480, 1457, 1473, 1474, Pl. vin Wiirttemberg, 1130, 1237 Wiirttembergische Naturaliensammlung, 13, 1253, 1609, 1620 Wiist, Ewald, 4, 802, 1050, 1058 wiisti (see Parelephas wiistt) Wulstkanten, spurs or arétes, 199, 201, 212, 219 Wyoming, 1003, 1012 xiphodonts, 34 Yabe, Hisakatsu, 5, 15, 802, 1300, 1305, 1420 yabet (see Palxoloxodon namadicus yaber) Yakutsk Museum, 1128 Yale University, 14, 183-185, 526, 527, 731, 745, 760 Yamawaki, 908 Yang Mu Kou, 699 Yangtze River, 835, 859 Yarmouth, 671, 682, 725, 726, 1033, 1510, 1512, Pl. v1 Yates, Lorenzo G., 3, 487, 802 Yatsu, Naohide, 15 Yaunde, 1193 Yedo, Japan, 1303, 1334 Yedobashi, 907, 13833 Yenangyaung, 737, 824, 825, 827, 842, 843, 845, 861, 863, Pl. 1v Yen-ching-kao (see Yenchingkou) Yenchingkou, 817, 818, 875, 1453 yoke-toothed mastodonts (see Zygolophodontinz) Yokohama, 897, 1188, 1289, 1301, 1417 yokohamanus (see Palxoloxodon yokohamanus) Yokosuka, 906, 908 yokotwi (see Bunolophodon yokotii) yorki (see Archidiskodon yorki and Palzxoloxodon york) Yorktown formation, 733 Yoshima-mura, 893 1675 Young, Chung Chien, 802, 807, 816, 894, 897, 899, 1362, 1418, 1452, 1488, 1594, 1623, 1630 Yuan Chii Hsien, 1487 Yii She Hsien, 721, 1418 Yiinchu basin, 1482 Yiishe, 837, 897, 899, 1402, 1483, 1484 ytishensis (see Stegodon ytishensis) Yukon, 1088, 1134, 1185, 1145, 1161 Yuma County, Colorado, 307-310, 460, 739, 1500 Yunnan, China, 115, 349, 355, 739, 817, 1451 Zacapt, 1013 Zacualtipan, Mexico, 1506 zaddachi (see Zygolophodon borsoni zaddacht) Zalensky, Vladimir Vladimirovich, 4, 802, 931, 1127, 1129-1131 Zanesville, Ohio, 1047, 1084, 1097-1099, 1409 Zanzibar, 1204 Zdansky, O., 899, 1480 zdanskyi (see Stegodon zdanskyt) Zebbug Cave, 1182, 1187, 1257, 1262, 1263 Zen Range, 1440 zeuglodont, 36, 37, 53, 400, 1424 zeylanicus (see Elephas indicus ceylanicus) Zeylonae, India, 1310 Zingiberaceae, 718 Zittel, Karl Alfred von, 4, 93, 802, 912, 1118, 1630 Zoological Garden, Berlin, 1401 Garden, Rotterdam, 1314, 1329, 1331, 1609, 1620 Gardens, Egypt, 1194 Institut en Museum, Amsterdam, 15 Museum of Academy of Sciences, Leningrad, 1128, 1148, 1387, 1388 Park, Washington, 1314, 1609, 1621 Society of London, 15, 745, 760, 1199, 1200, 1332, 1336, 1609, 1615 Zoologische Staatssammlung, Munich, 14, 759, 1609, 1615 Zuffardi, P., 4, 802, 1049, 1055, 1123, 1137, 1404 Zukowsky, Ludwig, 4, 802, 1193, 1221, 1409 zukowsky?i (see Loxodonta africana zukowskyt) zulu (see Loxodonta zulu) Zululand, 1187, 1286, 1402, 1437 Zumpango, Mexico, 554, 1013, 1080-1082 Zurich, 134, 212, 216, 218, 745, 760, 1464 Zwenigorod, Russia, 1136 Zygolophodon, 10, 11, 43, 124, 132, 162, 166, 191, 192, 194, 195, 197-206, 211, 212, 223, 254, 525, 686, 688, 690, 694, 718, 736, 819, 822, 902, 903, 1401, 1545, 1546, 1575-1577, 1602, Pl. 1 borsoni, 10, 30, 124-126, 132-134, 139, 164, 166, 191-195, 198-200, 203- 205, 207-211, 215, 283, 532, 633, 634, 699, 736, 748, 758, 763, 963, 1183, 1377, 1388, 1393, 1472, 1482, 1483, 1605, PI. 1 borsoni affinis, 192, 193, 207, 1397, 1464 borsont buffonis, 124, 192, 193, 207, 209, 1472 borsoni vellavus, 124, 192-194, 207, 209, 631, 632, 1393, 1472 borsoni vialetii, 124, 192, 194, 207, 209, 631, 632, 13938, 1472 borsoni zaddachi, 192, 193, 208, 209, 736, 1397, 1472 pyrenaicus, 10, 30, 115, 192, 193, 199, 200, 203, 205-207, 217, 736, 821, 902, 1394, Pl. 1 pyrenaicus aurelianensis, 115, 192, 193, 201, 2038, 205, 207, 736, 821, 903, 1410, 1460, 1464, 1605 zy golophodont, 1545 ; Zygolophodontine, 27, 30, 32, 119, 120, 124, 126, 128, 132, 191-195, 200, 689, 690, 698, 699, 734, 736, 13870, 1528, 1575-1577, Pls. 11, 11, x ERRATA Page 4: Middle column, 36th line under heading 4, for Dietrich, Wolfgang O. read Dietrich, Wilhelm O. Page 14: Left-hand column, 50th line, for Wolfgang O. Dietrich read Wilhelm O. Dietrich. Page 773: Right-hand column, first line, for Dietrich, Wolfgang O. read Dietrich, Wilhelm O. Page 841: Caption to Fig. 712, 5th line should read ‘5; 1884.3, Pl. xvi, fig. 2 (as M. perimensis);’’ ete. Page 1022: Page 1382: Page 1393: Page 1401: Page 1514: Page 1520: Footnote 2 refers to Loxodonta africana oxyotis, the upper-case 2 belonging at the end of the 2nd line of the right- hand column. Under the genera Hypselephas and Platelephas, for 1941 as the date of this volume read 1942. Right-hand column, 20th line under the heading, for Anancus arvernensis read Anancus arvernensis macroplus. Right-hand column, 4th and 5th lines under the heading, for Loxodonta capensis cyclotis read Loxodonta africana cyclotis. To the list of Proboscideans on this page add ? Rhynchothertum francist. 16th line, for Cordillerion chilensis read Cuvieronius chilensis. SRM, t ot hg re es ee 0 Wate 90E y INL LEAL A ‘ D1", Ye Vip leo f (iL fa ‘ 4 My, 744 7 ¢ 44,