HENRY FAIRFIELD OSBORN
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OTHS, LOxODONTS, AND ELEPHANTS
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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 <stc 5s sees. 9 eee bi ne eee oe eats eee eee 1327 indicus: bengalensis de Blainville; US845)...)¢ choirs as cessive sess viel ciao Sacra eee ere aero ae 1327 mdicus sumatranus Temminck, 1847. iis 55 eteye ents, oo eee end Gee Oe oe eee ee eener 1329 indicus lursutus Viydekker; VOV4 \ o 1) 5:cfastecrete cys vee sbs, HRT er eRe eee ere 1332 mMnaicus Buske |=~thaleolocodon buska|) Matsumoto, O2/ame meester eae ee inet einer 1333 Distinctions andsmeasurements ol theplncdianveleplrarityem eric reiterate tern tee ene enna seen ee 1334 Characters of the Upper Pliocene and Lower Pleistocene species Hypselephas hysudricus and Platelephas plat- YCOPNGUUS? sab de, evden HAs Aa ends mld ebbolors awed DORE OC SCTE ie CraerC ORI e ree ieee eerste 1339 Hiypselephas| Osbor\, 1936, generic definitione err reeiy- er aerate eer eee eee 1340 hysudricus Malconer and iCautley, ql S45 el S4 Gre ier yan rete Creer tec eet eee 1340 Observations of Osborn on fourteen specimens collected by Barnum Brown in the Siwaliks......... 1345 G@ranial characters: and ‘affinities? smn a.m acionenies «cis cro eee IIe ee INE ELS ree 1348 Piatelephas) Osborn, 91936; ceneric detinitionyars tarot ere atte lee ere ate erence ee 1358 platycephalus Osborny 1929). «<0: sake vaya otesstelateishoiaie forseiol|skai-/ <P ake fe Ee ee nde ene rete ae 1359
JXpayosra bbs way {Olney rye -O.6- goqcogqcoso sod soocndacasasen sna ain olatalioci iaatid Merete Stewie s eruaqu rome 1362
xil CHAPTER XOKTE OME
OSBORN: THE PROBOSCIDEA
PAGE
NOMENCEATURE ORCEEYPROBOSCIDBAS eo atte akee nena eee eee eee eae 1363 dhevAmenicans Viastodons (Miastodoniamentcants) peerieaeteier eerie mei reier i are ere 1363 The Northern or Woolly Mammoth (Mammonteus primigentus)... 2.0... 000 cee ee cece cece eees 1365 Mistiofisupertamillestr. 2 sxc veryerae esciye ata aera eee ee PC A TSE CREE OE Tea ERR ee 1367 LGTES] C0) O41 EW 00) SST es ede Dea RA CRIN A ELIE AN aati PIA eo dics Am Nag ah am one hala foc 1368 Iastrotsublamiliess, 32215 vc sete eee co ls SPS aero Ss OLR eT Ce Ee TS 1369 TAS HIOM GENES hie chasse eciteie «soo ssl ctemele wit oat ae ae Ravane Aint eee eat atte teers er a ee Re cielo eee 1371 Ihistiof«species; subspecies, andivanleties.. «sacri peter ote aavieck coe eee oe PO rae ner Poker oeea ie ee ee 1382 RAE IGHOLOGIC SUCCEHSSIONOH HES ROB OS Cl HA eee eer ee eae 1421 U.N a (Cr: eee Ee ee SR en eR Re ee Nr ee ns PEs Le Men Rea ae aS fag ee a air remae eeg oe T - 1422 iHoceneyand:Olicocene/of WNortheAtrica sn eet ae ee ee er oe eee eee eee 1423 Miocene ON Orth: Atri cas. eae ces aychs sei Ess A SL Mk SNES Oa ee 1426 Miocene; of (CentralvandsWastvAtnicas ias.5cnececk <n eeetaeeetok we ceva eae oe ee 1428 Pleistocene of North: Afni as .05.c. Aotrieueoie Rake any ask Aa Oe oe EIS Can ee 1429 Pleistocene,of¢Centraliand’Mast wAtricay..14a05 anc herd oh eee a Cee eee 1432 Pleistocene of South Adri case ys sik aes ek tice ssavoceaauses etek sehen Dra ee ee 1437
dj WWeYeW Ohute al teeny erencarpe tere MeeIats teno eter Cra ca ern Mee eee Ene Tee Ce ennirreer eS fp Preheat een ac 1439 IMiocene*of¢BaluchistanvandeSindlys on -raci ie rave ke era ace Nee CE EC eee ee eee 1439 The Siwalik series (Miocene—Pleistocene) of North India.................... 0. cece cee cee eee neeaes 1442 Bleistocenexofs@entral lari as pieces: sore Se techies ote Sates PRON ey ELE ere ae 1447 Pleistocene oli @eyloni naa erte noe elevator nc> (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. <cctate re onu A MSPS Na el see eG enn eB ooh eae Taare en ene Nahe ee 1516 Pleistocene vo Amgen tin ayes. eke oties cnc latere sine eet ate Sra toro ee ee rere eee cen ete rece TnI 1516 Pleistocénevof-the Andeanvvalleys 0s 455-850 matin eure skivnadis Broce serials Bereta tent ee NSE eRO tee eee etees it ue ectets 1519 Pleistocenexol- Brazil and HrenchiGuwisinay, or. etveo nso oeye ee et etenste csv eto ool naae ee EI erotic 1521
CONTENTS xiil
CHAPTER PAGE XXIII. AFFINITIES, MIGRATIONS, AND PHYLOGENY OF THE PROBOSCIDEA: A SUMMARY............. 1523 HAV OLSUD OT SINT VES mye gea- crane p sa erste ee te MONA ere oy esos RC ave eee TR A tc Sat ec Ea ee O 1524 IDSC] SN RAINE CEUET SSI 5) eSNG Shap eae etre ects & tr AE SPR RN rae eT D y 5, tO aR con Dre ree maa NE ee 1525 pihwenty-OneisU bam lies sey srr cy sere ere ede ees ace Gre we CEN ANE Pee ovcas cdot) Secs tay ae cioee ae ee 1526 HH Ontiyat OUT Le ENOL yeweuen ete Ee ear en ie CASE SUSES one sa AIS oh pce era oa ein ual rcee sare as aia cao te ANE 1526 Walidtspeciesn (Gaz) memaratrrctre emer ercrs 6 caterer socio IMe cari res ane ai cP yay Uh SEEN Rep ke ee a 1527 Osborn’s final (1935) classification of the Mceritherioidea, Deinotherioidea, and Mastodontoidea............ 1529 Osborn’s final (1935) classification of the Stegodontoidea and Elephantoidea..................2.2200 000008 1539 Explanation of terms used throughout the text of the present Memoir...................... 0... Boreas cits 1545 Characters; atinities sandimigrationsortheseroboscidesaen aaniee eee eee eee eee eee 1552 IMicerit hres ates ye trate eevee cece me Ce eet ACA a Senco UPC TTT ae (REE Re a 1552 ID GINOLMETES steers reste se ace ero econo ROPE Gaerne toe aan AaRcPT CN Ce ne ee 1553 Longirostrines (genera, T’rilophodon, Megabelodon; subgenera, Genomastodon, Choerolophodon, Tatabelodon)..1555 Gnethabelodonts(Grathabelodon) mew cic isk ore ee etn Teta nies ee tre ee ee 1557 Amenelodontss CAuvebelod one htOnlea) Renee enna eer a earner 1558 Tetralophodonts (genera, Tetralophodon, Morrillia; subgenus, Lydekkeria) ................ 000.0000 1559 Notorostrines:(Cordullerion) rs ivr icin crn aoc Ooo On eee ae ee ne ee eee 1560 Rhynchorostrines (Rhynchotherium, Blickotherium, Aybelodon)..... 0.0.00... 00 0000 cee cece cee ee 1561 Brevirostnmess (ANanicus wenialopnodons Sy Nconolopiiuls) Meniniareeeeeiaen eran ener 1563 Humboldtines (Cuveronius, Eubelodon, Stegomastodon)). ..1....252+05s0505s ss. 58+ +oesse esses ee 1566 Serridentines (Serridentinus, Ocalientinus, Serbelodon, Trobelodon).... 2.0... .0 0c cee ccc eee v cece eee 1568 Rlatybelodonts,(Blatjbelodon.-horynovelodon) ameeeeioee eee een eee 1570 Notiomastodontsy(Notomastodon) eater eee eee een een nn eee 1572 iRalscomastodonts! (Palvomastodon) rar aici een ae OE nn Pee oa Cee eee 1572 Mastodonts| (Mastodon, Maomastodon: Pluomastodon)s 0... 5.02 eo ceeee ee ode ee eee 1574 ZV <OLOpHOGonts|(ZYGolopRodon whurici.s) ee eee Cee ecient eee oeeeorie 1575 Stecolophodontsil(Stegolophodon) isn. se cian cele Oe ania is ee ea oe ic ee Cee aoe 1578 Stegodonts(Stegodon) sire scree svsns over oe heey oere eer Oke eee On a EE Ee eee 1579 Mammontines (Archidiskodon, Metarchidiskodon, Parelephas, Mammonteus)............ 0000000000000 1582 Loxodontines (genera, Loxodonta, Palxolorodon, Hesperoloxodon; Sivalikia, Pilgrimia, synonyms of PAleOlOr OM ON)! oscesnsycat sos: spat tuaisiehs aeeea.8 trator steaks Sa eee Ee Oe 1590 Elephantines (Hlephas, Hypselephas, Platelephas)........... 0.0.00 000 005 abe fa, dish. oy tape senile ee 1595 Skeletaltmatertale to.c'sos 25.0 cece dhsus actus clio tueeto ante oni ae Or een IST nec eae eee 1600 iHeightsiofsproboscideans.estimatedvandtactualea-emeeeaeeeeee eee ebioeace reer renee 1604 APPEND TO VOLUME IT RROBOSCIDEAN DENTAL TNISTOLOGYAee eee eee eee eee 1607 LIST OF CATALOGUE NUMBERS OF SPECIMENS MENTIONED IN THE PROBOSCIDEA MEMOIR, NO} TAGIM 0 20 0 (ee re ee eo ean atl wine Kine at orp Une timc aos Ooreoe css bo0 boo c000508 1609 SUPPLEMENTARY BIBLIOGRAPREY. .....6 0. ccceen lene ear sna scent eee eee Oe eee 1623
INDEX TO VOLUMES. T AND Tl. .cc52.e.ccntedideted Gedenst a ctai cece ons ne Aetna ei eer aera 1631
LIST OF ILLUSTRATIONS
PLATE PAGE Frontispiece. Restoration of the Jeffersonian Mammoth (Parelephas jeffersonii). After painting by Knight. PRUNE SLOOP HOcOntine ss iSteqOlophOdONer® cs mindy. acai ista Siar caieie aes bls ecw ace tus ba aise e Mela Oe Link oe AS Beretta 850-851 XIV. Migration and evolution of Meritherium and Deinotherium.........0. 00.0.0 cece eee cee cece cess seseess 1552-1553 XV. Migration and evolution of Trilophodon, Megabelodon, Gnathabelodon, Phiomia, Amebelodon, Tetralophodon, and LH PLR Sere TAR Ces Tok cee Oe Oe are OOOO ere ae eta a aha es 1556-1557 XVI. Migration and evolution of Cordillerion, Rhynchotherium, Blickotherium, Aybelodon, Anancus, Pentalophodon, and SSVRTDTUE/D ACE cca TEN AERPET oD rCoe hNe AE NE RHO TERRE CCN ae an a gO ae aa SA 1560-1561 XVII. Migration and evolution of Hubelodon, Cuvieronius, and Stegomastodon..... 2.0.0... cee cc cece eee ee ve eeeees 1566-1567 XVIII. Migration and evolution of Trobelodon, Serbelodon, Serridentinus, Ocalientinus, Platybelodon, Torynobelodon, and INOLCOTR ASLO LOT ine Nahas SNe este NOT ara Fey RUA oa he og, ES SPL Sloss er peo at nai e VORA eee et re ea 1568-1569 XIX. Migration and evolution of Palzomastodon, Miomastodon, Pliomastodon, Mastodon, Turicius, Zygolophodon, and SUA QUT ODD 5 See CRG SG SOIT Some CAG SRM Ne OEE CO eT ee ee SE 1572-1573 NOXCoe Miprationzan devolution Oly Stegod ord act s.e.oaicerstercte te cassie raver stcys SENSI hous, SE yorsie ieee oes eer ERIE nee ioe 1578-1579 XXI. Migration and evolution of Archidiskodon and Metarchidiskodon............. ccc cece euveeecceeeeeeeeesen 1582-1583 XXII. Migration and evolution of Parelephas, Mammonteus, and Elephas.......... 2.0.0.0 200 ccc ceeeee cece eeeeeeeee 1584-1585 XXIII. Migration and evolution of Loxodonta, Palzoloxodon, and Hesperolorodon.......... 0.00200 ccc ce eee eee 1590-1591 POG Osborm=neccs)! Comelationmibableron 1922; and 1929" ease oes eae ees oaoe oer ane ee anes 1606-1607 XXV. Northwestern India and adjacent territory, showing especially the Siwalik Hills where Falconer and Cautley made their classic collections and Dr. Barnum Brown recovered in 1922 the fine Proboscidea collections for the ANSoaVST eC ai CTA RUISTAYDI toed arenes a eaten oe ol arch aN aeRO Rene tn Se tiven ae Rec = on Were tn Merce Lhd Are ce WEL Ye 1606-1607 XXVI. Elephas indicus: (Fig. 1) Complete transverse section near tip of unerupted tusk. (Fig. 2) Area of thin section within circle on Fig. 1. (Fig. 3) Same as Fig. 2, photographed between crossed nicols...................... 1630-1631 XXVIII. Elephas indicus: (Fig. 1) Area within circle on Pl. xxvi, Fig. 2. (Fig. 2) Transverse section of about half of a small, mature tusk about 14 inches long. (Fig. 3) Part of section within circle on Fig. 2. (Fig. 4) Same as Fig. 3, with crossed nicols. (Fig. 5) A small area near outer edge of cement band of section shown in Figs. 2-4. (Fig. 6) Phiomia wintoni: Transverse thin section about one-half inch from tip of small, worn tusk............... 1630-1631 XXVIII. Phiomia wintoni: (Fig. 1) Same as Pl. xxvny, Fig. 6, with crossed nicols. (Fig. 2) Area within circle on Pl. xxvun, 18h Bassa ds AS eR Ree cere aE td eRe eae rene a Ee Se She edie oe acd do cok tus 6 1630-1631 XXIX. Trilophodon obscurus: (Fig. 1) Transverse section of part of tusk. (Fig. 2) Area within circle on Fig. 1. Crossed nicols. (Fig. 3) Same tusk as Figs. 1 and 2, thin section across enamel cut in a plane vertical to surface and parallel to longitudinal axis of tooth. Crossed nicols. (Fig. 4) Trilophodon (Megabelodon) sp. Transverse thin section of tusk near edge of enamel band. (Fig. 5) A different part of same thin section as Fig. 4. Crossed TANKG) Sioa anes CS Cee ee eae eR See i BO EE eA mee Goa Peer U cccb ade oles e hae hbakcosos 1630-1631 XXX. Trilophodon (Megabelodon) sp.: (Fig. 1) Vertical thin section of broken molar tubercle. (Fig. 2) Part of section in circle on Fig. 1. Crossed nicols. (Fig. 3) Transverse thin section of a molar cusp, from same tooth as Figs. 1 and 2 but a different cusp. (Fig. 4) Part of section in inner circle on Fig. 3. (Fig. 5) Part of section in outer CUP CIE LOM BIO ors fos cscs rm wv o0 le y's Susans oe VIEWED OL OY RTT OTe ee RO CC Ee eee 1630-1631 FIGURE 681. Scene on the ancient Solo River, illustrating the Dietrich-Osborn theory of the Middle Pleistocene age of Pithecanthropus erectuswelhestoration by sFlinseht.;< 4.04. = 5 3s sey Gewese See ee Cae Er REE DO See ee eee 804 6825) Wamuily,crouploh Stegodon omentalis grangenz. Restoration by, Hlinsehis--. 24 -).2 4-460 2222-1) eid aise eee 804 GSaumstegadomelepnantordes(—clifiz) cotype molars Atter Wy dekKeree trie eater ile ii erie neta eee 808 GSAaameSLegadononentalis qnangen) relerred mirsh ebb SuUpenor m0] aie gene arene e eee ee cee ene 808 685. Structural evolution of the cones, conelets, and ridge-crests in the Stegolophodon phylum, in comparison with Palzomastodon. 810 686. Structural evolution of the cones, conelets, and ridge-crests in the Stegodon phylum..............-.....02-.00...0000- 811 OS7ae Gradualsprogressive hy psodonty im) superior grind ersiolSteqodona ne lee eee eee eee eee 813 688. Gradual progressive hpysodonty in superior and inferior grinders of Stegodon.... 2.1.2... cece ee eee 813 GSoReiunccrusicompareduwithinlophodon molarsy After Viay chm arrri ie eisai iste iii erence eee 819 C90Reelunccrsandystegolophodon tormrot, grinding tectht ws eee rae eee sear eee ee eee 821 691. Map showing geographic distribution of types and referred specimens of Stegolophodon and Stegodon.................. 823 6925eeHossil-beaningshorrzonsral ong wtihe wars wad clive Eun erg U0 2 aye aes eevee ete area akg en eae 824 Goss steqolopnodonilatidensplectotype palates After. C lites esmenier rere eer ete nese ie ey eee 826 694. Stegolophodon latidens, cotype third right inferior molar. After Clift................................ 222s eee eee 826
XV
Xvl
FIGURE
695. 696. 697. 698. 699. 700. 701. 702. 703. 704. 705. 706. 707. 708. 709. 710. 711. 712. 713. 714. 715.
716.
OSBORN: THE PROBOSCIDEA
PAGE Elephas cliftzi Falconer and Cautley equals Stegodon elephantoides (= cliftii), cotype first left superior molar. After Clift... 826 Stegodonselephantoides ecto typellower jaws eAtter Chitty ctr ee tee ernie nen eee 826 Stegodon insignis, lectotype and cotype molars. After Falconer and Cautley.................... eee eee cee ween eee 829 Stegodon ganesa, lectotype third superior molar. After Falconer and Cautley..................2 20 cece cece eee e eee ee 829 Steqodon bombifronscotype skull. Atten Halconenand @autleyzeeiermmian cee aoe ieee ere eerie 830 Elephas cliftii, original type figure of first left superior molar, equals Stegodon elephantoides (=cliftii). After Clift......... 831 Elephas cliftii, new type figure, equals Stegodon elephantoides (=cliftiz). After photograph of type cast............... 831 Stegodon sinensis, type third superior deciduous premolar. After Owen...............2.6.2.- esc e reste ween eee tees 831 Stegodon orientalis, type. Portion of true molar and posterior end of milk molar. After Owen....................... 832 Stegolophodon cautleyi, lectotype third left superior molar. After Lydekker..................... 50. cee e eee e ees 832 Sieqodonnmgonocephalus. type momature) skull eAtbenrgy lantimere terse eel sire ist eileen eee eet 833 Stegodon (Archidiskodon?) mindanensis, type molar. After Naumann......................... see c este eee eee es 833 Siegodonaimawana sy pelowelaw. Auver Mantineemasrera ae rac science eee irc cacti oe cit eric ener ree 834 Stegolophodon stegodontoides, type right third superior molar. After Lydekker..............0..0 0.0.0 c cece eee eee 834 Stegodon aurorz, type right second superior molar. After Matsumoto..................... eee e eect eee eee 834 Sfeqolophodon sublaiidens sty pe molar. At ter p NOLO RTA le ror nyse rere ya teins eet 835 Sicgadon punjorensis aby pe) palate: Alter PAOtORTap Ors: er sae se lel eine) ieee er nstere neler teen ee 835 Stegolophodon cautleyi, lectotype left third superior molar. After Lydekker..................... 0... c eee eeeeeeeees 841 Stegolophodon cautleyi, lectotype left third superior molar. After Falconer and Cautley.....................2...20.. 841 Stegolophodon cautleyt, cotype first superior molar. After Lydekker..................-.-- cece eee e teen etre eee 842 Stegolophodon cautley1, referred left second superior molar. After Lydekker................................--00-00 842 Stegolophodon latidens, lectotype (new figure) right second and third superior molars. Orthogonal projection after cast.. 848 Siegolophodonmatzdenscotype tbninduntenior molar sAtter Cliftee ee eerie ree aeeecetr eae aa eee eee 843 Stegolophodon latidens, lectotype right second and third superior molars. Perspective. After Clift................... 844 Stegolophodon latidens, lectotype molars. Section after Falconer and Cautley, in comparison with Mastodon americanus,
Ching tSUPETIOnMMOLAR ss sysnersge eee ier cere oo A iam savant ele ye CERNE cee een Gh Sines sn eee ne 844 Stegolophodon latidens, lectotype third superior molar. After Gaudry................---:-ee esse eee cette eee reese 844 Stegolophodon latidens, referred third right superior molar from Japan. Primitive stage. After Matsumoto............ 845 Stegolophodon sublatidens, type. Posterior half of a third right superior molar. After photograph.................... 846 Stegolophodon stegodontoides, type third right superior molar. After Lydekker.......................--..--000 eee 847 Stegolophodon nathotensis, type fragmentary molars. American Museum (Barnum Brown) collection................. 848 Stegolophodon cautleyi progressus, type cranium. American Museum (Barnum Brown) collection..................... 849 Stegolophodon cautleyi progressus, type right superior dentition. American Museum (Barnum Brown) collection........ 849 Stegolophodon cautleyi progressus, type skull, left lateral, palatal, and right lateral aspects. After photograph.......... 850 Stegolophodon lydekkerz, type third left superior molar. After Lydekker....................2... 0 ee sete esse eee eee ee 851 HaAlconensinaplomtiner cologysolmlndlammerer pric: nice ce Tacit SE ciel ays cic ois ccc oases Ree ee ae aCe ara 852 Map of chief Miocene and Pliocene fossil mammal deposits of Asia. After Osborn...............0000 0200s eee eee ee 853 Species of Stegodon from India, China, and Java. Restorations by Flinsch...................... 2c eevee teens 855 Stegodon ganesa, S. insignis, and S. bombifrons crania. After plates by Falconer and Cautley.....................5.. 856 Stegodon ganesa, cranium with tusk extremities turned inward, in contrast to Falconer and Cautley’s restoration (Fig. 732)
WithMusksaurmedmounwardeands closely, appressed 1-1s<ci1s)ocrcic cine eater iereicenl eee Ice arene ence erates eh teat erate 857 Stegodon bombifrons, lectotype. A generalized cranium. After Falconer and Cautley.........................2.20.. 858 Stegodon insignis, a specialized cranium. After Falconer and Cautley.............2.... sees teers 858 Stegodon ganesa, a specialized male cranium. Restoration after Falconer and Cautley..........................000. 858 Stegodon sinensis, type third right superior deciduous premolar. After Owen................2220 cece ee eee eee eens 860 Stegodon elephantoides, lectotype second and third left inferior molars. After Clift............... 0.00002 cece eee 861 Stegodon elephantoides Clift (=cliftii Falconer). Cotype first left superior molar. After cast....................0-5. 862 Stegodon elephantoides Clift (=cliftii Falconer). Cotype first left superior molar. After Falconer and Cautley......... 862 Stegodon elephantoides (=cliftii), referred third left inferior molar. After Falconer.....................020-.0000-0-5 863 Stegodon bombifrons, lectotype, cotype, and referred crania. After Falconer and Cautley......................00-05, 864 Stegodombombijronss” Restoration by-blinschy aie. ecmeciss acts ccc ee oe tne oe ee ae alee ne ne nee ep reer 864 slegoaon vombijnons) cotype skull: yAfter Haleonerand:Cautley eee ener cece ane eee eco elscietetasity tae iris 865 Stegodon bombifrons, referred third right superior molar. After Falconer and Cautley.................-.....+-.005-. 866 Stegodon bombifrons, referred fragment of third right superior molar. American Museum (Barnum Brown) collection... 866 Stegodon insignis, lectotype and cotype molars. After Falconer and Cautley.................. sce e ects eect eeee 867
Stegodon insignis, referred superior and inferior molars. American Museum (Barnum Brown) collection.............. 868
FIGURE 749. 750. 751. 752. 753. 754. 755. 756. 757. 758.
Cm oO mH res
LIST OF ILLUSTRATIONS xvil
PAGE Stegodon insignis, referred second right superior molar. American Museum (Barnum Brown) collection. .............. 869 Stegodon insignis, referred inferior mandible. American Museum (Barnum Brown) collection. ....................... 869 Stegodon insignis, referred juvenile and young adult lower jaws. American Museum (Barnum Brown) collection. ...... 870 SLeqodonmnsigmasy reterred craniay »AtverHalconerandsCautleyaqe-) aa ens see eas eaee ee seeneee eee eens 870 Stegodon insignis, referred third left inferior molar. American Museum (Barnum Brown) collection.................. 871 Stegodon insignis, oft-reproduced referred skull and tusks. After Falconer and Cautley.................0.0.000020.. 871 Sieg Od Oneg GILES nme NV CSLOLA WOME Ya H INSehie pees snr ASAE teL eka erates ica Mee nea ae PSP ae er) eee ae 872 SGOT, CASIO, INET OREN AON lon TMNT. oo gooovoeavt sugeoouoweoeeaeeuspodgeoconuossonenneonemanonndunpune Aue 872 Sectioned molars (lectotypes) of Stegodon ganesa and Stegolophodon latidens. After Falconer and Cautley............. 874 Stegodon insignis birmanicus, type ramus with third left inferior molar. American Museum (Barnum Brown) collection. 875 Stegodon orientalis grangeri, type and referred molars. American Museum (Walter Granger) collection. ............... 876 Stegodon insignis-ganesa ref., S. insignis birmanicus type, and S. ordentalis grangeri type. Comparison of left third inferior TXNO | US PROME RN ote Piya eC Tes SN cs yoy EA Le REP ONGP oc foy sc RU ah an ct I yc IGA Rae Ee Sari ren en ne a 877 iSteqadannomentaus grangenz, relerred superior and interiom molars. soe. sencilla aac cere te eve cele ee eee eens 878 Stegodon orientalis grangeri, type and referred superior and inferior molars. After photographs....................... 879 Stegodon orientalis grangeri, infantile, juvenile, young adult, and mature adult crania...... 0.0.0.0... 0.00.00 eee ee eee 880 Stegodon pinjorensis, S. insignis, S. airdwana, and S. orientalis grangeri, sections of third left superior molars.......... . 881 Stegodon pinjorensis, type skull. American Museum (Barnum Brown) collection...................00.....0 02.0000 882 Stegodon insignis, referred lower jaw and right superior tusk (American Museum, Barnum Brown, collection), compared with S. ganesa, referred skull with tusks, after Faleoner and Cautley...................00. ceeeee ete ee cee eeee 882 Stegodon pinjorensis, type skull, also same skull superimposed on referred skull of S. ganesa...............-..-..-00.. 883 SATU. (HOTTA, INORG Loy MINN Nog go nacponcocgooncbecobatencovencecoduneenedoveesanusessoeuauaover 883 Stegodon orientalis, type. Portion of true molar and posterior end of milk molar. After Owen....................... 884 Stegodon insignis(?) = orientalis(?) ref. and Serridentinus lydekkeri type. Molars, after Schlosser...................... 885 SATO CUMODGNG, WHATS TOMER ee “Ne INGRAM Goo nacho pa poo+ na oeandmbannneocecdsoraonoodouuncaeencvnconovad 886 Via prot kendengshon zona brim yavaseAticerel Ub Oise yews prci rere eeraic riers vey ieee enya ee 887 Swajcin quran, waicrnxel Sawlll, AMET AEN: ono ooe nod doaoosonowssoodsungaccns snoenenadsongoducaanoovapes 888 SAO GORDO INGTON HOM OZ IMIS; oo oo bingognoaddoopdoonebuddeds adgoonbosanacdseassongenbosadanncous 889 Sia Odlon, Hr teOC TOGO, INEWONoN lon NMAC so ocnoscucsucconoancozphyponoumonhsansohSnnenoosoucosenuanvose 890 Stegodon trigonocephalus, type skull, after Martin, and single ridge-crest of molar, doubtfully referred by Naumann to S. UST TUPSHOTA LOIS #4 TUES CU 2 chen, etn BE eee aM ait SNOT eo Tire So aC re PoE eve ae 890 Stegodont crania: Stegodon pinjorensis type, S. bombifrons cotype, S. orientalis grangeri ref., S. trigonocephalus type, S. HIROKO KURI) yok es ee ae ee aE ee ee Oe ir coe ona ite ain avis © Aeon ois Geos Shas cob mace oeoo6 O04 891 Stegodon (Archidiskodon?) mindanensis, type inferior molar (incomplete). After Naumann.....................-.... 892 Stegodon airdwana, section of referred second molar, and section of ridge-crest of third superior molar. After Janensch.. 892 Stegodoniaurore) ype second might superior molars Atte Matsumoto reser eee eiieie ciel eee Nari elaine eee 893 Stegodon aurore, type second right superior molar (vertical section). After photograph............................. 893 Stegodon bondolensis, type mandible with third molars zn situ. After van der Maarel.................. pe oie sec eel) Stegodon trigonocephalus praecursor, type third left inferior molar. After von Koenigswald.......................... 896 Parastegodon? kwantoensis, type lower jaw with right second molar. After Tokunaga.........................0000-. 897 Stegodon yitishensis, type third left superior molar. After Young....... Seb JR a i eae ee eee 897 Stegodonmopicinalisy typermolan tragment se Atiter lel OP W.OOCle jessie relia i ser rites tener 898 Stegodon officinalis, referred molar fragment. After Hopwood................................. Schr {Bee 898 Steqodon edansky7 «type thirdnightanterion molars eAtiter Elopwoodeass ee aoe aaa eee aera 899 Parastegodon [Stegodon?] sugiyamaz, type, probably a left second superior molar. After Tokunaga.................... 900 Thirteen fossil mammal-bearing formations of Japan. After Matsumoto.................. 000. cee cece eee ress teeee 902 MheoretickphylozenysomthelMfastodontidsess Afters Matsumotomeeeres teeter eee einai einen 903 iMheoretieyphylogemys ofthe Stezodonts:, “Aiiter Matsumotomaarreeiciente eileen 904 Theoretic phylogeny of the Elephantide of Asia and Europe. After Matsumoto....................-.0eeee ee eeeee 905 Elephantide: Primitive, intermediate, and progressive mandibles and grinding teeth. After Falconer and Cautley..... 910 General climatic distribution of the subfamilies of the Elephantoidea and Stegodontoidea, including theoretic migration [nme sa OSS) eee eae cekke ceases of SO CE ae ere Seo 2h PPS Ge TS PE Ben 914 Asiatic lephant a juyenile cranium. eAtter, Os bOnmran ds Ge SOL V7. genteel erate rar tte eee o-%. AGG Asiatic elephant, juvenile cranium, also orbitosphenoidal region, left sce. vies Osborn and Gregory........ Foes ee ee Asiatic elephant, infantile cranium, basis cranil..................... HT e dip alels Or CARERS GL One ee 917
Asiatic elephant, infantile cranium, occiput and jaws............ Sit EL Ge EAE Ae 4. ye RL HOE htop 21S
XViil
FIGURE 800. 801. 802. 803. 804. 805.
806.
807. 808. 809. 810.
811. 812.
813. 814. 815. 816. 817. 818.
819.
820. 821.
822. 823. 824. 825.
826.
827. 828. 829. 830. 831. 832. 833. 834. 835. 836.
837. 838.
$39.
840.
841.
OSBORN: THE PROBOSCIDEA
PAGE Asiatic elephant (Hlephas indicus bengalensis), adult cranium, palatal view... ... 2.0.0... 000000 e ee cece ee eee 918 Five wire section lines. Key to sections. Young Hlephas indicus bengalensis cranlum.................-0.00. 000000 919 Asiaticrelephant py oungs slither oraviewsOlskullsatter: Greg orysery menor riences ketene everett aie raters earner ae 920 Elephas imdicus cranium. Fronto-occipital growth curves of vertex...-:2....--5---.--+ss+saceseeaes sens seeen ees: 921 Elephas indicus cranium. Growth curves of vertex: midfrontal vertical section. ................... 00: s sees cece eee 921 Preliminary study of fronto-occipital-basilar planes: Elephas indicus, Loxodonta africana (adult skulls), and Parelephas jeffersonii (young male skull) subsequently referred to Archidiskodon imperator.. 0.0.0... ce cee 922
Later study of fronto-occipital-molar-3 planes. Para-occipitofrontal section of Loxodonta africana, Parelephas jeffersonii male and female (the latter subsequently made the type of Mammonteus primigenius compressus by Osborn), and
ePRG siti Cus ee Se Ge A ec RE Ate i a eo as Pahoa tala raage Ne yen Stora te ae eae Ce RE 922 Blephasanaicusysuperiornmasal crow hi stages) (Sections) /aeemee aaa ee oor kre tee ee aera 923 Mlepiasmndccus sranstrontaligrowubstazesi (Sections) years aa aces) cnet eee 923 Hlepraswnaccius .occipitohonzontalagTowthistagess(SCClOUS) seat een aie eee eel 923 Mid-occipitofrontal sections, vertical longitudinal: Crania of Loxodonta africana, Elephas indicus, and Parelephas jeffersonit
[=Alrchediskodon1mperaton) rn oats ee ree ORT NG PIS Sieyatlecs coke outta n oReTagU UN es Ue RSE ee OUR Re RE eae te 924 Frontal sections: Crania of Loxodonta africana, Elephas indicus, and Parelephas jeffersonii |= Archidiskodon imperator].... 924 Nasal contours (sections): Loxedonta africana and Elephas indicus compared with Parelephas jeffersonii and Archidiskodon
TT OIE ee RE RC eR ah ca aed Lae ea a en Pt cc Tat Oe RL rico, IRN cS 925 Midfrontal or intertemporal forehead (Loxodonta africana, Elephas indicus, and Parelephas jeffersonii)................ 925 Occipitohorizontal sections through back of occiput (Lorodonta africana, Elephas indicus, Parelephas jeffersonii)........ 925 Map. Successive habitats and world migration routes of the archaic-toothed mammoth Archidiskodon............... 934 Mid-cranialsaxess ooxodontines sHlephantines =\Vlammontines) see eeer eee iac iene elena eee 937 Archiaiskodon andy Stegodon comparative) proiiles Oli Crania.cese cere een eee sete piel rae eiealet aenea erie eens 938 Hypsicephalic crania of the Mammontine (Archidiskodon imperator, Mammonteus primigenius, Parelephas jeffersoni, and
PP DASNUNG LOND) bers, reeves eA el aE es GE aa Ss aa ae OS ee Man a een ED oy ol ST ea AST Te 939 Comparative series of superior molars showing evolution of the ridges in the Elephantoidea and Stegodontoidea (Mam-
monteus primigenius compressus, Archidiskodon planifrons, Stegodon aurore, S. ganesa, S. insignis)... 2.2... 20. 939 Map of central region of the Siwalik Hills, 200 miles south and north of Simla. ...........00.0 0.00.00 e eee eee eee ees 940 Map of chief Lower and Upper Pleistocene localities of western Eurasia in which occur species of Archidiskodon, Parelephas,
iMammonteus Loccdonia anauraleolocoaon., AtterOsbornta.e aneaser ee ee eee aici ae ere ee 941 Map showing geographic distribution of the principal species of Archidiskodonts..................50 000000. c eee eee 942 VWaalehiveronavelmerracess SOULMBATTI CAR. mttoe ).tencuelstnee aa et ei ir Meiers on Paeie hte St case sis Crea ronan ee Raee ante ime rrere 945 Archidiskodon imperator maibeni, family group along the Platte River, Nebraska. Restoration by Flinsch............. 947 Archidiskodon planifrons, lectotype right second superior molar, and cotype left third inferior molar. After Falconer and
(CHITIN seca eicuchc todo. ocx tcc, Onc er ee Cle gcc eee Cee SPE cere cath ere tei Rea err eearee ars eecicies imac oiehos ao a,c onto 951 Map of favorable exposures, southwest of Simla, of the Archidiskodon planifrons life zone, Upper Siwaliks, India, chiefly
IEHaUCole- atl eos Naearen es: Anat taro Sih Oeota to acne eo aE ae eRe OOo oman a Av 6 Gontthg ENA. y Or 6 952 Archidiskodon planifrons, new lectotype figure of right second superior molar (Miss Woodward’s drawing) ............ 952 Archidiskodon planifrons, lectotype right second superior molar. After Lydekker..................000 020 eee eee. 953 Archidiskodon planifrons, referred right third superior molar. After Faleoner and Cautley.....................5555. 953 Archidiskodon planifrons, profile of skull. Reproduced from Gaudry after Falconer and Cautley..................... 953 Archidiskodon planifrons, referred first left superior molar. American Museum (Barnum Brown) collection............ 956 Archidiskodon planifrons, referred first left and right superior molars. American Museum (Barnum Brown) collection... 956 Archidiskodon planifrons, referred second left superior molar. American Museum (Barnum Brown) collection.......... 956 Archidiskodon planifrons, referred third left superior molar. American Museum (Barnum Brown) collection........... 956 Archidiskodon planifrons, referred left third inferior molar. American Museum (Barnum Brown) collection............ 956 Archidiskodon planifrons, portion of referred first and second right inferior molars. American Museum (Barnum Brown)
POI UCEVAN 0) 0 etree, cas RE oe Oe en Lei eth, ae POA Creo ut oR ni S 3.06) o.8 pip ware .0 GD 957 Archidiskodon planifrons, referred fourth inferior deciduous premolar. American Museum (Barnum Brown) collection. . 957 Archidiskodon planifrons, fourth left inferior deciduous premolar and first left inferior molar. American Museum (Barnum
Bitch, gad ele) Veena Leh Sie iG ae ko ac ren Cee nn ee oi elder hie We Oe ae Me aire odnioG oc 5'6.0.0.0 Yo ic 957 Archidiskodon planifrons, portion of lower jaw with right third molar in situ. American Museum (Barnum Brown) col-
IVYci 00) ena es ee NM le ROR Rn Le le ee hd ee re RAN aA D Bae A) 0 ido 04. cor 957 Archidiskodon planifrons, portion of lower jaw with right third molar in situ. American Museum (Barnum Brown)
COLLET OTA S vz ehh eee a ag Be Poe Le A PAS ENE in) AA they eh SS SONS or AE ATG EEG I oe eee 957
FIGURE 842. $45. 844.
845. 846.
847.
LIST OF ILLUSTRATIONS XIX
PAGI Archidiskodon planifrons, referred third right inferior molar. American Museum (Barnum Brown) collection........... 958 Archidiskodon planifrons, referred third left inferior molar. American Museum (Barnum Brown) collection............ 958 Archidiskodon planifrons, portion of referred lower jaw with left third molar in situ. American Museum (Barnum Brown)
Cale Gnpe mest ak atten Ake ork) RAE PONG c.hbloliy yeah We RM 2 ote lah el cic 1 al sai Unease 958 Archidiskodon planifrons, referred right third inferior molar. American Museum (Barnum Brown) collection........... 959 Archidiskodon planifrons, referred left third inferior molar, also transverse vertical section. American Museum (Barnum
STOWE) ECOlECHIOTE ies eth tare Re pe Penh red! a Bh oe expr shee Med tated Pumas | Ulver (ante dan geile 959 Leith-Adamsia siwalikiensis, type third right superior molars [=synonym of Archidiskodon planifrons|. After Falconer
mine! (CRITTHLE ew aie Maree Rees aiold Sand ice oe ees col enn cen eee ara” Ce eee OR ES SS NP fe ele 960 Archidiskodon planifrons, referred cranium (supposed female). Redrawn after Falconer and Cautley................. 960 Archidiskodon planifrons, three referred primitive mandibles from the Siwaliks, India, Chagny-Bellecroix and Senéze,
HPA Cewe Atitete Via yetvam ds Ror amie 202 2 cous toc on eS USPS oN te Ga ale Me Tee res So ee 962 Archidiskodon planifrons of Chagny, referred tusks, maxille, condyle, and atlas. After Mayet and Roman............ 963 Map of type locality of Archidiskodon proplanzfrons, and type and referred localities of A. planifrons; also theoretic range
from supposed African center northward to France and Britain and eastward to India.......................... 964 Archidiskodon planifrons of Piltdown, England. Molar fragments, after Smith Woodward.......................... 965 Archidiskodon planifrons, molar fragments from Piltdown, England. Redrawn by Miss G. M. Woodward........... . 966 Archidiskodon planifrons, lectotype sixth, seventh, and eighth superior ridge-plates. After drawing by Miss G. M. Wood-
VC eee cre eG by Ored Lao n veA a: 1 herd oh os iA Oh SA Ty ad 9 Nee hae Sc te ae Cee dE Raa an als oo ee 966 Archidiskodon planifrons of Piltdown, England (sectioned molar), in comparison with lectotype and referred molar sections. 967 Scene on the ancient river Ouse, illustrating the Osborn theory of the Upper Pliocene age of Hoanthropus dawsoni. Restora-
{RYOEA Lonie | RINGANS(C) OVE ks anenat ee Aenene ReniMt odes ne eae MPN ERP A el sy ocean starr! a Run ten A) aera ire ean Mee ied ht we es hala gh CL. 6 Bat 968 Archidiskodon planifrons rumanus, type third left inferior molar and portion of referred third right inferior molar. After
SLCTAMES CU wR oor Nee MCA atts SPY cs8 RRA he rh aia an, Aan naroe he ti eo eee cy ok DU eure ee ee go Le ze = 969 PAN CIAUsKOdOnmmenvaconaus: Lectotype cranium (©)s AttiersNestin snes si cictcrirtter este) sees ieee 970 Anchidiskodonmnendionaliscovype cranium (A))s eAtiter NestiC gare cere aan ae ecient aca eee 971 Archidiskodon meridionalis, restored cotype cranium (A). After Weithofer................0... 00000. c eee eee eee eee 971 Archiduskodonimentaronalus, lectotype cranium) (©). Atiter: Weithofer..= sae. ae se es ee ee eee 972 Archidiskodon meridionalis, referred third inferior molars from the Val d’Arno and Norwich Crag. After Falconer and
(CHinUleNTs 62 det eee Ore a ee ee meee ot Ren One ait Oh Aa A Bn BAT hE Aa Ges osDeobdt 973 Archidiskodon meridionalis, referred third superior molar from Chagny, France. After Gaudry...................... 974 Elephas lyrodon [= Archidiskodon meridionalis, female], type skull. After Weithofer.......................0....00.. 975 Crania (19) of the Mammontine (Mammonteus primigenius, Parelephas trogontherii, Archidiskodon meridionalis, A. impera-
toppandeAnnlanifrons)» sAiter PohligsHalconer; Weilthofersy ance cee eee cette eee eee eee ere 976 Archidiskodon meridionalis of Durfort, skeleton, largely restored. After photograph..........................0..0.. 978 Archiaduskodon mernidionalis of Durtort. Restoration by, Plmsch:-...s2.4-022--2--- ees coe aces aa eee 979 Archidiskodon meridionalis of Durfort. Skeleton redrawn after photographie plate in Gaudry........................ 979 Archidiskodon meridionalis, referred third left (?) superior molar from Essex, England. After Lydekker............... 980 Archidiskodon meridionalis cromerensis, type third left superior molar. After Depéret and Mayet.................... 980, Archidiskodon, Parelephas, Mammonteus: Primitive grinding teeth from the Forest Bed and Red Crag, England. After
peneilusketches bythe presentyaubhorj ya ..c\-)- cea craieieaetekcr st err ae Irene oe ete ee tel ie eee eee 981 Map showing fossil Proboscidea route from southern equatorial Africa... .......2..... 05220. c ese e eee e sees eet seeess 983 Archidiskodon proplanifrons, type right third superior molar. After original......................0.. eee eee e eee ee 987 Archidiskodon subplanifrons, Osborn’s original type figure of third right inferior molar...........................-.. 987 Archidiskodon subplanifrons, type right third inferior molar (new figure)....... 22.22... 5... s tees eee cece e tees 988 Archdiskodonyplaninons, reterred third might inferior mola. seer eet eee eee eee eee 988 Archidiskodon broomi, Osborn’s original type figure of third right inferior molar......................00..0..0052... 989 Archidiskodon vanalpheni, type third left superior molar. After Dart....................... 00s see eect eee eee ee 990 Archiduskodonimalletty. stype thind left superior molar) Atiten Dante cree sete eric eesti ie acer heres 991 Archidiskodon loxodontoides, type third left superior molar. After Dart............................ eee eeee eects eee 992 Archidiskodomuyorkoatypemolariragment. Atiter Dante t = rtrersee ie eee eee er eet renee eee 993 Metarchidiskodon griqua, type third left superior molar. Original figure of Haughton and new figure after Osborn... ...... 995 Metarchidiskodon griqua, referred fragmentary molar, from Kaiso Bone-beds, Africa. After photograph. ..... nee eee LS) Archidiskodon imperator, type fragment of a third right superior molar. Leidy’s original type figure.................. 999 Archidiskodon imperator, type third right superior molar. After Osborn...................0..20.00e eee eee eee eee .. 999
Archidiskodon imperator, Leidy’s type molar and Osborn’s neotype combined. After Osborn......................... 999
XX
FIGURE 887. 888. 889.
890. 891. 892. 893. 894.
895. 896. 897. 898. 899. 900. 901. 902. 903. 904. 905. 906. 907. 908. 909. 910. 911. 912.
913. 914.
915. 916.
ONT.
918. 919. 920. 921. 922. 923. 924. 925. 926. 927. 928. 929. 930. 931. 932.
933. 934. 935.
OSBORN: THE PROBOSCIDEA
PAGE Archidiskodon imperator, type molar compared with type molar of Parelephas columbi. After photograph.......... ... 1000 Archidiskodon imperator, neotype third right superior molar. After Osborn........... 2.6.6.0... 0s eee, 1000 Archidiskodon imperator, referred third superior and inferior molars of two individuals, showing mechanical reversal of the
convexvandsconcavesunacess) -Atitier: Osborne: ornare tier selec tse ecole eT oh Rec even ected terete econ tote 1001 Map showing distribution of Archidiskodon imperator west of the Mississippi River. After Hay....................., 1003 Archidiskodon imperator, referred cranium of young male from Texas. American Museum Cope Collection. ........... 1004 Archidiskodon imperator, referred mandible, in comparison with mandibles of Parelephas jeffersonii.........-.....55-. 1006 Mandibles of Elephas indicus, Loxodonta africana, Parelephas washingtonii, and Archidiskodon hayi..... 2... ...0.5055. 1006 Archidiskodon imperator, right superior tusk of record size, from Post, Texas, combined with a superb tusk formerly in the
National Museum of Mexico, now destroyed. After photograph......................-... see esse eee e eee eee 1007 Archidiskodon imperator, referred young male(?) skull from tar pools of Rancho La Brea, California. ................. 1007 Archiduskodonmauniperators revered skull strony Vale hora) yp WeKAS eyes eye eyepe eye tees aes een vey ete Ne eset eet tse ret ee etter 1008 Archidiskodon imperator, referred skull from Victoria, Texas, as mounted in the American Museum................... 1009 Archidiskodon imperator, referred jaw from Tule Cafion, Texas. ...-. 0,-0.0. 5 66 ete ee ee oe eee 1010 Archidiskodon imperator, crushed skull of aged male (Nebraska Museum) from Hay Springs, Nebraska. ............... 1011 Archidiskodon imperator, crushed skull of adult male in the American Museum, from Hay Springs, Nebraska.......... 1011 Archidiskodon imperator, referred third right superior molar, from Zumpango, Mexico. After photograph... .......... 1013 Archidiskodon imperator, referred mature male cranium, from Tepexpan, Mexico. After photograph. ................. 1014 Archidiskodon hayi (2), referred mandible in Geological Institute of City of Mexico. After photograph............... 1014 Archidiskodon imperator silvestris, type third left superior molar. After Freudenberg. .......-..-..-+.-- 25-2020 s200- 1015 Archidiskodon imperator falconeri, cotype jaw after Freudenberg. Originally figured by Villada...................... 1016 Archidiskodon imperator, referred. Aged male. Reconstruction.............-.---+2+ ese ee eee eee 1017 Archidiskodon imperator, referred right forelimb, as mounted in the American Museum. ... 0.2... 60.00.0000 0 0200 sees 1018 Archidiskodon imperator (young adult) and Loxrodonta africana oxryotis (““Jumbo’’), referred limb bones... ...........-. 1018 Archidiskodon imperator. After restoration by Osborn and Knight, 1908... 2... 2.6.0... 0. eee eee eee 1019 Archidiskodon imperator maibeni, forelimbs of type skeleton, also cranium and tusks of Parelephas jeffersoni........... 1020 Archidiskodon imperator maibeni, mounted type skeleton in Morrill Hall, University of Nebraska..................... 1021 Shoulder heights of living and extinct elephants (Elephas indicus, Loxodonta africana oxyotis, Archidiskodon imperator, A.
WIVARULEP TOL ANCH) oto ou o gbadhe oopen eo Ud dou deocn ceo noadeecr UaDo Ga cApdd son UBOSS oO monn pCoDmnEdoucaGDocouC 1022 ANadhaitetontia an innoaignn LNvuee ieee ggoooedscocaseosauaeosooece sudanoeacooosec0aD DD CRod IoD COS anor 1024 Archidiskodon planifrons, primitive mandibles from the Siwaliks, India, Chagny-Bellecroix and Senéze, France. After
PEATE aS Ray ae aV eieera Ste Ne cb itl ca MRR coo on Cae on ena aan bokeas Daeg Nato o> 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<re.. sich, ant ateccie Pichu ute e, tie eet ae at cole te ge aan a Mey UP ROMO eh uecotna cua Rog Mey cera: 1166
Mammonteus primigenius, referred skull and half-grown tusks of male specimen found on the Yukon River, Alaska........ 1166
FIGURE
1033. 1034. 1035. 1036. 1037. 1038. 1039. 1040.
1041.
1042. 1043. 1044. 1045. 1046. 1047. 1048. 1049.
1050.
1051. 1052. 1053.
1054.
1055. 1056. 1057. 1058. 1059. 1060. 1061. 1062. 1063. 1064. 1065.
1066. 1067. 1068.
1069.
1070. 1071. 1072. 1073. 1074. 1075. 1076. 1077. 1078. 1079.
LIST OF ILLUSTRATIONS Xxiil
PAGE Circilarguis srOfhenvoollyaVianmnothiotsibenianneeen eee eae eee eee eee eee nee eee 1166 Wroollyeviammo thy soname saver wean cen sets =. his ce ei es gh a alee An oy ne ees GE | 1166 Simataomuneseelkarma Cavey Moravia eAtter photographs. 4444.0 0quateeasness ss acces oeeye loeeee cee see fon 1168 MEmTIGIN OE OF WEKO mes, Mion, Laie ja eVooyae ON, posaneudsbaaveaduacecodovescddaugcoadsancasecousausass- 1168 Giantekallimg stone of the Moravianjhunters, After photogtaphh..eese)sssso lee eeesnecee assesses senses snes. 1168 Ivory figurine of a woman’s head, from Brassempouy. After Pilloy in Piette’s ‘“L’Art pendant l’Age du Renne”.......... 1168 Equine ivory statuette from Lourdes. After Pilloy in Piette’s “L’Art pendant Age du Renne”...................... 1169 Loxodonta africana, young adult bull of the Lake Paradise region, east central Africa. Photograph by Mr. and Mrs. Martin Johnson, and shown in film “Simba.” Courtesy of Mr. Daniel E. Pomeroy...................0000---eeeeeeeee 1170 Crania of Loxodonta africana, Palxoloxodon namadicus, and Hesperoloxodon antiquus. After Falconer and Cautley, Pilgrim, Wrelthorer manda olli gy. Saga steer he... tewere te POAT oe ae A ht AOE ue ae he a sh aot 8 alse ape a ee al 1172 Cuvier’s figures and definitions of Hlephas primigenius (Messerschmidt’s cranium), H. indicus, and EF. africanus........ 1173 Loxodonta africana and Elephas indicus, third inferior molars. After Owen..............0..0000cc eee ceeceeeeeeees 1175 Loxodonta africana and Elephas indicus, second superior molars. After Lydekker...........0..... 0.00000 ce eee eee 1175 onoaontaafrmcancaonyous(« Jumbo:2), eranitim and jaws... <2506 shane ee dee atend. See eee A 1176 Palzoloxodon namadicus, type cranium, from the Nerbudda. After Faleoner and Cautley......................--... 1176 Palzxoloxodon and Hesperoloxodon as depicted by the cave men of North Africa and Spain. After Pomel and after Breuil... 1184 Map showing geographic distribution of the principal species of the Loxodontine...........................--..--. 1186 Habitat of the African elephant (Loxedonta), forest and savanna of the Uasin Gishu Plateau, Kenya Colony. Calf of old fe- malercharsinexelephant. eAtiterphotopraphibys CarlihwAkeleys meee oe ee ees Gene deen eee 1189 Habitat of the African elephant (Loxodonta). Same region as previous figure. Females and young bulls in forest. After PhotographubyeWermilt NOOSE Vel bee iresstiucsayske Rue es APR Io a ey SecA moe CIOs care Re Cn 1189 Small herd of African elephants (Loxodonta). After film photograph by Martin Johnson............................ 1189 Alkelevaorouploreaimcantelephantsant thesAmenican! Museum 44 see eer eee eae detec aie ee eee reine 1190 “Khartum”’ (Loxodonta africana oxyotis), formerly living in the New York Zoological Park. Two growth stages. After jo) AYO YAO Ye) 0) 01 re ah pe eae ait AEE rina ise ae arts ee Re ae EL ood fon, oe ene Aeon Pe RR te ence I eats ola ae 1194 Remarkable cave paintings of white rhinoceros and African elephant, discovered in South Africa. Courtesy of London Hibs trreu Ge cING WS he dejesucscoen, ox best oe Sev as Saco cpecd lay shone ek be Mens Jester cree eee | EP eh ARID Peer Ay gr OS Re 1194 Mapishowinerdistnbution) of existing Atricanvelephant) (orodonta)masa ose oes aoe ae ae ee ae eee ae eee 1195 Loxodonta africana oxyotis and L. africana pumilio. Restorations by Flinsch.....................0..00 cee eee eee 1196 Loxodonta africana, Blumenbach’s original figure of type, a right second inferior molar, described as Hlephas africanus..... . 1197 Hlephasvmniscus) |= Loxodontarajmeana|. “hype molaratter Goldiussse-yanee eee eee ee coe eee eee nee eee 1197 Loxodonta africana peeli, referred skull and tusks of adult male, from Mt. Kenya. After photograph................. 1198 Loxodonta africana oxyotis (‘“Jumbo’’), referred middle-aged skull (24 years)............. 000. cece cece eee eee 1199 Loxodonta africana oxyotis (Jumbo’’), superior and palatal views of cranium. Age twenty-four years................... 1200 Comparison of tusks of Loawodonta africana oxyotis and Mammonteus primigenius. After photographs................. 1201 Loxodonta africana albertensis and L. africana peeli, full-grown heads (male and female).....................-..-.... 1202 Lozxodonta africana (?)cottonz, foetal cranium, jaw, and milk dentition. After Eales.............................00.. 1203 Tusks of African elephant, believed to be the heaviest in the world, shown in front of a typical Arab door at Zanzibar. One tusk now in British Museum. After Kunz, courtesy of executors of his estate.................00. 00020 ee cee ee 1204 Horodoniaconnaliaes typemightsuperiormmolary eAttersAradasemere eter areas eee nee nee ee 1205 Young Addobush elephant (Loxodonta) from Cape Colony. After photograph..................... 0.000 eee e eee eee 1205 Hesperoloxodon of Europe, Palxoloxodon of India and of the Mediterranean Islands, compared with drawing of Hesper- olonodonm byscave men or northern Spain’) Restorations by Mlinschle eee eee e rere leer erent iene 1206 Comparison of crania of Palxoloxodon namadicus, Hesperoloxodon antiquus italicus, H. antiquus ausonius, and H. antiquus platyrhynchus. After Faleoner and Cautley, Pilgrim, Weithofer, Graells, and Pohlig............................ 1208 Palzoloxodon namadicus, type female(?) cranium. After Falconer and Cautley..........................---0-e-- 1211 Channellof the Godavarinear Nandur) Madméshwar Indias After eilorimm js -eee eee aie eet einen 1213 Comparison of Hesperoloxodon and Palzxoloxodon (syn. Sivalikia) superior and inferior molars........................ 1214 Three progressive broadening stages in the Palzoloxodon (syn. Sivalikia) and Hesperoloxodon superior grinding teeth. ... 1215 Hesperoloxodon antiquus (Upnor elephant). Restoration by Wlnsch..2.-....05.2-.....2-2-5.--2- 2s ses eee eee 1216 Hesperoloxodon antiquus, lectotype left second inferior molar. After Falconer and Cautley.......................... 1218 Hesperoloxodon antiquus, referred left second inferior molar from Gray’s Thurrock. After Falconer and Cautley........ 1219 Hesperoloxodon antiquus, referred third right superior molar. After Falconer and Cautley........................... 1220 Hesperoloxodon antiquus (Upnor elephant). Referred second superior and inferior molars. After photographs.......... 1220 Hesperoloxodon antiquus of Upnor. Mounted skeleton in British Museum. After photographs....................... 1223
XXiV
FIGURE
1080. 1081.
1082. 1083. 1084.
1085. 1086.
1087. 1088. 1089. 1090.
1091.
1092. 1093. 1094.
1095. 1096. 1097. 1098. 1099. 1100. 1101. 1102. 1103.
1104. 1105. 1106. 1107. 1108.
1109. 1110. 1111. 1112. 1113.
1114. 1115. 1116. 1117.
1118. 1119. 1120. 1121.
1122. 1123. 1124.
OSBORN: THE PROBOSCIDEA
Hesperoloxodon antiquus of Upnor, skeleton in the British Museum. Redrawn to show original and restored parts......... Scapule of Hesperoloxodon antiquus of Upnor, Loxodonta africana (“Jumbo”’) of the Sudan, and Hlephas indicus. After
INNS eevee TOG 01010) 015) teach cage cae Gente can aca a eackoel Nini 0 hich 01d. 0.9,0 6 orn oman Go ditoo om cocoonentg adn aou mS Dorsolumbar vertebre of Hesperoloxodon antiquus of Upnor and Elephas indicus... 2.2... - 0. ee cee eee
Vertebral columns of Hesperoloxodon antiquus, Loxodonta africana oxyotis, Elephas indicus, Mammonteus primigenius, and
ITA NMEA ile 6 paso 00cm eBons nog odgucUmooos sco DSDGyboooUHEGa HOU RDCOOUACDC LD can ugOoMOnapoo oC Hesperolorodon antiquus nanus, type, possibly a left second superior molar. After Acconci.....................-.-5. Hesperoloxodon antiquus platyrhynchus, type premaxillary rostrum with tusk and portion of right superior maxillary with
SECONGHMNOLALIUN SHU PATUCTGTELIS® acizche octet trols Tet teree esteem ee Pen en eee ne ested eee ea oy oie Reema eR Cees ele Hesperolorodon antiquus ausonius, type third right and left inferior molars. After Depéret and! Mayet: la es anes Hesperoloxodon antiquus, H. antiquus germanicus, H. antiquus italicus, progressive stages in evolution of grinding teeth... .. Hesperolorodon antiquus germanicus, type. Portion of right second inferior molar. After Stefiimescu.................. Hesperoloxodon antiquus germanicus, right tusk in Field Museum of Natural History, from Steinheim, male and female
tusks in Gotha Museum, from Tonna, right male tusk in Stuttgart Museum, from Steinheim..................-. Hesperoloxodon antiquus germanicus, right tusk excavated at Steinheim. Diagrammatic sketch reproduced through courtesy
ai DY BKaae I CG Sis ny ace cen cicens apes ohms ba Coeds Can Rn DOA TAD OOGE Doin CBGOmC Momma SOD too ec Lanoe 90.8004: Hesperoloxodon antiquus italicus. Restoration by Flinsch... ......... 0.0.65. 0 cee ee eee eee eee ees Lozodonta africana albertensis. Restoration by Flinsch.......... 22.20... 6 eee eee eee te eee eee Map of valley of the Liri, Italy, showing the location where Hesperoloxodon antiquus italicus was found, and other exposures.
Miter be sorenzoiand MJB asia = sc crn ey rwn ctes Cvencatts Dae c= PD ch cit iar lege epee en cee oie Pekar ese Mammalian fossils associated with type cranium of Hesperoloxodon antiquus italicus. After Osborn...........-....-. Hesperoloxodon antiquus italicus of Pignataro Interamna, type cranium before; removal! ae eae hee eee Map showing location of Pignataro Interamna, region of valley of the Liri. After Century Atlas Q138 2 ee ae emer Hesperoloxodon antiquus italicus, type cranium. After De Lorenzo’s original sketch and measurements................ Hesperolorodon antiquus italicus, type cranium. After De Lorenzo and D’Erasmo.......-.-.- 2-20.06 esses seen eee. Hesperoloxodon antiquus italicus, front view of cranium. After De Lorenzo and D’Hrasmoe ayes ees ones eae Hesperolorodon antiquus italicus, type superior grinders. After photograph. ............ 02... 2050s e sees e eee eee Hesperoloxodon antiquus italicus, type right second and third superior and inferior grinders. After Osborn............. Hesperoloxodon antiquus italicus, type mandible, right lateral view with second and third superior teeth superposed on cor-
responding inferior teeth, and superior view. After Osborn. ......... 0.22.00 605 e eee eee ee eee eee ees Hesperolorodon antiquus italicus, type third right inferior molar. After Osborn... ... 0.0... - 0.050 s serene cere eens Hesperoloxodon antiquus italicus, type cranium, three aspects... .. 1... 0-65-50 e eee ee teen eee eee Hesperoloxodon antiquus italicus, type cranium as reconstructed and mounted in the American Museum. After Osborn... . Hesperoloxodon antiquus italicus type and Loxodonta africana peeli ref. crania. After Osborn® she soar cette piace a Comparative bathycephaly of the Loxodontine: Loxodonta africana, Palzoloxodon namadicus, and Hesperoloxodon
ann iidicok, INuoe Oki, oy acedas secaobonduboo COs dU eroGuopODGuavOn Usb cooAD Too 9Ud peaedaOs ase Gods Comparison of scapula of Hesperoloxodon antiquus italicus, Loxodonta africana, and Elephas UNAUCUS Nereida estes Palxoloxodon namadicus, referred cranium, of the Godavari Alluvium at Nandtir Madméshwar, India. After Pilgrim. .. Hesperolorodon antiquus italicus, referred femur, with Dr. Pohlig standing beside it. Miter photog rap hine ise eee ln Intracranial brain casts of Hesperoloxodon antiquus italicus, Loxodonta africana, and Elephas indicus. After Osborn... .... W. Bauer Quarry at Steinheim on the Murr, showing the site of the 1928 discovery of the cranium of Hesperoloxodon
Hesperoloxodon antiquus germanicus, referred third right superior molar from Steinheim: space cree eect er aterm here Elephas antiquitatis Kriiger [= Hesperoloxodon antiquus germanicus ref.], type molar. After Breislak:. erge cress wet Bathymetric map of the Mediterranean Islands. By permission of Longmans, Green and Company, from map edited by
@bshiolnand ee te cacy. es occ ace os sees Bier oars needed chou.s co tebe aap ote setter on cnet sel see beeen cd Meera enerrWarspeyen cchaters.cnteG-batet eternal Dwarfed elephants of the Mediterranean Islands. Diagrammatic representation. .........- 55-20-50 0s scenes eee eens Dwarfed elephants of the Mediterranean Islands. Restorations by Flinsch. .............0 0-000 -s00e sees seen ees Loxodonta africana pumilio, young “pygmy” elephant passing beneath adult Elephas indicus, female. After photograph. . . Palzoloxodon mnaidriensis, referred, the ‘“Elephas (antiquus) Melite”’ of Pohlig. Fully adult cranium from the Grotta di
te Chianti salhy, wevep lillie oo ee cccosadu sondpovagd Oued gaccuugouOdc aD LOO obo BDA DOH ua ace no Ue Palzoloxodon mnaidriensis, referred juvenile cranium, the “Hlephas (antiquus) Melite” of Pohlig. After Rohligneeentr a Palxoloxodon melitensis, type third left superior molar. After Falconer... ........-. 000s esse eect eee etter eee Palzoloxodon melitensis, referred mandible from the Grotta di Pontale, Sicily... .. 0... 6.5.5.2 ee eee eee eee eee ee
FIGURE 1125. 1126. 1127. 1128. 1129. 1130. 1131.
1132.
1133.
1134. 1135. 1136. 1137. 1138. 1139. 1140. 1141. 1142. 1143. 1144. 1145. 1146. 1147. 1148. 1149. 1150. 1151. 1152.
1153. 1154. 1155. 1156. 1157. 1158.
1159. 1160. 1161. 1162. 1163. 1164. 1165. 1166. 1167. 1168.
1169.
LIST OF ILLUSTRATIONS XXV
PAGE Palzoloxodon mnaidriensis, third right inferior molar. After Leith Adams.....................0 0 eevee eeeeeeeeeeee. 1264 Palzoloxodon mnaidriensis, type and paratype molars. After Leith Adams....................00 000000 eee eee eee. 1264 Denizens of ancient Malta. Restoration by Leith Adams as dwarfed African elephants...........................-- 1265 ACL EOLOLOMONECY DRLOLES i COLY Pe nOlarsamAtvera Bb aAlcerya er er een + Ae ee elie tere ier eae eee 1266 Palkaglonoilap: GHEE, CONTSD WON, ING BENG 5 506 na dabagnseo556 540056 seegus sdeouesaboossaaeoGugaeeonase sot 1267 Sectionsois Grottorok Muparellopealermoysicilys) Atiten Vauireys). 4. 44). 58 see ose Jee oe aoe Jeelaeees Saee enee 1268 Tusks in the three dwarfed species of the Mediterranean Islands: Palzoloxedon falconeri, P. melitensis, P. mnaidriensis. ANTES VENOMS oe Se Pater cote: CES rae ict mcEA er oe Aan nae year aan Poe ee a ae ee are OEE, Te Pa Oe 1271 Two types of molars belonging to Hlephas [= Palzolorodon] mnaidriensis according to Vaufrey, namely, ‘type endioganal’ and ‘type pachyganal,’ from Shantiun and Puntali. After Vaufrey...................0-00-0eeeeeeeeeceecesss. 1271 Molars referred by Vaufrey to Elephas [= Palzoloxodon] melitensis, from Luparello, Sicily, and Benghisa, Malta. After WERT 0.4 che Bh ne eh eRe Choe rae LEE ar etry ih ert Tae a een de OO eae Tree geeery ree | Ae 1272 Ulne of Palxoloxodon mnaidriensis, P. melitensis, and P. falconeri. After Vaufrey................20000000-0 cee eee 1272 Palzxoloxodon atlanticus, cotype right second inferior molar. After Pomel.................00. 0000 cece eee cece sees 1274 Palzoloxodon atlanticus, referred third left superior molar. After Pomel.......................0000000eeee eee eee eee 1274 iPaleolozodenwolensis uy pe lett third interionmolar Atter homeless. asses eee oe ee eases eee eae Ae. 1275 Palzoloxodon recki, lectotype left second inferior molar. After Dietrich....................0.00eceececeeceesseeeee 1276 Palxoloxodon(?) andrewsi (Dart’s type of Archidiskodon andrewsi). Left third inferior molar, restored. After Osborn.... 1278 Palzoloxodon hanekomz, type ?third right superior molar. After Dart...-.................00-+..---++00--:22-eess 1279 ale oloxodonMorkemuypeiehte,ubirdsnten ormolaneALter» anieeeeeneree eee teen sien ase eee ee eee eee 1280 ELcomxodonmnilmaniaty pes (inirdinieniormolars AtteL Dante eee eee en eee Le ee eee eee eee eee 1281 OLeolOxOCOTmKILAN? atv per ower leitnmolaraeAtterD ania aeneene eee ene aoe eee ae eee 1281 Palzoloxodon archidiskodontoides, type ?second superior molar and part of right humerus. After Haughton............ 1282 Palzoloxodon transvaalensis, type right third superior molar. Modified after Dart’s photographs..................... 1284 Palzoloxodon sheppardi, type left third superior molar. Modified after Dart’s photographs.......................... 1284 Koxodontavuluesty pewhirdlettinteriommolaryyAtter scottes sone ee eee ee oe een ease eee 1286 Loxodonta zulu, referred third left inferior molar, from Kaiso Bone-beds, near Lake Albert, Africa. After photograph... ... 1287 Goxodonianprimamsy Peaunird lefty imienorimolaneeATters Dante eee eee eee eee ee eee eee eee 1287 Loxodonta africana var. obliqua, type third right inferior molar. After Dart...................0.0. cee cece eee ee eee 1288 Loxodonta subantiqua, type ‘possibly a right lower molar, probably the second.” After Haughton................... 1288 Palzxoloxodon namadicus naumanni, type, of Makiyama, in comparison with Hesperolorodon antiquus germanicus ref. of oblige Diagrammatic outline sketchy. :2ccs0i= sete aeeeace eee eee 1294 Palxolorodon namadicus namadi, type right third superior molar. After Makiyama................................ 1296 Palzoloxodon protomammonteus, type left third inferior molar. After Matsumoto.......................-...-..02.. 1297 Palxoloxodon protomammonteus proximus, type fragment of left third inferior molar. After Matsumoto............... 1298 Palzxoloxodon namadicus yabei, type right ramus of mandible, containing third molar. After Matsumoto.............. 1299 Palxoloxodon (Archidiskodon?) tokunagai mut. junior, type right second inferior molar. After Matsumoto............ 1300 Parelephas protomammonteus matsumotoi, type. Portion of left mandibular ramus with third molar zn sztu. After Saheki. INotidetermined: bythe: presenti atithor arc scars ee Ae PSD 1300 Palxoloxodon yokohamanus, type second right superior molar. After Tokunaga.......................0.00020-0005- 1301 IPalzoloxodomnhysuarinaieus, cotype molars;) Attern photographseen eer eae eee eee ene eee 1302 Map showing Japan as part of the Asiatic continent in Plio-Pleistocene time. After Longmans’ New School Atlas...... 1304 Map showing Japan as part of the Asiatic continent in Plio-Pleistocene time. After Yabe.......................... 1305 Referred Elephas indicus. Male and female Ceylon elephants. After photograph by Plate Ltd...................... 1306 First definition of the genus Elephas, bracketed with ?Rhinoceros. After first edition of Linneus’ ‘Systema Nature,” | Oe) | | Pe a ee errs ne Aki oe Pr nmneerLs rice ty oma DE eC O Gn ad MOORE aG om td ad 1aa0%9 OLE 1309 Facsimile of portion of page 11 of Linnzus’ Memoir of the Museum Adolphi Friderici Regis, Stockholm, 1754, in which first appears, theyspecies: name: Hlephas wndicus:., cio = odo ase ane de cm roe He ee oe coe eee e cae Sele see ee eee 1309 Facsimile of page 33 of Linnzus’ original tenth edition of the “Systema Nature,” 1758, in which Elephas maximus is sub- shitubedstorvMlephas endieusss... rule SG ate Pe ae re eee ee el Sc ee ea Pe 1310 Indian elephant group in the American Museum of Natural History. Specimens from the hills in the Province of Mysore, shot! 923 bynMir: Arthur Si: Verm ay sc scot sh ees a he custo eos oor Pade eke eet os a sae oti nce caval ee ee 1311 Elephas indicus sumatranus, pair of young elephants from Sumatra captive (1921) in the Zoological Park of Washington. Aftenphotogtaphy. «x.y. Secreto oe eta ath A SR es eu cailn LPR ira etry aI = Se PS 1314 The Sumatran elephant, apparently a female, living in the Amsterdam Zoological Gardens, August, 1913. After photo-
(35) 0) | eae Nea ee eee ee SAL cars hued Mi Oa OAR ME eG Clb nice ocIORb 4 cas S 5 Giyolow-cic kid < ohn c 1314
XXV1
FIGURE 1170. 1171. 1172. 1173.
1174. 1175. 1176.
MAUZ Pe 1178. 17/9: 1180. 1181. 1182. 1183. 1184. 1185. 1186. 1187.
1188.
1189.
1190.
1191. 1192.
1195. 1194. 1195. 1196. WIE 1198. 1199. 1200. 1201.
1202.
1203. 1204.
1205. 1206.
1207. 1208. 1209. 1210.
1211. 1212. 1213.
OSBORN: THE PROBOSCIDEA
PAGH Elephas indicus ceylanicus and E. indicus bengalensis, erania. Types. After de Blainville...............4......00.5. 1316 Crania of Elephas indicus (Dauntela var.) and E. indicus (Mukna var.). After Faleoner and Cautley................ 1317 Elephas indicus bengalensis, type, and E. indicus ceylanicus, referred, crania. After photographs. .................... 1317 Map showing geographic distribution of the principal species and subspecies (living and extnct) of Hlephas, Hypselephas,
Enrol Etat tlio) 1k eee pies ato Gon tte reer non en aoaut oA Non co Oso demo Gosia ponU OG 66 utd 1318 Platelephas platycephalus, Hypselephas hysudricus, and Elephas indicus. Restorations of heads by Flinsch............. 1320 Elephas indicus bengalensis. A herd of wild elephants in a bamboo jungle of Mysore. After photographie see eecseeeels 1322 Elephas indicus, referred third inferior molar (vertical section) of an unusually large specimen from Assam. After Falconer
Gravel (CONN ca oa Moot stare aes Parmer o NEOs oan enon Ma U ULC OOO MEO co con O boom nbd rdonon Su ao Ba 1324 Elephas asiaticus. Blumenbach’s original type figure of a first right superior molar. .... 0.0.0.0. .6 60. c cece eee 1325 Elephas indicus ref. Section of a partly worn third inferior molar. After Gaudry............ 0.0.0... eee eee eee 1325 Elephas indicus bengalensis, known as the “giant tusker of Udiapur,” showing abnormal length of tusks. After photographs. 1326 Comparison of crania of Elephas indicus ceylanicus and EF. indicus bengalensis. 2... 0.0.66 eee 1327 Elephas indicus sumatranus formerly living in the Rotterdam Zoological Gardens. After Lydekker................... 1329 Elephas sumatranus, cotype male and female crania from Palembang, in Leiden Museum. After photographs.......... 1330 Elephas sumatranus crania (adult and infantile), in Munich Museum. After photographs... ...................405. 1330 Sumatran elephant from Batang Serangan. Mounted specimen in Munich Museum. After photograph.............. 1331 Sumatran elephant (infantile) in Munich Museum. Mounted specimen. After photograph. ....................505. 1331 MountedsBurmeseelephants, -Atiter:photoprapliecerr acbseeis seein eer ae ietel otalekate oles ecco ier eke tee ete alee eset 1332 Elephas indicus hirsutus, type, formerly living in the Gardens of the Zoological Society, London. Mounted specimen in the
Britishy MiuseunayONeturaleen stony, sa eAtberyly.cl eke Taal retriever tere totter ete eee 1333 Elephas indicus buski Matsumoto [=?Palzxoloxodon buski], type first superior molar of the left side, from Japan. After
IVER [Ub TOL Oe See ee ee Cent ees Retreat ire. tre Sn hone Minne canny aS ana De athe aieG SAAS 1333 Second right superior molar from between Kanagawa and Tokio (Yedo), Japan, referred to Palzoloxedon by the present
authors After Iiy deleker icy eharce ss levees desma clasts oe yeu satay: Bina ene ea ee ee eon 1334 Superior view of heads of a young African elephant and of an adult Indian elephant. After Geoffroy St.-Hilaire and
Hired Gri ci@uvieErs 2. shaves. el ee che dhs aerate eae eth aoe ee Go ei ie ee a etre: irene 1335 Loxodonta africana and Elephas indicus, crown views of third right inferior molars. After Owen..................65. 1335 Comparison of low-browed African cranium (Lorodonta africana) and high-browed Indian cranium (Hlephas indicus),
showinexdeeplyzembeddediraim belowacramialvaircellsiyereyaey si tret-t- te ter tele tenet te else te ete epee eee eee ate 1335 Unguligradism. Radiograph of right foot of a young Indian elephant...............-- 0.0 e eee eee eee eee ees 1336 Hstimated shoulder heights of Indian elephant, skeletal and flesh.............. 22.0. .05 20. cee as eee cere eens 1337 “Elephas planifrons” and ‘‘Elephas hysudricus” life zones. After Pilgrim............ 0... 0:0. eee cece nee e eee eee eee 1338 Map showing Upper Siwalik exposures of the Simla foothills, India. ..............6. eee e eee e eee eee es 1339 Hypselephas hysudricus, type and paratype molars. Sections after Falconer and Cautley...............0...0 000.005. 1341 Hypselephas hysudricus, paratype third right inferior molar. After Falconer and Cautley...................50.00555 1342 Hypselephas hysudricus, referred third left inferior molar. American Museum (Barnum Brown) collection............. 1342 Hypselephas hysudricus, referred second left inferior molar...........--. 0... s sees eee ee eee eee ees 1343 Hypselephas hysudricus, referred third left superior and inferior molar sections. American Museum (Barnum Brown)
GOLlEC HONE eras ci asec ites Slane AE ek Aull Corie cei boots DSW pee Anis comnts ced Ae eos Si 1344 Hypselephas hysudricus, veferred third right superior molar, inner view, also transverse section, and photograph of occlusal
gies, JNmrertteanal Ihren Asian 1B5x0),\aa)) COME, oo oc ocongecconenoensoconnonpansaneseoencuddo0Gs00GE 1345
Hypselephas hysudricus, referred superior and inferior molars from India. American Museum (Barnum Brown) collection. . 1346 Comparison of two crania of Hypselephas hysudricus with Mukna and Dauntela varieties of Elephas indicus. After Faleoner
rintol (Chim sini saan nol beret Onno aa tn cane center inka Clr tan coe bicte to Puna no Ganuano Enso eect adc. uri ce 4 OMe 1349 Hypselephas hysudricus, referred adult male cranium. After Falconer and Cautley.... 2.0.6... 5000. e eevee eee 1350 Hypselephas hysudricus, portion of young jaw with greatly elongated rostrum. American Museum (Barnum Brown)
seyaNA Pci toy aves oe Rg a eae EE ee eae ee PPh eee Menem er ae ie tanlehen aia eoen, volar sath of SUH Adios aigec bane 1351 Platelephas platycephalus, type cranium (palatal view). American Museum (Barnum Brown) collection. ...........--. 1352 Archidiskodon planifrons, referred adult cranium of supposed female, with small tusks. After Falconer and Cautley..... .. 1352 Hypselephas hysudricus, male cranium (palatal view). After Falconer and Cautley..........6 6.000000 1352 Hypselephas hysudricus, referred female cranium in Amherst Museum, collected near Kullu, a district of the Punjab, by
IMI OER Ey cojaie ony ee eto. Rianne Lee AL ea amen Doe iced Ge Moi ea ernie cient castor rica nr cas n.or6 Gecuole.4 onc tarts 1353 Hypselephas hysudricus, restored juvenile skull. American Museum (Barnum Brown) collection. After photograph....... 1354 Hypselephas hysudricus, referred cranium. After Falconer and Cautley.............06 60 ceee eee eee eee ees 1354
Hypselephas hysudricus, referred juvenile crania in British and American Museums. ... 0... 6.6.00. 0 0 essere es 1355
LIST OF ILLUSTRATIONS XXVil
FIGURE PAGE 1214. Hypselephas hysudricus, referred juvenile cranium and jaws. American Museum (Barnum Brown) collection... . . 1356 1215. Hypselephas hysudricus, referred right second superior molar, originally selected by Osborn as the type of Hlephas plates e-
phalus angustidens. American Museum (Barnum Brow Collectionvga ersten te Ar ro ee nye ee ea ee 1357 1216. Upper Pliocene and Lower Pleistocene strata near Siswan, India, showing site where type cranium of Platelephas platyce-
LOCUS WS TOUUGl, Avie ol OUC NN e4 cog sdewe oD ono eddy ue cbdn aco uenoop ee orb er uobmeanan: Se ree ee 1358 1217. Comparison of the types of Platelephas platycephalus and Stegodon pinjorensis. Cranial sections......... eee ove _ 1360 1218. Platelephas platycephalus and Hypselephas hysudricus, right cranial profiles............................ Tey hee: . 1360 1219. Platelephas platycephalus, four aspects of type cranium. American Museum (Barnum Brown) collection. ........... 1361 1220. Geological relationships of African Proboscidea. Columnar section by Colbert....................... Bae tari aie aL 1221. Geological relationships of Oriental Proboscidea. Columnar section by Colbert..........-........ Anos eran ee) 1222. Geological relationships of European Proboscidea. Columnar section by Colbert.............0..00..00 0000 ee 1457 1223. Geological relationships of Asiatic Proboscidea. Columnar section by Colbert. ..............0.....002220000000. sy MET 1224. Geological relationships of North American Proboscidea. Columnar section by Colbert. ................... en Lao 1225. Geological relationships of South American Proboscidea. Columnar section by Colbert......................... o-» L516 1226. Models of Recent and extinct Mammoths and Mastodons. After Knight......................00.00ee see. ee lo22 1227. Map showing geographic distribution of the Mceritherioidea, Deinotherioidea, and Mastodontoidea............. 1528 1228. Map showing geographic distribution of the Stegodontoidea and Elephantoidea (including the Stegolophodontine if the
TNs Levers Pana 3 0 CEE) oy cae een —= ee ac RRP CRORE ICP eR A ne Pe Pe es aM teed RNR ge od 1538
1229. Four-coned ancestral grinders of the Proboscidea (e.g., Marither‘um) compared with the six-coned Paleomastodon molars... 1544
1230. Molar diagrams showing typical crown pattern (median sulcus, median conules, double trefoils, serrate spurs) in each of the LOUrprAniiliesrotetherViastodomtoideat |. snv-s.cah a «Walt cssoeepsucsrenceeeay cert Seve Pate ae tassel yea ey A rier 1546
1231. Accelerated evolution of ridge-plates from Archidiskodon planifrons into A. imperator; also Stegodon granger? molar, mail enamel foldings, inserted to show V-shaped valleys of the stegodontoid molar as compared with U-shaped valleys of the
Gleyaowriaortel iio) Bik: eA ancnde a cent oat Rees tet Pen aaa On oncud at oashrsd dame canoe Saopes oe Te Ea Oa 1232S Brevirostrns-s broversion ol rdee-crests in-Amancusm...--- + oases eee eee ee aeons ea errata 1548 1233. Humboldtine: Retroversion and centroversion of superior ridge-crests in Cuvieronius and Stegomastodon............ _ 1548 1234. Crown view of third inferior molars of the right side of Loxodonta africana and Elephas indicus. After Owen... ... 1549 1235. Archidiskodon subplanifrons, type third right inferior molar. Section showing cement, dentine, and enamel. Drewine By
DE Pelbevicttmb ra dle yc reste oticirs hots A alhcciels betne Son cheer oa teade et ete) Te ees esas aoc e ere erence oe ee eee 1549 1236. Chief head and dental forms of four of the superfamilies (I IV) of the Proboscidea (Mceritherioidea, Deinother ‘ides a, Masto-
dontordearmbileplnan toes’) rae sacs ave asi exeqsssrevec ares etch ay one the ete eee cca em eet Pome ae ea aaa eer ce 150) 1237. Divergent adaptive radiation of crania and incisive tusks in six bunomastodont subfamilies............... poh eee a5 5T 1238. Deinotherium giganteum, juvenile jaws showing replacement molars. After Lartet........................2..5.. 1554 1239. Geneplasmic evolution of the archaic-toothed mammoths during a three-million-year period so far as known to April, 1935.
Ps NTGTOCK OFS] 000 1 Ne er en RE Ue Res obit. on iciciats: cen etna} Sita bu Soncrotgen.o OS -a.a 4% agutrens Soe eevee . 1581 1240. Alloiometrons: Naaetine sieeily and weight proportions. wAtiterl sb Oloir iret teeter eee eee . 1581 1241. General climatic distribution of the subfamilies of the Elephantoidea and Stegodontoidea including theoretic simieion
lines (1938). After American Geographical Society North Polar Projection...................-........+-.--:-- 1589 1242. Worldwide distribution of the Proboscidea in past and present time. Same as figure 6 of Volume I, with modifications... . . 1594 1243. Foot trail of Indian elephant ‘‘Gunda,”’ formerly living in the New York Zoological Park, taken in sand.............. 1598
1244. Elephas indicus ref., showing contrast in proportions between adult and young. After photograph by Underwood and \ Bf aves 7(070ys Dee er ene eateries Olin mache 6 cob mi ome ecm Molo O olas.o See © bane 599
Hi bere om, if oe Ay’! nado
CHAPTER XIV
THE ROOF-TOOTHED STEGODONTS, SUPERFAMILY STEGODONTOIDEA
PossIBLE ANCESTRY IN THE MIOCENE ZYGOLOPHODONTS OF WESTERN EUROPE. PRIMITIVE FOREST-BROWSING ELEPHANTS OF THE ORIENTAL REGION. SLOWLY PROGRESSIVE FROM MIDDLE MIOCENE TO MIDDLE [UPPER]
PLEISTOCENE TIME.
MIDDLE [UPPER] PLEISTOCENE TIME.
I. INTRODUCTION.
Is 2. 3.
4. 5.
Ge
History of classification. Habits and general characters. Approximate descending ridge formule, after Falconer,
Lydekker, Martin, and Osborn.
Geologic and diphyletic order of the Stegodontine. History of discovery of the subfamily Stegodontine.
Principles of type revision of the species.
The Stegodontinz and Mastodontine of China. Pliocene to Pleistocene Proboscidea of Japan. Phylogenetic discussion of the thirty described species
of Stegodonts and Stegolophodonts. Probable African-European-Asiatic origin and migra- tion of the primitive Stegodonts.
Il. Type RevIsION OF THE SPECIES IN ORDER OF ORIGINAL
DiscoVERY AND DESCRIPTION.
1. The first two Stegodonts, discovered in Burma, 1828.
Mastodon latidens Clift. Mastodon elephantoides Clift.
2. Discoveries in India and Burma (1845, 1846).
Third species, Hlephas insignis, India. Fourth species, Elephas ganesa, India. Fifth species, Hlephas bombifrons, India. Sixth species, Hlephas cliftvi, Burma.
3. The Stegodonts of China, India, Java, the Philippine
Islands, Austria, Japan, and Burma.
Seventh species, Stegodon sinensis, China.
Kighth species, Stegodon orientalis, China.
Ninth species, Mastodon cautleyi of Perim Island.
Tenth species, Stegodon trigonocephalus of Java.
Eleventh species, Stegodon mindanensis, Philippine Islands.
Twelfth species, Stegodon airdwana of Java.
Thirteenth species, Stegodon ganesa var. javanicus of Trinil, Java.
Fourteenth species, Mastodon stegodontoides of Lehri, India.
Fifteenth species, Hlephas (Prostegodon, Parastegodon) aurore of Japan.
Sixteenth species, Mastodon (Bunolophodon) longi- rostre Kaup forma sublatidens of Austria.
Seventeenth species, Stegodon orientalis shodoénsis of Japan.
Kighteenth species, Stegolophodon nathotensis, India.
Nineteenth species, Stegolophodon cautleyi progres- sus, India.
Twentieth China.
Twenty-first species, Stegodon insignis birmanicus, Burma.
Twenty-second species, Stegodon pinjorensis, India.
species, Stegodon orventalis grangert,
BRACHYODONT TO SUBHYPSODONT, BUT PROGRESSIVE RIDGE FORMULA.
Ill.
IV.
805
EXTINCTION IN
Twenty-third species, Stegodon bondolensis, Java.
Twenty-fourth species, Stegodon trigonocephalus prae- cursor, Java.
Twenty-fifth species, Parastegodon? kwantoensis, Japan.
Twenty-sixth species, Stegodon yiishensis, China.
Twenty-seventh species, Stegodon officinalis, China.
Twenty-eighth species, Stegodon zdanskyi, China.
Twenty-ninth species, Parastegodon [Stegodon?| sugi- yamai, Japan.
Thirtieth species, Stegolophodon lydekkeri, Borneo.
SysTEMATIC ARRANGEMENT OF THE STEGOLOPHODONTS AND
STEGODONTS IN PHYLOGENETIC ORDER.
1. Characters of the subfamily Stegodontine. 2. History of the generic names assigned to the Stegolopho-
donts and to the Stegodonts. Generic characters of Stegolophodon Schlesinger. Systematic description of species of Stegolophodon. Stegolophodon cautleyi of the Upper Miocene [| Middle Pliocene], Perim Island.
Stegolophodon latidens of the Lower Pliocene {?Lower Pleistocene] of Burma and [Middle Pliocene] of India.
Stegolophodon sublatidens of the Middle(?) Pliocene of Austria.
Stegolophodon stegodontoides of the Upper(?) Plio- cene of India.
Stegolophodon nathotensis of India.
Stegolophodon cautleyi progressus of India.
Stegolophodon lydekkeri of Borneo.
SUCCESSION OF SPECIES OF THE GENUS STEGODON. Genus Stegodon Falconer and Cautley, 1847, 1857. Skulls of Stegodonts in the British and Indian Museums. Characters of referred skulls of Indian Stegodonts,
after Falconer, 1868.
Stegodont crania of China and of the East Indies. Systematic description of species of Stegodon.
Stegodon sinensis of the Yangtze River, China. Stegodon elephantoides of Burma. Stegodon cliftii of the Irrawaddy River, Burma, and of the Dhok Pathan horizon, India. Falconer’s Notes of 1868 on Stegodon cliftii. Stegodon bombifrons of the Lower [Middle] Pliocene, Dhok Pathan horizon. Falconer’s Notes of 1868 on Hlephas | = Stegodon| bombifrons. Lydekker’s Notes of 1886 on Elephas [= Stegodon| bombifrons. Stegodon insignis of the Upper Pliocene [to Upper Pleistocene] of India. Stegodon ganesa of the Upper Pliocene [to Upper Pleistocene] of India.
806 OSBORN: THE PROBOSCIDEA
Faleoner’s Notes of 1868 on Stegodon insignis Middle Pleistocene, Java (synonym of S. and S. ganesa. ardwana or S. trigonocephalus). Lydekker’s (1886) comparison of Stegodon in- Steqgodon (Archidiskodon?) mindanensis of Minda- signis, S. ganesa, and S. bombifrons. nao, Philippine Islands. Stegodon insignis-ganesa a collective species. Stegodon aurore of Mt. Tomuro, Japan. Stegodon insignis birmanicus of Burma. Stegodon orientalis shédoénsis of Japan. Stegodon orientalis grangert of China. Cranial characters of S. orientalis grangeri. V. ReceNTLY DESCRIBED STEGODONTS FROM JAVA, CHINA, AND Stegodon pinjorensis of India. JAPAN. Stegodon orientalis of Szechuan, northwest China. Stegodon bondolensis of Java. Lydekker’s Notes of 1886 on Stegodon orientalis. Stegodon trigonocephalus praecursor of Java. Stegodon airdwana of Trinil, Pithecanthropus erectus Parastegodon? kwantoensis of Japan. zone, Java. Stegodon yiishensis of China. Stegodon airdwana fauna, Kendeng-Schichten Stegodon officinalis of China. layer of Trinil, Java. Stegodon zdanskyi of China. Comparison of Stegodon airdwana of Java with Parastegodon [Stegodon?] sugiyamai of Japan. Stegodon insignis-ganesa of India. Stegodon trigonocephalus of the vicinity of Surakarta, Appenpix: Matsumoto on the phylogeny and classification Java. of the Japanese Mastodonts, Stegodonts, and Elephants Stegodon ganesa var. javanicus of the Trinil horizon, (1924-1927).
I. INTRODUCTION 1. HISTORY OF CLASSIFICATION [The name Stegodontoidea first appeared in a diagram by Professor Osborn in his article of June, 1935, entitled, ““The Ancestral Tree of the Proboscidea. Discovery, Evolution, Migration and Extinction over a 50,000,000 Year Period”? (Osborn, 1935.937, p. 407, fig. 2), in which superfamily he included both Stegolophodon and Stegodon under the family Stegodontide of Young-Hopwood, thus removing them from the Elephantoidea.
In Volume I of the present Memoir (published Aug. 15, 1936) Professor Osborn confirmed his separation of the Stegodontoidea from the Elephantoidea (pp. 22, 25) but he withdrew the Stegolophodonts, placing them in the superfamily Mastodontoidea, family Mastodontide, and creating a new subfamily, the Stegolophodontinz (see pp. 700, 737, and PI. rv), to embrace the various species, owing to the “intermediate position of [the] molars between the true Mastodontide ... and the true Stegodontoidea,”’ and suggesting (p. 191) the “possible derivation of the grinding teeth of the Stegodontoidea from those of Stegolophodon.”” The true Stegodonts he retained in
the superfamily Stegodontoidea, subfamily Stegodontine.
As early as 1857 Falconer observed (1857.1, p. 314) that “The 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.’’ Later Falconer (in Murchison, Pal. Mem., 1868, Vol. II, footnote, p. 268) remarked that ‘The Indian fossil species, which have been ranged under the designation of Stegodon, establish, through their molar teeth, a manifest and nearly unbroken passage from the Mastodons into the true Elephants.” Also, as recently as 1932, van der Maarel (1932.1, p. 162) expressed the opinion that “all the species of Stegolophodon .. . being all very primitive forms ... may as well be reckoned to the family of the Mastodontide.”’
Therefore, while various scientific observers have regarded certain of the Stegodonts as transitional between the Mastodontide and the Elephantide, it remained for Professor Osborn to assign the superfamily name Stego- dontoidea to the true Stegodonts and to remove the Stegolophodonts to the superfamily Mastodontoidea, under the new subfamily name Stegolophodontine, the members of which he designated (Vol. I, p. 690) as “pro-stego-
donts.”’
The regrettable death of Professor Osborn in November of 1935 precludes the full treatment of these groups as contemplated by him (see Vol. I, p. 197); it is deemed best, therefore, to allow the present chapter to remain as
THE STEGODONTIN:: HISTORY 807
first written by him, making such changes (either in square brackets or in editorial notes) as are consistent with the knewn opinions of the author, namely, as deduced from published statements, accumulated notes, and interlined text.
The classification of the genera Stegolophodon and Stegodon, now to be described, would appear, therefore, to be as follows:
SuPeRFAMILY: STEGODONTOIDEA Osborn, 1935, 1936 Separated by Osborn (Vol. I, 1936, pp. 22, 25, of the present Memoir) from the Elephantoidea Osborn, 1921, as a distinct stock, but without diagnosis. However, according to his observations on the sectioned molars, the valleys separating the adjacent ridges are closed or V-shaped at the bottom in the Stegodontoids (Fig. 764) and open or U-shaped in the Elephantoids (Fig. 1231). He also considered that the extremely short face of the Stegodontoids could not have given rise to the longer face of the Elephantoids.
FamILy: STEGODONTIDA Young-Hopwood, 1935
This name appears in Young (1935.1, p. 5) but without definition. Later in 1935, Hopwood, in his Memoir on the “Fossil Proboscidea from China,’ defines the family as follows (p. 71): ‘‘The animals included in this family have skulls which resemble those of the true elephants but which are more primitive. They have very long sockets for the tusks, and the grinding teeth and palate are well below the plane of the occipital condyles. The grinding teeth remain brachyodont throughout the whole of their evolutionary history, but they parallel the teeth of the true elephants in showing a progressive increase in the number of ridges. This is especially true of the third molars. With this increase in the number of ridges, and its accompanying increase of length, there is an ever increasing curvature of the occlusal surface which reaches its maximum in certain specimens referred to S. airawana and S. insignis. Owing to the short palate, it was impossible for the whole of the very long teeth to be accommodated in the upper jaw at one time. The curvature of the crown allowed the tooth to follow a more or less circular path which brought it from a position practically parallel to the plane of the occiput to the correct position for mastication. There is an ever increasing amount of cement, and the ridges show an increasing number of mammille on their crests.”
“This sub-family comprises two groups" of animals. One, with compressed, tectiform, ridges is given the generic name Stegodon Falconer & Cautley. The other, in which the ridges are blunter, and composed of round- ed conules, is known as Stegolophodon Schlesinger. Both genera occur in India, but, so far as is known, Stegodon is the only genus found in China.”
SUBFAMILY: STEGODONTIN Osborn, 1918, 1921
Original reference: Bull. Geol. Soc. Amer., 1918, Vol. XXIX, pp. 135, 136 (Osborn, 1918.468); Amer. Mus. Novitates, 1921, No. 1, pp. 12, 18 (Osborn, 1921.515).
SuBrAMILY Derinition (Osborn, 1918.468, p. 136): “The Stegodontine may be distinguished as a phylum confined to Asia, in which the grinding teeth remain brachyodont, short-crowned, although a very large number of cross crests evolve, especially on the posterior grinding teeth. From an early member of this subfamily, perhaps of Middle Miocene time, were given off one or more branches of the elephant and mammoth phyla.”’
(Osborn, 1921.515, pp. 12 and 13): ‘‘We observe that the Stegodonts are persistent browsers, probably tropical, forest-living proboscideans. According to Pohlig, from the skeleton discovered in
According to Osborn, 1936, Vol. I, p. 700, by removal of the Stegolophodonts to the family Mastodontide, the Stegodontide embrace the true Stego- donts (Stegodon Falconer and Cautley) only.—EKditor.]
808 OSBORN: THE PROBOSCIDEA
Trinil, Java, they have short, massive bodies like those of the Mastodontinz of the north temperate
forests. The skull and tusks do not lead into either the Elephantine or the Mammontine types . . . The
distinctive feature of the grinding teeth is the rapid multiplication of t ‘ansverse crests which rise from
the formula 4.5.6.6.7-8 in S. [Stegodon] cliftii (Lower Pliocene) to 5.9.10.12.13 in S. insignis (Lower Pleis-
tocene). Jaw rapidly abbreviated. Upper tusks straight, parallel, slightly upeurved (adapted to dense
forests). Grinders brachyodont to subhypsodont, crests breaking up into small mammille, valleys filling with cement.”’
In the first article, namely, ““A Long-jawed Mastodon Skeleton from South Dakota and Phylogeny of the Proboscidea,”’ 1918, Osborn placed the Stegodontine under the Elephantide (p. 135), and in an accompanying table mentioned the species Stegodon ganesa, S. cliftii, S. bombifrons, and S. latidens. ‘The last-mentioned species was made by Schlesinger the genotype of Stegolophodon.
The above definitions were based on both Stegolophodon and Stegodon. After the removal of the former genus to the superfamily Mastodontoidea, family Mastodontide, subfamily Stegolophodontinz, no revised definition of the subfamily Stegodontinz was given by the author.
LATE TERTIARY GEOLOGY OF INDIA The latest opinion regarding the geology of India (the Siwalik Hills and Perim Island) will be found in Edwin H. Colbert’s Memoir of 1935 on ‘‘Siwalik Mammals in The American Museum of Natural History,” pages 6 to 55; in a summary on page 21 it will be noted that Doctor Matthew in 1929 assigned to the Siwalik beds a somewhat higher position in the geologic time scale than Doctor Pilgrim in 1927. Compare also Chapter XXII of the present Memoir, “Geologic Succession of the Proboscidea,” which has been written by Doctor Colbert.
Throughout the present Volume, therefore, the later determinations will be inserted in square brackets or added in footnotes. It should be recalled that the chapters constituting this Volume were written about eight to ten years ago and were awaiting final revision by Professor Osborn.— Editor. |
2. HABITS AND GENERAL CHARACTERS
We observe that this subfamily [Stegodontinz’] includes forest-living browsers, which probably developed in a
Fig. 20. forested or semi-forested oriental region, ranging through India into Burma, China, Japan, and southward into Java, Borneo,” and the Philippines. The ancestors of the subfamily may be found in Miocene deposits of western Kurope®. A single tooth, named Mastodon (Bunolopho- don) longirostre Kaup forma sublatidens, has been de-
Elephas clifti.—The first (?) left upper true molar; from the Siwaliks of Burma. 4. The lower border of the figure is the inner border of the specimen.
(From. Gaudry’s ‘ Enchainements.’)
CoryPE OF STEGODON ELEPHANTOIDES Cure? (=cLirru FALCONER)
Fig. 683. Type of Elephas cliftii Falconer and Cautley, 1846, a first left REFERRED SUPERIOR MOLAR OF STEGODON ORIENTALIS GRANGERI upper true molar, 1.M!, one-half natural size. The same six crested tooth Pig. 684. Referred first right superior molar, r.M! (rev.) of Stegodon appears in figures 686, 700, and 701. After Lydekker, 1886.2, p. 81, fig. 20 orientalis grangeri (Amer. Mus. 18536—wrongly numbered 18580). Actual (taken from a woodcut in Gaudry, 1878, p. 176, fig. 232). median length 181 mm.
‘At the time this chapter was written the subfamily Stegodontinw was thought to embrace both the Stegolophodonts (now removed to the Mastodontoidea subfamily Stegolophodontine, p. 700) and the true Stegodonts.—Editor.]
*[Type locality of Stegolophodon lydekkeri Osborn, 1936 (see Vol. I, p. 700).—Editor.]
31See Vol. I, p. 197.—Editor.]
THE STEGODONTINA:: HISTORY 809
scribed by Schlesinger from near Teschen (Schlesien), Austria; this resembles Mastodon |= Stegolophodon| latidens of Burma and is referred in the present Memoir to Stegolophodon sublatidens (see Vol. I, p. 737; Vol. II, p. 846).
The skull is subelephantine in type, brachycephalic, brachyopic, the rostrum being elongated to support the tusks; the grinding teeth and palate are depressed far below the occipital condyles (bathycephalic). As in the elephants, the jaw is greatly abbreviated. The upper tusks are straight or shghtly upeurved, elongating, without trace of enamel band, to a length of about 10 ft. Considering the large size of the tusks the skull is relatively small. The grinding teeth are brachyodont to subhypsodont, yet the ridge-crests in the posterior molars, M8, multiply from five plus [Stegolophodon|] to fifteen plus [Stegodon], each crest breaking up into small nipples, mam- ille, or conelets. The lower incisors disappear very early.
As in certain of the Mastodontide and as in all the Elephantide, the grinding teeth increase the number of their ridges by adding crests both in front and behind, as first observed by Falconer. Thus in M 1 of Stegodon \s-6-7-¥
orientalis grangert (Fig. 684) the ridge formula may be written: *"“. The descending scale of the ridge formule in the premolars and molars of the principal species of Stegodonts is approximately as below.
3. APPROXIMATE DESCENDING RIDGE FORMULA, AFTER FALCONER, LYDEKKER, MARTIN, AND OSBORN
Observe that (1) the maximum upper ridge-crests, rising from 2le to 51e, are, so far as known, less numerous than the maximum lower ridge-crests which probably rise to more than 51; (2) this is compensated for by the fact that the upper molars are throughout broader than the lower molars; (8) the differences in width and in the number of ridge-crests of the wpper molars in comparison with the lower molars are beautifully shown in figure 687, also in figures 759 and 762, the type of Stegodon orientalis grangert. See also details of progressive ridge-crest formule, talons and half ridge-crests, under each species.
Maximum EstTiIMATED MAXIMUM CONELETS UPPER AND LOWER
PER RIDGE- RIDGE-CRESTS CREST Dp 2 Dr 3 Dp 4 M 1 M 2 M 3 Dp 3-M 3 Wey i Cah 2 Se 7-9 7-9 9- 2- Stegodon airdwana 2 = = Suat ean 13-20+ 5le Stegodon insignis il 2 5-6 7-14 716-8 _7ig-6-8 Yo-11-46 11 42 49 Stegodon ganesa_ 3 g ee GE EAE Oa. AMEE 2 . : : 6-9-4 5iin6 ¥-6-4 V-6-7-4 4-8-4 Me11-i6 Stegodon orientalis grangeri week ud a == ae ae -11 41 46 ° 5- 6 7 946 Stegodon bombifrons + a a SS = 11+ 34e ° 6M 6% cr Stegodon elephantoides SM ou 10 5-8 . Bradeies 6 61% 646 Stegodon elephantoides (=cliftii) 4 s si =a an 10-12 28e QY. o 614 ~ Stegolophodon stegodontoides oe 5-6 / ms payin 6-5 5 ~ ¢ Stegolophodon latidens 2 a Ane ee 4-5 23(?) - 6-3-6 aaa Stegolophodon cautleyi ae Bean a 4-5 2le
FormMuLaA: RipGe-crests oR Lopus (Dp 2—M 3).—The above ridge-formula table, assembled from several sources, iS approximate, first, because various observers differ in the method of counting the ridge-crests, second, because within each species the ascending mutations may lead up to the next higher progressive stage, as, for example, in the transition of Stegodon insignis-ganesa to S. airdwana of Trinil, Java.
810 OSBORN: THE PROBOSCIDEA
The individual number of ridge-crests from Dp 2 to M 3 is mainly assembled from a very careful collation of the ridge-crest formule given by Falconer (see below), supplemented by the observations of Lydekker, of Martin, and of Osborn.
ConELETS.—The conelets of Stegolophodon and of Stegodon arise chiefly by binary, rarely by ternary, fission of the primary cones. Starting with the original loph of the Palzomastodon stage, consisting of two cones, normal binary fission would produce:
2—4+ (NS. cautleyi)—8 (S. elephantoides)—20+ (S. airdwana), etc.
But the binary fission is not so regular as this. The newer anterior and posterior crests exhibit fewer conelets
than the older mid-crests, so that the highest number of conelets usually will be found in the third and fourth
erests.
OC LES
65 €7ests S77 Conwelecs cag STEGOLOPHODON STEGODONTOIDES
$+ conelets 3 STEGOLOPHODON LATIDENS
Zmz CAUTLEY!
5+crests 4-5 conelets 2
2-3 conules Zn?
2p crests 2.cones 1
Fig. 685. SrrucruraL Evoturion or tHE Cones, CoNELETS, AND RipGb-CRESTS IN THE STEGO- LOPHODON PHYLUM, IN COMPARISON WITH PALAXOMASTODON
(1) PaAtaomasropon, Primary Type. Four primary cones; ridge-crests, proto-, meta-, and rudimentary tritoloph. [See pp. 143 and 691 of Vol. I, for subfamily position of Palwomastodon.—Editor.]
(2-4) SrecoLopHopon Puytum. Binary fission of primary cones into four to five conelets, vestigial conules in S. cautleyi and S. latidens (protoconule =p.1., metaconule=m.1.); addition of trito-, tetarto-, penta-, and hexalophs. Gradual loss of median sulcus, presistent, however, in the first two anterior crests.
Stegolophodon latidens (3, left) is a third lower molar, r.Ms, inserted for comparison, and has seven plus (7'4) ridge-crests, four plus conelets, and median sulcus.
m 3°
/§ crests 20 or more Conelet 10 SISSUTECS
CR ee /4 Crests // conelets 9 Oo at A oO WO
/ /z2crests J or less conelets & STEGODON ORIENTALIS GRANGER!
ve a| 8 ie
2 —e eS rms 93 crests 11% conelets 7 STEGODON BOMBIFRONS
2mz Be Ace l.mgz Wo crests 5% &conelets 6 STEGODON ELEPHANTOIDES Zm- 72 = 85 crests 10+ conelets 5 STEGODON ELEPHANTOIDES @CLIFTII)
Fig. 686. SrrucruraL Evoution OF THE CoNnEs, CoNELETS, AND RIDGE-CRESTS IN THE STEGODON PuytuM, tN ASCENDING ORDER (5-10). (5-10) Srecopon Pxytum. Binary or ternary fission of the cones into conelets (5-20); addition of anterior and posterior ridge-crests (10-15 in M 8); addition of cement.
(5) Stegodon elephantoides (=cliftiz) with six and a quarter ridge-crests and ten plus conelets (M!). (6) Stegodon elephantoides with ten ridge-crests and five to eight conelets (Ms). (7) Stegodon bombifrons with nine and a half ridge-crests and eleven plus conelets (M®). (8) Stegodon orientalis grangeri with thirteen and a half ridge-crests and eleven or less conelets (M3). (9) Stegodon insignis-ganesa with fourteen ridge-crests and eleven conelets (M3).
(10) Stegodon arrawana with fifteen ridge-crests and twenty plus conelets (M°).
811
812 OSBORN: THE PROBOSCIDEA
Fiaures 685 ANp 686.—Particularly interesting and significant in the Stegodontine is the transformation of the original cones by fission into conelets. Thus in the archetypal Paleomastodon molar there are two cones in the protoloph, in Stegolophodon latidens and S. cautleyi each cone splits into two, making four plus conelets in the metaloph; in S. stegodontoides each of these four conelets tends to split into two, tending to form from five to eight conelets, but this splitting is not regular and no loph actually attains eight. In Stegodon elephantoides the equal splitting gives rise to from five to eaght conelets; in S. elephantordes (=cliftir), S. bombifrons, and S. insignis- ganesa the fission gives rise to from eleven to twelve conelets, hence each ridge is finally surmounted by twelve cone- lets which, when slightly worn, present eight loops. In Stegodon airdwana, the most progressive species, the conelets range from thirteen to twenty plus. Thus the maximum number of cones and conelets in each crest runs as follows:
Molar cones and conelets: Primitive (Palzomastodon) 2-4—6-8-12-20+ progressive (Stegodon airdwana).
Rince-crest Evotution.—The Stegodontine also furnish a beautiful example of the evolution through which each ridge passes in turn, from the primitive submastodontoid type seen in Palzomastodon into the highly progressive subelephantoid type seen in Stegodon insignis-ganesa and S. airdwana. The superior (-loph) and inferior (-lophid) ridge-crests may receive a brief numerical terminology, namely:
Pro-protoloph—id = One-half, anterior rudimentary ridge Pentaloph—id = Fifth ridge Post-metaloph—id = One-half, posterior rudimentary ridge Hexaloph—id = Sixth ridge Protoloph—id =First primary ridge=protocone and Heptaloph—id = Seventh ridge paracone of Ungulata Octaloph—id = Kighth ridge Metaloph—id = Second primary ridge = hypocone and Ennealoph—id = Ninth ridge metacone of Ungulata Decaloph—id = Tenth ridge Tritoloph—id = Third ridge Endecaloph—id =Eleventh ridge Tetartoloph—id = Fourth ridge Dodecaloph—id = Twelfth ridge
Crest Appition.—In Stegolophodon and in the Mastodontide the homology of the protoloph and of the metaloph is simple as compared with other ungulates, but since the increment of ridge-crests in the elephantine molar is by addition to both the anterior and posterior ridges, namely, the pro-protoloph and the post-metaloph, it soon becomes difficult to determine which ridge-crests correspond with the primary protoloph and metaloph of Palexomastodon and of other ungulates.
INTERMEDIATE Mouars Unrrorm.—A constant feature in the Proboscidea appears to be the uniformity of
the three ‘intermediate molars,’ namely, Dp 4, M 1, M 2, which tend to have the same ridge-crest formula in each species, for example:
Stegodon insignis-ganesa Dp 4 7% Mi 7% M 2 7%-8 =intermediate molars Stegolophodon latidens Dp 4 4 Mi 4% M2 4-5 =intermediate molars Trilophodon angustidens Dp 4 3 Mi1 3 M2 3 =intermediate molars
Consequently it may be difficult to distinguish these ‘intermediate molars’ from each other by the ridge formula alone; whereas they may be distinguished by the character of wear, by the width of the crowns, and by the condition of the fangs.
RipGe-crest ELevation.—The progressive elevation of the ridge-crests in the Stegodontine is illustrated in two diagrammatic figures (Figs. 687, 688), which demonstrate the constant progressive heightening of the ridge- crests as we ascend from the Lower(?) Pliocene Stegodon sinensis and [Middle Pliocene] S. bombifrons into the Middle Pleistocene S. airdwana stage. The early phases of ridge-crest elevation (Fig. 687) may be compared with the later phases (Figs. 688, 781) as follows:
Stegodon orientalis Owen Stegodon aurore Matsumoto Stegodon orientalis grangeri Osborn Stegodon airdwana Martin Stegodon bombifrons Falconer and Cautley Stegodon insignis Falconer and Cautley
Stegodon sinensis Owen
THE STEGODONTINA: HISTORY
813
DESCENDING ORDER OF SpEciIES.—(1) At the summit of the known Stegodontine is Stegodon airdwana, the most progressive both in ridge-crests and in conelets—a Lower to Middle Pleistocene stage. The following extract from a letter by Doctor Dietrich (March 10, 1924, and notes) confirms the Middle Pleistocene age of this species:
For a long time I have been trying to prove that the Stegodon |airdwana] species from the Pithecanthropus strata [Trinil, Java] are the very youngest, that is, young Pleistocene. In detail: ‘‘Geologisches Alter. Aus morphologischen Griinden muss die javanische Art jiinger sein als die chinesische und alle bekannten kontinentalen Stegodon-Arten. Pleistocin lebte, wird von fast allen Autoren angenommen; ihr plezstocdnes Alter diirfte gesichert sein. Erweist sich die chines- ische Stegodontenfauna als alt-pleistocin, dann ist die Trinilfauna jiinger als alt-pleistociin. Ich halte es aus geologischen und anderen Griinden sogar fiir wahrscheinlich, dass die Trinilfauna (und damit Pitheacanthropus) jung-pleistociin ist; das wird sich
mit Hilfe des St. Airawana bei besserer Kenntnis der kontinentalen Stegodonformen vielleicht erweisen lassen.”
4 STEGODON ORIENTALIS 1s LS eke
STEGODON ORIENTALIS GRANGER! A.M. 1/8705 Ys
STEGODON SINENSI!IS iy Type Ys Fig. 687. GrapuUAL PRroGRESsIvVE Hypso- DONTY IN SUPERIOR GRINDERS
Stegodon orientalis Owen, type. Superior deciduous premolar, r.Dp*.
Stegodon orientalis grangeri Osborn. Superior deciduous premolar, r.Dp*.
Stegodon bombifrons Falconer and Cautley. Third superior molar, M®.
Stegodon sinensis Owen, type. Superior deciduous premolar, r.Dp*.
All figures four-fifths natural size, excepting Stegodon bombifrons which is four-fifteenths natural size. Observe that the ridge-crests (1-5) in transition from S. sinensis type to S. orientalis type are broad, progressively elevated, and approximated.
tmner vzew All Ws nat, 325¢
727 F STEGODON AIRAWANA ref Cast. A.M. 6835
STEGODON INSIGNIS Type
Fig. 688. GRrapUAL PROGRESSIVE HyPpsopoNTy IN SUPERIOR AND INFERIOR GRINDERS
Stegodon airawana Martin. Middle Pleistocene of Java. Elevated and approximated ridge-crests (9-156), right third inferior molar, r.M3.
Stegodon aurore Matsumoto, type. Middle Pleistocene {?Upper Pliocene] of Japan. Elevated and closely approximated ridge-crests, right second superior molar, r.M?.
Stegodon insignis Fale. and Caut., type. Lower Pleistocene, Upper Siwaliks, Boulder Conglomerate, of India. Third superior molar, M°, with eleven +ridge-crests.
All figures to same scale, one-third natural size. Observe that in Stegodon atrawana and S. aurore the ridge-crests are much more elevated and approximated than in S. insignis, but that S. insignis is less elevated than S. orientalis (Vig. 687).
Dass St. Airawana im
TaB_e III
Upper PLEISTOCENE
MIpDLE PLEISTOCENE
LOWER PLEISTOCENE
Upper Pliocene
to Lower Pleistocene
Upprer PLIOCENE
MIpDLE PLIOCENE
Lower PLIOCENE
Mr0-PLIOCENE
STEGOLOPHODON PHYLUM
India, Burma
Stegolophodon stegodontoi- des type
\Stegolophodon | cautleyt | leetotype Stegolophodon | latidens ref. | [AUSTRIA ISteqalophndon | sublatidens type] Stegolophodon latidens lectotype and cotype! Stegolophodon latidens ref.!
Stegolophodon cautleyi progressus type
Stegolophodon nathotensis type
Japan
\Stegolophodon latidens ref.
STEGODON PHYLUM
Borneo | India, Burma | China, Japan Philippine Islands, Java |
Stegodon in- signis ref. |
Stegodon gan- |
| esa ref.
Stegodon ori- \Stegodon airdwana type entalis Stegodon ganesa var. shodoénsis javanicus type =S. airdwana or S,. type trigonocephalus Stegodon (Archidiskodon?) mindanensis type
Stegodon pin- Stegodon ori- _Stegodon trigonocephalus type
| jorensis entalis type
| type Stegodon ori-
| | entalis
| grangeri Stegodon in- type | signis lecto- type and ref.
Stegodon gan- esa lecto- | | type and ref. |
| |
Stegolophodon — Stegodon \Stegodon
lydekkert | insignis aurore type birmanicus | type
|| type!
| | Stegodon bom- | bifrons ref.
Stegodon bom-
| bifrons lec- totype
Stegodon bom- bifrons ref.
Stegodon bom- bifrons ref.
|
\Stegodon ele- phantoides lectotype'
|Stegodon ele- phantoides
| (=cliftii) || cotype'
| | | |
Stegodon sinensis
type
[See note on page 824 below, in which the geologic age of Stegolophodon latidens is given as Lower Pleistocene. As S. elephantoides and S. elephantoides (=cliftit) were found in the same locality and at the same geologic level, they should also be regarded as of Lower Pleistocene age. Note also that S. birmani- cus is placed in the Lower Pleistocene. Compare Colbert, Chap. XXII, pp. 1450, 1451.—Editor.]
814
THE STEGODONTIN: HISTORY 815
(2) Next in descending order are referred Stegodon insignis and S. ganesa, of Lower to Middle [Upper] Pleisto- cene age. (3) Third in descending order is Stegodon orientalis grangeri, slightly more primitive than S. insignis both in ridge-crest and conelet progression—probably of Lower Pleistocene age. (4) Next is the [?Upper Pliocene] Stegodon aurore of Mt. Tomuro, Japan. (5) Far more primitive both in ridge-crests and conelets are Stegodon bombifrons [of the Middle Pliocene] and S. elephantoides (= cliftii), known to be of Lower Pliocene (type) age.! It is probable that the referred S. bombifrons of the Middle [Upper] Pliocene, Tatrot horizon, will prove to be some- what more progressive than the lectotype of S. bombifrons of the Middle Pliocene, Dhok Pathan horizon.
(6) Representing the Stegolophodon phylum is the Upper(?) Pliocene Stegolophodon stegodontoides type, of approximately the same age as Stegodon ganesa lectotype and S. orientalis grangeri type. (7) Stegolophodon latidens ref. is recorded in the same Middle Pliocene (Dhok Pathan) geologic level as Stegodon bombifrons lecto- type. (8) The Upper Miocene [Middle Pliocene] yields Stegolophodon cautleyi lectotype of Perim Island (Dhok Pathan). (9) The Middle Miocene [Mio-Pliocene] yields Stegolophodon cautleyi progressus and S. nathotensis.
4. GEOLOGIC AND DIPHYLETIC ORDER OF THE STEGODONTIN
The geologic level of Falconer’s types of Hlephas |= Stegodon| ganesa and EF. |=S.] insignis, although unre- corded, is probably Pinjor (see Vol. I, Fig. 413). Pilgrim writes (letter of January 26, 1927): “I am now almost convinced that the bulk of the fossils were collected from that zone, i. e., Upper Pliocene [Pinjor zone, Moginand], but that some came from the Boulder Conglomerate zone Falconer distinctly states.’’ As observed by Barnum Brown, the exact geologic horizon of the type and referred specimens of S. insignis and S. ganesa is uncertain, because apparently in the Pinjor horizon are obtained jaws and skulls redeposited from the Boulder Conglomerate above. 5. HISTORY OF DISCOVERY OF THE SUBFAMILY STEGODONTIN. PRINCIPLES OF TYPE
REVISION OF THE SPECIES
GENERA.— Up to 1924 sixteen species of [Stegolophodonts and] Stegodonts had been discovered and described between the years 1828 and 1917, in the order shown in the list in Section II below. In 1924 Matsumoto described a new subspecies from Japan, namely, Stegodon orientalis shodoénsis. In 1929 Osborn described Stegodon orientalis grangert from the pits near Wanhsien, Szechuan, China, Stegodon insignis birmanicus from the Pliocene of Burma, Irrawaddy River, and Stegodon pinjorensis from the Lower Pleistocene (?Pinjor) of India; also two Stegolopho- donts (Stegolophodon nathotensis and S. cautleyi progressus). All the early species were described either as Mastodon (e.g., Mastodon latidens, Mastodon elephantoides) or as Elephas (e.g., Elephas insignis, Elephas ganesa). Although the name Stegodon appears as early as 1847 (‘Fauna Antiqua Sivalensis,’’ Faleoner and Cautley, PI. XLII), it was not until the year 1857 (pp. 314, 318, table opp. p. 319) that Falconer ventured to select Hlephas cliftui, E. bombifrons, E. ganesa(?), and FE. insignis as representing a new subgenus, Stegodon.
It was in 1922 that Matsumoto in a letter to the present author announced his intention of making Mastodon latidens the type of a new genus, Prostegodon. The name, however, is preoccupied by the Stegolophodon of Schle- singer, 1917 (genotype Mastodon latidens Clift), in which genus Osborn united (1929) the four species M. latidens Chft, M. cautleyi Lydekker, M. stegodontoides Pilgrim, and M. (Bunolophodon) longirostre Kaup forma sublatidens Schlesinger, also Stegolophodon nathotensis Osborn from the lower Middle Siwaliks and Stegolophodon cautleyi progressus Osborn from 2,000 feet above the base of the Lower Siwaliks, India.”
See footnote on preceding page (p. 814) regarding the Lower Pleistocene age of Stegodon elephantoides (=cliftii).—Editor.] *ITo these six species should be added Stegolophodon lydekkeri Osborn, described in Volume I of the present Memoir, page 700.—Editor.]
816 OSBORN: THE PROBOSCIDEA
|Recently six additional species of Stegodon have been described, namely, Stegodon bondolensis van der Maarel 1932, and S. trigonocephalus praecursor von Koenigswald, 1933, from Java, S. officinalis, S. zdanskyi Hopwood, 1935, and S. yiishensis Young, 1935, from China, also Parastegodon |Stegodon?| sugiyamai Tokunaga, 1935, from Japan. The generic determination of Tokunaga’s species Parastegodon? kwantoensis, 1934, has not been given in the present Memoir, owing to Professor Osborn’s views regarding the genus Parastegodon (see next paragraph).— Kditor.|
The genus Parastegodon of Matsumoto, 1924, belongs to the genus Archidiskodon or to a progressive Stegodon, as the genotypic species, Hlephas (Parastegodon) aurore, is slightly more primitive than Archidiskodon planifrons.
PRINCIPLES OF TyprE Revision.—The consideration of these matters of descent and phylogeny, however, must be preceded by a very rigid review of each species of Stegodont in the order of its original description, following the standard methods of type analysis established throughout this Memoir, namely:
1) Determination of the actual type specimen or the specimen first mentioned among a series of cotypes.
2) Fixation of the original type figure, the one first published by the author or selected by the author from other publications.
3) Enumeration of the type characters observed in the type specimen by the author or by subsequent observers.
4) Elimination from the type list of characters founded on referred specimens which do not actually belong to the same species as the type.
5) Determination of the type locality and distinction of the topotypes.
OriGInAL Descriptions.—The revision of these various species of Stegodonts, on the strict application of the five rules above, has been a long and very difficult task. In order to establish absolutely the original author’s intention, the author’s original description is cited in full. When too prolix, as in the case of many descriptions by Owen and by Lydekker, excerpts are made in the author’s own language.
This revision and establishment of type characters must be followed by a restudy and revision of referred speci- mens, which in most cases can only be done in future by monographie research directly upon the specimens them- selves, amplified by knowledge afforded by fresh materials. Consequently the present Stegodont chapter lays no claim to completeness or finality; it leaves many questions wholly undecided, for example, the sexual or specific dental or cranial distinctions between Stegodon insignis and S. ganesa.
In the meantime our present establishment of the types and of the type specific characters based upon the type specimens themselves as well as of characters derived from properly referred specimens in the same geologic horizon, as in the ease of Stegodon bombifrons, may lay a firm foundation for future monographic research.
6. THE STEGODONTINAZ AND MASTODONTIN OF CHINA Two species of Stegodonts described by Owen in 1870, namely, Stegodon orientalis and S. sinensis, were determined from collections brought in by dealers for medicinal purposes and not procured in situ by palzeontolo-
gists.
THE STEGODONTINA: HISTORY
0/4) —_ ba |
a) Koxen’s Notes oF 1885 AND SCHLOSSER’S Nores oF 1903 Koken, ‘‘Ueber fossile Siugethiere aus China,” 1885, pp. 31-44; Schlosser, ‘Die fossilen Sdugethiere Chinas,’”’ 1903, pp. 43-49 Koken’s specimens were also probably collected from the caverns of Yunnan by dealers for von Richthofen; his list is as follows (Koken, 1885, p. 33):
Proboscidia. ieee VCSLOCONEDEnUINENSTSUVAINISUILEILSTS MEE ee nea eeeeaen eee Yiinnan. eee VC stod onatiep le Qnasonismermagray- tere icericis Seine ae esses och enh Meee Yiinnan. “Bl SOCIO. UG L0 Be tactician caniotn bic Societe CORE Eee ern ee? Shanghai; oberer Hoangho (West-Kansu). AMM SECO OOOTIBLICSLO USER mera a tere een syst ree SAUL n eS Nore Ry eeteo eee Yiinnan; Szechuen. GS, SWOT CH WOT ROSS ait binw 6 Anais Ht HOES CE MEE Dee Re OE no nee Yiinnan.
Schon frither beschriebene oder erwihnte Arten sind durch *ausgezeichnet.
The type fragments all appear as though they had been brought in by native collectors, for medicine dealers, since they consist of broken teeth only; the same is true of the Schlosser collection. Consequently we regard the specific and generic determinations made from these imperfect materials as of provisional value. The species of Proboscidea contained in Schlosser’s article of 1903, following Koken’s review of 1885, are as follows:
REFERENCE IN PRESENT Memorr
Fokien, Kansu Stegodon orientalis Owen, 1870=Stegodon insignis Falconer (fide : Koken, 1885, fide Schlosser) Stegodon orientalis Shanghai Stegodon sinensis Owen, 1870= Stegodon clifti Falconer (fide Koken,
1885), = Elephas clifti (fide Lydekker, 1886) — Stegodon sinensis Stegodon bombifrons Falconer, 1846=Stegodon aff. bombifrons (fide Koken, 1885),=Hlephas bombifrons (fide
Lydekker, 1886) 2Stegodon bombifrons Rothe Thone Mastodon latidens Clift, 1828 = Mastodon aff. latidens (fide Schlosser) ?Stegolophodon latidens = Shansi Rothliche Sande = Mastodon Lydekkeri Schlosser, 1903, related to Mastodon latidens Tientsin, Honan, etc. (fide Schlosser, 1903), to Serridentinus (fide Osborn) ?Serridentinus lydekkeri Mastodon perimensis var. sinensis Koken, 1885= Tetralophodon (Ly- dekkeria) sinensis (fide Osborn) Tetralophodon (Lydekkeria) sinensis Rothe Thone Mastodon pandionis Falconer (fide Koken, 1885) = Incerte sedis (fide = Shansi Osborn) Incertzx sedis
b) AmMericAN Museum Discovery OF STEGODON ORIENTALIS GRANGERI NEAR THE YANGTZE River, CHINA The first scientific party to collect fossils 7n situ in China, with records of the actual locality and geologic level, was that of the American Museum, under Walter Granger, in 1920-1921.
In a preliminary notice of a collection secured during the winter of 1920-1921 by Dr. Granger of the Central Asiatic Expedition, Matthew and Granger (1923) described the material as occurring in a series of pits or fissures at the village of Yenchingkou in the vicinity of Wanhsien, province of Szechuan, about one hundred and forty miles distant in an air line from Chungkingfoo, the type locality of Stegodon orientalis Owen. This Yenchingkou material includes a fairly complete adult skull, two young skulls, a series of palates and lower jaws, and many teeth, which Matthew and Granger figured and partly described (pp. 567-571, figs. 8-6). They remark (p. 567): “Stegodon orientalis Owen. Schlosser regards this species as identical with S. insignis of India, basing the reference upon the fragmentary teeth described by Owen. Matsumoto regards it as distinct, upon the evidence of the referred material which he describes and figures. The Yen-ching-kao material includes a fairly complete adult skull, two young skulls, a series of palates and lower jaws and many teeth. It should enable us to estimate the affinities of the species more exactly when it has been cleaned up and studied.”’ They deferred further description of this fine material to Osborn (see below, pp. 875-881).
818 OSBORN: THE PROBOSCIDEA
GroLogic AGE OF STEGODON ORIENTALIS AND S. ORIENTALIS GRANGERI Typps.—Matthew and Granger (1923, pp. 563, 565) observe:
The Chinese fossil mammals described by Owen in 1870 [Footnote: ‘Quar. Journ. Geol. Soe., London, XXVI, pp. 417-436, Pls. xxv1I-XxIx.’] came from ‘a cave near the city of Chung-king-foo in the province of Sze-chuan.’ Chung-king is on the Yang-tse-kiang above Wan-hsien, about one hundred and forty miles distant [southwest from the Granger locality of Yenching-
kou] in an air line. . . . Possibly the Chinese informants of Consul Swinhoe, who sent the fossils to Owen, misled him, uninten- tionally or deliberately, as to the locality... . Owen regarded the [Chungkingfoo] fauna as Pliocene and described the follow-
ing species:
[Chungkingfoo] [Yenchingkou] Stegodon orientalis [type]. Parts of molars. [S. orientalis grangeri type Rhinoceros sinensis {type}. Parts of 4 upper and 4 lower molars. R. sinensis ref.
Tapirus sinensis |type]. Parts of 3 upper and 4 lower teeth. _T'apirus sinensis ref. Chalicotherium sinense [type]. Part of an upper molar. Chalicotherium sinense ref. Hyena sinensis [type]. Canine, 2 premolars. Hyena sinensis ref.|
Owen’s descriptions and figures accord very well with some of the species in our collection [the Yenchingkou collection of the American Museum], so that we have referred them to his species, whether or not later investigation proves them to be exact topotypes.
7. PLIOCENE TO PLEISTOCENE PROBOSCIDEA OF JAPAN
Discovery.—Matsumoto’s discoveries and descriptions of the Japanese Proboscidea up to 1924 are recited in more detail at the close of this chapter (pp. 901 to 909). His Elephas (Prostegodon, Parastegodon) aurorex, 1915, 1918, 1924, from a comparison with Stegodon airdwana of Java, appears to belong to Stegodon aurorex rather than to represent a distinct genus, 1. e., Parastegodon. Matsumoto (1918, pp. 51, 52) gives the history of discovery of Stegodon and Elephas in Japan, beginning with Leith Adams (1868), Naumann (1881), Brauns (1883), Lydekker (1886), Martin (1886 [1887]), Tokunaga (1906), Sato (1914), Kato (1914), Matsumoto (1915, 1918). The Japanese species of Stegodon referred by Naumann to S. cliftii and S. insignis have been transferred by Matsumoto to Owen’s Chinese species Stegodon sinensis and S. orientalis. Matsumoto remarks as to sexual characters (op. cit., 1918, p. 52):
Thus, the present writer’s opinion [Footnote: ‘This Vol., p. 10.’] that, St. sinensis, as well as St. orientalis, is geologically younger than St. cliftii, evidently holds true also in the Japanese specimens. One evidence noticeable is that the Stegodont species are usually found in couples. For example, St. cliftii and bombifrons are found associated with each other from the Dhok Pathan to the Tatrot horizon, St. ganesa and insignis from the Boulder Conglomerate horizon and the Lower Pleistocene of Narbada, Sf. orientalis and sinensis from the Uppermost Pliocene to Lower Pleistocene of China and Japan, and St. airdwana
and trigonocephalus from the Lower [to Middle] Pleistocene of Java. One may imagine the possibility, that each couple of species represent sexual dimorphism of one and the same species.
Matsumoto (1924, 1926, 1927) continues the history of discovery of the Stegodonts and Elephants of Japan up to the year 1927, as set forth in detail in this chapter, and presents his recent views as to the phylogeny of these animals as shown in figures 791, 792, and 793. To the fossil fauna of Japan he adds the following five re-
ferred and new species (op. cit., 1926.1, p. 1):
Genus Stegodon Valeoner & Cautley. 4. S. clifti Falconer & Cautley, ibid. [Journ. Geol. Soc. Tokyo, Vol. XX XI], 1924 [1924.3], p. 327. Akira-mura, Kage District, Province of Ise—possibly Plaisancian-Astian. 5. S. sinensis Owen, ibid., 1924, p. 328. Island of Shodo (Shédo-shima or Shozu-shima), Inland Sea—Milazzian-Tyrrhenian. 6. S. orientalis Owen, ibid., 1924, p. 330. (Also as S. bombifrons, ibid., p. 329.) Nagahama, Minato Town, Province of Kazusa; Togane Town, same province— Calabrian. Riuge, Ikadachi-mura, Province of Omi—Calabrian or possibly Cromerian. Okimisome, Ube Coal-Field, Province of Suw6—Cromerian. 7. S. orientalis shédoénsis, nov., ibid., p. 333. Islands of Mitsugo (Mitsugo-shima) and Island of Shédo, Inland Sea; off Nagasaki, Eastern Sea—Milazzian-Tyrrhenian. Kashiwazaki, Province of Echigo (?this form).
Of the above the genus Parastegodon Matsumoto, 1924, is regarded in the present Memoir as belonging to
the genus Archidiskodon or to a progressive Stegodon.
THE STEGODONTINA: HISTORY 819
8. PHYLOGENETIC DISCUSSION OF THE THIRTY DESCRIBED SPECIES OF STEGODONTS AND STEGOLOPHODONTS
DovusLE or MuuriepLe PHyLA OF THE STEGODONTS. SCHLOSSER, 1903.—Schlosser (1903, p. 191), in his revision of the Stegodonts of China, doubtfully suggested that west European species, originally described as Mastodon turicensis [= M. tapiroides, M. pyrenaicus, ete.], separated as Zygolophodon by Vacek in 1877 and (1926) by Osborn as Turicius (M. turicensis, M. tapiroides) and Zygolophodon (M. pyrenaicus), may have given rise to such Upper Miocene [Middle Pliocene] Stegodonts of India as Stegolophodon cautley?.
Schlosser was also the first to suggest (op. cit., p. 191) that Mastodon turicensis |= Turicius tapiroides| of the Lower Miocene of Europe may have given rise to the Mastodon [=Stegolophodon| latidens of the Lower Pliocene’ of Asia from which in turn sprang off the true Stegodonts, such as Stegodon insignis.
More in detail, Schlosser, who was the first to discuss the double phylogeny of the Stegodonts (op. cvt., p. 206), separated them into two divisions as follows:
Mastodon latidens Clift Stegodon ganesa Falconer and Cautley Mastodon cautleyi Lydekker Stegodon insignis Faleoner and Cautley
Stegodon cliftii Falconer and Cautley Stegodon bombifrons Falconer and Cautley
Two Puyta SuaaEstep BY Pinerim, 1913.—The second to discuss the phylogeny of the Stegodonts was Pilgrim. The foundations of a diphyletic arrangement of the Stegodonts were laid by his observation (1913,
Lower Miocene. Turicius (4,5) COMPARED WITH TRILOPHODON (2)
Vig. 689. Left third inferior molar, 1.M3. Comparison of Turicius turicensis [=tapiroides] (upper 4,5) with Trilophodon pontileviensis (lower 2), one-half natural size. After Mayet, 1908, Pl. x1, figs. 2, 4, and 5.
These molar teeth exhibit the profound difference between the zygolophodont molar (above), resembling that of a primitive Stegodont with uninterrupted, widely open valleys, and the bunolophodont molar (below), in which the valleys are closed by the central conules.
(Upper) Figs. 4, 5.—‘‘Mastodon turicensis [= T’. tapiroides). Pontlevoy. Derniére molaire inférieure. Paris Muséum. Grandeur naturelle.” Fig. 5.—‘‘Td. vue d’en haut.”’ See also figure 138C, Cl.
(Lower) Fig. 2.—‘‘Mastodon angustidens. Valun de Pont- levoy. Derniére molaire supérieure. Communiquée par M. Jean de Bodard, Pontlevoy. Grandeur naturelle.’ [= T'rilophodon pontileviensis. |
pp. 293, 294) that Mastodon [= Stegolophodon] latidens occurs in the same Lower [Middle] Pliocene beds with Stegodon bombifrons. Pilgrim’s discussion of the phylogeny of the Stegodonts may be paraphrased as follows:
(a) In the lower deposits of Perim Island there appears a species Mastodon cautleyi representing a line of evolution which in all its earlier stages is entirely unknown in Europe [see Turicius (?) and Zygolophodon (?)]. The lectotype of M. cautleyi is a last upper molar, figured by Lydekker in 1886 (1886.1).
See note on page 824 below where Stegolophodon latidens is given as of Lower Pleistocene age.—Editor.]
820 OSBORN: THE PROBOSCIDEA
(b) In the higher Perim levels, as well as in the Middle Siwaliks of Lower [Middle] Pliocene age, is found a further development of this type in the species Mastodon latidens, with an increased number of ridges. The larger form of MW. latidens is to be regarded as a direct descendant of Mastodon cautleyi. The last species of masto- dont to be referred to this line is represented by a tooth from Lehri, for which Pilgrim proposed the name Mastodon [Stegolophodon] stegodontoides.
(c) The type tooth of I. stegodontoides, of which the horizon is uncertain but may possibly be Upper Siwalik, is distinguished from that of M. latidens by the almost entire absence of accessory columns [conules of Osborn]; M. stegodontoides, like M. latidens, carries on each of its ridges four columns [=conelets of Osborn], while the anterior ridges of the next higher stage, Stegodon elephantoides (=cliftiz),’ carries nine or ten mamille [=conelets of Osborn].
(d) So close is M. stegodontoides to S. elephantoides (=cliftii) that it is hard to separate the two genera.
(e) The true Stegodon type represented by S. bombifrons appears in the Lower [Middle] Pliocene, Dhok Pathan horizon, before the extinction of the Mastodon [= Stegolophodon] cautleyi-latidens-stegodontoides phylum which is parallel with it.
Pilgrim accordingly divides the Stegodontine into two generic phyla: to the first he applies the name Masto- don |=Stegolophodon|, to the second he applies the name Stegodon. This may be graphically represented as follows:
Masropon Serius [=STeGoLorpHOpON] STEGODON Series [=SreGopon] Upper Pliocene Mastodon stegodontoides Stegodon bombifrons [Middle Pliocene] Lower Pliocene! Mastodon latidens Stegodon cliftii [= Stegodon elephantoides (=cliftiz), Upper Miocene [Middle Pliocene] Mastodon cautleyi Lower Pliocene]!
ScHLESINGER, 1917.—Schlesinger recognized the distinctness of Stegolophodon, basing his type on Mastodon latidens Clift.
Marsumoro, 1922.—The fact that Mastodon latidens appears in the same Lower [Middle] Pliocene geologic horizon as S. bombifrons is very significant; it tends to support the idea that the Stegodonts were diphyletic. This idea is perhaps carried a step further by Matsumoto who writes (letter, Nov. 20, 1922): “In my report just in preparation on the Japanese ‘Mastodonts,’ I follow you to refer ‘Mastodon’ latidens to the genus Stegodon creating however a subgenus Prostegodon for it. Prostegodon is the primitive representative of the Stegodon- phylum, representing half bunomastodontine and half stegodontine dental characters. Schlosser’s opinion, that Prostegodon might be ? a descendant of ‘Mastodon’ turicens, does not appear to be correct at all.”
OsBorN, 1923.—Osborn (1923.601, p. 2) erroneously adopted the generic name Prostegodon Matsumoto, based on the genotypic species Mastodon latidens Clift. Prostegodon, however, is preoccupied by Stegolophodon Schlesinger, 1917. Thus the diphyletic arrangement of the Asiatic and European species and genera would appear as follows:
Stegolophodon stegodontoides Stegodon ganesa
Stegolophodon cautleyi Stegodon insignis
Stegolophodon latidens Stegodon bombifrons
Turicius (?) sp. Stegodon elephantoides (= cliftiz)
Zygolophodon (?) sp. Marsumoto, 1924, 1926.—A more recent step is that of Matsumoto in a diagram embodied in a letter dated Sendai, November 20, 1924, in which the polyphyletic Stegodontinz are divided theoretically into five
'[See footnote on page 824 regarding the Lower Pleistocene age of Stegodon elephantoides ( =cliftii).—Bditor.}
THE STEGODONTINA:: HISTORY 821
phyla (Fig. 792), and Trilophodon [= Zygolophodon| pyrenaicus (Fig. 791) is placed as ancestral to Tetralophodon falconert, Parastegodon |= Stegolophodon| latidens, P. [=S.] stegodontoides, and Stegodon.
In other words: (1) The type of Mastodon |=Zygolophodon| pyrenaicus of the Middle Miocene or its an- cestors Zygolophodon pyrenaicus aurelianensis Osborn of the Lower Miocene of western Europe may be ancestral to the Stegolophodon and Stegodon species of southern Asia.
Fic. 230. — Derniére molaire inférieure du Mastodon luricensis (tapiroides), aux 2/0 de grandeur. — Miocéne moyen de Simorre, Gers. (D’aprés Lartet. )
Turicius (C) anp SteaoLtopHopon (A, B) Form or Grinpina TEETH CoMPARE WITH STEGODON Mo ars (Fic. 686)
Fig. 690. (C) Type third right inferior molar, r.M3, of Turicius turicensis simorrensis Osborn, 1926, erroneously determined by Lartet (1859, Pl. xv, fig. 3) as Mastodon tapiroides, two-fifths natural size. Upper Middle Miocene of Simorre. After Gaudry, 1878, p. 174, fig. 280. Reversed in drawing. See also Vol. I, pp. 207 and 220 of the present Memoir.
Cotype and lectotype of Mastodon [=Stegolophodon] cautleyi Lydekker, 1886.
A (Cotype). First superior molar of the left side, 1.M!, one-third natural size. After Falconer and Cautley, 1846 [1847, Pl. xt, figs. 3, 3a], as “Mastodon latidens.”’ Length 4 inches, width 2.3 inches. Brit. Mus. M.2817. Cast Amer. Mus. 26965. Perim Island. See Lydekker, 1886.1, p.xv, fig. 5.
B (Lectotype). Third superior true molar of the left side, 1.M%, one-third natural size. After Falconer and Cautley, 1846 [1847, Pl. xxx1, figs. 6, 6a] as “Mastodon latidens.”” Length 8.5 inches, width 4.5 inches. Brit. Mus. M.2705. Cast Amer. Mus. 26966. Perim Island. See also Lydekker, 1886.1, p. xv, fig. 6, and 1886.2, p. 73, fig. 18. Same as figures 141 and 142 of Volume I of the present Memoir.
(2) Matsumoto held (1924) that the Stegodonts are not merely diphyletic (e. g., Stegolophodon and Stegodon), but polyphyletic by subdivision of the species of Stegodon into five distinct lines of descent, as clearly displayed in figure 792.
OsBorn, 1927.—The present author (1927) takes the more conservative view that there are certainly two distinct phyla, namely: (a) Stegolophodon cautleyi, S. latidens, S. stegodontoides, and (b) Stegodon elephantoides (=cliftii) to S. airdwana. The latter appear to present a progressive series in the increasing number of conelets and ridge-crests or lophs, but without very marked divergence, as shown in the geologic succession table (Table III, p. 814).
822 OSBORN: THE PROBOSCIDEA
SuGGEsTED EvROPEAN-ASIATIC ORIGIN AND MIGRATION OF THE PRIMITIVE STEGODONTS
The suggestion by Schlosser (1903), rejected by Matsumoto (1922), that the species of Stegolophodon may be derived from mastodonts related to the Turicius or Zygolophodon of western Europe, should not be lightly dismissed. These west European animals have been grouped in the genus Zygolophodon by Vaeck (1877), based on the three species Mastodon borsoni, M. turicensis, and M. tapiroides (=M. pyrenaicus). The genus Zygolo- phodon embraces a type molar tooth fundamentally homologous and analogous, as shown in the accompanying comparison of the lectotype of Mastodon |= Stegolophodon| cautleyi from the Middle Pliocene of Perim Island, with a third inferior molar erroneously referred by Lartet (1859) to M. tapiroides |=type of Turicius turicensis simorrensis Osborn, 1926—Fig. 690] from the Middle Miocene of Simorre. Molars of the Turicius turicensis | =tapiroides| type occur in the Lower Miocene, Burdigalian, of the Falun de Pontlevoy, contemporaneous with the referred Trilophodon angustidens |= T. pontileviensis—Fig. 689, 2].
Actual relationship to primitive Stegodonts [Stegolophodontine] of Asia is represented by the Mastodon (Bunolophodon) longirostre Kaup forma sublatidens of Schlesinger from the Pliocene of Teschen (Schlesien), Austria (Fig. 722). A striking analogy to the Stegodon type is seen in the molar teeth referred by von Meyer to M. {|Turicius| turicensis from the lignites of Elgg and Kapfnach, and finally in the strictly stegodont Mastodon |T.| virgatidens of von Meyer (Fig. 168). The only way to test this theory, however, is to place models or casts of the teeth of Zygolophodon, of Turicius, and of Stegolophodon side by side to see whether they compare in close detail, in which case the genus Zygolophodon Vacek would replace the genus Stegolophodon Schlesinger.
ConcLuSION: PROBABLE AFRICAN-HUROPEAN-ASIATIC ORIGIN AND MIGRATION OF THE PRIMITIVE STEGODONTS Osborn, 1927': If, as now appears probable, (1) the Trilophodon phylum first arrived in southern Europe and migrated eastward into India, (2) it is also probable that certain primitive species of the forest-living Zygolopho- don or Turicius phyla gave rise in Lower Miocene time to forest-living animals which spread into the forests of southern Asia and developed into the Stegodon series, as first adumbrated by Schlosser (1903).
The European species actually resembling these animals is the Mastodon (Bunolophodon) longirostre Kaup forma sublatidens Schlesinger from Schlesien, Austria (Fig. 722), as described below.
II. TYPE REVISION OF THE SPECIES IN ORDER OF ORIGINAL DISCOVERY AND DESCRIPTION In the previous Section I of the present Chapter XIV we have discussed the habits and general characters, the ridge formule, the geologic order, the history of discovery, the principles of type revision of the species, the Stegodonts of China, Japan, and the East Indies, the phylogenetic succession, and, finally, the probable origin of the Stegodontine [and Stegolophodontine] in western Europe, and hence more remotely in Africa.
We now pass in Section II to the very complicated subject of the type revision of the species on the principles enumerated above (p. 816). For this purpose we will review the thirty species described in the years between 1828 and 1936 in the order of their description, quoting extensively from the original type descriptions and re-
producing every available type or lectotype figure directly after the original author.
"Compare Vol. I, pp. 195, 197, also Pls. 1 to 1v.—Editor.]
THE STEGODONTINA: HISTORY 823
EGODONTINA
LEE
Fig. 691. Geographic distribution of the principal species of Stegolophodon and Stegodon. The white dots within the black areas represent the approximate localities where the types of these thirty species were discovered. Numbers 1,9, 14, 16, 18, 19 and 30 are Stegolophodonts (see also Fig. 1228). The white crosses represent referred specimens.
STEGODONTS [AND STEGOLOPHODONTS] IN ORDER OF THEIR DISCOVERY AND DESCRIPTION See Figure 691 Speciric REFERENCE
OrIGINAL NAME IN PrEseENT Memoir 1. Burma 1828 Mastodon latidens Clift, Irrawaddy River, Burma = Stegolophodon latidens 2. Burma 1828 Mastodon elephantoides Clift, Irrawaddy River, Burma = Stegodon elephantoides 3. India 1846 [1845] Elephas insignis Falconer and Cautley, Siwalik Hills, India = Stegodon insignis-ganesa 4, India 1846 [1845] Elephas ganesa Falconer and Cautley, Siwalik Hills, India = Stegodon insignis-ganesa 5. India 1846 Elephas bombifrons Falconer and Cautley, Siwalik Hills, India = Stegodon bombifrons 6. Burma 1846 Elephas cliftii Faleoner and Cautley, Irrawaddy River, Burma = Stegodon elephantoides (=cliftiz) 7. China 1870 Stegodon sinensis Owen, vicinity of Shanghai, China = Stegodon sinensis 8. China 1870 Stegodon orientalis Owen, Chungkingfoo, province of Szechuan, China = Stegodon orientalis 9. India 1886 Mastodon cautleyi Lydekker, Perim Island, India = Stegolophodon cautleyi 10. Java 1887 Stegodon trigonocephalus Martin, ?vicinity of Surakarta, Java = Stegodon trigonocephalus 11. Philippines 1890 Stegodon mindanensis Naumann, Mindanao, Philippine Islands = Stegodon (Archidiskodon?) mindanensis 12. Java 1890 Stegodon Airdwana Martin, Alas-Tuwa, Java = Stegodon airdwana 13. Java 1908 Stegodon Ganesa var. javanicus Dubois, Trinil, Java = Stegodon airdwana [or S. trigonocephalus] 14, India 1913 Mastodon stegodontoides Pilgrim, Lehri, Punjab, India = Stegolophodon stegodontoides 15. Japan 1915, 1918, 1924 Hlephas (Prostegodon, Parastegodon) aurore Matsu- moto, Mt. Tomuro, Kaga, Japan = Stegodon aurore 16. Austria 1917 Mastodon (Bunolophodon) longirostre Kaup forma sublatidens Schlesinger, Teschen (Schlesien), Austria = Stegolophodon sublatidens 17. Japan 1924 Stegodon orientalis shodoénsis Matsumoto, Island of Mitsugo (Mitsugo-shima) and Island of Shédo, Inland Sea;_ off Nagasaki, Eastern Sea, Japan = Stegodon orientalis shodoénsis 18. India 1929 Stegolophodon nathotensis Osborn, near Nathot, India = Stegolophodon nathotensis 19. India 1929 Stegolophodon cautleyi progressus Osborn, near Chinji Bungalow, India =Stegolophodon cautleyt progressus 20. China 1929 Stegodon orientalis grangeri Osborn, Yenchingkou, China = Stegodon orientalis grangert 21. Burma 1929 Stegodon insignis birmanicus Osborn, Mingoon opposite Man- dalay, Burma = Stegodon insignis birmanicus 22. India 1929 Stegodon pinjorensis Osborn, near Siswan, India = Stegodon pinjorensis 23. Java 1932 Stegodon bondolensis van der Maarel, Bondol, near Kuwung = Stegodon bondolensis 24. Java 1933 Stegodon trigonocephalus praecursor von Koenigswald, Bumiaju {Not determined 25. Japan 1934 Parastegodon? kwantoensis Tokunaga, Kakio, Kanagawa Prefecture} by the 26. China 1935 Stegodon yiishensis Young, Yushe | present author 27. China 1935 Stegodon officinalis Hopwood, Szechuan(?) = Stegodon officinalis 28. China 1935 Stegodon zdanskyi Hopwood. Exact locality unknown = Stegodon zdanskyt 29. Japan 1935 Parastegodon sugiyamai Tokunaga, Iruhi in Saida Village, Shikoku Not determined by the present author 30. Borneo 1936 Stegolophodon lydekkeri Osborn, near Bruni = Stegolophodon lydekkerv
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(Left) Map illustrating J. Crawfurd’s journey to Ava and Martaban in the years 1826 and 1827. After William Clift, 1828, Pl. xiv. (Right) Map illus- trating the explorations of Barnum Brown for the American Museum of Natural History in the year 1923. Based on the official Indian Survey Gazetteer.
3-3. Ava, upper levels of the Irrawaddy Series (600 fect in thickness) = Upper Pliocene, Upper Siwaliks of India, containing Stegodon birmanicus Osborn and undetermined Bos. [Lower Pleistocene (cf. Colbert, chap. XXII, pp. 1450, 1451 of the present Memoir.—Hditor.} 2
“
Pondaung Clays (50 feet in thickness) = Upper Eocene, containing the suilline Anthracotheriide; also of the Order Perissodactyla: Fam. Titano-
theriide, Gen. Sivatitanops, sp. S. cotleri, S. birmanicum, S. rugosidens, Gen. Eotitanotherium, sp. 2. lahirii; Pam. Amynodontide, Gen. Paramynodon, sp. P. cotteri, P. birmanicus;
Fam. Tapiride [=Lophiodontide|, Gen. Indolophus, sp. I. guptai, Gen. Chasmotherium, sp. C. birmanicum. [For additional members of the Pondaung fauna, see Colbert, 1938.1, pp. 255-398.—Editor.]
1. Yenangyaung (250 miles south of Ava), lowest levels of the Irrawaddy Series (estimated at 1500 feet in thickness), base of the Middle Siwaliks, con- taining Slegolophodon latidens type, Stegodon elephantoides type, S. cliftit type [Stegodon elephantoides (=cliftii)).
[Nore sy Epwin H. Corserr:—The total thickness of the Irrawaddy Series is estimated at 5,000 feet, of which the upper levels only (about 600 feet), so far as known, are mammal bearing and are now regarded by recent geologists and paleontologists as of Lower Pleistocene age. Stamp (1922, pp. 497, 498), for example, has shown that Mastodon [Stegolophodon| latidens and Hippopotamus irravaticus are probably limited to the upper Irrawaddy beds (Lower Pleisto-
cene), He considers that, Pilgrim erred (1910, p. 196) in placing these species in the lower Irrawaddy fauna (ef. Colbert, Chap. XXII, pp. 1450, 1451, of the present Memoir). Colbert places all the proboscideans discovered in Burma up to the present time, namely, Stegolophodon latidens, Stegodon elephantoides, S.
clephantoides (=cliftii), and S. birmanicus, in the upper Irrawaddy Series, thus assigning them to the Lower Pleistocene. The lowest levels of the Series are considered of Middle to Upper Pliocene age.
Likewise the Pondaung clays are of a total thickness of 6,500 feet, the mammal-bearing portion apparently not exceeding 50 feet,—Editor. |
$24
THE STEGODONTINA: HISTORY 825
1. THE FIRST TWO STEGODONTS, DISCOVERED IN BURMA, 1828
Gro.ocy, IRRAWADDY River, BurmMa.—In the years 1826 and 1827, J. Crawfurd, F.R.S., while on an em- bassy to Ava, Burma, discovered an extensive deposit of organic remains in that unknown and distant region. On the Irrawaddy River, 250 miles below Ava, a gravel and sand deposit contained fossil bones, as mapped and described by Buckland (1828). This is the type region of Mastodon latidens Clift, 1828, and of Mastodon elephant- oides Clift, 1828, also of ‘‘Hlephas cliftiv”” Falconer and Cautley, 1846. According to Buckland (1828, p. 378) the exposure is an extensive one:
These plants were found most abundantly in the same region with the fossil bones, but occur also along nearly the whole course of the Irawadi from Ava to Prome. They were principally collected from a tract of country [Footnote: ‘See annexed map, Plate xuiv.’] extending over a square of more than twenty miles on the east bank of the Irawadi, near the town of Wetma- sut, about half-way between Ava and Prome, between lat. 20° and 21° N. The occurrence of bones was most abundant in a small space near the centre of this district, occupying about one third of the above-named area, the surface of which is com- posed chiefly of barren sand hills mixed with gravel; beneath these are strata containing shells and lignite, through which they sink wells about two hundred feet to collect petroleum.
This indicates that the cotype specimens came from lower and higher geologic levels, a fact not realized in Buckland’s paper, nor in subsequent descriptions excepting those of Pilgrim.’ Referring to Clift’s paper of 1828, Buckland mentions (p. 380):
... two new and strongly characterized species, one of which, from its approximation to the elephant in the structure of the teeth, Mr. Clift proposes to designate by the name of Mastodon elephantoides: to the other he has given the name of Mastodon latidens.
Lydekker (1886.2, p. 81) states that the type of H. cliftii, 1.M"', ‘‘was obtained near Yenankhoung, on the left bank of the Irawadi in Upper Burma, by Crawfurd in 1826, and is preserved in the Museum of the Geological Society.’”
D. N. Wadia (1919, p. 213) includes this Burmese |Yenangyaung] deposit in the ‘Irrawaddy system” [series], which combines marine and fluviatile strata. The upper part (= 2,000 feet), composed of ‘‘sands and clays with abundance of fossil wood and mammals,” is of fluviatile origin and corresponds to the Manchhars of Balu- chistan and to the Siwaliks of the sub-Himalayas.
Barnum Brown visited this region in 1923 for the American Museum of Natural History and prepared a new map (Fig. 692, right) of this classic collecting ground which should be compared with Crawfurd’s original map (Fig. 692, left). The three geologic levels, discovered along the Irrawaddy River, Burma, are shown in these maps.
HISTORY OF DISCOVERY OF BURMESE STEGODONTS Curr, 1828.—The first proboscidean described from Burma was Mastodon latidens Clift (1828, p. 371), the second species was Mastodon elephantoides Clift (1828, p. 372), both species founded on excellent cotypes with the excellent figures which are reproduced herewith (Figs. 693, 694, 695, 696).
MASTODON LATIDENS.—The cotypes of WM. latidens (Figs. 693, 694) were found in 1826 on the Irrawaddy River, 250 miles below Ava, near Yenangyaung, Burma. Clift’s figured specimens include: Plate xxxvi, upper jaw (palate with anterior molar teeth, M’ much worn), which rightly (Osborn) belongs to Mastodon | = Stegodon| elephantoides; Plate xxxvu, upper molar teeth of the right side, M’, M* (a younger animal), also Plate xxxvill, fig. 1, anterior part of the lower jaw; these certainly are the cotypes, because they conform with Clift’s deserip- tion—all from the left bank of the Irrawaddy.
See this Volume, Chapter XXII, p. 1450, by Edwin H. Colbert.—Editor.] *[Now in British Museum (Brit. Mus. M. 10520)].
FOUR CLASSIC STEGODONT AND STEGOLOPHODONT TYPES OF CLIFT AND FALCONER AND CAUTLEY
Corrre Turrp INFERIOR MOLAR OF STEGOLOPHODON LATIDENS Fig. 694 (right). Cotype third right inferior molar, r.Mg, of Mastodon latidens Clift, 1828, Pl. xxxvu, fig. 1, one-third natural size. Jaw omitted.
From near Yenangyaung, Burma.
Fig. 695
Lrecroryer LowrrR JAw or STEGODON PELEPHANTOIDES
Vig. 696 (right). Lectotype second
and third left inferior molars, 1.M3-2, of
Mastodon elephantoides Clift, 1828.
Pl. xxxviu, fig. 2, one-third natural
size. From near Yenangyaung, Burma,
WAG de 6 Scharf Lahey
Laecrotyph PALATE OF STEGOLOPHODON LATIDENS
Fig. 693 (left). Lectotype palate (perspective of r.M®*) of Mastodon latidens Clift, 1828, Pl. xxxvn, fig. 1, with r.M? in situ, about one-third natural size. Inverted to show natural position of molars. From near Yenangyaung, Burma.
Observe beginnings of binary fission of cones in tritoloph. An accurate scale drawing of these teeth is shown in figure 716 from a cast (Amer. Mus. 21978).
Coryrk OF STEGODON ELEPHANTOIDES Curr (=cLieru FALCONER)
Tig. 695 (left). Type first left superior molar, 1.M!, of Elephas cliftii Paleoner and Cautley, 1846, after photograph of cast Amer. Mus. Warren Coll. 10382, one-half natural size. Inverted to show the molar in natural position. From near Yenangyaung, Burma. See also figure 683. Compare Clift, 1828, Pl. xxxrx, fig. 6. Original in British Museum (Natural History) M.10520.
Printed by C.
1 Right Lower Jaw, Mastodon latdens. 2 Lge Lower Jaw, Mastodon Elophanindes. t Nak size
Mules below Ava.
Found: by J Crawfurd Esq] 250
Fig. 696
826
THE STEGODONTIN: HISTORY 827
MASTODON LATIDENS (oP. crv., Pp. 370).—On comparing the teeth of our Mastodon latidens with those of the Mastodon of the Ohio (M. giganteum), we shall find the elevated points or ridges in the tooth of the former more numerous, less distant, and the interstices less deep than in those of the latter; in short, we shall observe that the teeth begin to assume the appearance of those of the elephant. On advancing to Mastodon elephantoides, we shall find all these features of similarity more strongly developed ;—the points and ridges are still more numerous, and the structure, were it not for the absence of erusta petrosa, becomes almost that of the tooth of the elephant.
MASTODON ELEPHANTOIDES.—This species (Fig. 696) was found in the same locality as the cotypes of M. latidens. Clift’s figured specimens include: Plate xxxvir, fig. 2 (lower jaw with My, M, in situ); Plate xxxv1 (palate figured by Clift as M. latidens), and Plate xxxrx, fig. 6 (a first superior molar, 1.M!) afterward made the type of Elephas cliftii by Falconer and Cautley, referred in the present Memoir to Stegodon elephantoides (=cliftii).
MASTODON ELEPHANTOIDES (OP. crT., Pp. 372).—The tooth [Fig. 696], which is eleven inches long and three inches and a half broad, has no less than ten denticules [i.e., ridges], and each of these denticules is mammillated with small points; five being the smallest number, and eight the greatest on any one denticule. In front of this beautiful tooth we have a remnant of the preceding one,...
Osborn, 1924: Pending final revision, which ean only be made by a reéxamination of the specimens and the localities from which they came, we may designate these Stegodonts from Burma as follows:
Mastodon [= Stegolophodon| latidens Clift, 1828, founded upon a palate from near Yenangyaung, Burma, with five and a half ridge-crests in the third superior molar, four and a half to five in the second superior molar, and with four to five mamillz (or conelets) on each crest. See Clift, 1828, Pl. xxxvu, fig. 1 (Fig. 693 of the present Memoir). Also a lower jaw (see Clift, Pl. xxxvu, fig. 1), r.M3 with seven ridge-crests (Fig. 694 of the present Memoir).
Mastodon [= Stegodon| elephantoides Clift, 1828, founded upon a third inferior molar of the left side, 1.M3, from near Yenangyaung, Burma, with ten ridge-crests, five to eight conelets on each. See Clift, 1828, Pl. xxxvuit, fig. 2 (Fig. 696 of the present Memoir). Also a palate showing r.M? and |.M?, with six and a quarter ridge-crests (see Clift, 1828, Pl. xxxvr), erroneously marked “Upper Jaw of Mastodon latidens.” (Not figured in the present Memoir.)
Stegodon elephantoides (=cliftiz) Also a left first superior molar, |.M!, with six and a quarter ridge-crests, from near Yenangyaung, Burma. See Clift, 1828, Pl. xxxrx, fig. 6, marked ‘Upper molar of M. Elephantoides.” Afterward made the type of Elephas cliftii by Falconer and Cautley. (Figs. 683, 695, of the present Memoir.)
ORIGINAL CorypPEs OF MASTODON LATIDENS Curt (Fries. 693, 694)
First Species.—Clift’s designation (1828) of Mastodon latidens as including a lower jaw (Pl. xxxvui, fig. 1—Fig. 694 of the present Memoir) and a palate containing M’, M? (Pl. xxxvu, fig. 1—Fig. 693 of the present Memoir) establishes these specimens as the cotypes. The same superior teeth, r.M?’, r.M’, were sectioned and figured by Falconer and Cautley (1846, p. 48 [1845, PI. 1m, fig. 8]):
The last tooth shows five principal ridges with a posterior talon ridge and a subordinate ridge in front. The ridges are transverse, and divided by a longitudinal cleft into two pairs of principal points without intermediate mammille in the hollows. The enamel is very thick, and the cement is reduced to a thin layer which is only observable in the bottom of the hollows. . . . The anterior tooth [M?’] had been a long time in use, and the ridges are nearly all worn out. They were four in number, in this as well as in the two teeth which preceded it in the jaw. We believe this to be a small or dwarf variety of M. latidens. . . .
(Clift, 1828, p. 371): Dentition—HEach tooth of the lower jaw [Fig. 694] consists of seven denticules, which are elevated, rounded, and mammillated: the mammillz being from three to four in number. The dentition both in this species and in M. elephantoides, very much resembles that of the elephant. We have the molar tooth gradually protruded forward, and rising as the fangs are added, according to the demand made by the abrasion of the exposed crown, and the consequent absorption of the anterior fang; the posterior part of the tooth not having cut the gum, while the anterior portion is completely worn away. The relics of the preceding tooth, the place of which the tooth in use was progressively supplying, are plainly to be seen [Foot- note: ‘See Plate xxxvul. fig. 1. Plate xxxvu. fig. 2.’].
828 OSBORN: THE PROBOSCIDEA
According to the above, the ridge formula is as follows: Ridge formula of Mastodon |= Stegolophodon| latidens: Dp 4; M 1; M 2% M 3°". Osborn, 1927: The above is the main ridge formula given by Falconer, omitting the anterior and posterior talon ridges (compare tables above, Section I, No. 3, and below Section IV of this chapter, also full description).
ORIGINAL CoryPEs OF MASTODON ELEPHANTOIDES Curt (Fig. 696) Srconp Specres.—The second species is Mastodon elephantoides Clift.
Curt, 1828, p. 372.—Mastodon elephantoides.—M. dentibus molaribus latis, denticulis numerosis, compressis. This species must have been smaller than the last; and though we have one fine example of the lower jaw, showing the tooth in the highest degree of perfection, that is the only portion of the animal from which we can safely draw any inference as to its struc- ture and habits. The tooth, which is eleven inches long and three inches and a half broad, has no less than ten denticules li. e., ridge-crests], and each of these denticules is mammillated with small points [conelets]; five being the smallest number, and eight the greatest on any one denticule. . . . The denticules of the tooth are much more compressed than those in the species last described; they are closer together [Footnote: ‘Eight denticules of M. elephantoides occupy the same space as five denticules of M. latidens.’|, and the enamel appears to be not so thick. They form a series of plates mucronated with small points. There is no apparent commissure, neither is there any central depression: on the contrary, the plates rather rise in the middle.
Clift uses the word denticules in the sense of ridges, or crests, or lophs; subsequently Falconer uses the word denticles in the sense of ‘“‘mammille” or conelets. Osborn introduces the word conelets, because these small, rounded “mammille”’ appear on the summits of the primary cones, as explained above (p. 812).
(Clift, 1828, ‘Explanation of Plates,’’ Pl. xxxrx, fig. 6): ‘Upper molar tooth of Mastodon elephantoides.”’ [No mention of this tooth is found in the text.] As reproduced herewith (Fig. 695) this molar corresponds closely in scale and structure with Clift’s type of M. elephantoides (Fig. 696).
Osborn, 1927: From these two teeth, figured and described together by Clift, 1828, not improbably represent- ing the same species, also from the palate with r.M? and 1.M? in situ (Clift’s figure, Pl. xxxvr), the following ridge formula may be written:
Ridge formula of Mastodon [= Stegodon] elephantoides: M 1° M 2°* M3;5.
The above ridge formula is similar to that of Stegodon bombifrons and much more primitive than that of S. insignis-ganesa. Consequently Mastodon elephantoides Clift, 1828, of Lower Pliocene age,’ based upon two figured specimens (Figs. 695, 696), appears to be well established as the second species of Stegodon described from Burma in 1828. The name elephantoides was dropped, however, by Falconer and Cautley in 1846 and a new name, Hlephas cliftii, was applied to the second specimen (Fig. 695 of the present Memoir) figured by Clift, Pl. xxxrx, fig. 6.
Species M. ELEPHANTOIDES DroppED BY FALCONER AND LyYDEKKER.—In 1846, Falconer and Cautley erroneously alleged that Clift had confused the remains of the two species Mastodon latidens and M. elephantoides under the name M. elephantoides; they accordingly (op. cit., p. 47) dropped the name elephantoides and proposed a new name for the species of “transitional Mastodons,” which had been partly confused with Mastodon latidens. Thus to the Stegodonts with six ridges on the intermediate molars they gave the name of Hlephas cliftii (Figs. 683, 695), and to the Stegodonts with a greater number of ridges the name of FH. insignis (Fig. 697). In a sub- sequent paper (Falconer, 1857, p. 314), these and other species appeared under the subgeneric name of Stegodon Falconer. Lydekker (Palaeontologia Indica, 1880, pp. 256, 257) also set aside the prior specific name elephantoides Clift, 1828, and made it a synonym of Stegodon cliftii Fale. and Caut., 1846. In subsequent literature (e. g., Pilgrim) the Faleconer-Lydekker usage is followed, i.e., M. elephantoides is dropped.
M. eLEPHANTOIDES Revivep.—In the present Memoir the specific name elephantoides is revived by Osborn for reasons given in the systematic revisions above and below.
‘See note on page 824 above regarding the Lower Pleistocene age of Stegodon elephantoides.—Editor.]
THE STEGODONTINA: HISTORY 829
2. DISCOVERIES IN INDIA AND BURMA (1845, 1846) Tuirp Species.—The third species of Stegodont (Hlephas insignis) was described (Falconer and Cautley, 1846, p. 37) as follows (see Fig. 697):
... the four anterior ridges being affected by wear, and the six posterior ridges entire, ... The white mass in the centre [dentine] represents the body of ivory, which is projected upwards in ten angular lobes terminating in a sharp edge. .. . The interspaces of the five posterior ridges of enamel are completely filled up by a mass of cement, or ‘cortical,’ much exceeding the enamel in thickness; and in quantity in nearly as great an amount of development as the ivory core of the ridge.
Fic. 697. Lecroryrr anp CoryPr or STEGODON INSIGNIS (Left) Lectotype superior molar, 1.M%, of Elephas insignis Falconer and Cautley, 1846 [1845, Pl. 1, fig. 6a], one-third natural size. Probably Pinjor horizon, Upper Pliocene [Lower Pleistocene], Upper Siwaliks, India. Inverted to show natural position of molar.
(Right) Elephas [=Stegodon] insignis cotype. Anterior portion of a third inferior molar, M3. After Falconer and Cautley, op. cit., Pl. u, fig. 6b.
Falconer distinguished this species as possessing a ridge formula of M 3 2°, with valleys between acute ridges deeply filled with cement, as compared with Stegodon elephantoides which has ten less acute ridges without cement (Fig. 696). Lectotype ridge formula (Fig. 697) of Elephas | = Stegodon] insignis: M 3 1°.
FourtH Specres.—The fourth species of Stegodont (described by Falconer and Cautley, 1846, p. 45) was named Hlephas ganesa:
Vig. 7a, pl. 3., represents a section of the last uppér molar of an undescribed Indian fossil species, named #. Ganesa, in this work. The crown consists of ten principal ridges, with a subordinate ‘talon’ ridge in front and behind [i.e., M 3%1°]. 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. A small portion is broken off at the anterior end. The disposition and relative propor- tions of the ivory, enamel, and cement, bear the closest resemblance to those of the corresponding tooth of H. insignis (pl. 2, fig. 6a), and the number of ridges agrees. The section presents the same chevron-formed character in the ridges, but the interspaces are narrower, the cement is in less quantity, and the layer of enamel is thicker.
Fig 7a
Lydekker observes (Lydekker, 1880.1, p. 268): ... “these latter teeth, however, cannot be distinguished from those named
Stegodon insignis, and as we shall see subsequently, it is only the
LECTOTYPE OF STEGODON GANESA F a Fig. 698. Lectotype M? of Elephas Ganesa Faleoner and 2dult skulls of these two very closely allied species that can be
Cautley, 1846, [1845, Pl. m1, fig. 7a], one-third natural size. — istinguished.’’ This has led to the opinion that S. insignis may Probably Pinjor horizon, Upper Pliocene [Lower Pleistocene].
Upper Siwaliks, India. represent a female and S. ganesa a male of the same species.
830 OSBORN: THE PROBOSCIDEA
The ridge formula of the leetotype third superior molar of Stegodon ganesa (Fig. 698) appears to be practi- cally the same as that of the leetotype of S. insignis (ig. 697). Other more progressive ridge formule are shown in the comparative ridge formul tables above and below in this Memoir.
Ridge formula (Fig. 698) of Hlephas [= Stegodon| ganesa: M 3 *?*°*.
Fier Species.—Faleoner described (Falconer and Cautley, 1846, p. 46) Elephas bombifrons, the fifth species
of Stegodont, as follows:
This species, of the distinctness of which we are assured, by possessing several crania containing perfect teeth, belongs to the same group [Stegodon] as the two species last described. The crown is divided into similar transverse ridges, composed of numer- ous mammill, which yield a corresponding chevron-shaped section, and the interspaces are occupied by a thick coat of cement; but they differ, in being broader and less elevated, with more open hollows. ‘The principal ridges of the last molar [M*] do not exceed eight in the upper jaw, and nine in the lower [M3]; while in #. znsignis they amount to ten in the former [M*], and reach as many as thirteen in the latter [M3]. The last tooth of the upper jaw measures eleven inches in length, by four and
a half in width. Falconer and Cautley (1846 [1847, Pl. xxvm1]) assign to M®* nine ridges and a heel.
Lectotype ridge formula of Stegodon bombifrons: M 35.
SrxTH Specres.—A sixth species of Stegodont (Figs. 683 and 695) was erroneously proposed by Falconer and Cautley in 1846 (1846, p. 47) under the name of ‘Elephas cliftii.. They selected the type of this species as
follows:
Lg #
CoTyrPr OF STEGODON BOMBIFRONS
Vig. 699. Cotype skull of Blephas bombifrons Falconer and Cautley, 1846 [1847, Pls. xxvu, xxvii], one-sixth natural size. Brit. Mus. M.2979; cast Amer. Mus. Warren Coll. 10378. From the Siwalik Hills, India, probably the Dhok Pathan horizon, Upper Siwaliks, Middle Pliocene. As a view of the third molars was not given in the lectotype of Lydekker (Ialconer and Cautley, op. cit., Pl. xxvi, Brit. Mus. M.2978), the cotype skull is figured here.
THE STEGODONTIN#: HISTORY 831
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.
This was an unfortunate error. As explained above, the tooth in Pl. 39, fig. 6, of Clift’s Memoir, measuring 155 mm. in length, 83 mm. in breadth, has a ridge formula of M 1 **; the palate specimen (PI. 36), erroneously identified by Clift as “Mastodon latidens,” also has a ridge formula of (?)M 2°*. Consequently they belong to the same species, namely, Stegodon elephantoides, but to preserve the name cliftii, which runs all through the previous literature, they are designated in the present Memoir as Stegodon elephantoides (= cliftiz).
STEGODON ELEPHANTOIDES (= CLIFTII)
Fig. 700. Original type figure of Elephas cliftti Falconer Fig. 701. New figure of type of Elephas cliftii Falconer and and Cautley, 1846, a first superior molar of the left side, 1.M?, Cautley, 1846, a first superior molar of the left side, 1.M', de- one-half natural size. From near Yenangyaung, Burma. scribed and figured as M. Elephantoides by Clift, 1828, Pl. xxxrx, After Clift, 1828, Pl. xxxrx, fig. 6, figured as an “Upper fig. 6. Subsequently selected by Falconer and Cautley as the type molar of M. Hlephantoides.”’ Original in the British Museum of Elephas cliftii. After photograph of type cast (Amer. Mus. (Brit. Mus. M.10520); cast Amer. Mus. Warren Coll. 10382. Warren Coll. 10382). One-half natural size. Observe that the
Inverted to show natural position. molar is here placed in its natural position. British Museum (Brit. Mus. M.10520).
Original in the
3. THE STEGODONTS OF CHINA, INDIA, JAVA, THE PHILIPPINE ISLANDS, AUSTRIA, JAPAN, AND BURMA
SEVENTH SpEcies.—The seventh species of Stegodont described (Fig. 702) was the Stegodon sinensis of Richard Owen (Owen, 1870, p. 417), alleged to be (p. 421) “from marly beds in the vicinity of Shanghai,’”’ China, which has been mistakenly regarded by some authors as close to, or as a synonym of, Stegodon cliftii (Owen, op.
Tig. 702. Type r.Dp* of Stegodon sinensis Owen, 1870, Pl. xxvu, figs. 1, 2, natural size. From “marly beds in the vicinity of Shanghai,’’ China, of Upper Miocene [Lower Pliocene] age.
832 OSBORN: THE PROBOSCIDEA
cit., p. 418; Brauns, 1883, p. 44). This synonymy is not borne out by careful comparison with the type figures, because S. s¢nensis is much more primitive than S. clifti.
Eraurx Species.—In the same paper Owen (op. cit., 1870, p. 421) describes Stegodon orientalis, the eighth species of Stegodont (Fig. 703), “from a cave, near the city of Chung-king-foo, in the province of Sze-chuen,” ‘more resemble the teeth of Stegodon Cliftii, St. insignis,
‘
based upon molar fragments which he rightly observes and St. ganesa of Falconer than does the St. sinensis; and in the apparent quantity of coronal cement... as well as in the evidence of a hinder talon... they are more like Sé. insignis than St. Cliftii.”
Fig. 703. Type of Stegodon orientalis Owen, 1870, Pl. xxviu, figs. 1-4. “Portion of true molar” (figs. 1, 2) and “hind end of milk-molar” (figs. 3, 4). From near Chungkingfoo, China, probably of Lower Pleistocene age.
Nintu Species.—Mastodon cautleyi Lydekker, 1886 (1886.1), was founded on five teeth of the upper jaw, of which we select as the type (Fig. 704) the third left upper true molar, from Perim Island, India, a tooth first figured Fig. 18. by Falconer and Cautley and referred to the species
Mastodon latidens Clift (Pl. xxx1, figs. 6, 6a Vol. I, fig. 142 of present Memoir, the caption of which is erroneous—corrected in present chapter, p. 821). The four cotypes, obviously smaller and much more primitive teeth than the type of Mas- todon latidens, are also from Perim Island and are recorded, like the lectotype, from an older geologic horizon, namely, Upper Miocene [now (1935-19388) regarded as Middle Phocene, Dhok Pathan]. Ac- cording to these specimens, the ridge formula of Mastodon {|= Stegolophodon] cautleyi is about the same as that of Mastodon | = Stegolophodon| latidens.
see
Mastodon cautieyi.—Vhe third left upper trve molar, in an unworn condition; from the Siwaliks of Perim Island. 4. The lower border of the figure is the inner border of the specimen.
LECTOTYPE OF STEGOLOPHODON CAUTLEYI Mastodon [ = Stegolophodon| cautleyr: M2* Mises
Tig. 704. Third left, superior molar of Mastodon cautleyi Lydekker, 1886, Trento Sprecres.—In 1887 Martin described selected as the type (see Pilgrim, 1913, p. 294). Reproduced after Lydekker, 1886 4 j as i 5 (1886.2, p. 73, fig. 18), one-half natural size. (“Fossile Siiugethier-reste von Java und Japan’’)
THE STEGODONTINA: HISTORY 833
from Java a species to which he gave the name Stegodon trigonocephalus (Fig. 705), in reference to the triangular shape of the head. The geologic level is regarded by Matsumoto as equivalent to the Lower Pleistocene, Boulder Conglomerate beds of India. It may be of the same geologic age as the type of Pithecanthropus erectus. The juvenile type does not admit of giving the mature ridge formula; the ridges are closely compressed, buried in cement, and each ridge is surmounted with ten to twelve conelets, the exact number being indeterminable from the figures.
Fig. 705. Type of Stegodon trigonocephalus Martin, 1887, immature skull, one-eighth natural size. From vicinity of Surakarta, Java, probably of Lower Pleistocene age. (Right figures) Tab. 11, figs. 1, 1a; (left figure) Tab. 1m, fig. 1.
ELEVENTH Specres.—The eleventh species (Fig. 706) was described by Naumann in 1890 from Mindanao, Philippine Islands, as Stegodon mindanensis. It was first referred to Stegodon trigonocephalus by Naumann in 1887, but subsequently was made the type of a distinct species. The compressed cement-covered ridges with multiple conelets indicate a Lower to Middle Pleistocene stage of evolution, similar to that of S. trigonocephalus.
TWELFTH Specres.—A twelfth species, Stegodon airdwana (Fig. 707), was described by Martin in 1890 from Alas-Tuwa, Java, based upon a type jaw containing the right and left third inferior molar teeth, Ms. In its high ridge formula the author compared it to Stegodon insignis and S. ganesa. In a later paper by Janensch (1911, p. 187), he distinguished S. airdwana from both S. insignis and S. ganesa and wrote the ridge formula as follows, according to the present writer’s understanding of the Janensch system [ef. Table V]:
Ridge formula of Stegodon atrdwana: Dp 372 Dp 4°" M 1 ** D '4-9-16 YY \g-1 1-12-16 M 2 #24 Mf 3 Heute
This species is important because it is highly characteristic of the
Trinil Pithecanthropus erectus beds.
Typx or Stecopon (ARCHIDISKODON?) MINDANENSIS THIRTEENTH Spectes.—In 1908 Dubois described from the Trinil Vig. 706. Type of Stegodon mindanensis
Naumann, 1890, molar tooth. Mindanao, Philip-
pine Islands, Lower to Middle Pleistocene age. ganesa [which proves to be a synonym of either S. airdwana or S. tri-
After Naumann, 1887, Taf. 1, figs. 1 and 2, under the ; as
name Stegodon trigonocephalus. gonocephalus (see p. 889 below)].
Kendeng-Schichten, Java, Stegodon ganesa javanicus, as a variety of S.
834 OSBORN: THE PROBOSCIDEA
FOURTEENTH SPECIES.— Very important is the Mastodon stegodontordes of Pilgrim, 1913, from Lehri, Punjab, India (Fig. 708), which Pilgrim described as follows (p. 294):
The last species of Mastodon which can be referred to this line [the W. cautleyi-M. latidens line] is the tooth from Lehri, of which the horizon is uncertain but may be possibly Upper Siwalik, figured by Lydekker in Pal. Ind. ser. 10, Vol. I, Plate 39, as M. latidens but which is better recognized as a new species, owing to the almost entire absence of accessory columns, for which I propose the name Mastodon stegodontoides. So close is this to Stegodon clifti that it is hard to separate the two genera. It will be seen that Mastodon stegodontoides carries on none of its ridges more than the usual four columns while anterior ridges of Stegodon clifti carry nine or ten mammille. FIFTEENTH SpEcIES.—Very progressive is the ei Elephas (Prostegodon, Parastegodon) aurorze of Mat- sumoto, 1915-1924, from Mt. Tomuro, Kaga, Japan
(Fig. 709), which Matsumoto first regarded as a stage
TYPE OF STEGOLOPHODON STEGODONTOIDES
Fig. 708. Type r.M® of Mastodon stegodontoides Pilgrim, 1913. After Lydek- ker, 1880, Pl. xxxrx: “Mastodon (Tetralophodon) latidens, Clift. The third right upper true molar: from Lehri, in the Punjab. The specimen is drawn of the natural size, and is viewed from the inner [outer] side.”” Ind. Mus. A.86. Reduced to one-half natural size. Provisionally placed in the Upper Pliocene, Pinjor formation (see Fig. 413, also Pl. xt).
Vig. 707. Type of Stegodon Airawana Martin, 1890, Tab. 1, figs Kendeng-Schichten horizon (Pithecanthropus erectus zone), Middle Tyre OF STEGODON AURORA 1, 2, lower jaw, one-fourth natural size. From Alas-Tuwa, Trinil, Java, Vig. 709. Type r.M? of Blephas (Prostegodon, Parastegodon) aurore Pleistocene. Matsumoto, 1918, Pl. xx, figs. 1 and 3, one-half natural size.
THE STEGODONTINA: HISTORY 835
in Elephas close to the Upper Pliocene Hlephas [= Archidiskodon| planifrons of India, but which he subsequently made the genotype of a new genus Parastegodon. With our fuller knowledge of Stegodon atrdwana, the present species may be placed in the true Stegodon phylum, distinguished by cranial characters from species of the Ele- phantidz, as Stegodon aurore.
SixTEENTH Spectes.—The subspecies Mastodon (Bunolophodon) longirostre Kaup forma sublatidens Schle- singer, 1917, from near Teschen (Schlesien), Austria (Fig. 710), appears to be very close to the Mastodon [ = Stego- lophodon| stegodontoides of Pilgrim; consequently it is removed to this genus, namely, Stegolophodon sublatidens.
SeVENTEENTH Species.—The subspecies Stegodon orientalis shodoénsis of Matsumoto, 1924, named from the Island of Shédo, Inland Sea, is regarded by its author as a descendant of S. orientalis Owen, of China, contemporaneous with the referred S. insignis of the Middle [Upper] Pleistocene, Narbada of India. Not figured in present Memoir.
EIGHTEENTH SpEciEs.—The type of Stegolophodon nathotensis Osborn, 1929, consists of fragmentary molars (Fig. 724) collected by Barnum
Brown in 1922 in the Lower Chinji horizon, near Nathot, India.
Typr OF STEGOLOPHODON SUBLATIDENS
Fig. 710. Type of Mastodon (Bunolophodon) longirostre Kaup forma sublatidens Schlesinger, 1917, one-half natural size. Compare fulllegend equtleyi of the Upper Miocene [Middle Pliocene] is the Stegolophodon caut- below of figure 722.
NINETEENTH SPECIES.—Somewhat more advanced than Stegolophodon
leyi progressus Osborn, 1929 [of the Mio-Pliocene], the type of which (Amer. Mus. 19446) is a complete cranium (Figs. 725,727) collected at the summit of the Lower Chinji horizon, 2,000 feet above the base of the Lower Siwaliks, India. This is a young individual in which the molar ridge-crests are intermediate in formula and pattern between the ‘Mastodon’ cautleyi of Lydekker and the ‘M.’ latidens of Clift.
TWENTIETH SpectEs.—Somewhat more primitive and an- cient than the Upper Pliocene [Lower Pleistocene] Stegodon in- signis Faleoner type is the subspecifie stage Stegodon orientalis grangeri Osborn, 1929 (Fig. 762) of the Yangtze River region near Wanhsien, Sezchuan, 140 miles northeast of the type locality of S. orientalis Owen. This subspecies is now very fully known and is amply illustrated and defined in the present Memoir. See type figures on pages 876, 877, 879, and 881 below.
TWENTY-FIRST SPECIES.—Stegodon insignis birmanicus Os- born, 1929 (Fig. 758), from the Upper Pliocene of Burma. See type figure on page 875.
TWENTY-SECOND Species.—Stegodon pinjorensis Osborn, 1929, from the Lower Pleistocene of India, upper levels of the Pinjor horizon (Figs. 711, 765, 767).
|The following species have been described since the author’s
Fig. 711. Photographie reproduction of type palate of
intensive review of the true Stegodonts, consequently the de- Stegodon pinjorensis Osborn, 1929 (Amer. Mus. 19772), collected - ‘ ; A : by Barnum Brown three miles north of Siswan, India. About terminations of Professor Osborn cannot be given and the spe- one-eighth natural size. See figure 765 below.
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THE STEGODONTIN: HISTORY 837
cies are listed here according to the nomenclature of the various authors, without comment, with the exception of Parastegodon? kwantoensis and P. sugiyamai.
TWENTY-THIRD SPECIES.—The type of Stegodon bondolensis van der Maarel, 1932, is a fragment of a mandible with what is inferred to be the third molar of each side, found at Bondol, Java (Fig. 782 of the present Memoir).
TWENTY-FOURTH SpecrES.—In 1933 von Koenigswald described from Bumiaju, Java, a lower jaw with third molar of both sides complete, which he named Stegodon trigonocephalus praecursor (Fig. 783 of the present Memoir).
TWENTY-FIFTH SPECIES.—The species Parastegodon? kwantoensis Tokunaga, 1934, Pl. 1x, a portion of a jaw with second right molar in situ, is indeterminate, owing to the fact that Professor Osborn regarded Parastegodon as either a progressive Stegodon or a primitive Archidiskodon. See type figure 784 on page 897 below.
TWENTY-SIXTH SpPEcIES.—Stegodon yiishensis Young, 1935, from Yiishe, China. Type, a well preserved upper left third molar, PI. v, fig. 1 (Fig. 785 of the present Memoir).
TWENTY-SEVENTH SprEciEsS.—The type of Stegodon officinalis Hopwood, 1935, is a fragment of an unworn lower molar, said to have come from Szechuan, China. This is figured in Hopwood, 1935, Pl. vu, fig. 3 (Fig. 786 of the present Memoir).
TWENTY-EIGHTH Spectes.—The type of Stegodon zdanskyi Hopwood, 1935, is a fragment of a right third lower molar, consisting of the first four ridges, and figured in Hopwood, Pl. vu, fig. 5 (Fig. 788 of the present Memoir).
TWENTY-NINTH Spectes.—The type of Parastegodon sugiyamai Tokunaga, 1935 [Stegodon? sugiyamai] is either a first or a second molar, probably of the upper left side, found in Shikoku, Japan (Fig. 789 of the present Memoir).—KEditor.|
TuirtTieTH SpEcrEs.—In Volume I of the present Memoir (p. 700, fig. 660) will be found the type descrip- tion of Stegolophodon lydekkeri, dedicated by the present author to his friend Richard Lydekker. The type is a third left superior molar from Borneo, ‘“‘much more progressive than the S. latidens type.”
III. SYSTEMATIC ARRANGEMENT OF THE STEGOLOPHODONTS AND STEGODONTS IN PHYLOGENETIC ORDER
+1. CHARACTERS OF THE SUBFAMILY STEGODONTIN 4’
SUBFAMILY CHARACTERS.—(1) Habits chiefly browsing; tropical forest living; crushing of coarse leafage, herbage, and wood fiber; not progressing to the grazing type. (2) Cranium relatively abbre- viated, mesocephalic, bathycephalic; grinding-tooth plane deeply depressed to occipital condyles; occipitofrontal plane neither elevated nor expanded, non-acrocephalic, non-hypsicephalic. (3) Tusks straight or slightly curved, horizontal or subhorizontal in direction, continuously serving in browsing habits. (4) Grinding teeth brachyodont to subhypsodont, ridge- plates of M 8 increasing from 3% Bceerador to *=* [Stegodon].
‘In Volume I (1936) of the present Memoir, the author separated the Stegodontoidea (true Stegodonts) from the Elephantoidea (pp. 22, 25), also the genus Stegodon from Stegolophodon, placing all the Stegolophodonts in the superfamily Mastodontoidea, family Mastodontide, new subfamily Stegolophodon- tine (p. 700) and the true Stegodonts in the superfamily Stegodontoidea, family Stegodontide, subfamily Stegodontine (see pp. 806-808 above).—EKditor.]
838 OSBORN: THIE PROBOSCIDISA
These characters are observed in the species and subspecies discovered since 1828, many of which were origi- nally referred to the genus Mastodon and the genus Hlephas and are now referred to the primitive genus Stegolo- phodon and to the more progressive genus Stegodon.
2. HISTORY OF THE GENERIC NAMES ASSIGNED TO THE STEGOLOPHODONTS AND TO THE STEGODONTS
The history of the generic term Stegodon Falconer is fully given in Chapter X. XI, also in the present Chapter XIV, which may be summarized as follows: (1) The name Stegodon applies chiefly to the more progressive Stegodonts. (2) To the more primitive Stegodonts, e.g., the Upper Miocene [Middle Pliocene] Mastodon cautleyz, the Lower Pliocene’ M. latidens, and the Pliocene M. stegodontoides, the name Stegolophodon Schle- singer 1s applicable. The name Stegolophodon also applies to the Mastodon (Bunolophodon) longirostre Kaup forma sublatidens Schlesinger of Austria. The name Stegolophodon Schlesinger, 1917, preoccupies the name Prostegodon Matsumoto, 1922-1924. (8) The name Parastegodon Matsumoto, 1924, based on the genotypic species Hlephas aurore, appears to be in part a synonym of Archidiskodon. These interpretations of the generic names adopted in the present Memoir are displayed in the following table.
PHYLOGENY. STEGOLOPHODONTS AND STEGODONTS ARRANGED BY COUNTRIES IN APPROXIMATE ASCEND- ING ORDER OF EVOLUTION, THE MOST PRIMITIVE FORMS BELOW, THE MOST PROGRESSIVE FORMS ABOVEE |The species described by various authors since 1929 are omitted here owing to the fact that they had not been studied in detail by Professor Osborn.—Editor.]
STEGODON Japan 1924 Stegodon orientalis shodoénsis Matsumoto, Island of Shédo, In- land Sea, Japan = Stegodon orientalis shodoénsis 1915-1924 Hlephas (Prostegodon, Parastegodon) aurore Matsumoto, Mt. Tomuro, Kaga, Japan = Stegodon aurore Philippines 1890 Stegodon mindanensis Naumann, Mindanao, Philippine Islands =Stegodon (Archidiskodon?) mindanensis Java 1908 Stegodon ganesa var. javanicus Dubois, Kendeng, Trinil, Java | = Stegodon airdwana or S. trigonocephalus | 1887 Stegodon trigonocephalus Martin, vicinity of ?Surakarta, Java = Stegodon trigonocephalus 1890 Stegodon Airadwana Martin, Alas-Tuwa, Kendeng, Trinil, Java =Stegodon airdwana China 1870 Stegodon orientalis Owen, Chungkingfoo, Szechuan, China = Stegodon orientalis 1929 Stegodon orientalis grangeri Osborn, Yenchingkou, Szechuan, China = Stegodon orientalis grangeri 1929 Stegodon pinjorensis Osborn, near Siswan, India = Stegodon pinjorensis India 1846 [1845] Stegodon Ganesa Fale. and Caut., Siwalik Hills, India = Stegodon insignis-ganesa 1846 [1845] Stegodon insignis Fale. and Caut., Siwalik Hills, India = Stegodon insignis-ganesa Burma 1929 Stegodon insignis birmanicus Osborn, Mingoon opposite Man- dalay, Burma = Stegodon insignis birmanicus India 1846 Hlephas bombifrons Fale. and Caut., Siwalik Hills, India = Stegodon bombifrons Burma 1828 Mastodon elephantoides Clift, near Yenangyaung, Irrawaddy tiver, Burma = Stegodon elephantoides 1846 Elephas cliftdi Fale. and Caut., near Yenangyaung, Irrawaddy River, Burma = Stegodon elephantoides (= cliftir) China 1870 Stegodon sinensis Owen, near Shanghai, China = Stegodon sinensis
STEGOLOPHODON
India 1913 Mastodon stegodontoides Pilgrim, Lehri, Punjab, India = Stegolophodon stegodontoides Austria 1917 Mastodon (Bunolophodon) longirostre Kaup forma sublatidens Schlesinger, Teschen (Schlesien), Austria = Stegolophodon sublatidens 3orneo 1936 Stegolophodon lydekkeri Osborn, near Bruni, Borneo = Stegolophodon lydekkeri Burma 1828 Mastodon latidens Clift, near Yenangyaung, Irrawaddy River, 3urma = Stegolophodon latidens India 1929 Stegolophodon cautleyi progressus Osborn, near Chinji Bungalow, India = Stegolophodon cautleyt progressus 1886 Mastodon cautleyi Lydekker, Perim Island, India = Stegolophodon cautleyt 1929 Stegolophodon nathotensis Osborn, near Nathot, India = Stegolophodon nathotensis
[See note on page $24 aboye regarding the Lower Pleistocene age of Mastodon [ =Stegolophodon| latidens.—Editor.]
THE STEGOLOPHODONTINA: STEGOLOPHODON 839
SuPERFAMILY: MASTODONTOIDEA Osborn, 1921 FaMILy: MASTODONTID Girard, 1852 SUBFAMILY: STEGOLOPHODONTIN® Osborn, 1936
Genus: STEGOLOPHODON Schlesinger, 1917
Original reference: Denk. Naturhist. Hofmus., 1917, I, p. 115.
Genotypic species: Mastodon latidens Clift, 1828.
Compare Stego (lopho) don Pohlig, 1888, p. 252.
Syn.: Prostegodon Matsumoto MS., in Osborn, 1923; Matsumoto, 1924, p. 325.
GENERIC CHARACTERS.—(Schlesinger, 1917, p. 115, footnote): “Ich schlage fiir M. latidens, das sich durch seine kurze Symphyse von dem Subgenus Bunolophodon, durch seinen Molarenbau von Dibunodon 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.”’
Osborn, 1926: (1) Six! species of Stegolophodon, namely, Mastodon latidens, the more primitive M. cautleyi, and the more progressive M. stegodontoides; also S. cautleyt progressus, S. nathotensis, and Mastodon (Bunolophodon) longirostre Kaup forma sublatidens. As defined by the teeth, these species have in common the following generic characters: (2) Lophs as in Mastodon and Zygolophodon, tendency to form from four to six transversely arranged cones and conelets (conelets somewhat irregular) and to con- solidate into ridge-crests; (3) molar pattern transitional between the Zygolophodon type and the Stegodon type; (4) ridge-crest formula known in Stegolophodon latidens as follows, Dp 3° Dp 4+ M 1 ““*** M 2422 M3 i=. [Anterior ridge-crests with persistent median sulcus (see Vol. I, p. 700).—Kditor.]
This genus resembles Trilophodon and Tetralophodon in the retention of a broad enamel band on the straight superior incisive tusks (Figs. 725 and 727); it differs widely from T'rilophodon in the presence of four ridge-crests on the intemediate molars (Fig. 726); it also differs from T'etralophodon in the absence of trefoils and in the progressive tendency of the cones and conelets to form regular transverse ridge-crests surmounted by regular conelets, as seen in the genotypic species Mastodon [= Stegolophodon|] latidens, also in Mastodon | = Stegolophodon| stegodontoides.
Four of the known species are provisionally distinguished as follows:
Middle Pliocene Lower Pliocene? Middle(?) Pliocene Upper Pliocene Stegolophodon cautleyi Stegolophodon latidens Stegolophodon sublatidens Stegolophodon stegodontoides M2* M3 M2425 M34 (2)M3% M3 4-5 conelets on each ridge- 4-5 conelets on each ridge- 4-5 conelets on fourth, 5-6 conelets on each ridge- crest, very irregular. crest, more regular. fifth, and sixth ridge-crests. crest, very regular. Posterior
Posterior ridge-crests arched ridge-crests arched or convexo- or slightly convexo-concave. concave. Stego(lopho)don (see Pohlig, 1888, p. 252). By strict rules this term may be regarded as a nomen nudum and should not stand in the way of Schlesinger’s excellent name Stegolophodon.
Prostegodon. (1) For the first printed use of the name Prostegodon, see Osborn, 1923.601, p. 2: “‘Prostegodon, new genus, Matsumoto. Ina letter from Dr. H. Matsumoto, dated November 20, 1922, from Sendai, Japan, he writes: ‘In my report just in preparation on the Japanese ‘Mastodonts,’ I follow you to refer “Mastodon” latidens to the genus Stegodon, creating however a subgenus Prostegodon for it. Prostegodon is the primitive representative of the Stegodon-phylum, representing half bunomastodontine and half stegodontine dental characters. Schlosser’s opinion, that Prostegodon might be ? a descendant of “Mastodon” turicens, does not appear to be correct at alll? Genotypic species Mastodon latidens Clift. This genus should be credited to Doctor Matsumoto.”
‘Seven species including Stegolophodon lydekkeri described in Vol. I, p. 700, of the present Memoir.—Editor.| *[See page 824 above where it is stated that S. latidens is limited to the upper Irrawaddy beds (Lower Pleistocene).—Editor. |
840 OSBORN: THE PROBOSCIDEA
(2) In 1924 Matsumoto’s report on the Japanese Mastodonts, referred to above, appeared in the Journal of the Geological Society of Tokyo, Volume XXXI, in which he defined the genus Prostegodon (p. 325). This,
however, was published in the Japanese language.
(3) In 1926 Matsumoto published his English text on this genus (““On Two New Mastodonts and an Arche-
”)
typal Stegodont of Japan,’’ 1926, p. 9), from which the following is a direct quotation:
Skull and mandible only imperfectly known, brevirostral. Lower incisor-tusks might be absent, or abortive if present at all. Intermediate molars four- or five-ridged, last molars five- or six-ridged. Grinders essentially lophodont, though their first and second ridges may show a slight tendency of bunodonty and of trefoil pattern of cusps; mesial longitudinal cleft evident; inner and outer cusps opposite, instead of being alternate; valleys widely open, free of cement.
(4) Thus Prostegodon Matsumoto-Osborn, 1923, 1924, 1926, becomes a synonym of Stegolophodon Schlesinger , 1917.
(5) The genus Parastegodon Matsumoto, 1924 (1924.2), founded upon the genotypic species Hlephas (Prostego- don) aurore Matsumoto (Fig. 709), also probably [in part] becomes a synonym of Stegodon, because as shown below the genotypic species 2. (P.) aurore {= Stegodon aurore of the present Memoir] is somewhat more primitive than Stegodon airadwana Martin of the Middle(?) Pleistocene and quite distinct from H. [Archidiskodon| planifrons. The name Parastegodon was originally published by Matsumoto in 1924, pp. 256, 257: Parastegodon gen. nov. = Stegodon mindanensis-Elephas aurore group; type Elephas aurore; it was subsequently cited by Matsumoto (1926.1, p. 1). Elephas aurore was originally figured in 1918, Pl. xx, as reproduced in figure 709 of the present Memoir. From the isolated type second superior grinder of the right side, r.M’, it was difficult to determine whether this was a progressive Stegodon like S. airdwana or whether it was transitional to a