Ar = Sans ‘ 741.2 | aed 81 ‘ANNALS OF THE NEW YORK ACADEMY OF SCIENCES SCA | Be Vol. XXVI, pp. 215-315 Editor, Epmunp Otis. Hovey “REVIEW OF THE PLEISTOCENE OF EUROPE, ASIA AND NORTHERN AFRICA BY Henry Farrrietp Osporn NEW YORK PUBLISHED BY THE ACADEMY 30 Juty, 1915 THE NEW YORK ACADEMY OF SCIENCES — ) (Lyceum or Naturat History, 1817-1876) OFFICERS, 1915 President—GrorGr FREDERICK Kunz, 601 West 110th Street Vice-Presidents—CHARLES Be BERKEY, RayMonpD C. OsBurn, CHARLES BASKERVILLE, CLARK WISSLER Corresponding Secretary—Henry E. Crampton, American Museum Recording Secretary—Epmunp Otis Hovey, American Museum “ Treasurer—Huxnry J. Cocturan, 389 Fifth Avenue Inbrarian—RaLpeH W. Tower, American Museum Editor—Epmunp Orts Hovey, American Museum SECTION OF GEOLOGY AND MINERALOGY Chairman—CHARLES DP, BerKey, Columbia University Secretary—D. W. JoHNson, Columbia University . | SHCTION OF BIOLOGY Chairman—RayMonp ©. OspurN, 557 West 124th Street Secretary—WiLL1AM K. Grecory, American Museum SECTION OF ASTRONOMY, PHYSICS AND CHEMISTRY Chairman—Cuartes BASKERVILLE, College of the City of New York Secretary—Ernest EH. Smiru, 50 Hast 41st Street SECTION OF. ANTHROPOLOGY AND PSYCHOLOGY Chairman—CLark WISSLER, American Museum Secretary—Rosert H. Lowiz, American Museum The sessions of the Academy are held on Monday evenings at 8:15 o'clock from October to May, inclusive, at the American Museum of Natural History, 77th Street and Central Park, West. [ANNALS N. Y¥. Acap. Scr., Vol. XXVI, pp. 215-315. 30 July, 1915] REVIEW OF THE PLEISTOCENE OF EUROPE, ASIA AND es NORTHERN AFRICA? By Henry FAarrriretD OsBorn (Presented by title before the Academy, 12 April, 1975) CONTENTS Page MIMO CMIELLOM aera se iy ses = eraieteusieacleletsnene) RedoodebugnE@mocacwodunogandcc ot 217 Sevenuhstaunall phase——-OwWatierm aryiayecuas 4onnc00gbosbacboKdGoGs OR Sern atc 60.0 255 Hioraviot the irsteinterclacral periodic crise nett eetee 256. Mammals tof) theshrst: Imterclieiallestacee sirens rciieneleyeneleeneneneiene neal 256. 125-4 OVC] Koy ols eae eM eREa o eeara enone or meoramri oN tetas gioco. 00 257 SHRP VV ISS 5 FeO aE BS HAA FOU eyed oa) ete NO 25% GaN el aA (Se PL) MUCK ORNT ST Nn None Part ak CRRA LS ao oc 258 Minsteinterclacial (oie iran Cen accomere nile seine een eee 260 Southern elephant (Hlephas meridionalis) ...........0.002000008- 261 ELI pPPOPO CAAT A wee ine eve ouaveavateuoio ele menaneeas PARENT AON bt ne UREN Ah 0.15 263 SA DERE PCOOUIS eee rs a ase aay erin Bae ae cana Dele Ves Pie Reape 2265) MOOSE) CATCES ee 5 Stier Se Bowel pela Sie 2 ale eae nee: ee le ACO 264 Problematic evidence of man................... eee SRT ota 5:6 o 264 EDOM bn AU KERN AY=Y0) Li; Den men MU ON hanera yea) NUE Lt MAMI HE ee Ee so Oe 265 IPILNECANtNTODUS CLRECUUS» aoe wee ee 1S RO Siarses ae 266 Life of the Mediterranean islands.............. {ioaoay ead, 2 cee ee 267 Second Glacial stage—-Saxonian, Mindel, Kansan....................0-0- 268 Second Interglacial stage—Mindel-Riss.............. TELE ee ool 269 Moisture followed by aridity................... Rae rae Aas ciSic- O10. 3 269 CO) Dia aeh psa icine Roget a Abeea tau CEPA NUR CRC Nw uae ae aC ADE rN Oe oo oo 270 a4 Ee yea a ce2 keane RAO Eels ae Al an ern Ne NR AME SIE DI a ENN eS Go 0 271 SUIT VAV ATS Sree aes Ce Ue fe al ck nn Ul lg at aca A ore Ors ae Zl 1 Dp. 78 KCI KO OTS rr m saeey ce oe UR aie naan em en REMAN Sunt o Gla dao 6 272 aN il AYES SIRE SI RR a AUPE NE a Noe RN eR GS JESS SER a eee 272 LLOMONVCTCELO CHO CHSTSs al ec eee ie NERO es aut Sa chatc 274 Chellean culture and ancient Interglacial fauna..................--.. 276 Fauna of the Pyrenees, Cantabrian Alps, Spain and Portugal........ 276 Second and third Glacial and Interglacial epochs....:.............-.6+--- 207 AQHA ONIENMO IRAN es 6 a ddoocodanacbodoocoboe. deine ter uatene aoe Rota OIRO PACT Old elephant (Hlephas antiquus)............ HE RIP e Eee ch W185, 5;0 (5-0 PACTS FUMIO CER OSES Ls Sie eS RE erm reee Se ah sn a eRe rey a moat 278 RO VATIOES eos clan A US Sa oh Shee Vm eee ee Coa ah aaa Ae ere OaQcrey cA a ane a De ea 279 Mecaceros ciganteussacim ade eee oe MMM ee OIoeb.O O.0 0 280 Hap Minekorereda deen eee nese nse Sos AOR IT ee eels 280 N axe boo eye y armen NMA ERIN R Apc Ee oi RL IH on @ Davee to CITE COG 280 Garni VOres oe Veen Nance PIO ae MEUM aie Se SRE ee a ere ee 281 third (‘Glacial sstase—lilinolam. Polamd iam iRiSsh act suelerwecis lets eicielaretelevenenel= 283 4 Ravieeh cea teri ante pnaalal ae Ala ike eeu aia S Dale Bic rors ah ealaaearcuetenstelere kere 283 Third Interglacial stage—Riss-Wtirm, Sangamon...............-+-+-+++0:: 283 Climatic changes during the third Interglacial stage......... Rear ec eASy: | 2 Kohat: RN etna aL aati one LRTI Anais Mn EW Urat A a Gig Ab\o-a a0 6 0 0 285 TAMIA S55 LS os Ral) Shae Ree RR Ree re Ge a ino ee toe) act i ic ea a 286 Piltdown man, Hoanthropus dawsoni............6 slaistors modaiatiereke nee eee 286 Bre-@hellean and -Chelleamtaumalis oes ce cee eile eee 287 Chellean culture on the “low terraces”...............-+++----- 288 Grays-Thurrock, Ilford (Hssex, Bngland)................+.---++: 288 OSBORN, REVIEW OF THE PLEISTOCENE als Page ANCHeUeAMEXGUIEUTE) LAs: eee n re ass: ous ieensr oe casmeba ms cc ramen a reiomrs iets ioccbebenare 289 AWFAUIETUDERS EUS Chics ctsretenen siebeus pens tetots meerausie eae tetiters Sites frescos oceperace al Gta man slattelele 289 (Cavell SSPE Giese WGC SLO ROIS CTE eNO SEENON els «cS MERN re ana chin ue aw 289 Krapina Neanderthaloid race.............--. ats oS ete a cpevar anand eneaiilns 290 Mousterian, culture; temperate: fawieie se ces ce eice eiclele sv etelclere seers 290 Tourth Glacial stage—Wtirm, Mecklenburgian, Wisconsin......... cesta ene 291 Becinnines-of the remdeenr amd (cave period. 2 4. see see es ee elo e eoele)« 291 Periodeot che, fnaleclacial maximum teers cic ieee ee ete cere ee 291 Fauna of the fourth glacial stage........... aliaen aiavabasianen real ai Sataeet te 292 Mp PEt Sans vO LP MANE Nerve een apy si ey cen tals ears ROR a CR Peiarg tee 294 Mousterian Paleolithic culture.............. EERE BO AS 294 INGA d enbinaleracestraramtete siyeraeveistere classe ener ira eU PM Ca et evra ttLy crateecrene 294. Postglacial stage—continuation of upper Paleolithic, reindeer or cave FOVSTENCOYD Foe Sean hagtS RN Src cee roe Pn aE inst rar rei) AERA SAM: NUNS TNAB 296 COPED AAA TS ces cee ee eT TIS NG IEE es ies tate Ia URIS A OH UTA AE 296 Uppers LaleoMchics tourvor tive human ralceSiyaeescicve slats lara terest ale 297 Aurignacian, first upper Paleeolithic culture stage................ 298 SolutreansssecondeemliGuresstagseemiay sec cessor tele ia ice eretehe sie 298 Magdalenian, third culture stage and fauna..................... 298 OS CST ACTA Sa AUT ls repey mireyretayete rete eea Sie eel Tone al Se PTS es eee, GST cares Lg itreT MANO: Aaah 299 ScehweiZersbilde cawiereiccee nels rors ie Se ieee sao oil KeSSeCrlOcGh i Gaviesiewicieic cee i evenelavel d avelecene stele cas oes UN SU RIN RU ATA Is DeaLara cy 302 Migrations of the large mammals of the fourth glacial and post- CEUIEWENEE IL. OYE) AKG Ss rcpt SCONCE RRO ie CICTER CHER SOI U TT ETHIE NRE eee NON EA AUS aan acy 303 AY Meh aa AVOYPLOYS See tcl GROSS CCST NRC SISO CL CRUG TIRE ie OC bai ay MMR 304 VVOOL liye TIMO COTO SHAT scare tre eaueia Ne ere Saas VST ae aes EEE UME 305 HT ASTIO THEO tees esc crsea ayetctitetciate stores tole wire fer er onan eeeereoiee ratte RIOR ale AA 307 HOrsSesxots they Pleistocene@nn nice cic one PARE ELD AEN SORE R E Ie 308 ETS each a ci a tional sick oy cat are eure ons Siena SCN Rabat Tae MSO eA var sO see ire Fae Ae 310 Transition to the European forest stage......... HRA Butea neth Ae OSA Te 311 Micration of the funda fauna. os eaoe ss ce ces MEATS ey Min ntevenad ere eae: sere 311 Retreator the steppestaumae tere oversea e cilia ireraticls epee aro staratatius 311 Survival of forest and meadow fauna........... Eee ict en cee ten 312 Azilian-Tardenoisian, final upper Paleeolithic culture............ 313 HOLES eran Ayres wee oc ss eV estos Aosta ac sraaeeet ian PE ES Pea aT ates UDA NA 313 INTRODUCTION We observe that the Upper Tertiary closes with a Pliocene northern world rich with life, a world replete with Asiatic and African influence. The Tertiary Period is followed by the Quaternary, or Age of Man, a time of transition and of vast extinctions in Hurope and North America through natural causes, as well as of the geographic redistribution of life and establishment of the modern zodgeographic regions. Toward the close of the Quaternary Period man becomes the “destroying angel” and very nearly completes the havoc which Nature has begun. 218 ANNALS NEW YORK ACADEMY OF SCIENCES C®NOZOIC QWURNKAANIAT GosdasocduooudgduGod Age of Man MOT EUAIV its 5 eoeteeue erebenerse octoton eee Age of Mammals We thus enter a new Cenozoic faunal phase, the Seventh. When its transitions are complete the world wears an entirely new and somewhat impoverished aspect: the North has banished all the chief southerly forms and established the five modern zodlogical regions of the Old and New Worlds, namely: Palearctic, Nearctic, Oriental, Ethiopian, Neo- tropical. SEVENTH FAUNAL PHASE—QUATERNARY IN THE NORTHERN HEMISPHERE THE GLACIAL EPOCH. VERY GRADUAL EXTINCTION OR EXPULSION OF SOUTHERN TYPES OF AFRICAN, SOUTH ASIATIC AND SOUTH AMERICAN ORIGIN FROM THE NORTHERN, OR HOLARCTIC REALM. FIRST APPEARANCE IN CENTRAL EUROPE AND NORTH AMERICA OF THE CIRCUMPOLAR TUNDRA FAUNA, IN EUROPE OF THE STEPPE FAUNA. IN NORTH AMERICA EXTINCTION OF THE REMAINING LARGE ENDEMIC QUADRUPEDS. THIRD AND FINAL MOD- ERNIZATION OF EUROPE AND NORTH AMERICA BY THE HARDY FOREST, MEADOW AND MOUNTAIN RUMINANTS AND THE CARNIVORKES. The grand geologic divisions of the Quaternary in the New and Old ' Worlds are the same, namely, beginning with the Pleistocene and closing with the Holocene. times. Domestication. I. PLEISTOCENE, or GLACIAL HPocH. 3. PosTeLactaL. Mammals of tundra and steppe type gradually disappearing or retreating. Mammals of existing north tem- perate type multiplying. 2. GLACIAL. Period of successive glacial advances and interglacial retreats. Mammals of extinct and existing species com- mingled. 1. PREGLACIAL. Period of the lowering of temperature in the northern hemisphere and modification of plant and animal L life. Ie HOLOcENE, or RecENT HrocH. Mammals of prehistoric and recent QUATERNARY MEANS OF ESTABLISHING THE TIME DIVISIONS OF THE QUATERNARY The fluctuations of climate and of the plant and animal life of the Pleistocene are so numerous, so widespread, and so profound that it seems best to introduce the subject by a review of the great time divisions OSBORN, REVIAW ChE THE PLEISTOCEN L 219 together with some discussion as to the period when we should consider that the Quaternary proper begins. The fullness and precision of Ku- ropean faunistic Investigation is in very strong contrast to the prelim- inary results of American work, and in no other period may we anticipate more weighty inductions from correlation between the history of the Old and New Worlds. It is absolutely clear that a final and positive time scale and subdivision of the early Age of Man are not far distant and that the vast labors of European and American geologists, botanists, zoodlogists, paleontologists and anthropologists will finally be rewarded with a harmonious theory of all the phenomena of the Quaternary Period, the determination of the chronology of the various races and an approximate estimate of the duration of the entire Quaternary Period itself. The reader will observe that this correlation, derived from at least five distinct branches of natural science, is based on evidence of four kinds. 1. Geological: glacial deposits and erosions, which furnish the chief data for estimates of time. 2. Botanical: plant deposits, alternations of northern, arctic, steppe, temperate and southern floras, which furnish the chief data for estimates of tem- perature. 5. Palzeozoélogical: evolution and extinction of mammal and bird life, which furnish the chief divisions of the Quaternary time scale and afford sup- plementary knowledge of conditions of moisture, temperature and foresta- tion. . Anthropological: the successive stages of human culture and implements, the skeletal remains of man, which combined furnish the minor sub- divisions and correlations of Quaternary time. 6 pb PLIOCENE AND PLEISTOCENE LIFE oF AstA AND NortH AFRICA LIFE OF ASTA The region of the rich Tertiary flood plains of India? was one of the main sources of the large mammals which wandered into northern Africa and southern Europe in Pleistocene times; in other words, the large mammals—the elephants, the rhinoceroses, the hippopotami—were all in- vading forms from Asia and Africa. The relations between these three geographic regions are, in fact, so close that they might be embraced in a single zodgeographic realm were it not that throughout the Pleistocene the forests and meadows of southern Europe also maintained a northern Hurasiatic fauna which is entirely absent from southern Asia. >See p. 323, English edition of “The Age of Mammals.” aS 7 i ANNALS NEW YORK ACADEMY OF SCIENC 220 Ghee a Oiee 6 Sia Baines aye ewe ee QUdIOL IN jeddy) “uemuog 13aM0T auad -Oly teddy “(tue xg jo esvys) ueguog seddQ QUIIO![q JSIMO'T 10 uviuog ysow.edd () aUusd0I[q JOMO'T Q8Uu00 “og teddy 0} 2a[ppry ausv01[q ysoutsadd -) 000 ‘F 00g 00¢'6 -00¢'8 000° 01-00¢'6 00& ‘TI-000' OT 000 ‘FI-00G" IT 000 ‘91-000 FT SOON GeO 8th G Cas OL suozLioy uvedoing SHIPBATS JO 9S¥q VAOGB JOT WoO Oo 0 -O-O Ot) 0800 0 0b teb00 0 OF IUNGDILOT ‘xisayOL{NU) PIOFBIIS IsIVl ‘odofeJUB Vssiel “wnorp Ww wnmsayroh ‘thaowuay ‘wor.wddry IWASSOLYOS “JV snr posayyov ue DID wayyoaph, woliovydwuy — ‘nuaslio iT “wnuowndday vnualiywg ‘sptoy -JBILS OSIVl PUB WNWUAYLOpHIIAH ‘Sodoy -9JUB DULIOdISdITS a81Bl ‘snsa00b0.47, ‘(9D L908) snupjododdiy ‘sniunzod -OIiMA PY “UDP. SNK ‘(BdTBOS) UOPOISVT ‘UOLDAM “ha yypoph) = UWIAAOYYD.190 ‘UOPODIALS ‘SngvDbn4409 PUB sIswaIg -plund U0pojagn.4a,, ‘VYousnYy UWOpogsn fi Sy Ie AL 1S @[PPI JO eseg “** UOZLMOY LSB N — SHTTBANTS O] PPL uOZ “L104 UBY Id YOU “*'*spaq aBpueyg GIGL “Wiig soqojdaT ‘soquuay ‘(yuepungqe ) ) snumpododdiyy ‘tsnhiyoddryeA ‘snip -yodoohvayy ‘wowwddiy ‘sisuaypars Wop -O18SD Jy ‘suotfiqwuog pus Wy4Jyo wopotoly “** TOZLLOY JOLT, soquula HH ‘s0qgojda'T ; “ sypemig aeddq ‘uo.wddry ‘suosjvunid snydariy ‘Uuopobaiy ** uoziaoy aofurg snjoaiing ‘sog ‘snjawnyg ‘“vsauoajn{ sng ‘snnbm ‘sniuysiyoid snurvyso1a0vuy QU0Z 9}B19 ‘snowpnshy spydargy ‘nsaunh wuopobazy | -Wo[suoo Jeprnog |} S[ISSOO, OLST1ogOVIBYD UOLVOYISSB[D “‘pypuy fo spsodoq Wn) g-poo,y inysa gt, oY, fo Woypoyissp]Q—* | ATAV TL, 221 OSBORN, REVIEW OF THE PLEISTOCENE ‘9u90051[9 edd Q ‘uviuejinby sddq JO UvI[BsIpIng JaMO'T O00 0°08 So O08 O80 oD 050 OO O00 0-080 sUs00IJ JOMOT ‘uvTy -OAJOH «0 «UR ITRSIDING UBIUOJOT, 0F UBIJ@ATOFT 9U90 “OT O[PPHAL *( ear0ur -Ig JO 9dB4s) ueIuO VIO], (suopnes) JUIBS JO 96x48) URTZBUTARG SU9DOTIA, [PPTL ‘uBIyeueg ysoursedd o00'T ‘ssouyoryy CCC PO) 000‘8 ‘XB ‘SSoUyOIYy °XB I ese er eres eee 004° [-eseq 002 ‘8-002 ‘T (000 ‘F-002'¢ WNIVPUL WNVWIYLOIANDD YQ ‘SUS -uayjong uopo,nwjay, ‘ypsofunjq sD120 “0910, “Whiway wap “Wynosndaia wop -O1SDUNWAH ‘snunoii{p pue snazunbr6 SNPONYIDLT ‘aSUat{ONg wWiNilaYyLOIDLYyQUP SnUuDdIA{D snpohyonig ‘asuabunlyaqw $04990910,[, ‘asuai6ng OTE S OST IEDC COU GOOD GD slinan aod sees s*Sureured JuB[d pue wofpwm ywQnKy DNDILI “JO LhAIWOD VAN ‘OsUuaIpws wWwniwaYyJORH ‘uUopoznw)d,y, ‘Upsofun)g Lhiounuay ‘sdooqohyA ‘snp -OhYODAG “ABA asualDh wnrisay way ‘SIUOLpUDd PUB stapYsSNnHuy UOP0]IQD99,], snaay qd “Oh ‘synsdossig ‘ajnbojdn zy ‘snazunb Ww Ye Uollavyduy ‘sapro1say,ooDsyzUn IUNQDILO GY ‘vhsawUwonndorig ‘ehsay Opn) ‘Sodojajyue jpeurs pure sn.vad ODDMOLT ‘snzvsnd ehaowunuay ‘sng ‘UWOpOLUsvy ‘asuaipurs “yo WNWMOY LON FT “IVA “dS pur svyJNUusOwonu UOPOJIQD.149, J, (sepoeds rflurmgo TIMOT OT} JO Jsour ATquqoad) snoayqed DUS ‘nuMhTT ‘snovpiajynd uohaydup ‘HNAO OL DMV) ‘SNMOOODDM ON ‘LRLODID I “UNIMOYIORFL “Sng “(a0aRdS) worwnddr yy WNIMOYLOIDAY IU Geo20 0 ‘Spoq Tyong see eee BuRpIny Teysseq, , LO ILIN GL IOMO'T T[NBSB SY 10 satan, dodd gq uozloy IVyYyouByL Laao'T uoz -140Y 1LULY OQ tao] u0Z oy ify tedd Le : | Seo OS SED UTA: “** SHIPBAIG 1dM0'T 999 ANNALS NEW YORK ACADEMY OF SCIENCES The precise researches of Pilgrim,* published subsequent to the corre- lation proposed by the author in 1910 in the Pliocene chapter of the “Age of Maminals,”* have resulted in a new classification and correlation of the Tertiary flood-plain deposits of India which are of the utmost im- portance and interest to students both of phylogeny and of geographic distribution and migration. In geologic time the Indian series extends from the Bugti beds, which are of Aquitanian or Upper Oligocene age, to the uppermost Pliocene of the Upper Siwaliks. The correlation with the successive intermediate phases of European life appears to be quite close. As regards the origin of the Proboscidea, the author discovers in the Upper Oligocene of India animals which he believes resemble the Lower Oligocene Palwomastodon as well as the Maritherium of the Fayim de- posits of northern Egypt. These animals are referred to respectively as Hemimastodon and Meritherium. This discovery would favor the hy- pothesis that the Proboscidea may have originated in southern Asia rather than in Africa. Pilgrim believes that the Mastodon cautleyi of the Upper Miocene of Perim Island gave rise to the Stegodon types of the uppermost Miocene of Pikermi age, namely, to S. cliftt and S. bom- bifrons, from which originated the genus Hlephas which first appears in the dominant type Hlephas planifrons of Middle to Upper Phocene times in strata 3,000 feet above those in which the earhest forms of Stegodon occur. The species #. planifrons is especially important because it has re- cently been recognized by Schlesinger® in the Pliocene “Belvederschot- ter” north of Dobermannsdorf near Vienna. The horizon is regarded as of Middle Pliocene or even earlier age. Pavlow® has also recorded the occurrence of H. planifrons from beds in Bessarabia which are regarded as of Lower Pliocene age. In the Upper Pliocene of Europe occurs Elephas meridionalis which is regarded as a descendant of 1. planifrons, while in the Upper Pliocene of India occurs the Hlephas hysudricus, which Pohlg considers as a geographic variety of the Huropean #. merid- ionalis. In the uppermost Phocene of India also occurs the Dicerorhinus platyrhinus, which is believed to be closely related to the D. etruscus of the Upper Pliocene of the Val d’Arno of Italy. Some authors mistakenly regard the “Altelephant” (7. antiquus) of 8 PILGRIM, Guy H.: “The Correlation of the Siwaliks with Mammal Horizons of Hu- rope.’”’ Records, Geol. Surv. India, Vol. xliii, Part 4, pp. 264-326, Pl. 26. 1913. +See Life of Southern Asia, pp. 323-332, ‘“‘Age of Mammals.” 5 SCHLESINGDR, PAUL: “Studien tiber die Stammesgeschichte der Proboscidier,’ Jahrb. d. k. Geol. Reichs., Vol. 62, pp. 87-182. Vienna, 1912. 6 Pavtow, MARIE: “Les éléphants posttertiaires de diverses localités en Russie,’ Ann. géol. et mineralog. de la Russie, vol. xi, pp. 171-174. Moscou, 1910. OSBORN, REVIEW OF THE PLEISTOCENE 293 a the European Pleistocene as related to H. hysudricus of India, but Pil- erim and Pohlig rightly compare H. antiquus with the Narbada elephant (EZ. namadicus), which first occurs in the Pleistocene of Asia. In fact, it is not known whether the phylum of H. antiquus, which is quite dis- tinct from that of #. planifrons-H. Meridionalis, originated in Asia or in Africa. To sum up, among the contributions of southern Asia to the Pliocene and Pleistocene fauna of Europe are the following: Elephas planifrons, entering Europe in the Pliocene, related to the Hlephas meridionalis, the southern elephant. Elephas hysudricus. Hippopotamus, H. javadicus, related to H. major. Bison sivalensis, the short-skulled bison, related to Bison priscus. The long-skulled Leptobos. related to the LZ. etruseus of the Val @Arno and Bos primigenius of the First Interglacial Stage. The Sumatran type of rhinoceros, Dicerorhinus platyrhinus, related to D. etruscus and D. merckii. The hyzenas, related to H. crocuta, the spotted hyzena, and A. striata, the striped hyzena. The horse. Equus sivalensis, related to the Arab, or desert type of Hurope. Among the mammals which did not find their way from Asia into western Hurope are the camels and the various giraffoids. The absence of the antelopine members of the Bovide is also a very characteristic feature of the Pleistocene of Hurope as contrasted with their abundance in Asia and their presence in diminished form and numbers in the Upper Pliocene of Hurope. LIFE OF NORTH AFRICA It would appear that in Lower Pleistocene times when there were broad land connections between Kurope and Africa the latter continent contributed to Europe some of its indigenous mammals and others which had been derived originally from Asia. It is natural to suppose that the hyzena and hippopotamus, now so characteristic of Africa, entered Hu- rope either from Asia or from the north African region. With these ani- mals may have come the lion (Felis leo) and the “old elephant” (#. antiquus), which is a primitive offshoot of the same stock that gave rise to the African elephant (Loxodon africanus). We observe that in Lower Pleistocene times north Africa is still dis- tinctively a part of the Ethiopian Region, closely connected with central and southern Africa in its fauna. Throughout the Lower Quaternary the fauna of north Africa is also closely related to that of Asia. More- over it has a number of species in common with the Quaternary fauna of 204 ANNALS NEW YORK ACADEMY OF SCIENCES Europe, including those noted below which came into Europe from Africa. The contrary theory of the relative geographic isolation of Africa and Europe in Quaternary times originated with Pomel’ as the result of his exhaustive review of the entire fauna of north Africa. He concludes that since the resemblances between the European and north African faunas are rare and often doubtful, the two continents were for long periods separated by the Mediterranean Sea and Straits of Gibraltar. ( C : — os ———— eS ere vl ee q Nh NLT E \ ) | 0 Ni 7 ATS q f° (lll nem, l i WUYy Nt za G Wu wily li, CCUM TITEL 7p erry CHA, oF, we, Lorman iy —_— SS» 7 PLEISTOCENE LUT Fie. 1.—Pleistocene, or Ice Age A period of maximum total elevation facilitating free migrations and invasions of life, culminating in the Glacial epoch, and followed by a prolonged depression. Portions of northern Burope and the coasts of North America greatly depressed. Then a period of reélevation. Rearranged after W. D. Matthew, 1908. Climate.—At the beginning of the Quaternary Period north Africa was characterized by abundant rainfall which led to the formation of great alluvial or flood-plain depositions. In the Barbary and Sahara regions the life was closely similar to the grand plateau life of equatorial Africa at the present time, including elephants, rhinoceroses, zebras, wild asses, giraffes, wild cattle, buffalo, antelopes, gazelles, gnus, elands, hippopotami, wart-hogs, lions and hyenas. The presence of these ani- ™PomEL, A.: “Les bléphants Quaternaires.’ Carte Géol. Algérie, Paléont. Monogr. Algiers, 1895. OSBORN, REVIEW OF THE PLEISTOCENE 995 mals is consistent with the climatic theory of subtropical temperature and alternate dry and rainy seasons. Various phenomena point to increasingly long periods of drought and progressive secular desiccation of this great region as the Pleistocene advanced, resulting in the partial extinction and partial migration of the great equatorial life into central and southern Africa. Eurasiatic Invasion.—At the close of the Quaternary the bear (Ursus), as a characteristic forest-dweller, requiring a moist climate, became ex- tinct, while the Eurasiatic deer, wild sheep, wild boar, smaller mammals of European type, survived and established for this region its present affinity with Europe and its Palearctic fauna. We must account for this northerly, or Eurasiatic, fauna of north Africa as having entered the continent during the latter part of the Pleistocene Epoch and as sur- viving in the forested regions of present and prehistoric times so as to unite northern Africa closely with modern and prehistoric Europe. North Africa thus becomes a part of the Palearctic Region. Thus in no region of the world have more profound changes occurred during and since Pleistocene times than in Africa north of the Sahara Desert. Sources of the Pleistocene African Life-——It is premature to attempt to ascertain the original sources of all the various members of this im- posing assemblage of mammals. There remains always a great element of doubt which can be eliminated only by the discovery of the complete Czenozoic history of Asia and Africa. It would appear probable from our previous studies that the several continents contributed to the remote original ancestry of the African fauna somewhat as follows: Africa or Asia, elephants and mastodons. Northern Eurasia, deer and bear, wild sheep, wild boar. Southern Eurasia, wild cattle and buffalo. North America or northern Eurasia, rhinoceroses, various Hquide, camels. The most comprehensive comparison of the fauna of Africa and Europe is that of Stromer,* in which the entire fauna of Europe and north Africa, including the Reptilia and Mammalia, is compared from Lower Hocene to Pleistocene times. This author observes? that during the middle period of the Tertiary the mammal fauna of southwestern Eu- rope, western Asia, India to China, partook of the tropical or subtropical ®STROMER, ERNST: “Uber die Bedeutung der fossilen Wirbeltiere Afrikas fiir die Tiergeographie.” Verhandl. d. Deutsch. Zoél. Gesellschaft, pp. 204-218. 1906. ®STroMeER, Ernst: “Die einstige Verbreitung afrikanischer Siiugetiere.”” Naturwis- senschaftliche Wochenschrift, N. F., X Band; der ganzen Reihe, XXVI Band, No. 51, pp. 814-816. Dec. 17, 1911. 226 ANNALS NEW YORK ACADEMY OF SCIENCES character of the African high plateau fauna, rich in antelopes, giraffes, zebra-like ancestors of the horse, elephants, rhinoceroses, hyenas and apes. Late in Diluvial times in Europe numerous representatives of what we now consider a tropical African fauna, including hippopotami, lions, hyenas and apes, were widely distributed. Asiatic and European Affinities——The total assemblage of the Pleisto- cene life of north Africa may be summarized as follows from Pomel: REE adHomocl ate Pleistocene) A noteworthy distinction between Mastodon (Early Pleistocene only) orth Africa and Hurope is the sur- Blephants (several species related vival in north Africa of the masto- both to Lorodon and to Elephas) dons throughout early Pleistocene Rhinoceroses (two species of the times; also of several species of hip- African, or Diceros type) Beira odueactana eases parions side by side on the plains of ipparions, Numidia with the early north Afri- amels ae (Libytherium, Giraffa) can horsest® or zebras. Both the Wild cattle (Bos), three species mastodons and the hipparions are Buffalo (Bubalus) absent in the Pleistocene of Europe. Dwarf abel Opes: gazelles, gnus, oryXx, We may now review thous Seen a north Africa itself in Pleistocene Wild boar (Sus) times. Six species of elephant Wart hogs (Phacocherus) occur, including the mastodons, the Lions (two cavern species) southern mammoth (HH. meridiona- Hiya =) (spotted and sued) lis), and the “old elephant” (1. an- Jackals (Canis aureus), India Marauce: (Ol northern one) tiquus). ‘The most characteristic Deer (of the Cervus type, one and widespread elephant (L. atlan- species ) ticus) belongs to the African sub- Bear (of the Helarctos group) genus Loxodon while differing from Wild sheep and goats (Ovis' palwo- the recent Atrican/elephaniaanel™ tragus, O. promaza) Pee : : 4 africanus) im several poimts. The latter species only occurs in the recent deposits of the latest prehistoric period. Similarly the two species of rhinoceros (D. mauritanicus, D. subiner- mis) resemble the modern African types, but there is nothing to indicate the presence either of the modern African “black” (D. bicornis) or “white” (D. simus) species. Among the Pleistocene horses, in addition to the surviving hipparions and the species (H. numidicus) related to the Val D’Arno type of Hu- rope, there is a third species (#. mauritanicus) which exhibits tooth characters of the recent zebra. Thus there is every reason to believe that 10 BouLb, M.: “Observations sur quelques Equidés Fossiles.’’ Bull. Soc. Géol. France, Ser. 3, vol. xxvii, pp. 531-542. 1899. OSBORN, REVIEW OF THE PLEISTOCENE 297 in Pleistocene times ancestors of the zebras, which are now confined to equatorial Africa, extended to the extreme north of the continent. To the same period belongs a wild ass very similar to the Ethiopian ass (#. asinus), an animal which survived in this region until exterminated by the Greeks and Romans, and is now confined to the highlands of Abyssinia. Among the Artiodactyla the presence of camels’? (C. thomasiw) in Paleolithic Pleistocene times and even in deposits of Neolithic age (C. dromedarius) is extraordinarily interesting. There is no evidence as to domestication. ‘The earlher of these two camels of ancient Libya had longer legs and was of heavier build than the dromedary. The rare re- mains of the Jater form, probably identical with the recent dromedary, may be those of a race which was already emigrating or becoming extinct. The presence of the camel is one of the most convincing proofs of con- nection of this fauna with that of the Upper Siwaliks of southern Asia, and thus of North America. Especially significant of Asiatic and Siwalik affinity are the Pleisto- cene cattle and buffaloes of north Africa, including contemporary species of Bos, all belonging to late Quaternary or to the Neolithic age, partly domesticated, and with remote resemblances to the Pleistocene cattle of France and Spain. Similar Asiatic affinity is found in the remains of a buffalo (Bubalus antiquus) allied to the existing Indian form; this was a powerful beast which presumably lived in herds, frequenting grassy plains and swampy districts, and in its presence here we seem to find con- firmation of what geology teaches us in regard to the dampness of the Quaternary climate. The disappearance of the buffalo from north Africa at the commencement of the Recent Period was no doubt due to the in- creasingly dry conditions, and partly to destruction by man. The great number and variety of antelopes is most astonishing in this region, which at present is inhabited only by the gazelles (Gazella), the hartebeests (Bubalis)- and addax antelopes (Addax). he Pleistocene fauna includes gnus (Connochetes), several species of Bubalis still rep- resented in the Barbary States, an aberrant form (Oreonagor), related to the nilgai of India, nine species of gazelles (Gazella), the oryx (Oryx), the nabor (Cervicapra redunca), several large elands (Oreas), such as now inhabit south Africa, as well as dwarf antelopes (Cephal- ophus). Beside these plains and desert types of ruminants, the hills were covered with wild sheep (Ovis palwotragus) very similar to the ex- isting Barbary sheep, as well as goats (Ovis promaza). 2 PomeEL, A.: “Caméliens et Cervidés,” Carte Géol. Algérie, Paléont. Monogr. Algiers, 1893. 998 ANNALS NEW YORK ACADEMY OF SCIENCES In the rivers there lived in early and later Pleistocene times a series of species of hippopotami (H. hipponensis, H. sirensis, H. icosiensis) leading to a form (H. annectens) related to the existing Nile hippopota- mus. ‘There are also two types of wild boar (Sus), and more abundant than these were the wart-hogs (Phacocherus) found in the caves and alluvial deposits of Barbary. Preying upon these Herbivora were lions, leopards and hyznas, which are compared by Pomel with Pleistocene cave forms of Europe. There are also jackals, wolves, the ichneumon and, possibly, a polecat. Fic. 2.—Skeleton of the Pleistocene pigmy hippopotamus of Madagascar, Hippopotamus madagascariensis, together with a skull of the recent hippopotamus, H. amphibius In the American Museum of Natural History. African-European Distribution.—Of this imposing list the following types occur both in Africa and in the Lower and Middle Pleistocene of Europe, the species being similar if not in some instances identical. Southern elephant (H. meridionalis). which is also found in Pliocene and early Pleistocene deposits of Europe. Hlephants similar to HE. antiqwus of Europe and its dwarf representatives in Malta and other Mediterranean islands are found in the Upper Pleistocene deposits of north Africa. Long-headed rhinoceroses. It would appear probable that the woolly rhi- noceros (D. antiquitatis) which is closely related to the “white” rhinoceros (D. simus) originated in Africa, but no animal resembling it has been discovered in the African Pleistocene. OSBORN, REVIEW OF THE PLEISTOCENE 299 One of the Pleistocene horses of north Africa (H. numidicus) is closely similar to the Upper Pliocene H. stenonis of Europe. With these animals may have come the lion (Felis leo) which was widely spread over southern Hurope. Hyenas. The striped (H. striata) and spotted hyzenas (H. crocuta) are com- mon to Hurope. Bears. The bear (Ursus lybicus) found fossil in Algeria seems to belong to the Helarctos group, possibly derived from the small U. etruscus of the European Pliocene and now represented by the Malayan sun bear. Machsrodonts. Recently (Stromer) sabre-tooth tigers have been discovered in Pleistocene Egypt. Primates. The primates are represented by the macaque (Macacus), not very different from the existing forms which frequent the region of the Straits of Gibraltar. In Pleistocene times the macaques ranged northward into southern France (Harlé). Suillines. Wild boar (Sus) may have affinities with the Pliocene types of Europe. With the exception of the above list, there is little in common between the large fauna of north Africa and that of Europe in Pleistocene times. AFRICA IN PALZOLITHIC AND NEOLITHIC TIMES. Giraffes very similar to the recent African giraffe (C. giraffa) have been found in mid-Pleistocene deposits associated with Paleolithic stone implements of the Chellean type.1?_ Industry’ similar to the Chellean but not necessarily of the same age is found in Africa from Egypt to the Cape. Giraffes are also depicted in rock drawings of Neolithic age in Algeria. , In Neolithic times there existed at least one species of deer, whereas at present there are two kinds of deer, the red and the alloy, in north Africa, both undoubted Hurasiatic migrants. The prehistoric men of the Barbary States apparently obtained and domesticated the horse, species of sheep and several dogs, and left many sketches of animals on the rocks of the region.*® PLEISTOCENE oR GLACIAL HrocH IN EUROPE After the establishment of the single glacial theory by Charpentier and Louis Agassiz (1836-1840), there gradually developed in Europe and 122 PaLLARy, P.: ‘Note sur la Girafe et le Chameau du Quaternaire Algérien.” Bull. Soc. Géol. France, Ser. 3, Vol. XXVIII, pp. 908-909. 1900. 1 OBERMAIER, HuGo: ‘‘Der Mensch der Vorzeit. Band I. of Der Mensch aller Zeiten.” Alleg. Verlags-Gesellsch. m. b. h. Berlin, Munich, Vienna, 1912. 14 See LYDEKKER, RICHARD: Deer of all Lands. The North African red deer (Cervus elaphus barbarus) is smaller than the European race. EHyidence on the range of the com- mon fallow deer (Cervus dama) in northwestern Africa is not very full. % See PoMBL, ’93, ’94, ’95, ’96, ’97, 798. 930 ANNALS NEW YORK ACADEMY OF SCIENCES America the hypothesis of several glacial advances of varying duration and severity alternating with interglacial temperate periods during which the ice retreated and conditions of climate prevailed which in some in- stances were even milder than the present in the same latitudes. As early as 1856 Morlot observed a relatively warm flora between two Swiss glacial deposits at Diirnten, and he subsequently advanced a theory of three glacial stages. James Geikie (1871-1894) developed the hypoth- esis of a succession of six glacial and five interglacial stages and climates. In 1883 Boule from his observations along the Mediterranean coast main- € ] Young Drift Limit —- Middle Drift Limit ——. Old Drift Limit ++++ Scale 1:5,000.000 0 2 SO 100 Km te a/+4,) 7 Fic. 3.—Glacial map of northern Germany and the Netherlands This map shows the drift and terminal moraines of Glaciations I-II, Scanian + Saz- onian (old drift), III Polandian= Riss (middle drift), IV Mecklenburgian = Wiirm (upper drift). After Leverett, 1910. tained that there is evidence of three great glacial advances, the first fall- ing within the close of the Pliocene Epoch, the second falling properly within the Pleistocene. he firm foundation of the quadruple theory in Europe was laid, however, by the researches of Penck and Briickner*® in the Alpine region, published in 1909. According to this classic work the entire Glacial Epoch is assigned to the Pleistocene or Quaternary Period. Its deposits include the entire “Diluvium” and “Drift” of earlier geolo- gists. All the river gravels, boulder-clays and moraines of the Glacial Epoch *¢ PENCK, ALBRECHT, and BrickNrR, HpouarD: Die Alpen im Wiszeitalter. III. Die Hiszeiten in den Sudalpen und im Bereich der Ostabdachung der Alpen. Leipzig, 1909. OSBORN, REVIEW OF THE PLEISTOCENE 231 are of later date than the marine Pliocene deposits of southern Europe. Before any of these glacial deposits occurred there was an elevation of the marine Pliocene strata along the southern Alpine borders from sea level to a height of from 300-500m.; there also occurred erosion of these marine strata by rivers. Thus in the valley of the Po there is a consid- erable time interval between the closing marine conditions of the Plio- cene and the opening Pleistocene conditions. In the valley of the Rhone also there is a marked interruption between the strata of the Pliocene and of the Glacial epochs, the latter overlying the strata recognized as Upper Pliocene, which in turn overlie the marine Pliocene.*” This interval between the Pliocene and Pleistocene corresponds with very important changes which occur in the mammalian life of Europe, namely, in the extinction of many characteristic Pliocene mammals, such as the anthropoid apes, the antelopes and the mastodons. 4 Sierra de Gredos Alps Mts ae, [S18 Atlas Mts Bere Mts. | seeel Ol = ! | pangs Germany Scandinavian Plareau Denmark t Se er Strait of Gibra/tar Garonne Phone North Skager Valley Valley Sea Fak PROFILE OF PAST AND PRESENT SNOWLINES AND CLACIERS OF EUROPE Fic. 4.—Theoretic snow levels during the Glacial Epoch Prepared under the direction of the author by Chester A. Reeds from data given by J. Geikie, Penck, Briichner, Leverett and Stieler’s Hand-Atlas, December, 1914. The traces of four different glacial advances and retreats observed around the northern slopes of the Alps by Penck and Briickner?® were fol- lowed with their “river drifts” and moraines down the Danube to the neighborhood of Vienna; they were found to be clearly marked in the re- gion of the upper Rhine and of the Rhone around Lyons, and distin- guishable both by the greater or the less extension of their borders and by the greater or less erosion which has occurred in the intervals between their successive depositions. These four advances were named _respec- tively the Giinz, the Mindel, the Riss and the Wiirm. As an instance of the disparity between the duration of these several glacial advances with the accompanying descent of the ice and snow line, the old moraines of the Riss or third glaciation form a girdle around the more recent Wurm or fourth glaciation, proving that the Riss was not Op. cit., pp. 654-655. ZOpy Cite Ds Ade 239 ANNALS NEW YORK ACADEMY OF SCIENCES only a more extensive glaciation but that the snow line was 100m. lower. It is also estimated that the climate of the Riss was one-twelfth more severe than that of the Wurm. In northern Germany only three great glacial advances are recorded, while still farther north, in Scandinavia, there was in a sense only one Glacial epoch, since the ice cap never retreated so far as to permit of interglacial deposits. ‘This is in accordance with the anthropological fact that only toward the close of Postglacial times does Scandinavia show traces of human habitation in the arrival of the Neolithic men; whereas in France and Germany there is evidence of human habitation as early as the Second and Third Interglacial Stages. In the meantime American geologists have also discovered similar proofs of four successive glacial advances and more temperate inter- glacial stages. The correlation of these conditions in the New and Old Worlds suggested by Penck, Chamberlin and others has recently been reviewed with great precision by Leverett,1® to whose work we shall fre- quently refer. The most recent results of geologic and anthropologic correlation with some original modifications are graphically presented in the accompanying diagram (Fig. 5) by the author and Reeds.”° The river terraces are of great importance both in geology and in an- thropology. In general the “high terraces” belong to the earlier glacia- tions and the “low terraces” to the latest. Thus the “high terraces” of the Alpine region belong to the Riss or glaciation III; in the valley of the Rhine the “high terrace” is visible near Basle; the “low terrace” of the Wiirm or glaciation IV occupies vast surfaces on the upper Rhine and contains a mammoth (FZ. primigenius) fauna. The “high terraces” in the Paris basin reach 30m. above the level of the Seine, while the “low terraces” are only 5m. above the level of the Seine and subject to floods ; the “high terraces” in the valley of the Seine contain the First Inter- glacial (H. meridionalis, H. stenonsis) fauna, while the “low terraces” of the Seine and of the Somme contain the Second and Third Interglacial fauna (LH. trogontheru, H. antiquus, and D. merckit). DURATION OF THE PLEISTOCENE The Pleistocene was estimated by the American geologist Dana (1874) to be equal to about one-fourth of the entire Cenozoic Era, 1. e., 700,000 years. By Ward (1885) and Williams (1895) it has been estimated at 19 LEVERETT, FRANK: ‘‘Comparison of North American and Huropean Glacial Deposits.” Zeitschr. f. Gletscherkunde, Vol. iv, pp. 241-316. 1910. 20 REEDS, CHESTER A.: Dr. Reeds has prepared the climatic curve from data furnished by Penck, Leverett, Taylor, Chamberlin, Salisbury, Geikie, Schmidt, Coleman and Osborn. Dated October, 1914. OSBORN, REVIEW OF THE PLEISTOCENE CORRELATION Of, CLIMATIC, RACIAL, CULTURE & LIFE STAGES (9/4 | PREHISTOR? VEOLITHIC POST GLACIAL 2DAUNSSN\ it} prrran-rammenos he ptee ——oREN ELLE | SE CENT FOREST, MEADOW, ALPINE: “Newer Loess’ ZGSCHNITZ/7S,! ; 1) | 2 MacoALEN/AN PALAEO- | GRO-MAGNON ZLLEEELES SSL BUM ER \ |, SURIGNACIAN._J_LATHIC._| Grimato | REINDEER PERIOD, ARCTIC IV. GLACIAL LA ae 23,000 YEARS WEANDERTHAL| TUNDRA, STEPPE , ALPINE WR, WISCONSIN yi eA:| | 4 MOUSTERIAN : FOREST, MEADOW Vawext Terraces’ E A) 2150000 YEARS COLD FAUNA pi ARRIVAL: STEPPE, TUNDRA, FAUNA iti r Satan cae a a aa 5 1 | eee | LOWER |” (KRAPINA), rAasT WARM. AFRICAN-ASIATIC 3.INTER- Alii P|” (cratnae Ps GLACIAL Allitii | |2cHeLLEAN LITHIC E.ANTIQUUS, HIPPOPOTAMUS: RISS -WURM itil 4 41/00000 YEARS D.MERCKI!, E.TROGONTHERI/ itt; ALSO FOREST, MEADOW SANGAMON Ait | | 1 PRE-CHELLEAN P/LTDOWN : Middle Loess 2Ai1\'\ | stre500 rears EURASIATIC FAUNA jj) Bi URGE ie Se ese easaie Zo AGN °6{/50000__» Z Le | ee WI. GLACIAL % ee COLD TUNDRA FAUNA RISS, POLANDIAN ZB Ze, 15S ) WOOLLY MAMMOTH & Middle Drift” ef! 7 AASASSS SSSA SS, 7 jl RH/NOCEROS. FIRST A LZ Hili! | | STEPPE & REINDEER nace UiNiue (eteagouo weak meet Nips t's iN Venn eu uih s NS oee = Wee GLACIAL ni MINDEL-RI liji! [9229000 , WARM AFRICAN: ASIATIC LRISS ZAI iii)! FAUNA HELVETIAN Hi YARMOUTH hij ‘i 10:250000 E.ANTIQUUS , E. TROGONTH- Long Warm tii! HEIDELBERG) py p. MERCKII, HIPPO- ifjl: Stage lijitg wh275000 , POTAMUS “Older Loess’ A i jij iii! /24300000 Gog he ce Sa igee a oP ea Wi ci lt eden Cac Hea ar aay ca i! 13+325000 Gj HA Mi A I!) “2 i 144350000 Lise Rican ae Wer eC Aarne el IL GLACIAL LA ys| FIRST COLD MINDEL , SAXONIAN 25 oe FAUNA on Hey rift A] 161400000 YEARS ZAG SA Aas ae pee ne ISINTER- 2A | FR WARM Oh GLACIAL I Rane? ah AFRICAN - ASIATIC. FAUNA ele Myf GUNZ-MINDEL ZM\''\i |etasqo00 HIPPOPOTAMUS CAN VE y YYj#; SA iN MACHAERODUS ZR Vsherg000 7 Khe RRA PaO REV TT ai L.GLACIAL 7h. COLD FORFa BED Go | GUNZ,SCANIAN ZA | THROPUS /NEBRASKAN ir oe 501 ae ae pee” (TRIN/L) , "Old Terraces A | ia ee eee | Br Ra Ie ON SNCs | Vidiya PLIOCENE PLIOCENE Zp epee WARM FOREST GLACIAL —|2zeel88| STONE CULTURES | HUMAN | STAGES OF MAMMALIAN © INTERGLACI/IAL AND COLD FAUNAS RACES AND PLANT. LIFE Fig. 5.—Divisions and contemporaneous cvents of the Glacial Epoch Prepared under the direction of the author by Chester A. Reeds from data observed and correlated by J. Geikie, Penck, Chamberlin, Salisbury, Leverett, Schmidt, Coleman and Osborn. October, 1914. Yo 4h LE MY OF SCIEN( / 7 vy) W YORK ACADE y u NALS NE AN 34 2 000°0% 000 “02 2UNx) *[BIOBLH) ISA | 000‘091-000‘00L [EPUL-2ZUNy ‘TeIoKR[oLezUy 4S 000° 02 [OpUL “[Blov[E) puooes ]] 000 ‘00F-000‘0FZ SSIY-[OPULpY “[Blov[sie}UT PUODY deBY S19qQ[eploH 000 ‘0Z ssly “Bovis PYLE TIT UBI[[9YOD-ed 000°001-000'09 uve[[eyO Tin A -ssiy ‘[eOovpsiajyUy pry uvolnetaVy UMULXB IL T qeorqod uafnnT UBIIOISNO [A 000 ‘02 WunA\ [BIovpy YWnoy AT UINWIXBIT JT] UBIOBUSLIN VY : : z JVIM AI UIYo P uvdIJN[OG uelua|epse [| 000 ‘FZ-000' 91 THAG 000‘FS 4 0000% UBITIZV ZLIN HOSE) [e10R [5380 OLY P[OON 000 *2 xavgd iladdog jo a8 y sainjwiedwey, laysty ‘sqvorjey Moug ‘saBvq4g [vloels.104u] se8¥)}g a1nj[np UeWN A SoPVWIIS GUILT, 9ATJLIOY 000 “8-000 ‘F Aylpluny puv sainjetodwe J, MOT ‘S9OUvAPY MOUgG ‘seSvIg [BI0B|yH aUnjoA, sry) UW poydopy sabnjiy [pion{hsazuy pup [DI9v)H JO SayDUltps yy GU J—T1 FTaVL OSBORN, REVIEW OF THE PLEISTOCENE - 9395 cne-third the entire Cenozoie, t. e., 1,000,000 years. If with Wallace we accept Croll’s theory and estimate, the last glacial advance would date back to the last period of great eccentricity of the earth’s orbit, namely, 200,000 years, but this we now consider excessive. The following figures show the variations of opinion on this subject and the two opposite tend- encies of greatly expanded or greatly abbreviated estimates of Pleistocene time: Lyell, “Antiquity of Man”..... 1863 800,000 years ipo nana eee cyet ewe cep ss econcnsicrecs wetshste 1893 100,000 “ Wialeottnaec cre cee soe 1893 400,000 ‘* SQUIER ES aR aoe neat tae 1900 400,000 “ IEEVIOS< Ros ee ew enc lanp Spier at ts 1909 520,000 to 840,000 years The very high estimate of 840,000 years made by an eminent and usually conservative authority such as Penck appears excessive unless we are to expand our estimates of Tertiary time (see p. 63) to 20,000,000 years and of the pre-Tertiary into hundreds of millions of years. All the arguments for the briefer estimates of Pleistocene time have recently been brought together by Wright.?1 Antiquity of Man.—Vast interest attaches to this duration problem in connection with the antiquity of man. In the calculations of Penck*’ the time since the Fourth or Wiirm glaciation has been used as a measure- unit to calculate the length of the previous glacial and interglacial periods. It is believed that since the climax of the Wiirm glaciation from 20,000 to 34,000 years have elapsed. Geologic, prehistoric and historic events since the close of the last glaciation make this estimate appear not excessive. In regard to the previous time intervals, the au- thor does not pretend to give an absolute age estimate, but simply a survey of the relative magnitude of the time periods with which we are déaling. The unit of measurement is Postglacial or post-Wiirm time which Penck** estimates at 20,000 years. On the basis of this estimate the time (520,000 years) covered by the whole Glacial Epoch is relatively distributed as follows: 21 WRIGHT, G. FREDERICK: The Ice Age in North America and its Bearings upon the Antiquity of Man. 8yvo. Bibliotheca Sacra Co. Oberlin, 1911. 2 PENCK, A.: “Das Alter des Menschengeschlechts.” Zeitschr. Ethnol., No. 3, pp. 390-407. 1908. ~ PENCK, A., and BricKNer, EpovuarD: ‘‘Die Alpen im Wiszeitalter. Dritter Band, Die Hiszeiten in den Sudalpen und im Bereich der Ostabdachung der Alpen.’”’ S8vo. Tauch- nitz, Leipzig, 1909. (pp. 1153-1176, “Chronologie d. Eiszeitalters in d. Alpen.’’ Penek.) 236 ANNALS NEW YORK ACADEMY OF SCIENCES Relative , Descent of duration Totals snow line. Units Years Years Meters Postglacial, post-Wtirm hemicycle... 1 20,000 20,000 tetas LW. OL AWiOR MAG EAGTAMTONS« sey e eiersieteeienne 1 20,000 40,000 1,200 3rd or Riss-Witirm Interglacial Stage. 3 60,000 100,000 seats PASOL RIGS GUACTATION sree iene hee 1 20,000 120,000 1,250 2nd or Mindel-Riss Interglacial Stage 12 240,000 360,000 Sie INL Gre Whaxpoyse, CAGWMMONTs 6oboonouucocbcos 1 20,000 380,000 1,300 ist or Gtinz-Mindel Interglacial Stage 5 100,000 480,000 Sagat It ore Grusps: (GHOACIPMIBKOIN. Gogg odooaodcoudb DoS 1 20,000 520,000 1,200 The three chief conclusions of Penck are as follows: 1. If the whole Ice Age extended over a period of 500,000 to 1,000,000 years, the Second very long warm Interglacial Stage, also known as the Mindel-Riss or Helvetian, is reckoned at more than 200,000 years, while the final relatively short interglacial stage, the Riss-Wiirm, is reckoned at 60,000-—100,000 years. 2. The duration of the Lower Paleolithic culture periods, the pre- Chellean, Chellean and Mousterian, would by this reckoning be much longer than that of the Upper Paleolithic culture periods, the Aurig- nacian, Solutrean and Magdalenian. Penck estimates that since the be- ginning of Magdalenian times 24,000 years may have elapsed and since its close about 16,000 years. 3. Compared with these prolonged Paleolithic divisions the Neolithic Stone and Metal periods have occupied an almost unappreciable length of time. If the beginning of the Neolithic lake dwellings is dated about 5,000 to 7,000 years ago we estimate that the beginning of the Copper Age in Europe dates back between 3,000 and 3,500 years; in Africa it is much more ancient. The human culture stages are arranged above not according to Penck but according to the more recent correlations of Obermaier, Breuil, Schmidt and others. GEOLOGIC AGE OF THE CULTURE STAGES The trend of Paleolithic research lately has been to draw all the human culture periods from the pre-Chellean to the Magdalenian closer together and to reduce the time assigned for their evolution. All the French authorities, led by Boule, Cartailhac, Breuil and Obermaier, are now agreed in assigning the earlier Paleolithic cultures, the pre-Chel- lean, Chellean and Acheulean, to the Third Interglacial Stage and not to the Second. Schmidt has also lately declared himself in favor of this view after a most exhaustive and valuable investigation of this problem. OSBORN. REVIEW OF THE PLEISTOCENE 937 A summary of the correlation presented in this chapter is embodied in the Table of Osborn and Reeds above. A summary of the very diverse opinions on this subject is embodied in the Table of Wiegers below. A yery strong reason for abbreviating our estimates of the period which has elapsed since the appearance of man of the pre-Chellean cul- ture stage in Europe is found in the relatively unchanged condition of the river valleys of the Somme in northern France and of the Vézére in Dordogne, in which the earliest human cultures occur. The Vézére has not materially changed since Acheulean times. The pre-Chellean, Chel- lean and Acheulean specimens found in the Somme valley are also con- nected with the present river system. Both on the Somme and the Marne the Chellean and pre-Chellean cultures occur on the “lowest ter- races.” Again, there is no faunal break between late Chellean and early Acheulean times nor between late Acheulean and early Mousterian times. The first great faunal break is that produced by the Fourth glaciation. In favor of Penck’s contention as to the earlier geologic age of the Chellean is the occurrence of pre-Chellean and Chellean paleoliths in association with a very primitive mammalian fauna such as is character- istic of Second Interglacial times. ELEVATION AND SUBSIDENCE OF LAND IN QUATERNARY TIMES The relations of the mammals of Europe with those of Asia on the east and Africa on the south were profoundly affected in Pleistocene times by the periods of elevation of the continental shelf, resulting in the creation - of new land connections which facilitated migration, or of subsidence which cut off and isolated many migrating forms from their centres of origin and dispersal. The maximum elevation, as represented in the accompanying diagram (Fig. 1), never occurred in all portions of the continent of Europe at the same time, because there were oscillations both on the northern and southern borders of Europe and Asia. The beginning of the Pleistocene Epoch is one of elevation and is re- markable for the broad land connections between Europe, Africa and Asia. It represents the last stage in that vast community of mam- malian life which during Pliocene times distinguished the entire region of Europe, Asia and Africa. ; The theoretical relation which elevation and subsidence respectively bear to the glacial and interglacial stages and phenomena is, broadly speaking, as follows: ELEVATION, emergence of the land from the sea, broad land connections facilitating migration, retreat of the ice caps, periods of erosion of the river valleys and formation of terraces. S ’ u MY OF SCIENCE Al 4 ACAD K > 7 YOR jW fi 4 NE ANNALS “ET6I ‘FET-90T1 ‘dd ‘| 139 ‘GF “Saqee “unig J ‘S187 -Qijor(ed ueap nz asteiuerpnig,, HWANITHZIIE, “LAUVHHONHOS ‘Suma 2 aa “SUADATA\ “SDOUVHHONKOS “SUADAIA “TANVYD ‘: TOISSnOSTG .eUsOp10Og 1ap UaTfeIspuny oeyost ‘SOSRIS OANJTNI OTGINORTRA puR d[so]oas Jo suo] vJeI1400 snoyABA Udg][Ayoul { waa[[e4O uda]NayO VW [BlOv[S19]UT IST MOIS) IIE useT[2qo Ude[Neyo V UdI[[P4D - UAB AL Udg[NIYOV :4[BM [Blov[siequy pug — [] UseMIYOY | Uatsaysnoypy so}B yy WdTL9}SNO IAL [BRD TIT asvyd ple Ay 8} OISSBUIOL) wogT|eyO ite Ay J Uelte{sno]y See UdTI9JSNO TA Berle Oy ae [BOB] Suejuy pag Was | tas $9489 : asvydueddays LESSEN NO) eee Ud1IOBUSLINW oy W Use14N[OS4LV [elze[sq nay J] Uellaysnoyy [BIZBLSy oH - UdTOBVUSLIN Y udeIJNTOY UdadjNJOS-sun ¢ uaaIN[OS UdLI9ISNO JA, UdlI9ISNO J] [BOB AT ]BIzBy.ojsog | “MZC JLO7Z-BPl[OK UdIUg[Vpsvyl tuni pes qng qiez-snjAouy { = sg ney a udlOvUSIINV udlOvUsIINVy : . Z > Q : yez-vuoyy | qrezuieyg exa8uny' UdIU9[BP9v [AL UdlUI[VPse JA] = venules U9pAINIOS ere EIGBL ‘S1aSOrM rakvy OL6L ‘yous ZIGL “‘Iprmyos ZLAL ‘IAlVULLaqG SUOISIAIC [BOISO[Oa4) suoyiny quasaffig fiq fibo1010n1) pun fibojonyoIspy {0 W0144D]AL40Q—TIT FISVL OSBORN, REVIEW OF THE PLEISTOCENE 939 SUBSIDENCE, submergence of the land and advance of the sea, land con- nections interrupted, advance of the glaciers, periods of deposition and filling of the valleys. Thus Boule and Geikie consider the glacial stages as mainly periods of continental subsidence and filling of the valleys, the interglacial as times of elevation and erosion of valleys and terraces. Penck estimates the elevation of southern Europe at 100m. in the beginning of Pleistocene times; he speaks of the elevation of the Alps during the Second Inter- glacial Stage. Geikie describes the southern half of the North Sea as dry land during the First Interglacial Stage traversed by a northern ex- tension of the River Rhine, while the approach of the Second Glacial Stage was heralded by a submergence of this area of the North Sea. Again (Geikie) during the Second Interglacial Stage the English Chan- nel and probably a large part of the North Sea became dry land. Finally (Pohlig) during the Third Interglacial Stage there was a period of continental elevation and a dry, cold steppe climate in western Europe. Consistent with this hypothesis is the deposition of loess during the Second and Third Interglacial Stages, also during the Postglacial Stage, because loess deposition is characteristic of dry and elevated continental climates with winds prevailing in one direction. Final Subsidence.—Consistent with this hypothesis also is the fact that general and local subsidence in the northern hemisphere was the chief feature of closing Pleistocene times or the very cold Postglacial Stage ; all the old continental connections which had been characteristic of the Tertiary were cut off; in the northwest the English Channel was formed, Great Britain became isolated from Europe, Ireland first lost its land connection with Wales and then with Scotland ; to the eastward the Medi- terranean Sea extended into the Aigean region and cut off the old land connection between Greece and Asia, which had so long served to connect Greece with the mammalian life of southern Asia. During a period of extreme subsidence, the Black Sea, the Caspian, the Sea of Aral formed a large single sheet of water known as the Hyrcanian Sea. In southern Asia similar subsidence and separation phenomena were in progress; the islands of the East Indies, Sumatra and Java were cut off from the Malay Peninsula. The separation of the Japan and Philippine archi- pelagos probably occurred in late Postglacial times. To the far north- east late in Pleistocene times Asia lost its connection with America, Bering Straits were reopened, and the so-called Nearctic region of North America was completely isolated from the Palearctic region of Hurasia after a long period of community and free intermigration of Holarctic life. 940 ANNALS NEW YORK ACADEMY OF SCIENCES Final Elevation—This extreme subsidence was followed in Jate Post- glacial times both in Europe and America by reélevation which gave the continents their present contours and climates. ALTERNATIONS OF CLIMATE AND FLORA Fluctuations of temperature and of moisture in Pleistocene times are indicated first by the advances and retreats of the ice caps, second by the , presence of arctic or temperate Mollusca in the coast waters, third by the variations in the flora in glacial and interglacial times, fourth by the alternate appearance of the northerly or southerly types of mammals and birds, fifth by the nature of the geologic depositions, sixth by the nature of the land Mollusca in the loess. Combining the evidence derived from these various sources the theoretic broad divisions of the climatic se- quence are as follows: (1) the cold and moist phases connected with the successive glacial advances and retreats; (2) the warm to temperate climates of the First and Second Interglacial Stages and first half of the Third Interglacial; (3) the dry and cold climate of the second half of the Third Interglacial Stage and early Postglacial times; (4) the damp and cool climate of late Postglacial times favorable to forests. The theoretic alternating conditions. of each complete glacial cycle are as follows :?4 Subsidence: Glacial Maximum: Tundra flora and fauna Glacial Retreat: Cool and moist forest flora and fauna Elevation : Interglacial : Dry conditions, flora and fauna. Steppe Glacial Advance: Cool forest fauna and flora Subsidence: Glacial Maximum: Cold tundra flora and fauna Low Glacial Stage Temperatures——Low temperatures during the pe- riods of glacial advance are attested both by the advent of northern Mol- iusca, marine and terrestrial, northern flora, and the repeated arrival in Europe of members of the cold fauna of the arctic tundras, including both the smaller and the larger mammals and the birds, as well as the cold fauna of the high, arid steppes of western Asia. Low temperatures are attested also in early Postglacial times during the great Aurignacian- Magdalenian art period by the heavy covering of hair indicated on all the animals depicted by the Upper Paleolithic artists. This hairy cov- ering coincides exactly with that of the extreme northern tundra types of reindeer, woolly rhinoceros (D. antiquitatis) and woolly mammoth (2. primigenius) found imbedded in the ice or frozen soil. *% Compare Wiist. OSBORN, REVIEW OF THE PLEISTOCENE oAT Moderate Estimates of Temperature-——TVhat the advancing glaciers alone do not constitute proof of very low temperatures is observed in Alaska, where very heavy snowfall or precipitation causes the accumula- tion of great glaciers, although the mean annual temperature of the glacier region is 40°-45° F. (4.44°-7.22° C.) as compared with that of northern Germany, 45°-50° F. (7.22°-10° C.), 4. e., from mouth of Rhine S. E. along source of northward flowing rivers, e. g., Elbe, Vistula, ete. Neumayr estimated that during the Ice Age there was a general lowering of temperature in Europe of not more than 6° C., and held that even during the glacial advances a comparatively mild climate prevailed in Great Britain. Martins estimated that a lowering of temperature to the extent of 4° C. would bring the glaciers of Chamonix down to the level of the plain of Geneva. Penck estimates that all other.atmospheric conditions remaining the same as now a fall of temperature to the extent of 4° to 5° C. would be sufficient to bring back the Glacial Epoch in Europe. Perhaps the strongest proof that Europe was not refrigerated during the first, second and third glacial advances is the survival of the African-Asiatic fauna throughout the whole period until the fourth glaciation, which was accompanied by widespread severity of climate. Warm and Temperate Interglacial Stages—Similarly the early hy- potheses of extremely warm or subtropical conditions, based chiefly upon the northerly distribution of hippopotami and rhinoceroses, animals which we now associate with tropical conditions, are not supported by the study of the interglacial flora. It is quite probable that both the hippo- potami and rhinoceroses of the so-called “warm, fauna” were covered with hair although by no means so thickly covered as the woolly rhinoceros and elephant of the arctic tundras. There is evidence that in the First Interglacial Stage southern England and France enjoyed somewhat warmer and moister conditions of climate than the present. The Second Interglacial Stage also, which is commonly distinguished as the “long warm” Interglacial Stage, was of somewhat higher temperature than the present. The general evidence is that both in Europe and North Amer- ica, especially in the First and Second Interglacial intervals, the climate in the northern hemisphere was somewhat more equable and milder than at present, with a higher mean temperature, at certain intervals with greater precipitation of moisture, at other intervals much more cool and arid. There was perhaps more sunshine than now. As a result of favorable interglacial conditions arboreal vegetation flourished to the far north along the Arctic ocean, and the present tundra regions of Siberia and British America then supported forests which 949 ANNALS NEW YORK ACADEMY OF SCIENCES have long since been extirpated, the northern limit of similar living trees now lying far to the south. -Alternations of Flora.—lt is clear from these great successive fluctua- tions of temperature, moisture and aridity during Pleistocene times that the flora cannot be treated as a unit nor as progressing in a single direc- tion like the flora of preceding epochs; the flora as well as the fauna presented alternations of arctic, boreal and temperate species which migrated southward and northward following the advances or retreats of the glacial cap. ‘Thus we may observe evidences of changes of climate and flora from forested conditions to steppe conditions and back to for- ested conditions. From the beginning of the Fourth Interglacial in- terval to the present time, the Alps region (Penck, Brtickner, 1909) has apparently gone through a cycle of changes such as the following: VEGETATION CLIMATE Erocu Fourth, forest conditions Western Huropean, oceanic Modern Third, steppe conditions Western Asiatic, continental ITourth Glacial and Postglacial Second, tundra conditions Northeastern-European, Fourth Glacial subarctic First, forest conditions Western-Huropean, oceanic Third Interglacial The elephants (Hilzheimer, 1913) in the structure of their grinding teeth afford clear indications of the plant life, whether consisting mainly of grasses or forests, but not of climate except in so far as vegetation is dependent upon moisture. The advance and retreat of the ice is de- pendent both upon moisture and extreme cold and involves the frozen subsoil conditions of the tundras which are fatal to forests. Cases of alternation of conditions favorable to Hlephas trogontherw, which is be- heved to be a grass-eating form, and of Hlephas antiquus, which is be- lieved to be a forest-living form, are observed in Taubach by Wist.?° This author observes in the lower layers Hlephas antiquus succeeded in the middle layers by H. trogontherw and then in the upper layers again by EF. antiquus, and deduces from, this succession a change of conditions from forest, to steppe, to forest. FAUNAL LIFE ZONES OF EUROPE Tn the whole history of the Mammalia in various parts of the world we know of no conditions so unusual and complex as those which prevailed in Europe in Pleistocene times. These conditions were the product of | > WustT, EwALp: “Die plistozinen Ablagerungen des Travertingebietes der Gegend von Weimar und ihre Fossilienbestiinde in ihrer Bedeutung fiir die Beurteilung der Klimasch- Wwankungen des Wiszeitalters.”” Zeitschr. Naturw.. Bd. 82. pp. 161-252. Leipzig, 1911. OSBORN, REVIEW OF THE PLEISTOCENE 943 cycles of environment and of life which have never prevailed before and will never recur even if the world were to enter a fifth glacial stage, for besides the extraordinary geographic and climatic changes which have been outlined in the previous pages there was the prodigal profusion of life which survived from Pliocene times and has since become extinct. The result of these complex conditions was the assemblage in Europe of animals indigenous to every continent on the globe except South Z sts oe ch mak ae te a i ee ne / 7 YY zh : mae gd Ly oy RS) yy C Pie. 6.—Pive chief zodgeographic regions of Europe, Asia and northern Africa from which the mammals migrated into western Europe during the Pleistocene America and Australia, and adapted to every climatic life-zone from the warm and dry plains of southern Asia and northern Africa to the tem- perate forests and meadows of Hurasia, from the alpine heights of the Alps, Pyrenees and Altai Mountains to the high, dry steppes of central Asia with their alternating heat of summer and cold of winter, from the tundras or barren grounds of Scandinavia, northern Europe and Siberia to the mild climate of southern Europe. All these animals had been evolving during the Pliocene Epoch in these various habitats and they 244 ANNALS NEW YORK ACADEMY OF SCIENCES also underwent a very considerable evolution during Pleistocene times in their respective habitats. Owing to successive migrations and invasions into Europe of these exotic types of the north and south we should not expect to find a con- tinuous phyletic evolution or transformation such as we have observed in the earlier epochs, excepting only that which occurred among the Hura- siatic forest and meadow types which appear to have been native or in- digenous in Europe from the earliest Pleistocene to prehistoric times. These Hurasiatic forest and meadow mammals were continuous residents, retreating in the coldest periods to the shelters on the east and south. Cervus elaphus, for example, passed through several subspecific stages of evolution. The invading hordes from the tundras, the steppes, from northern Africa and from Asia represent branches which had their evolu- tion elsewhere. This is true both of the mammals and of the races of men which had their genesis in the far east and southeast and arrived in Kurope when it was a fertile peninsula, a region too small to be the seat of a continental evolution or adaptive radiation. The five great regions which contributed to the European Pleistocene were as follows: AFRICAN AND ASIATIC, PLAINS AND FOREST TYPES EUROPEAN AND ASIATIC, ForREST AND MEADOW TYPES EUROPEAN AND ASIATIC, ALPINE TYPES STEPPE REGIONS OF SOUTHERN HURASIA AND WASTERN SIBERIA Arctic TUNDRA REGIONS OF NORTHERN HURASIA Interature—The African-Asiatic element in these Pleistocene faunas was the first to be recognized and commented upon by the early writers; it is commonly known as the “warm fauna.” We owe especially to Neh- ring the discrimination between the tundra and the steppe faunas. Gaudry, Harlé, Woldrich, Studer and Boule have added to our knowl- edge of these faune. Other contributors are Pohlig, Soergel, Forster, Hilzheimer, Wiist and Dietrich. A strict systematic revision and intro- duction of the trmomial system is greatly needed. The most complete recent faunal lists of the late Pleistocene deposits in which traces of man are found are those of Koken and Schmidt (1912), who have also insti- tuted the closest correlation between the migrations of the Mammalia and the successive stages of human culture. African—A siatic Mammals, Warm Fauna.—Vhese mammals include those which first appear in the Upper Pliocene and survive into Lower Pleistocene times; also those which first appear in the Second Inter- glacial Stage and constitute the so-called “warm fauna” which survived in Hurope until the middle or close of the Third Interglacial Stage. The OSBORN, REVIEW OF THE PLEISTOCENE . 45 principal members of this list together with the probable continental cen- tres of their origin are as follows: Macacus ? sp. (Africa) Macaque baboon Elephas meridionalis (Asia) Sudelefant Southern mammoth Blephas antiquus (Asia- Altelefant Straight-tusked elephant Africa) Dicerorhinus etruscus (Asia) Nashorn Htrusean rhinoceros 5s merckii (Asia) Mercksches Nashorn Broad-nosed rhinoceros Hippopotamus major (Asia) Flusspferd Hippopotamus Bos primigenius (Asia) Urochs, Auerochs Urus Bison priscus (Asia) Wisent, ‘‘Auerochs”’ Primitive bison Equus stenonis Upper Pliocene horse Macherodus latidens Sabre-toothed tiger Hyena spelea (Asia-Africa) Hodhlenhyiine Cave hyzena “striata (Asia-Africa ) Canis aureus (Africa) Schakal Jackal Felis leo antiqua (Africa) Altlowe Lion “ pardus Leoparde Leopard The remains of these animals play a very important part both in the subdivision of the geologic horizons of Hurope and in theories regarding the alternation of climates, as well as in the determination of the an- tiquity of man. They are found chiefly in the river sands, river deposits and “High Terraces” of the First and Second glaciations and “Low Ter- races” of the Third Glacial and Interglacial Stages. Some survivors are found in the shelter and cavern deposits of the Third Interglacial Stage as objects of the chase. Only two of these animals, the urus and the bison, survive to become members of the Prehistoric Forest and Meadow Fauna of Europe. The lion also survived into Postglacial times but dis- appeared in Kurope before the Prehistoric and Neolithic periods. Although originally derived from southern Asia or from Africa, the woolly elephant (1. primigenius) and woolly rhinoceros (Diceros antiq- uitatis) are not to be placed with the African-Asiatic fauna because they appear in Europe only with the northern Glacial or Tundra Fauna and are invariably indicative of cold climatic periods. Hurasiatic Forest and Meadow, Temperate Fauna.—These animals were resident in the forests and meadows of Europe during the entire Pleisto- cene Epoch and survived with a few exceptions into Postglacial and Pre- historic times. In fact, they are probably separable by differences of specific and subspecific value from their successors in prehistoric Europe, but many authors still embrace them within the existing specific terms. They are divided into Forest and Meadow types, the former with brachy- odont teeth adapted to browsing habits and forest environment, the latter with elongate or hypsodont grinding teeth adapted to meadows and a diet 246 . ANNALS NEW YORK ACADEMY OF SCIENCES of grasses. Some of them, like the beavers, are stream- and river-dwellers. Naturally there is not always a sharp line of division between the habitats of these Forest and Meadow types: the Carnivora especially wander after their prey from the forests into the meadows and along the stream borders. Sus scrofa ferus Cervus claphus Megaceros Germanie Cervus maral (Persia) Capreolus capreolus Alces latifrons Equus mosbachensis Trogontherium cuvieri Castor fiber Sciurus vulgaris Lepus cuniculus Arvicola amphibius sc ratticeps ie agrestis nA glareolus Myoxnus Mus sylwaticus Talpa europea Ursus deningeri ne arctos speleus “ arvernensis Felis catus Lynchus lynx Lynx cervaria Canis lupus “ neschersensis “ Suessi Vulpes alopex Meles taxus Mustela vulgaris ‘ martes Gulo luscus Fetorius putorius Lutra vulgaris Bison priscus Bos primigentus Equus caballus (? sp.) Forest MAMMALS Wildschwein Edelhirsch Riesenhirsch Reh Elch Biber Hichhornchen Wildkaninchen Wasserratte Nordische Wuhiratte Erdmaus Rotelmaus Deninger Bar Brauner Bar Hohlenbir gem. Wildkatze gem. Luchs Silberluchs Wolf gem. Fuchs Dachs gem. Marden Edelmarder. Baum- marder Iltis Fischotter Meapow MamMMALSs Wisent Urochs The cattle and bison frequent both the forests and meadows. Wild boar Red deer, stag Giant deer Maral deer Roedeer Moose (broad-faced ) Horse of Mosbach Giant beaver Beaver Squirrel Wild rabbit Dormouse Forest mouse Mole Brown bear Cave bear Val d’Arno bear Wild cat Lynx Eurasiatic lynx Wolf * Common fox Badger Marten Pine marten Wolverine Polecat Otter Bison Urus OSBORN, REVIEW OF THE PLEISTOCENE 244 Fetorius vulgaris Wiesel Weasel Sorex vulgaris Shrew Cricetus vulgaris Hamster Arvicola terrestris Schermaus arvalis gem. Feldmaus Meadow vole Of these forest-living animals the giant deer (Megaceros), the cave bear (Ursus speleus), the Maral deer (Cervus maral), the giant beaver (Trogontherium) and the early Pleistocene species of horse are among PREHISTORIC By] = ‘ AND | RECENT P| Post GLlacral FOURTH GLACIAL —————| faa it Third Interglac zal THIRD GLACIAL Second Interglacial GLACIAL UPPER PLIOCENE = sy ——_ Warm Temperate Cold Domestic Fic. 7.—Introduction, succession and extinction of the faune from the five chief zodgeographic regions African-Asiatic, Eurasiatic Forest and Meadow, Tundra, Steppe and Desert, Alpine. the few forms which became extinct during the Glacial and Postglacial epochs. The great majority of these species survived with successive subspecific modifications. It is a very remarkable fact that this true forest fauna of Europe is frequently found in the same deposits with the “warm fauna” of African- Asiatic origin. The bison and wild cattle appear in Europe from early 948 ANNALS NEW YORK ACADEMY OF SCIENCES DA Pleistocene times, and in late Glacial and Postglacial times they oecur as companions of the mammoth and the woolly rhinoceros. Alpine Mammals, Cold Fauna.—During the severe conditions of late Pleistocene times the Alpine mammals were driven down into the plains or to the lower mountains and hills, and their remains oceur principally during the last Glacial advance. They are represented both in the draw- ings and in the sculptures of the men of the reindeer or cave period. Ibex priscus Steinbock Primitive ibex Rupicapra tragus Gemse Chamois Ovis argaloides (Altai Mts.) Argalischaf Argali sheep Arvicola nivalis Schneemaus Alpine vole Lagopus alpinus Gebirgsschneehuhn Ptarmigan Steppe Fauna of Russia and Siberra.—sSteppe conditions of climate were rendered possible in Europe by the elevation and extension of land much farther to the north and northwest than at present. At such periods all the tempering influences of the Atlantic Ocean were cut off from northern Europe and helped to give ceutral Hurope a cold, dry con- tinental climate favorable to dust storms. Boule, Kobelt?® and Scharff?” have agreed in the opinion that the pres- ence of steppe mammals affords inadequate proof of steppe conditions in the country. Other authors (Hilzheimer, 1913), however, strongly sus- tain the steppe-climate theory. The evidence for steppe conditions of climate has been strengthened in recent years by the discovery of three successive loess deposits. The steppe regions of eastern Hurope around the Caspian Sea and of central Asia still maintain this highly characteristic steppe fauna. The climate is usually one of hot, dry summers with high winds and prolonged cold winters with sweeping snow storms. The animals are consequently very hardy. The fauna includes the jerboa, sushk, bobac marmot, dwarf pica, hamsters, northern voles, corsac fox, the manul, or Pallas’s cat (Felis manul). Covering the plains are the larger grazing animals such as the saiga antelopes, wild asses and wild horses (including the Equus przewalsku type). Another animal which probably belonged to the Steppe fauna is the Hlasmotherium. Elasmotherium sibiricum® Hlasmothere Equus przewalskii Wild Gobi horse “ hemionus Dschiggetai Dzeggetai, wild ass, kiang 8 KOBELT, W.: Die Verbreitung der Tierwelt. Gemiussigte Zone. Leipzig, 1902. 27 ScHarFF, R. F.: The History of the European Fauna. London, 1899. °3 GAUDRY, ALBERT. and BOULE, MARCELLIN: Materiaux pour l'Histoire des Temps Quaternaires. Trois. Fasc. L’Elasmotherium. 4to. Libr. F. Savy, pp. 83-104, pll. xyi- xix. Paris, 1876. OSBORN Saiga tartarica Alactaga jaculus Lagomys pusillus Spermophilus rufescens Cricetus pheus Arctomys bobac Myodes lagurus Arvicola gregalis Canis corsac Putorius eversmannt Tetrao tetrix , REVIEW OF THE PLEISTOCENE Saiga antilope Gr. Pferdespringer Zwergpfeifhase Rotliche Ziesel Kl. Steppenhamster Sibirische Zweibelmaus Birkhubhn 249 Saiga antelope Jerboa Dwarf pica, or tailless hare Suslik, or pouched marmot Steppe hamster Bobae, or Polish marmot Steppe lemming Steppe vole Corsac wolf Steppe weasel Moorhen Tundra or Barren Ground Fauna and Flora.°?°—Certain members of the Tundra Fauna adapted to the long cold winters and short summers of the lands bordering the Arctic Ocean appeared in Europe at the height of each of the great glacial advances. The remains of these animals are always found within or close to the glacial drifts until the Fourth Glacia- tion when they spread all over France, Germany and Austria. Thus the musk-ox (Ovihos moschatus)-is recorded in the (?) First Glacial ad- vance of the Forest Bed of England. A tundra fauna including the rein- deer (Rangifer tarandus) is recorded (Forster, 1913) with the Third Glacial advance (Upper Mauer sands). An extensive Tundra Fauna also appears with the Third Glacial, or Riss Stage, in the “Mammutlehm” of Cannstatt (Koken, Schmidt, 1912, p. 270); this is termed the “Older Primigenius’ Fauna and occurs on the “high terraces” with the older Diluvium ; it includes the woolly mammoth, the rhinoceros, the horse and the reindeer. The Tundra Fauna reappears toward the close of the Third Interglacial Stage (1. e. “Lower Rodent” layer), but the full series of species characteristic of the Tundra Fauna are not recorded in Hurope until the Postglacial Stage (1. e. “Upper Rodent” layer), when the entire Tundra lst given below is discovered either mingled with the culture . remains of the Neanderthal race of men in Mousterian times or is repre- sented in the art of the Cré-Magnon men of the reindeer period. The full or typical Tundra list of the Fourth Glacial Epoch is as follows: Elephas primigenius Mammut Mammoth Diceros antiquitatis Wollhaariges Nashorn Woolly rhinoceros Rangifer tarandus Ren Barren ground reindeer : Ovibos moschatus Moschusochse Musk-ox Lepus variabilis Schneehase Aretic hare Myodeés obensis Oblemming Obi lemming, or Siberian lemming. 2 NEHRING, A.: Uber Tundren und Steppen der Jetzt- und Vorzeit, mit besonderer Berucksichtigung ihrer Fauna, pp. 81-166. Berlin, 1890. 950) ANNALS NEW YORK ACADEMY OF SCIENCES Myodes torquatus Halsbandlemming sanded lemming Canis lagopus Hisfuchs Arctic fox Gulo borealis Vielfrass Wolverine (glutton) Fatorius erminea Gr. Hermelin Ermine Arvicola nivalis Arctic vole Lagopus albus Moorschneehuhn Ptarmigan Asio palustris Sumpfeule CYGNUS MUSICUS Singschwan Animals like the banded lemming adapted to extreme northerly condi- tions generally cling to these very obstinately and perish rather than con- form to an altered environment (Nehring). ‘This species dwells imme- diately to the north of the region of coniferous forests, among scattered shrubs of the common juniper (Juniperus communis), the dwarf birch (Betula nana), the polar willow (Salix polaris) and the mountain dryas (Dryas octopetala). ‘Thus we may be confident that the lemmings dis- covered in Pleistocene times in England, France, Belgium and a large part of Germany are proof of climatic conditions similar to those of the present circumpolar region. We must conclude that the borders of the ice caps were surrounded by tundra or barren ground conditions at sev- eral Pleistocene stages. The lemmings probably dwelt in the immediate neighborhood of the glaciers. The existing tundras are characterized by frozen subsoil and the absence of trees or shrubs except along the river borders. The reason for associating the woolly mammoth with this fauna is that the mammoth as depicted by the men of the Postglacial Stage agrees pre- cisely in its form, its proportions, and its hairy covering with the mam- moths which have been discovered in the frozen subsoil of northern Si- beria and are washing out in large numbers along the northern Siberian and American coast at Eschholtz Bay and elsewhere. At Thiede near Braunschweig, a classic locality, the lemming remains are associated with those of the arctic fox, arctic hare, reindeer, musk-ox and mammoth. Thus the comparison of certain regions of Pleistocene France and Ger- many with arctic Eurasia and the barren grounds of northern Asia and North America is based on the strongest evidence. MIGRATION THEORY OF FLORAS AND FAUNAS The principal contributors to the theory of northward and southward migrations and to the succession of faunas and floras are Nehring (1880- 1896), Woldrich (1882-1896) and Penck (1896-1909). Such alterna- tion is held by Penck to be demonstrated in Switzerland, where during the Third or Riss glaciation the woolly mammoth and woolly rhinoceros - Be Same generic si 3 REN, near ink iat Suna = iit roar roa the Pies {ie 4 by eee teyt Tt PASE Oy eee op aaioeiw oe SGICETES rs Bes sites THN eH Arr Pelyy Taiee a — ite ats ractec’ the tunes ie Sea is anc 2x ae fish G San oe CREE _ sigaossaie a “ LSiWADIA MGV Gboseay titan SF grsshia : 2 is “he Bie i hisfude: bit é varaiaktt ouchuless | os a “eTodgeimeH Taste 1V.—Distribution of Forest Meadow Mammals RECENT Return of Alpine Fauna to the Mountains. Wide dispersal III Pore Voresr g PREHISTORIC of Forest and Meadow Fauna over entire Northern Hemisphere MeEapow Fauna 3 i) Cervus elaphus a POSTGLACIAL ve ; (Severe climate) Retreat of Tundra and Steppe Fauna. wilder southward and “ capreolus IL enNDEEE PERIOD! 3 westward dispersal of Tundra, Steppe and Alpine Fauna in Burope Saf i ‘A, STEPPE, TV GuacraL to the Pyrenees, Alps, northern Spain and Italy. Arrival of Tundra Hoyle UYU Cer Os: NINE. AND Meapow 5 Fauna. Arrival of Asiatic Steppe Fauna. Ursus arctos AUNA a Bison priscus 8 (Steppe climate) Bos primigenius Return of Warm and Tem- Pxtinction or southward retreat PoHLiGc, H.: ‘Hine Elephantenhoéhle Siciliens und der erste Nachweis des Cranial- domes von Hlephas antiquus,”’ Abhand. kénigl. bayer. Akad. Wissensch., cl. ii, Bd. xviii, Abth. 1, pp. 75-108, pll. i-v, 1895. Sep. Munich, 1893. . 968 ANNALS NEW YORK ACADEMY OF SCIENCES The diminutive elephants of the Mediterranean islands were all de- scendants of the straight-tusked species 1. antiquus. 'The researches of Bate confirm this relationship. They attamed a height not exceeding five feet. The adaptability to which L. antiquus owed its wide geographic distribution and continued existence through a long period of time may account for its survival in the Mediterranean islands despite rapid dim- inution in size under adverse circumstances. The true African elephant (Loxodonta) never crossed the Mediterranean. The reduced existing fauna of the Island of Cyprus contains a min- cling of Eurasiatic and north African mammals and shows the effects of deforestation in historic times. Descendants of Eurasiatic ancestors pre- vail in the Mediterranean islands. The recently discovered Myotragus balearicus of the Pleistocene cave deposits of the Island of Majorca is now regarded as related to the Rupicaprine or Alpine chamois type (Andrews). SECOND GLACIAL STAGE—SAXONIAN, MINDEL, KANSAN The second glaciation was the greatest both i Europe and America. We observe that the most extended drift sheets are those in the Scandi- navian region, on the British Isles, around the northern Swiss Alps, and fo) 3 x) ») from the Keewatin center west of Hudson Bay in British America. ‘The whole rise and fall of the Mindel glaciation in the Alps is estimated by Penck as occupying a very long period of time. The snow line in the Alpine region descended 1,300m. lower than at the present time. The only notable exceptions are in the Labrador region of eastern North America where the main ice field was formed at a later stage, known as the Illinoian. It also appears that the Third Glacial or Riss drift of the western Alps is the greatest in that region and of similar age to the Illinoian. In this second glacial advance the Scandinavian ice field reached its farthest southerly limits. In northwestern Europe this main Saxonian (Geikie) glaciation extended to the northern slopes of the Carpathians, the Sudetes, the Erz Gebirge, the Thuringian and the Harz Mountains. From these ice sheets were given off the “Older Drift,” or “Lower Dilu- vium” of northern Germany, and in the Swiss Alps this Second glacia- tion sent out its Mindel drift as the most extensive fringe along the northern borders of the Alps; on the eastern and southern borders of the Alps the Second Glaciation was about as extensive as the Third glacia- tion; on the western borders of the Alps the Second glaciation was less extensive than the Third. Similar conditions prevailed in America; from the Keewatin Centre the ice cap extended its drift southward into OSBORN, REVIEW OF THE PLEISTOCENE 269 A Missouri, Iowa, Kansas and Nebraska beyond the limits both of the First and the Fourth glaciation. Thus the Saxonian drift of North Germany, the Mindel drift of the Swiss Alps and the Kansas drift of America are correlated (Penck, Lev- erett) both by their great antiquity and by their very wide extent. The eminent geologist Wahnschaffe, however, correlates the “Old Drift” of the north German lowland with the Third or Riss glaciation instead of with the Second. SECOND INTERGLACIAL STAGE—MINDEL-RIss Penck regards the Second Interglacial or Mindel-Riss Stage as by far the longest of the interglacial intervals in the Alpine region, estimating the period between the maximum Second glaciation (Mindel) and maxi- mum Third (Riss) as high as 360,000 years. In America also by com- paring the erosion of the Second Glacial (Kansan) drifts with those of the Third Glacial (Illinoian) drifts it would appear that the Second Interglacial Yarmouth Stage was of greater duration than the entire interval between the Third Glacial and present time. In the north Ger- man lowlands it is shown to be a long interval from the amount of sedi- mentation effected by the interglacial rivers and streams, but whether in this region it is longer than the First Interglacial Stage is doubtful (Lev- erett, 1910, p. 273). MOISTURE FOLLOWED BY ARIDITY In course of this long warm Second Interglacial Stage the climate again moderated, becoming slightly warmer than the climate of to-day. The chmate immediately following the retreat of the ice was cool and moist, then followed a long warm stage, but this stage was finally suc- ceeded by a period of aridity both in Europe and America in which the first loess deposits occurred. In Russia also the Second Interglacial seems to begin with a cool and moist phase followed by a more arid or steppe- like climate favorable to the deposition of loess. It would appear that the height of the interglacial aridity was reached during the deposition of the loess. The “Older Loess” deposition certainly began both in Hurope and America during the Second Interglacial Stage although in neither country is the “Older Loess” so continuous or so thick a deposit as the “Newer Loess.” In Europe the “Old Loess” lies between the “Old Drift” of the First and Second Glacial advances and the “Middle Drift” of the Third Glacial advance. At Mauer near Heidelberg the loess les imme- diately above the Upper Mauer sand layer which contains an arctic- tundra fauna (Forster, 1913). The various layers of loess are of the 270 ANNALS NEW YORK ACADEMY OF SCIENCES utmost importance both in Hurope and America in the correlation of human and mammalian life, also in their significance as to the climate of interglacial times. Loess consists of a fine, porous, silicious, calcareous silt, usually of light brown color, characterized by a peculiar competency to stand in vertical walls during erosion. Its origin and transportation are believed to have been partly sub-glacial, partly fluviatile, partly seolian. The fine mud carried by the sub-glacial streams in glacial times became desiccated and redistributed by the wind. Penck (1904) describes the Pleistocene loess as formed in districts traversed periodically by great streams leaving dry mud which in arid periods was redistributed by eolian agencies. Its Pleistocene distribution is quite independent of alti- tude since it occurs in the interglacial deposits of Europe from sea level to a height of 1,500m. CLIMATE A considerable part of the elevation of the Swiss Alps apparently took place (Penck, 1910) during the Second Interglacial Stage, and this in- creased altitude is considered by some European authorities to be the cause of the greater extent of the Third glaciation in the western Alps. The Hottinger breccia near Innsbruck is referred by Penck (1909, p. 1157) to the Second Interglacial Stage with its rich flora indicating a climate warmer than that of present times; this breccia lies on one of the old “High Terraces” of Second Interglacial times. The plants include the fir (Pinus sylvestris), spruce (Picea sp.), maple (Acer pseudopla- tanus), buckthorn (Rhamnus frangula), several willows (Saliva nigricans, S. glabra, S. incana, 8. triandra), the wayfaring tree (Viburnum lan- tana), yew (Taxus baccata), elm (Ulmus campestris), strawberry (Fra- garia vesca), self-heal (Prunella vulgaris), beech (Fagus silvatica), and mountain ash (Sorbus aucuparia), buckthorn (Rhamnus Hottingensis), related most closely to R. latifolia, now living in the Canary Isles, the box (Buzxus sempervirens), also a southern species; and most important of all a rhododendron (R. ponticum) which now lives in the Caucasus five degrees south of the latitude of Innsbruck and in a climate on the average 3° C. warmer. ‘Taking all the facts into consideration Penck concludes that the climate of Innsbruck in the days of the Hotting brec- cia was 2° C. higher than it is now. In correspondence with this the snow-line stood 1,000 ft. above its present level, and the Alps save for the higher peaks were almost completely denuded of ice and snow.® A picture of the flora of the Second long warm Interglacial Stage is also afforded in the Quaternary tuffs of Provence, where the remains of 5 SoLuaAs, W. J.: Ancient. Hunters and their Modern Representatives, p. 27. 8vo. MacMillan & Co. London, 1911. q , as OSBORN, REVIEW OF THE PLEISTOCENE Py rea | plants are associated with elephants of the LZ. antiquus stage. “The flora of the Quaternary tuffs,” observes Saporta,°® “is composed almost entirely of woody forms living in valleys and by the sides of streams.” It is for the most part analogous with the present flora of Provence. Of the thirty-seven species, twenty-nine still occur in this region. Among the forms which have since retreated to the south are the sweet bay (Laurus nobilis) and another species of laurel (LZ. canariensis) which is now con- fined to the Canaries. The greater humidity of the time is indicated by the presence of species of pine which require more moisture. As in the Norfolk Interglacial, the figs (Ficus) and the Judas trees (Cercis) flour- ished. The ash (Fracinus) is of a species now found in Corsica and Italy. On the whole, the forest trees and forest ground flora are surpris- ingly modern, including oaks, elms, poplars, willows, lndens, maples, sumacs, dogwood, hawthorn. Among the chmbing plants are the vine (Vitis) and clematis (Clematis). MAMMALS This life period was first observed by Lyell and Evans in Essex, Eng- land, and was subsequently recognized in Germany and France. Geo- logically the deposits are partly of fluviatile origin, consisting chiefly of river sands and gravels in which the remains of hippopotamus, elephants and rhinoceroses occur. These animals were formerly cited as proof of an almost tropical climate, but the evidence of the flora, enumerated above, and the equally numerous hardy types of animals tend to modify the former theories as to extremely warm Second Interglacial tempera- tures. The geographic connections of Europe with the south through the land bridges of Lower Pleistocene times still persisted in Italy in whole or in part, because the depression of the southern portion of the continent of Europe had not yet begun. Survivals.—The mammals occurring in these Older Diluvial sands and gravels include several Pliocene survivals from the First Interglacial Stage, associated with the etruscan rhinoceros (Dicerorhinus etruscus). Tf Montmaurin belongs to this stage we may include Macherodus. At Mauer two primitive types of bear, Ursus arvernensis and U. deningent, are recorded, also Trogontherium cuviert. The Mauer horse first identi- fied as #. stenonis is now referred to H. mauerensis. Among the chief localities where the river deposits containing the mammals referred to the Second Interglacial Stage occur are the fol- lowing : % Dr Saporta, G.: “La Flore des Tufs Quaternaires en Provence.” C. R. Sess. Congr. Sci. France. Aix, 1867. m9 ANNALS NEW YORK ACADEMY OF SCIENCES rand) HaARLY PHASE. WARM FAUNA. Mauer Sands (Lower), near Heidelberg, Germany. Warm Fauna. Montmaurin (Haute Garonne), Pyrenees, France. St. Roche, France. MIDDLE PHASE. Mosbach, near the Neckar in northern Baden (Fig. 9, 12). Siissenborn, near Weimar, Germany (11). LATER PHASE. COLD TUNDRA FAUNA. Mauer Sands (Upper), near Heidelberg. Germany (14). Cold Fauna approaching the Third Glacial Stage. Extinctions—The mammals of this grand life zone have lost nearly all resemblance to those of Upper Pliocene times with the exception of the survival of the etruscan rhinoceros and possibly of. the sabre-tooth tiger. The polycladine deer of Upper Pliocene times and of the Norfolk Forest Bed, or First Interglacial, have vanished; neither are there any traces of the axis deer (C. azis). Arrivals.—The Second Interglacial Stage is readily distinguished both in France and Great Britain by a number of important new arrivals, chief among which are the “old elephant” (#. antiquus) and the broad- nosed rhinoceros (D. merckw). Another very important arrival is the lion related to the African Felis leo. The southern elephant has now certainly passed into the Hlephas trogontherw stage of Pohlig for the type specimen of this intermediate species occurs at Stissenborn ; in fact, this is the “H. trogontherw stage” of Pohlig; it is also known as the “older H. antiquus” stage by Schmidt and other authors. The southern mammoth H. (meridionalis) trogontherw is replaced by the more pro- gressive and typical #. trogontherit. The broad-faced moose (Alces lati- frons), the giant deer (Megaceros) and the roe deer (Capreolus) are all present in the cooler and forested phases of this interglacial period. The true stag (Cervus elaphus) is certainly recorded. The cattle (Bos primi- genius) begin to be numerous and the bison (Bison priscus) also appear in numbers. Horses of larger size occur (H. mosbachensis, EH. siissen- bornensis) . Among the river-living forms are the beavers (Castor). The giant beaver (T'rogontherium) is by some authors said to make its last appear- ance in Europe in this sub-stage, but it is again recorded in the Third Interglacial at Chelles. Other rodents include the marmots (Marmotta) now found in the Alps, Carpathians and Pyrenees, whose remains may have been borne down by the streams. Beside the lions the carnivores include the typical Eurasiatic forest forms, namely, the lynx (Ff. lynz), two varieties of bear (U. deningeri, U. arvernensis), and the badger (Meles). The chief components of the fauna of the Second Interglacial Stage OSBORN, REVIEW OF THE PLEISTOCENE 9723 u are seen to belong, first, to the Eurasiatic Forest and Meadow Fauna, only separated by specific and sub-specific differences from the Prehis- toric Fauna of Europe; second, to the surviving African-Asiatic fauna, including the hippopotamus, two very distinct kinds of elephant, and two rhinoceroses; third, there is evidence in the late colder phases of this period of the first occurrence in Europe of the Tundra Fauna as repre- sented by the reindeer (Rangifer tarandus). This animal is recorded in the gravels of Siissenborn by Weiss. Hilzheimer®’ also speaks of the re- mains of reindeer as occurring both in Stissenborn and Steinheim in asso- ciation with the remains of HZ. trogontheru. This author regards FL. tro- gontherw from the structure of its grinding teeth as analogous in habit to the Asiatic elephant which inhabits the forests of India, and believes that the presence of this animal indicates a relatively moist climate and well forested country. In this assemblage it is noteworthy that the lEurasiatic Forest and oionnesdelvercensis hardy temperate types greatly pre- J GERG AS TRAE dominate over the African-Asiatic Straight-tusked elephant types. This is another indication EB. antiquus that the climate was of a warm-tem- peceoutueran imammoth perate character rather than such as EB. trogontherii 5 j Sa aia sO er now characterizes southern Asia and Africa. It follows that all the Afri- AFRICAN-ASIATIC TYPES Primates D. merckii Etruscan rhinoceros can-Asiatic mammals may have been D. etruscus well protected by hairy covering and SEU SIUC EHS adapted to a temperate climate. H. major Gay Sabre toorhetizes In the caverns near Montmaurin Minhorodie in the Pyrenees®® we find remains of Lion an early Pleistocene fauna which Felis leo spelea contains the sabre-tooth tiger (M. EURASIATIC Harpy FAUNA latidens), the broad-nosed rhinoc- Urus ; o* Boe mranigenils eros (D. merckw), the stag (C. ela- Bison phus), the brown hyena (fH. brun- Bison priscus nea striata). Stag, roe deer, moose, giant deer Bear, lynx, badger. wild cat (Late) Reindeer Rangifer tarandus The most typical fauna is that of Mosbach in northern Baden. Here there occur all the characteristic mammalian types of the period, the hippopotamus, the urus, the bison, 37 HILZHEIMER, Max: Handbuch der Biologie der Wirbeltiere, pp. 678-679. Stuttgart, 1912-1913.. 52 Bout, M.: “La Caverne A Ossements de Montmaurin (Haute-Garonne).” L’An- thropol., Vol. XIII, pp. 305-319. 1902. 24 ANNALS NEW YORK ACADEMY OF SCIENCES the broad-nosed rhinoceros, two species of mammoth (7. antiquus, HE. trogontherw), and the horse (#. mosbachensis). In the Lower sands of Mauer near Heidelberg there occur the first re- corded remains of man and a fauna including some primitive species. Homo heidelbergensis—To the faunal stage of Hlephas antiquus and the etruscan rhinoceros (D. etruscus) is to be added the Heidelberg man, determined from a lower jaw discovered by Otto Schotensack®? im 1907 in the Lower Mauer Sands at a depth of 24.10 m., one of the most im- - portant discoveries in the whole history of anthropology. The lower jaw is exceptionally massive, without chin projection, with a large but essen- Fic. 14.—Sand pit at Mauer near Heidelberg The lower jaw (Homo heidelbergensis) was found at the spot marked with a cross. After Schoetensack and MacCurdy. tially human set of teeth; in other words, it is a jaw in some respects similar to that of an anthropoid ape but containing the dentition of a man, namely, typically human canine and molar teeth. The jaw is now regarded by anatomists as resembling on a very massive and primitive scale the jaw of the neanderthaloid human type (Homo neanderthalen- sis) which first occurs in the Third Interglacial Stage. The fauna associated with Homo heidelbergensis is of an ancient char- 5° SCHOTENSACK, OTTO: Der Unterkiefer des Homo heidelbergensis aus den Sanden von Mauer bei Heidelberg: Hin Beitrag zur Paliiontologie des Menschen. Verlag yon Wilhelm Engelmann. Leipzig, 1908. oe OSBORN, REVIEW OF THE PLEISTOCENE O75 acter. Schdtensack likened it to that of the First Interglacial or Nor- folkian Stage. The presence of the etruscan rhinoceros would appear to justify this opinion, but it is overborne by the similarity to the fauna of Mosbach including the presence of Hqwus mosbachensis, a species highly characteristic of the Second Interglacial Stage. The entire fauna of these Lower Sands of Mauer is now identified (Schmidt, 1912), as fol- lows: Elephas antiquus, D. (Rhinoceros) etruscus,°° Equus mosbachen- sis, Sus scrofa fere, Alces latifrons, Cervus elaphus, Capreolus capreolus, Bison priscus, Bos primigenius, Ursus arvernensis, U. deningert, Felis leo, Felis catus, Canis neschersensis, Castor fiber. The enumeration of Fic. 15.—Heidellerg jaw The human lower jaw (%4 natural size) found near Heidelberg, on which is based the species Homo heidelbergensis. After Schoetensack. this fauna is very important as indicating the temperate climatic condi- tions which surrounded the Heidelberg man. Wurm observes that the Ktruscan rhinoceros only occurs in Mauer and that its variations indi- cate a transition towards the D. merckw which occurs at Mosbach and Stissenborn but not in Mauer. Above this layer occurs a deposit of the “Older Loess,” indicating an arid climate. The Upper Sands of Mauer contain a cold fauna which by some is referred to the close of the Second © WourRM, A.: “Uber Rhinoceros etruscus Fale. von Mauer a. d. Hlsenz (bei Heidel- berg).” Verh. d. Naturhist.-Medizin. Vereins zu Heidelberg, N. F. XII Band, 1 Heft, pp. 1-62, pll. I-IV. 1912. © Worm, A.: Beitrige zur Kenntnis der Diluvialen Siiugetier fauna von Mauer a. d. Elsenz (bei Heidelberg), I. Felis leo fossilis. Jahresberichte und Mitteilungen des Ober- rhein. geol. Vereins, N. F., Bd. II, Heft 1, pp. 77-102. 1912. 26 ANNALS NEW YORK ACADEMY OF SCIENCES Interglacial stage, by others (Wurm, 1913, p. 68) to the “Younger Loess,” that is, to the Fourth Glacial or Postglacial Stage. CHELLEAN CULTURE WITH ANCIENT INTERGLACIAL FAUNA In favor of the antiquity of the Chellean culture may be urged the fact of its association in several localities (Torralba, Abbeville, Piltdown) with the primitive mammals identified as Macherodus, D. etruscus, Equus stenons. The specific identifications may be incorrect, but these Pliocene species are characteristic of the Second Interglacial Stage and are not certainly recorded in the Third Interglacial Stage of northern Kurope at least. For example, at Torralba, Province of Soria, Spain, there has been discovered (Harlé, 1910, p. 75) an old typical Chellean camping site containing abundant remains of D. mercku and H. merid- ionalis (trogontherti) mingled with remains of other mammals of prim- itive type identified as Dicerorhinus etruscus and Hquus stenonis. These associations with Chellean remains tend to support the theory that the Chellean culture began during the Second Interglacial Stage. Another very ancient fauna associated with very primitive Chellean or pre-Chel- lean implements is that found near Abbeville, Gisement de Champ de Mars.°* Beside typical members (such as H. antiquus, H. meridionalis trogontheru, and D. merckw) of warm Second Interglacial times this fauna is said to contain such primitive types as Trogontherium, D. etruscus, Equus stenonis, also very numerous specimens of Machwrodus and Hyena brevirostris. We cannot fully agree with Schmidt (1912) when he observes that the faunal separation of the Acheulean and Chellean is not so marked that we are obliged to separate these cultures by a long period of time. FAUNA OF THE PYRENEES, CANTABRIAN ALPS, SPAIN AND PORTUGAL ° The entire warm fauna characteristic of Germany, Great Britain and France also penetrated the Cantabrian Alps, Spain and Portugal as far south as Gibraltar. A macaque (Macacus) related to the Algerian species occurs in the erotto of Montsauné (Haute Garonne) associated with the hyena (H. 82 WurM, A.: “iber eine Neuentdeckte Steppenfauna von Mauer an der Elsenz (bei Heidelberg).’”’ Jahresber. u. Mitt. d. Oberrhein. geol. Vereins, N. F., Bd. III, Heft 1, pp. 62-78, pl. vi. 1913. : 8 DP’AULT DU MESNIL, G.: “‘Note sur le Terrain Quaternaire des Environs d’Abbe- ville.” Revue Mensuelle de l’f&cole d’Anthropologie de Paris, VI year, pp. 285-296. 1896. 8 HARLE, EpouarpD: “Les mammiféres et oiseaux quaternaires connus jusqu’ici en Portugal. Memoires suivi d’une liste générale de ceux de la Peninsule Ibérique.’”’ Com- mun. du Service Géol. du Portugal, T. viii, pp. 22-85, pll. I-V. 1910. OSBORN, REVIEW OF THE PLEISTOCENE any striata), with a dhole (Cyon), and other members of the warm fauna of EF. antiquus and D. mercku. The porcupine (Hystrix cristata) also oceurs here. The striped hyena (H. striata brunnea) is associated with Machero- dus latidens in the cavern of Montmaurin (Haute Garonne). The striped hyena occurs at five other localities in the Pyrenees, Spain and Portugal (Furninha) ; it has also been recognized in Germany (Mos- bach), Austria (Hundsheim) and France (Harlé, 1910, p. 40); it dis- appears later or retires to the south, while the spotted hyena (H. cro- cuta spelea) becomes adapted to the extreme cold and survives with the reindeer to the end of Postglacial times. ‘he Carnivora of this region are Felis leo spelea, the panther (Felis pardus), the wild cat (Felis catus), and the lynx (Felis pardina). SrconD AND THIRD GLACIAL AND INTERGLACIAL Epocus AFRICAN-ASIATIC FAUNA Four great animals especially characterize this fauna: Hlephas trogon- therti and Hippopotamus major, Dicerorhinus merckii and Elephas an- tiquus. Old Elephant (Hlephas antiquus).°°—The “old elephant” or straight- tusked elephant (Hlephas antiquus) does not occur in France or Great Britain until the Second Interglacial Stage, but it is said to occur in the Arno valley of Italy during an earlier stage in which it is associated with a warm fauna including the southern mammoth and the hippopotamus. The typical #. antiquus is recognized by its narrow, elongated grinding teeth with comparatively few plates which, combined with its skull char- acters, suggest its affinity to the modern African (Lozxodonta) rather than to the Indian elephant (Hwelephas) group. While during the first, or Norfolk, interglacial period it is confined to Italy, in subsequent in- terglacial times it wandered into northern Europe as one of the grandest and most distinctive forms, attaining a very wide distribution. Pohlg certainly overestimates its size®® in assigning to it a height of 5 m. at the back (16 ft. 8 in.), or 1m. more than the mammoth, and with tusks also 5m. in length. In consequence of the size and weight of the tusks, the head, shoulders and fore legs were enormously developed. The same writer believes that the habitat of this mammoth retreated and advanced ® PoHLic, H.: “Dentition und Kraniologie des Hlephas antiquus Fale. mit Beitrigen tiber Hlephas primigenius Blum. und Hlephas meridionalis Nesti.”’ Nov. Act. Ksl. Leop.- Carol. Deutsch. Akad. Naturforsch., Vol. LIII, No. 1, p. 326. MHalle, 1888. *®% PoHLIG, H.: Hiszeit und Urgeschichte des Menschen. Leipzig, 1907. a"8 AVVNAES NEW YORK-ACADEMY OF SCIENCES with the successive ice waves and warm interglacial times. Because of the resemblance of the grinding teeth of H. antiquus to those of the African elephant (L. africanus) it has been assumed perhaps too readily that this ancient elephant was characteristic of a tropical climate. It resembles the African elephant in the prominence of the enamel bands of the grinding teeth, which are adapted to the comminution of twigs and woody food, which justifies the belief that this animal frequented the forests. For these reasons Hilzheimer regards 1. antiqwus as indica- tive of forest conditions. Rhinoceroses.—The three great rhinoceroses characteristic of the Hu- ropean Pleistocene are each of distinct geological value. In general D. etruscus belongs to the First Interglacial Stage, D. mercku character- izes the Second Interglacial Stage and most of the Third Interglacial, while D. antiquitatis is distinctive of the Fourth Glacial and the Post- glacial. The two species first named are apparently related to the Sumatran phylum (Dicerorhinus sumatrensis). The D. etruscus of the Val d’Arno, of the First Interglacial and of the early phases of the Second Interglacial is a relatively small animal, distinguished by brachyodont grinding teeth and long, slender limbs, a small anterior and a larger posterior horn. It is remotely related to the Sumatran rhinoceros but differs in the absence of cutting, or incisor teeth. It is essentially a browsing type. Its remains in Mauer are said (Wurm, 1912) to afford a transition to D. mercki. Related to this animal in the Second Interglacial Stage in Great Britain, Germany, France, Italy, there appears the broad-nosed rhinoce- ros known as D. megarhinus, or D. merckw. It resembles D. etruscus in its smaller anterior and larger posterior horn and in the elongation of its hmbs and feet, but differs from it in the possession of relatively long- crowned (hypsodont) grinding teeth adapted to grazing habits. This animal is very widely distributed geographically in the Second and the first half of the Third Interglacial Stage, and is in most localities asso- ciated with remains of the hippopotamus and “old elephant.” Quite distinct from these animals is the woolly rhinoceros (Diceros antiquitatis, D. tichorhinus) which belongs with the colder climates of tundra and steppe conditions and is almost invariably associated with remains of the true woolly mammoth (H. primigenius). Like the above described Sumatran species it lacks the front, or cutting teeth and has in consequence been improperly considered as related to Dicerorhinus, but really belongs to the modern African group of Diceros, resembling espe- clally the species D. simus, with which it closely agrees in its dolicho- OSBORN. REVIEW OF THE PLEISTOCENE 979 cephalic cranial proportions, its long-crowned teeth, and especially in the presence of a square upper lip and very large anterior horn and small posterior horn. It is thus distinguished both by the proportions of its horns and by the characters of its teeth and lips from the two species of Dicerorhinus. It is distinctively a grazing animal. SYNONOMY OF RHINOCEROS GENERA 1. Indian rhinoceros: R. indicus = RHINOCEROS Linnzeus 1758 2. Black rhinoceros of Africa: D. bicornis = DIcERos Gray 1821 &. Woolly rhinoceros: R. antiquitatis (Blumenbach 1799) — Ca@Loponta Bronn 1831 4. Sumatran rhinoceros: D. swmatrensis = DICERORHINUS Gloger 1841 5. White rhinoceros of Africa: D. simus = CERATORHINUS Gray 1867 The names of these three rhinoceroses are almost hopelessly confused in the early literature. The animals converge toward each other in sev- eral characters, namely, in the loss of cutting teeth and in the develop- ment of an osseous septum for the support of the nasal bones. The woolly rhinoceros (D. antiquitatis) is first recorded in Europe in deposits correlated with the Third Glacial or Riss Stage, the Mammut Lehm of Cannstatt (Koken, Schmidt, 1912). Bovines.—The bison (B. priscus) rivalled the mammoth as a wanderer and was able to adapt itself to wide diversities of climate in Europe, Asia and America. Originally of African-Asiatic origin it became thor- oughly acclimated as a Eurasiatic meadow and plains type and may have extended also into the forests like the existing woodland bison (B. atha- basce) of Canada. It is readily distinguished as brachycephalic while its contemporary, the gigantic urus, is long-headed (dolichocephalic), as well as less agile than the bison. In external appearance, as depicted in the very numerous engravings and paintings in the Font de Gaume and other caverns, this animal resembled the existing American bison (B. americanus) more than the still surviving Lithuanian and Caucasian form (B. bonasus). The animal appears in the First Interglacial, or Norfolkian Stage in France. In the Second long warm Interglacial Stage there existed a bison (B. priscus antiquus) which enjoyed a wide distribution. The animal found its way to the Mediterranean islands and gave rise to the pigmy varieties. The wild ox (Bos primigenius) also occurs in the First Interglacial Stage and survived the vicissitudes of the entire Pleistocene Epoch. The “nrus”’ of Czesar survived in its wild state in Europe as late as the seven- teenth century A. D., where it was still to be found in the forests of Poland and in a few game preserves. It then disappeared so completely 280 ANNALS NEW YORK ACADHMY OF SCIENCES that even its popular designation “auerochs” was transferred to the Lithuanian bison.*’ The designations of these two types are therefore very confusing and are distinguished by Kobelt as follows: Urus (Pliny), wild ox, urochs, auerochs (Old Ger- man), tur (Polish), urstier — Bos primigenius Bonasus (Aristotle) wisent or wisont, subr (Polish), auer- ochs (of recent date) — Bos priscus The relations of the wild cattle of Asia to domestication will be con- sidered on a later page. Megaceros giganteus.—During the first half of the Pleistocene this noble animal was widely distributed in Ireland, England, Scotland, the Isle of Man, France, Denmark, Germany, Austria, northern Italy and parts of Eurasia even into Siberia. The famous Megaceros beds of Ire- land are freshwater clays which frequently underly the peat bogs. As observed by Williams these are “boulder-clays” which were redistributed as lake sediments and accumulated under genial or temperate climatic conditions like the present. Owing to the similaritv in the palmation of its antlers the giant deer has been generally (Lydekker, Weber, Trouessart) placed within or very close to the subgenus Dama, the fallow deer; but Lonnberg®* regards the likeness between the giant deer and fallow deer as convergence and considers that the giant deer is more closely related phylogenetically to the reindeer, but it is nevertheless so specialized as to hold an independent place in the system of Cervide. The Megaceros last appears in early Postglacial times associated with the Aurignacean culture in Germany; it is not recorded (Schmidt) with the succeeding Solutrean or MagdaJenian culture. It thus became ex- tinct before the close of the Paleolithic. Elaphine or Red Deer.*°—Sir Victor Buck held that the Cervide orig- inated in Asia and from there spread westward into Europe or eastward into America. The Asiatic origin of the red deer race has since been ably maintained by Koppen. A very large race of late Pleistocene times has been compared by Nehring with the C. canadensis of North America. Reindeer—The reindeer of Pleistocene times are generally referred to the Barren Ground or Tundra type. In this type, which is typified by the existing Old World reindeer (Rf. tarandus, R. spitzbergensis) and by the American arctic forms (Rf. arcticus, R. Grenlandicus, R. granti, R. 8 POHLIG, H.: Eiszeit und Urgeschichte des Menschen, p. 131. Leipzig, 1907. RUTIMEYeER, L.: ‘Die Fauna der Pfahlbauten der Schweiz.’’ Neue Denkschr. schweiz. Ges. gesam. Naturwiss., Vol. xix, pp. 68-112. Ziirich, 1862. KOBELT, W.: Die Verbreitung der Tierwelt. S8vo. Leipzig, 1902. 58 LONNBERG, E.: ‘““Which is the Taxonomic Position of the Irish Giant Deer‘and Allied Races?’ Ark. Zo6l., Vol. 3, No. 14, pp. 1-8. Upsala, 1906. 6 ScHarrr, R. F.: The History of the European Fauna, pp. 246-251. London, 1899. OSBORN, REVIEW OF THE PLEISTOCENE 93] pearyt), the antlers are round, slender and long in proportion to the relatively small size of the animal, while the spreading beam and brow tines are as a rule but little palmated, although in some forms the brow tine is palmated. The woodland type, which is now extinct in Europe, is typified by sev- eral American species’? (2. caribou, R. montanus, R. osborni) in which the antlers are heavier, flatter, thicker, and more heavily palmated on the spreading beam and on the “brow tine” especially, while the tine above the brow, which corresponds to the bez-tine of the stag (Cervus), is elaborately developed and palmated thus contrasting sharply with the simple bez-tine of the Barren Ground group. Some writers’? (Scharff) maintain that the Barren Ground reindeer entered Europe first during the First and Second Glacial Stages while the woodland group first appears in the Third Glacial Stage. Others (Hilzheimer) maintain that all the known Pleistocene reindeer belong to the Tundra form and not to the woodland form. Again, Dietrich” recognizes a woodland caribou in the “high terrace” gravels of Stemheim in the valley of the Murr. Carnivores—The larger Pleistocene carnivores embrace the wolves (Canis lupus), the bears (U. arctos, U. spelea), the hyenas (H. crocuta spelea, H. (brunnea) striata), the leopards (Felis pardus of Spain) and the lions. The chief enemies of the wild horses and cattle of the Pleistocene were the hons (Felis leo spelea), descended either from the great cats of the Pliocene of France and Italy (Felis arvernensis) or more probably mi- grants from northern Africa. These lions are known from deposits in England, Belgium, Austria, southern Russia, France, Spain,” Italy, Sicily, Greece and Algeria.7* The fact that remains of this animal are so often associated with those of the cold Postglacial fauna makes Neh- ring’s’ suggestion seem plausible that the cave lion was a northern race of the recent African and western Asiatic lion adapted to a colder cli- mate and with a heavy coat. After examination of specimens from cen- tral and northern Europe Boule’ reaches the conclusion that these lions 70 GRANT. MADISON: “The Caribou.’’ Ann. Rept. N. Y. Zo6l. Soc., no. 7, pp. 175-196. New York, 1892. 1 Op. cit., p. 154. 7 DinTRicH, W. O.: ‘Neue fossile Cervidenreste aus Schwaben,”’ Jahreshefte des Vereins f. vaterlindische Naturkunde, Jahrg. 66, pp. 320-336. 1910. @ HARLE, EDOUARD: “‘Les mammiféres et oiseaux quaternaires connus jusawici en Portugal. Memoires suivi d’une liste générale de ceux de la Péninsule Ibérique.’’ Com- mun. du Service géol. du Portugal, T. VIII, pp. 22-85, pll. I-V. 1910. 7% BOULE, MARCELLIN: “Les Grands Chats des Cavernes.”’ Ann. de Paléont., Vol. I, pp. 20-27. Jan., 1906. % NEWRING, A.: Uber Tundren und Steppen der Jetzt-und Vorzeit, mit hesonderer Beriicksichtigung ihrer Fauna. Berlin, 1890. 989 ANNALS NEW YORK ACADEMY OF SCIENCES are not related to the tiger (/. ligris) as was supposed formerly by De Blainville and Lartet. While rich in individual variations Felis leo spelwa is nearer the lion than the tiger in most of its characters; it should, in fact, be considered a veritable race of the recent hon with the name Jelis leo spelea. It differs from both the recent lion and tiger In the more gentle and uniform slope of its facial profile and in its large, flat forehead, but its limb bones are longer and proportionately thicker. It sometimes equals and often surpasses the existing lions and tigers in size. It is represented in the cave engravings and drawings both of early and late Postglacial times. The cave hyena (Hyena crocuta spelea) is a variety of the living spotted hyena (Hyena crocuta) of Hast Africa, but it attained dimensions considerably greater than that of its living ally. It has the larger propor- tions, the heavier build, the broad skull, the powerful carnassial teeth which distinguish the spotted from the striped hyena (H. striata) of the present day. Although proportionately heavier the hind limbs may have been shorter than in the spotted hysena in adaptation to the cavern life which the inclement climate made necessary. The cave hyena was a very abundant type and is responsible for scattering of the vast num- bers of the bones of the contemporary animals in a manner not pleasing to the paleontologist. In the caves of southern France a variety (Hyena priscus) of the striped hyena (Hyena striata) also occurs and there are also discovered here additional remains of an animal (H. intermedia) resembling the cave hyena. Thus the Pleistocene species of Huropean hyenas under- went an evolution of their own. As a result the living African forms differ more from the Pleistocene hyenas of Europe than they do from those of Phocene times. Harlé’® records the striped hyena (H. striata) as characteristic of the earlier or warm Pleistocene of Spain and Portugal; the cave hyena (H. crocuta spelea) survived into the late Pleistocene through adaptation to the cold climate. The cave wolf (Canis lupus spelea), a member of the forest fauna, also attained dimensions greater than its living allies. According to Gaudry and Boule,“’ (1892) no constant osteological differences can be determined between the Pleistocene cave wolf and the modern wolf of western Hurope, although the cave form is of considerably larger size. This animal is represented in the Upper Magdalenian paintings of Font de Gaume. 7 Op. cit., 1910, pp. 46, 70. ™ GaupRY, A., and BouLr, M.: Matériaux pour l’Histoire des Temps Quaternaires, Fasc. 4, Les Oubliettes de Gargas. pp. 108-112. Paris, 1892. OSBORN. REVIEW OF THE PLEISTOCENE 233 THIRD GLACIAL STAGE—ILLINOIAN, PoLANDIAN, Riss In North America the Third Glacial Stage is heralded by the advance of a great ice cap radiating from Labrador which sent its glaciers to the south and far southwest, depositing the Illinoian drift which is regarded (Leverett, 1910, p. 315) as of an earlier period than the Polandian or “Middle Drift” of northern Germany or the Riss drift of the Alpine region. This Third glaciation of the Alpine region has a period of ad- vance and retreat which is relatively estimated by Penck at 20,000 years, the snow line descending 1,250m. With it are associated the “high ter- race” deposits of the Alpine region. The Third glaciation was greatly extended along the Rhine, in parts of Switzerland, in France, and in the valley of the Po (Fig. 9). In northern Germany the principal rea- son for separating the “Middle Drift” (Polandian) from the “Upper Drift” (Mecklenburgian) is the presence of loess deposits between them which seems to strengthen the evidence for a Third Interglacial interval. ‘These loess deposits are regarded by certain German geologists (Koken) as the continuation of the “Older Loess” but by Penck and Leverett they have been regarded as belonging to the “Newer Loess.” THIRD GLACIAL FAUNA The recurrence of a cold climate in Germany is heralded in the Upper Sands of Mauer by the arrival of the reindeer and other arctic types. In the Mammut Lehm of Cannstatt is found a fauna which is regarded by Koken and Schmidt (1912, op. cit.) as contemporaneous with the Third Glacial advance. It is noteworthy as containing two new arrivals from the tundras of the north, namely, the woolly mam- moth (H. primigenius) and woolly rhinoceros (D. antiquitatis), as well as the reindeer (Rf. tarandus). The other members of this fauna in- clude two species of horse, the giant deer, the stag, the bison and the urus. “Cannstatt,” observes Schmidt (1912, p. 270), “affords a geologi- cal and final connecting link between the Second Interglacial fauna of Mauer and the fauna of Early Paleolithic [or Third Interglacial] times.” If this fauna actually entered Germany during the cold period of the Third glaciation it returned to the north with the approach of the warm-temperate climate of the Third Interglacial Stage, because no trace of it is found until near the close of the Third Interglacial Stage. THIRD INTERGLACIAL STAGE—Riss-WURM, SANGAMON The Third Interglacial Stage is shorter than the Second, its geologic and faunal characters are more fully known, and it embraces the first 284 ANNALS NEW YORK ACADEMY OF SCIENCES undoubted remains of human stone industry in Europe as well as abun- dant remains of man. In the Alpine region this Riss-Wiirm Stage is indicated by “high terraces,’ which rise 25 to 50m. above the existing streams. ‘The Riss-Wtirm interval is evidently shorter than that be- tween the Second and Third glaciation. Penck considers that the depo- sitions of “Newer Loess” which occurred in the Alps near the close of Riss-Wiirm Interglacial times represent a cold stage, since the fauna which it contains is of the Tundra-Alpine type and the Paleolithic im- plements found in it are closely similar in workmanship to those found in deposits subsequent to the Wiirm glaciation (Penck, 1909, p. 1159). He regards this as the “Newer Loess” that was laid down prior to the Fourth glaciation. Koken and Schmidt, on the other hand (1912), regard the ‘“‘Newer Loess” as partly or wholly Postglacial, that is, as occurring after the Wiirm maximum. The Third Interglacial loess of northwestern Europe was comparatively scanty and discontinuous, from 1 to 5m. in thickness, and contains a terrestrial molluscan fauna as in America. Al] the indications are that this loess was deposited by pre- vailing westerly winds. Also along the Danube the loess is chiefly due to westerly winds. Penck attributes the scarcity of loess in the southern, eastern and western borders of the Alps to the presence of thick vegeta- tion even during the glacial stages, the moraines being pushed out into the forests. In America the Third Interglacial interval is known as the Sanga- mon: the deposits are composed of dark, black soil which is overlain by the main or thickest loess deposit of the central United States. There appears to have been a long interval between the melting of the Third Glacial ice and the deposition of the loess which contains a terrestrial molluscan temperate fauna, indicating climatic conditions not greatly different from those now existing in the same regions (Shimek, 1909). The geologic deposits of this stage are mainly of three kinds: first, fuviatile sands and gravels; second, loess; third, hearth or kitchen-mid- den deposits made by man toward the cold closing period of this stage. CLIMATIC CHANGES DURING THE THIRD INTERGLACIAL STAGE The Third Interglacial Stage opens with a renewal or continuation of climatic conditions favorable to an Asiatic-African fauna exactly similar to that of the warm Second Interglacial period. This warm fauna is known as the “Second #7. antiquus fauna.” It includes the hippopota- mus, the straight-tusked elephant (#. antiquus) and the southern ele- phant (2. trogontherii). This last species is even referred to by many writers as H. meridionalis. ; : OSBORN, REVIEW OF THE PLEISTOCENE I85 This is the Hlephas antiquus Stage of Pohlig, this animal being very abundant until toward the close of the Third Interglacial Stage when it makes its last appearance in Europe. The broad-nosed rhinoceros (D. merckii) is also abundant and appears in Europe for the last time. The successive climatic phases of mammalian life are most clearly recorded in connection with the culture stages of the Lower Paleolithic period, including the pre-Chellean, Chellean, Acheulean, and the begin- ning of the Mousterian cultures. As indicated on p. 233, the warm Asiatic-African fauna prevails from the pre-Chellean until toward the close of Acheulean times, when there is evidence of the advent of a cold dry continental climate, on the approach of which the hippopotamus, - Elephas antiquus, and Dicerorhinus mercku gradually retreat. Thus at Villejuif, south of Paris, the late Acheulean implements are found im- bedded in great drifts of loess, a proof that a cooler, drier climate which marks the transition from the last warm Interglacial Stage to the Fourth Glacial advance was prevalent. Chiefly in the southern parts of France we find the Hlephas antiquus fauna still persisting until the close of the Third Interglacial Stage or during the early Mousterian period, a sign that this old African-Asiatic stock did not become extinct but migrated from central Europe to warmer regions in the south and southwest. Flora—tIndications of changes of climate in the Third Interglacial interval are preserved in the tuf de la Celle-sous Moret (Seine-et- Marne) 7° which overlies Pleistocene river gravels near Paris. The lower levels contain the sycamore maple (Acer pseudoplatanus), willows (Salix), the Austrian pine (Pinus laricia). Higher up in the same de- posits we find the box tree (Burus) and not uncommonly the fig (Ficus) ; the canary laurel (Laurus nobilis) occurs less frequently; the canary laurel and the fig indicate that the winters were mild because both these plants flower during the winter season. The climate was more damp and somewhat milder than that of the present time in this region. The Mollusca of the tufa of La Celle also indicate that the cli- mate of northern France was more equable so as to permit species now widely separated to live together. The plants in the highest levels of the tufa, however, indicate a cooler climate and yield Acheulean flints. The tufa is itself covered by a sheet of loess corresponding to the return of a cool, arid period in late Acheulean times. Jn Lorraine below the level of the fauna of the Fourth glaciation there oceurs a flora in which the most northerly varieties of the larch (Larix) and the mountain pine (Pinus lambertiana) predominate. The lignites of Diirnten and of Utznach near Ziirich (Fig. 9, 18) contain fossil re- 78 Have, op. cit., 1908-11, p. 1812. 286 ANNALS NEW YORK ACADEMY OF SCIENCES mains of forests of Third Interglacial age similar to those which still flourish in the same region, consisting of spruce, fir, mountain pine, larch, birches, yews and sycamores with undergrowth of hazel. These lignitic deposits rest upon the remains of a retreating glacier and are in turn covered with those of another glacier and are therefore intergla- cial.*® With this hardy flora are associated remains of Hlephas antiquus, D. mercku, the urus and the stag. Fauna.—The mammalian fauna is broadly divided into: first, the warm African-Asiatic, which disappears from Europe at the close of the Third Interglacial Stage; second, the Hurasiatic Forest Fauna, in which we now include the urus (B. primigenius) and the bison (B. priscus) ; third, the Tundra Fauna, which retreats after the Third Glacial Stage to reappear with the approach of the Fourth Glacial Stage, when the full tide of Tundra life, including fifteen species of mammals and birds, and the advance wave of Steppe life, including nine species of mammals and birds, first arrive in Europe. The chief localities in which the fauna is recorded are the following: Warm Stage. CHeELLES,® St. ACHEUL, valley of the Somme, northern France. Warm Fauna. Grays THURROCK and ILForD, Essex, Hngland. Warm Fauna. Cool Stage. TAUBACH-WEIMAR-HHRINGSDORF-ACHENHEIM, Germany. Acheu- lean Stage. Temperate Fauna. DURNTEN, UTZNACH, near Ziirich, Switzerland (cool flora, fauna). LAVISTE, AYGELADES, travertines, Marseilles, France (flora). KRAPINA (cavern of), Croatia (fauna and human remains). Cold Stage. RixporF, near Berlin. Cold fauna. PILTDOWN MAN, EOANTHROPUS DAWSONI Fragments of a skull and jaws discovered by Dawson in 1911 near Piltdown, Sussex, have been described by Dawson and Smith Wood- ward.** ‘They were associated in a fluviatile sand layer with a single pre-Chellean flint and remains of deer (? deer), rhinoceros (D. etruscus or D. merckii), beaver (Castor fiber), and hippopotamus. The geologic age is not positively determined by the fauna nor by the nature of the river gravel deposits in which these specimens were found. The associa- 7 DAWKINS, W. B.: “Classification of the Tertiary Period by Means of the Mam- malia.” Quart. Jour. Geol. Soc., Vol. xxxi, pp. 379-405. Aug., 1880. 80 The Chellean culture is placed by Penck and Geikie in the Second Interglacial Stage. 81 DAWSON, CH., SMITH-WoopwarpD, A., Smiru, G. ELiior: “On the Discovery of a Paleolithic Human Skull and Mandible in a Flint-bearing Gravel Overlying the Wealden (Hastings Beds) at Piltdown, Fletching (Sussex). With an Appendix by Prof. Grafton Hlliot Smith.” Quar. Jour. Geol. Soc., Vol. 69, pp. 117-151, Pls. XV-XXI. London, 1913. OSBORN, REVIEW OF THE PLEISTOCENE 287 Ss tion of the pre-Chellean flint and of numerous Chellean flints in the overlying layer would tend to determine the age as either Third Inter- glacial or at the earliest Second Interglacial. The placing of the skull and jaws together as belonging to one indi- vidua! is not certain, but is highly probable. The cranial bones are extremely thick; the skull is not Neanderthaloid, but is of a high, dolicho- cephalic type, with a brain capacity variously estimated at 1,100ccm. (Elliot Smith, 1913) to 1,500ccm. (Keith, 1913).°°* The jaw resembles closely that of an orang (Simia satyrus); the two lower molar teeth preserved are more elongated than in any human type; the superior canine tooth (mistaken by the authors for an inferior canine) resembles that of the anthropoid ape. Thus the specimen may be concisely de- scribed as possessing the skull of a man combined with the jaw and the dentition of one of the higher anthropoid apes. A number of eoliths and one paleolith were also found in the same layer with the skull. PRE-CHELLEAN AND CHELLEAN FAUNA The dawn of the stone industry in Europe is known as the pre-Chel- lean. It is found at Chelles in France, in Spain and at Piltdown, Essex, England. It is important to note that the Chellean culture stage is regarded by Penck, Geikie and others as belonging to Second Interglacial times, or the Mindel-Riss, while Boule, Haug, Obermaier, Breuil and Schmidt assign the pre-Chellean-Chellean culture to the Third Interglacial Stage. While the latter opinion generally prevails among archeologists there is reason for further investigation before the geologic age of the pre-Chellean and Chellean cultures can be considered as definitely determined. The faunal period of the Chellean culture proper is shown in the val- leys of the Somme and of the Marne where mingled with the Pre-Chel- lean and Chellean flints are found the hippopotamus, the southern ele- phant (£. meridionalis trogontherii) with the straight-tusked elephant (Z. antiquus) and the broad-nosed rhinoceros (D. merckii). The typi- cal site of the Chellean culture stage is near the town of Chelles in the wide expanse of the Marne valley. The river deposits of the period of the Chellean culture in this valley are eight meters in thickness and contain beside the animals named above the giant beaver (Trogonthe- rium), species of bear, of hyena, various kinds of deer, larger and smaller kinds of wild cattle and a primitive wild horse. At Abbeville, at the mouth of the Somme, are found remains of the Sig, The corrected and final determination is at 1,300ccm., Smith-Woodward, McGregor. ANNALS NEW YORK. ACADEMY OF SCIENCES (94) 28 same animals associated with those identified as belonging to the sabre- tooth tiger (Machwrodus). Chellean Culture on the “Low Terraces.”—-\t appears that the Chel- lean culture stage In many regions was subsequent to the formation of the terraces; thus Chellean flints may occur in the superficial gravels both on the “middle” and on the “low terraces.” Haug (1912) sup- ports the theory that the Chellean culture belongs to the Third Inter- glacial Stage. This accords with the terrace chronology. The minute researches of Laville in the basin of Paris are confirmed by the observa- tions of Commont in the valley of the Somme. Along the Somme Chel- lean flints occur in the deep gravels overlying the middle and upper ter- races. ‘The fauna found in the “low terraces” of Chelles and of Grenelle is the same, namely, #. trogontheru, Trogontherium cuvieri, Hyena crocuta, D. merckii, Hippopotamus major, Elephas antiquus. All these animals found at Chelles occur in the gravels a few meters above the level of the Marne; they belong exclusively to the sands and gravels at the base of the diggings in the “low terrace.” Similarly in the valley of the Somme near Abbeville from the base of the “low terrace” are re- corded Acheulean flints with H. major, D. mercku, Hlephas antiquus, ete. lLaville collected at Arceuil in the valley of the Biéve, in gravels assigned to the “low terrace,” Chellean, Acheulean and Mousterian flints; these “low terraces” are only 5m. above the river level and are still occasionally flooded with the high waters of the Seine. It is hardly probable that the close geologic and faunistic association of the Chellean- Acheulean cultures in these “low terraces” could have been separated by a very long geologic period, amounting to a hundred thousand vears, as demanded by the theory of Penck. What is regarded as the typical Third Interglacial fauna of the more northern regions of Europe as found at Grays Thurrock, Ilford (Hssex, England) and at Taubach is as follows: Grays Thurrock, Ilford (Essex, Se aaeaeiey ean ee England). — The hippopotamus is antiquus He ; Southern mammoth, #. trogontherii here recorded by Dawkins. The ele- Broad-nosed rhinoceros, D. merckii phant of Essex is referred by Pohlig Hippopotamus, H. major to #. trogontherw. The horse Wild horse, £. AALS (probably (Ewart) is considered to belong to ie ae varie) the Forest or Nordic type. This Wild boar, Sus scrofa ferus : ees Bicone on rises Essex fauna is characteristic of the Urus, Bos primigenius river shores and of the neighboring Red deer, Cervus elaphus forests and meadows. The lions, Roe deer, ©. capreolus hvenas and bears which are re- OSBORN, REVIEW OF THE PLEISTOCENE 289 Giant deer, Megaceros corded here are not true cave types, Piyeenet a crocuid spelad but are in part ancestors of the cave Brown bear, Ursus arctos : : Tigh qe COT types which appear in the succeed- Lion, Felis leo antiqua _ Ing reindeer or cavern period. Wolf, Canis suessi Badger, Meles taxus Marten, Mustela martes Otter, Lutra vulgaris Beaver, Castor fiber Hamster, Cricetus vulgaris The water vole, Arvicola amphibius ACHEULEAN CULTURE FAUNA Warm Stage.—The early Acheulean culture as found at Taubach, Weimar, Ehringsdorf and Achenheim contains the hippopotamus and the straight-tusked elephant (H. antiquus). The principal feature of the early Acheulean culture stage seems to be the abundance of these African- Asiatic animals so that this is commonly known as the “warm Acheulean” fauna. Cool Stage-——The Acheulean culture endured for a long period of time and toward its close two typical members of the warm fauna, namely, the hippopotamus and F. trogontherti, disappear. Thus the late Acheu- Jean fauna does not include either hippopotamus or EH. trogontherii but there still survive the H#. antiquus and the broad-nosed rhinoceros (D. merck). These animals persisted in Europe for a considerable time and becoming adapted to a colder climate are sometimes found in asso- ciation with the advent of the true mammoth (F. primigenius) and the woolly rhinoceros (D. antiquitatis). During /Jate Acheulean times a dry, cool continental climate prevailed (Hilzheimer, 1913, p. 145) similar to that of the steppes of southern Russia between the Ural Mountains and the Caspian Sea. Evidences of this are observed even in the sheltered valley of the Vézére, a tributary of the Dordogne in southwestern France. Similarly as regards north- central France, Obermaier (1912, pp. 122-124) observes that while the climate was mild and temperate and the country still forested at the be- ginning of the Acheulean culture, in late Acheulean times the implements at Villejeuf south of Paris, are found embedded in drifts of loess, a proof that the colder climate which marks the transition from the Third Inter- glacial Stage to the Fourth Glacial Stage was now beginning to prevail. The fauna is still that of H. antiquus and D. mercku. More rarely (Schmidt, 1912) Acheulean paleoliths are associated even with remains 290 ANNALS NEW YORK ACADEMY OF SCIENCES of the woolly mammoth and the woolly rhinoceros, indicating that in northern localities the Acheulean culture reached the cold period of the Fourth Glacial Stage. Krapina Neanderthaloid Race-——TYo the Acheulean Stage there is re- ferred a human tooth found at Taubach. Of much greater importance is the presence of abundant skeletal remains of men of a primitive Nean- derthaloid race found in the cave-shelter of Krapina in Croatia. These remains are positively associated with the Acheulean stage by Schmidt (1912, p. 256) but they are regarded as more recent, of the late Mous- terian culture stage, by Breuil. The remains as finally described by Gorjanovic-Kramberger®* include hundreds of human bones intermingled in various separate strata with hundreds of stone implements and chips and thousands of animal bones. Of the contemporary fauna are re- corded Ursus speleus, Bos primigenius, Equus ? caballus, Dicerorhinus mercku, Megaceros euryceres, Castor fiber, Arctomys marmotta. The human racial type is unquestionably related to that found at Neander- thal and Spy. The race is somewhat dwarfed, of broader head form, with less prominent supraorbital processes. The species is Homo nean- derthalensis. Mousterian Culture, Temperate Fauna.—The earliest strata of the Mousterian culture stage in France show a fauna not differing essen- tially from that of the late Acheulean stage, namely, a fauna containing Elephas antiquus and Dicerorhinus merckii. Thus in La Micoque, one of the oldest stations in the Vézére valley, Dordogne, in which the cul- ture belongs to the transition between late Acheulean and early Mous- terian times, in the very lowest layers are found traces of the broad- nosed rhinoceros (D. merckw) associated with remains of the moose (Alces). But the last glacial stage is approaching and D. merckii gives place to the migrants from the tundra region of the northeast, covered with hair, adapted to an arctic climate, namely, the mammoth and the woolly rhinoceros. The main succeeding portion of the Mousterian cul- ture was contemporaneous with the Fourth Glacial Stage and the cold tundra, steppe fauna. 82 GORJANOVIC-KRAMBERGER, Karu: ‘Der Diluviale Mensch von Krapina in Kroatien Hin Beitrag zur Palaioanthropologie. Studien iiber die Entwickelungsmechannik des Primatenskelettes mit besonderer Beriicksichtigung der Anthropologie und Descendenz- lehre.” Herausgegeben von Dr. Otto Walkhoff. C. W. Kreidel’s Verlag, 4to. Wiesbaden, 1906. OSBORN, REVIEW OF THE PLEISTOCENE 2971 FoturTH GLActaAL STAGE—WURM, MECKLENBURGIAN, WISCONSIN BEGINNING OF THE REINDEER AND CAVE PERIOD The Fourth Glacial Stage, like the First, is believed to have been nearly contemporaneous in Europe and North America, consequently the estimates of Postglacial time in one country have an important bearing on the other. The First Maximum of the Fourth or Warm glaciation in the Alps is estimated by Penck as occurring 40,000 years ago. It was followed by the slight recession known as the Laufenschwankung, a tem- perate retreat followed in turn by the Second Wiirm Maximum, which is estimated as occurring 20,000 years ago. Similarly in America the “early Wisconsin” is followed by a recession interval (Peorian), and this in turn by the “late Wisconsin” which is the final great glaciation in America. The contemporaneous Mecklenburgian of the North German lowlands gave rise to the “Upper Drift,’ which in some respects bears a striking resemblance to the Wisconsin Drift of America both in its sys- tems of moraines and in its topography. This stage also includes appar- ently the “Upper Drift” of northern England with which the drift of the Alps correlates well. The Upper Drift of England covered also a large part of Wales. In Germany glaciation also occurred in the Rie- sengebirge and the Black Forest. In America part of the “Upper Drift” is loess-covered and: in the opin- ion of Koken and Schmidt the Upper Drift of Germany is also partly covered with the “Newer Loess.” The Postglacial Stage did not exhibit a steady amelioration of climate after the culmination of the Fourth Glacial Stage, but there is evidence of great oscillations and renewed glacial advances both In northern Britain and Scandinavia and in Ger- many. These Postglacial advances, as most clearly defined in the Alpine region, have been termed by Penck the Biihl, the Gschnitz and the Daun. They are correlated by paleontologists and anthropologists quite closely with the successive faunz and archeological implements of Postglacial time. Period of the Final Glacial Maximum.—The length of time which has elapsed since the close of the Fourth great glacial advance is estimated in America by the recession of the Falls of Niagara. This recession began with the end of the Wisconsin glaciation which is believed to have been contemporaneous with the Wiirm. As early as 1829 Bakewell esti- mated that since the Falls of Niagara were receding by the erosion of the Niagara gorge at the rate of three feet annually about 10,000 years had elapsed since the end of the Glacial Epoch. Lyell visited Niagara in 1841 and after consideration of all ihe data of erosion concluded that 999, ANNALS NEW YORK ACADEMY OF SCIENCES the time since the last great glacial stage was not less than 31,000 years. Gilbert, Upham and other geologists of the United States Geological Survey after pointing out many sources of error in all such calculations were inclined to the adoption of periods ranging from 6,000 to 10,000 years. Gilbert’s survey is the most careful which has been made; he estimated (1896) that the gorge of Niagara is not more than 7,000 years of age. The Canadian geologist Spencer arrived at a result almost iden- tical with that of Lyell, namely, 32,000 years. The most careful estimates on the subdivisions of Postglacial time in Europe are those of Penck (1909, p. 1168) which may be briefly sum- marized as follows: TV. 2nd Wtirm MAxIMUM, beginning of Upper Palo- lithieculturesisse oes se Oe sh oie ee cle shane bsioeine see ee OOOO On A OOO eats Achen retreat, period of Aurignacean and Solutrean CUO ee ae ero ee con ame ie net sktet ot eaeuloe eit ncrene homer Biihl advance, period of early Magdalenian culture (GONE ESKGI a) ieee Pn AO Rater NHLBI Aaa ins cairly Rana ate C 16,000 “ 24.000 * Post-Btihl, period of late Magdalenian culture..... 10,000 ‘ 16,000 ‘ Daun Stage, period of the close of the Upper Palzeo- Ihsan, AVAlbiENn OVligWee Gog ooA doo oodoaddoucsouGue 7,000 re ANAS OVE (COOOL) Ty IDWULO Ns cocosauoaboasccoduooSoce 4,000 “ 5,000 * According to Heim’s** calculations the period since the deposition of the Bihl moraines in the Lake of Lucerne amounts approximately to 16,000 years and if the Magdalenian culture deposits of this formation are of the age of the Bithl advance we my estimate with Netisch that the Buhl advance occurred at least 24,000 years ago. This advance is a very important period because it represents the last of the Arctic, Tundra and Steppe Faunas in central Europe prior to the establishment of the modern forested conditions and fauna. Fauna of the Fourth Glacial Stage.—The severe climate of the Fourth Glacial Stage is indicated by the mammalian life found at Achenhemm and Sirgenstein associated with what is known as the “full Mousterian” Paleolithic culture (Schmidt, 1912). Here for the first time there is a predominance of the Tundra Fauna (fifteen species) as well as the ad- vent of the Steppe Fauna (two species), while the Forest Fauna (seven species) and Meadow Fauna (four species) are still represented. The straight-tusked elephant (#. antiquus) and broad-nosed rhinoceros (D. merckw) have disappeared and are now replaced by the woolly elephant (EL. primigenius) and woolly rhinoceros (D. antiquitatis). The mam- 3S Heim, A.: “Uber das absolute Alter der Hiszeit.”’ WVierteljahrsschrift der natur- forsch. Ges. in Ziirich, Jahrg. 39. No. 2, pp. 180-186. 1894. OSBORN, REVIEW OF THE PLEISTOCENE 993 mal life of the Fourth Glacial Stage at Achenheim and Sirgenstein near the upper Rhine is divided among the various faunz as follows: TUNDRA Blephas primigenius Diceros antiquitatis Rangifer tarandus Vulpes lagopus Lepus variabilis Myodes torquatus STEPPE Equus germanicus (Steppe type) Spermophilus rufescens FOREST Cervus elaphus Lynchus lyn Canis lupus Vulpes alopex Arvicola amphibius Ursus speleus ALPINE Arctomys marimotta Ibex sp.? Lagopus alpinus MrEapow Bison priscus Bos primigenius ASTATIC Hyena spelea Felis leo spelea This “Lower Rodent” After the First Maximum of the Fourth glaciation the Laufen- schwankung may have temporarily favored the return of the Hlephas antiquus and D. merckii fauna as far as northern France because we occasionally find a glacial mixed fauna where H. antiquus and D. merckw occur in association with LH. primigenius. The close of the Fourth Glacial Stage or Second Wiirm Maximum is marked by the first appearance of very numerous arctic rodents, especially of the banded lemming (Myodes torqua- tus) type, which constitute the so- called “Lower Rodent” layer. The animals (Schmidt, 1912) character- istic of this Lower Rodent stratum as found at Sirgenstein, Wild- scheuer and Ofnet are as follows: TUNDRA Canis lagopus Fetorius erminea with a fauna such as we find at pres- ent in far northern Russia registers the coldest climate of Pleistocene times, corresponding to the Second Maximum of the Fourth Glacial Stage. It is well known that this Lower Rodent fauna is not local but rather a widespread phenomenon ex- Lepus variabilis Myodes obensis “f torquatus Lagopus albus FOREST Arvicola amphibinus MEADOW Talpa europea tending over northern and southern Germany and Belgium (Schmidt, 1912, p. 261). The presence of this Lower Rodent fauna at Thiede near Braunschweig in the border region of the Upper Glacial Drift of Germany is of special significance, as Wahnschaffe observes. This is the classical locality for lemming, the remains of lemmings being associated solely with those of arctic fox, arctic hare, reindeer, musk ox and mammoth. 994 ANNALS NEW YORK ACADEMY OF SCIENCES Upper Sands of Mawer.—In the Upper Sands of Mauer, Wurm (op. cit. 1913) and Férster** have observed the presence of a sub-Arctic cold fauna including the arctic reindeer (f. tarandus) and the banded lem- ming (Myodes torquatus), the steppe suslik (Spermophilus rufescens), also the steppe weasel (Ietorius eversmanni). This is positively corre- lated with the fourth glaciation by Wurm. The fauna of the Wildkirchli cave on Mont Centis in Switzerland shows no typical interglacial forms and may indicate either the approach of the Wiirm Glacial period or its Laufenschwankung (Schmidt, 1912, p93): : Mousterian Paleolithic Cultuwre-—The Mousterian Paleolithic culture ot the Neanderthaloid race appears to have begun toward the close of the Third Interglacial Stage and continued throughout the entire period of the Fourth Glacial, including the First Maximum, the Laufenschwan- kung, and the Second Maximum, ending with the “Lower Rodent” fauna in the deposits of the “Newer Loess.” Thus Koken and Schmidt speak of the Mousterian culture as found at the base of the “Younger Loess.” The early men of the Mousterian culture period witnessed in the north the Hlephas antiquus and broad-nosed rhinoceros, and the hippopotamus in the south, as proven in the Prince’s Cave of Monaco. The culture lasted through all the severe climatic conditions of the entire Fourth Glacial Stage. The men of the Neanderthal race in Mousterian times sought the rock shelters and grottos or entrances to caverns so that the greater number of paleoliths are found mingled with the remains of ani- mals of the chase around the old hearths. The commonest game evi- dently consisted of the wild horse, urus and reindeer. Less frequently the ibex and stag were objects of the chase. a Neanderthal Races.—To the Neanderthal race (Homo neandertha- lensis,*°) in addition to the remains of Krapina, belong many and widely distributed remains, including the classic type of Neanderthal discovered in 1856, the Gibraltar skull discovered in 1848, those of La Naulette, Spy (1887), Krapina (1902), Malarnaud, La Quina, Le Moustier (1908) and La Ferrassie. The skull of Neanderthal man was character- & WOrster, F.: “ther eine diluviale subarktische Steppenfauna aus den Sandhiigeln von Mauer.” Verhandl. naturw. Verein. Karlsruhe, 1913. s> The mid-Pleistocene man was definitely named Homo neanderthalensis by Cope in 1893 at the time of the discovery of the skeletons of Spy; it seems, however, that King had previously (1864) used the same term. Schwalbe (1901) remarks, . . . “the species of man which I, together with King (1864) and Cope (1893), designated as Homo neanderthalensis.” In the following year (1902) the same author introduces the term Homo primigenius, which is that generally adopted in Germany. Among French authors the same man is known to-day as Homo mousteriensis. This polynomial usage serves at least to emphasize the unanimous opinion as to the distinct specifie character of mid-Pleistocene man. OSBORN, REVIBW OF THE PLEISTOCENE 295 ized by an extremely receding forehead, by the great prominence of the supraorbital ridges, and by a rather slender jaw. The occipital projec- tion for the attachment of the superior muscles of the neck was large, indicating that these muscles at the back of the neck were strongly devel- oped, a character necessary to meat-eaters before the invention of knives 1G. 16.—Neanderthal skull Skull of a Neanderthal man from the cavern of La Chapelle-aux-Saints (Corréze), France. After Boule. and forks. This primitive type of man was shorter than the average European (that is, 5 feet 814 inches) ; he is estimated of as low stature as 5 feet 3% inches. His lower limbs were especially powerful, but his gait seems not to have been fully erect, for the knees are bent slightly forward. 296 ANNALS NEW YORK ACADEMY OF SCIENCES The most remarkable skull and skeleton of Mousterian age is that (Fig. 16) found by the Abbés J. and A. Bouyssonie and L. Bardon in the cavern of La Chapelle-aux-Saints (Corréze) in 1908, associated with stone implements and remains of the reindeer, urus, ibex and woolly rhinoceros. The cranium is dolichocephalic, with prominent supra- oR Ke? gat ig. 17.—Neancerthal man Reconstruction of the head of Homo neanderthalensis by Charles Rt. WKnight under the direction of the author. 1910. orbital processes and relatively short and broad nose, weak lower. jaw, lacking the prominent chin process. These characters, as well as the posterior position of the foramen magnum and the form of the palate, are distinctly simian or pro-human.*° POSTGLACIAL STAGE—CONTINUATION OF UPPER PALMOLITHIC, REINDEER OR Cave PERIOD CLIMATE For a long period the fauna of Postglacial time in Europe remained practically the same, namely, during the Mousterian, Aurignacian, Solu- trean and Magdalenian culture periods. The cold fauna is shown both in the animal remains and in the art, which is so characteristic of Aurignacian and Magdalenian times. As the “Lower Rodent” layer of Mousterian times is referred to the Second Maximum of the Fourth gfaciation and the period of most intense 86 BouLE, M.: “L’Homme Fossile de la Chapelle-aux-Saints,’ L’Anthropol.. Vol. MIX, pp. 519-525. 1909. OSBORN, REVIEW OF THE PLETSTOCENE 297 eold it is followed by an amelioration of climate (Achenschwankung) during Aurignacian and Solutrean times, which in turn is succeeded by a recurrence of a colder climate during the Buhl advance, at which stage it is believed the “Upper Rodent” laver of Magdalenian times indicates a fresh invasion of Arctic-Tundra types. This tundra “Upper Rodent” layer eccurs in the deposits of Sirgenstein, Wildsheuer and Ofnet; it is associated along the upper valley of the Danube with early Magdalenian paleoliths, and continues in Hohlefels, Schussenquelle, Andernach, Miin- zingen and Wildscheuer in association with the Middle Magdalenian cul- ture stage; there is, moreover, in Hohlefels, Schmiechenfels and Propst- fels an association of the cold steppe fauna with a late Magdalenian cul- ture stage. After this there was a decided amelioration of chmate, for in the final Azihan-Tardenoisian stage all the Steppe, Tundra and Alpine types disappear. UPPER PALBHOLITHIC, FOUR OR FIVE HUMAN RACES (1) The Grimaldi Race, a negroid type is found close to the warm faunal level at Mentone. (2) An initial feature of the long Postglacial Stage is the entire disappearance of the Neanderthal race of men and the invasion of Europe by a new race of the much higher Crd-Magnon type referred to the existing human species (Homo sapiens). This race was discovered in Dordogne at the hamlet of Cré-Magnon from which it takes its name. It has also been found in the caverns near Mentone lying above remains of the Grimaldi race. Créd-Magnon re- mains are also found at several localities in France in deposits of Aurig- nacian and Magdalenian times. In the south in Aurignacian times this was a tall, well-proportioned, hunting and nomadic race; in the more severe climate of the north it occurs as a dwarfed race. The skulls are dolichocephahc, with very broad cheek bones, hence designated as dishar- monic. The brain capacity is equal to that of the early Neolithic races and surpasses that of many modern races. It is probable but not posi- tively demonstrated that to this Cré-Magnon race we owe the early arts of engraving, sculpture, design and mural painting which are preserved in the Aurignacian and Magdalenian deposits of France and northern Spain. (3) In Solutrean times there was at Briinn and Predmost (Mo- ravia) another dolichocephalic race which is considered as distinct or transitional because unlike the typical Cré-Magnon race the cheek bones are narrow and the skull is thus harmonic. This is the Briinn-Prédmost race. (4) After the close of the Magdalenian culture stage, or during the so-called Azilian-Tardenoisian culture, the first brachycephalic hu- man races make their appearance in Europe. heir remains haye been 298 ANNALS NEW YORK ACADEMY OF SCIENCES found at Ofnet, Furfooz and at Grenelle. This is commonly known as the Grenelle or Furfooz race; it is very distinct from the preceding races in bodily structure and in culture. Aurignacian, First Upper Paleolithic Culture Stage-—This first of the Upper Paleolithic culture stages is widely distributed in western Europe. It takes its name from the small grotto of Aurignac (Haute Garonne) where the first discoveries of the culture and of a number of skeletons were made in 1852. The arts of engraving on bone and stone, of drawing and painting in single hnes, of sculpture of the human and of animal figures, all in bold but primitive forms, first appear mm Aurig- nacian times. ‘hus man through his art begins to make a permanent record of the contemporary mammalian life, especially of the mammoth, bison, reindeer and cave bear. With early Aurignacian times the cold climax is passed but we still find remains of the Arctic lemming (Myodes torquatus) fauna. The mammalian list of the Aurignacian stations both of the “Newer Loess” and of the caverns still gives a cold aspect with its Tundra-Steppe-Alpine types with which no warmer types are associated. In middle and late Aurignacian times the lemmings for a time disappear ; otherwise the fauna retains its northern character (Gulo luscus, Lagopus alpinus), which is not essentially altered by the presence of the hyena and stag. Solutrean, Second Culture Stage-——This stage, which takes its name from the type station of Solutré (Sadne-et-Loire) represents the climax of perfection in the Upper Paleolithic flint industry, which appears to represent partly a development of Aurignacian workmanship and partly a culture invasion. With Solutrean times Schmidt correlates the three Briinn skeletons and Prédmost (Moravia). It is noteworthy that no evi- dences of a Solutrean art have been discovered. The fauna like that of the Aurignacian represents an amelioration of the extreme cold of the Fourth Glacial maximum. ‘The wild horse and reindeer are abundant as well as the mammoth, rhinoceros, wolf and cave bear. Perhaps the somewhat more frequent appearance of such cold faunal types as the Alpine hare and grouse betoken the approach of the colder Bihl stage of Magdalenian times. Magdalenan, Third Culture Stage and Fauna.—This third Upper Paleolithic culture takes its name from the station of La Madeleine (Dordogne). It is distmguished by decline in the perfection of the flint industry as compared with the Solutrean stage, by a very decided deyel- opment of bone implements, and by a surprising advance in the arts of engraving and painting and the sculpture of animal forms in bone and ivory. The Magdalenian stage corresponds with the “Upper Rodent” OSBORN. REVIEW OF THE PLEISTOCENE ra) 99 strata which registers a period of extreme cold correlated with the Biihl glaciation (Penck, Schmidt), which again attracts the cold-loving ro- dents. The “Upper Rodent” strata associated with Magdalenian cultures in the shelters of Schweizersbild and Kesslerloch are of more recent date than the neighboring “Lower Terraces.” This “Upper Rodent” stratum contains an abundance of Tundra and Steppe types of smaller rodents and represents the last stage of extreme cold in Europe. Thus the banded lemming (Myodes torquatus) corresponds with early Magda- lenian times. In the upper levels of the Upper Rodent layer, which be- long to late Magdalenian times, the Tundra fauna gradually gives way to a more abundant Steppe fauna, the banded lemmings becoming less frequent while the jerboas (Alactaga jaculus), the hamsters (Cricetus pheus), the susliks (Spermophilus rufescens) become more abundant. The reindeer and the wild horse are very abundant. The mammoth, the woolly rhinoceros and the cave bear gradually retire from the middle and southern mountains of Germany, and in the very highest Magdalenian culture lavers the fauna begins to approach that of recent times, namely, the Eurasiatic Forest fauna. In Schmidt’s opinion there is no question as to the similarity of age of the Magdalenian layers of the Miinzingen loess deposits with the cave deposits of Schweizersbild and Thaingen. -From this evidence it can be positively determined that the chief deposits of the “Newer Loess” occurred after the Fourth glaciation. With Magdalenian times are associated the skeletons of La Madeleine, Laugerie Basse, Chancelade, La Hoteaux and Duruthy; all are regarded as of the Cré-Magnon racial type. POSTGLACIAL FAUNA ~ .The mammal fauna of this long Postglacial period is the same in tha upper valleys of the Rhine, the Danube, the Dordogne and the Garonne. Tt extends throughout the Pyrenees and the Cantabrian Alps of northern Spain. Even the reindeer myaded this region (Harlé) but the stag (C. elaphus) is more abundant. The moose (Alces) also invaded the Pyre- nees and northern Spain. The Saiga antelope (S. tartarica) occurs at thirteen localities in southern France (Harlé), and the steppe sushk (S. rufescens) is very abundant. It is a grand assemblage of the European Forest and Meadow types mingled with a few Eurasiatic-Alpine types, abundant Hurasiatic Forest types, but with the Tundra and Steppe types predominating numerically until the close of the Magdalenian period, when the Forest types again begin to greatly predominate. The numerical succession in Germany and Austria may be tabulated from the invaluable tables presented by 200 ANNALS NEW YORK ACADEMY OF SCIENCES Schmidt. We owe our knowledge of the cold fauna of southern France chiefly to Harlé (1871-1912). n n n n n am a @ Be om £5 oS) oo ue) FEMS) 8 88 820 See Ba Ou an Sn an ; Me: : ee ’ a m2 4 a i Vostglacial, Daun Stage, Azilian-Tardenoisian Cul- (HUURDS Olimar dCi Oi caine cla Han o-GRmiod oSloiee 6.aino-o 7 Postglacial, Gschnitz Stage, Hohlefels, Late Mag- Geukeamignie CmlnbiReynsoocosdso cass oomononooS ool Od 7 6 il 5 ae Fostglacial, Middle Magdalenian Culture, Upper RO CLETIU CNS UI DEE Ne ses octave cee eats RR arate n ep oneie esa eateries 10 a 6 Ss Postglacial, Early Magdalenian Culture, Biihl StacevsSiGeenshelnin yee iacri rei ico ieee ten: 9 5 ms 5 3 Postglacial, High Magdalenian Culture, Thaingen. 15 14 4 13 21 Postglacial, High Magdalenian Culture, Schwei- ZOTS WMG aN a ise crcdes icteric praia Gace aetna Tay ei eee RN aOR rae (G 2 ® 6 8 Postglacial, Solutrean Culture...............-.:. 6 3 2, 2 Postglacial, Late Aurignacian Culture........... 10 4 1 1 5 Postglacial, Aurignacian Culture, Sirgenstein, (OAS Mites grea ae ine rue erate Urea a a tOe nA Geena ahalenay tee rf i 1 1 3 LV. Glacial Maximum, Lower Rodent Layer..... 7 | as 2 3 IV. Glacial Stage, Mousterian Culture........... 8 5 1 2 4 The fauna of Voklnshofen includes a similar intermingling of tundra, steppe, mountain, and meadow-forest types. The same is true of the scattered deposits*’ in Thuringia near Saalfeld, Gera, Jena, Leipzig, ete. *“ also includes twenty species of mammals divided into typically modern tundra forms of northern The loess fauna near Wiirzburg, Bavaria, Asia, typical modern steppe forms of central Asia and Siberia, together with the four characteristic great mammals of the period, the mammoth, the woolly rhinoceros, urus and bison. ‘The arctic character of the fauna of Chateauneuf-sur-Charente*’ in central France is very conspicuous, most of the species belonging either to the tundras or the steppes of modern Europe. The bones of many young animals occur in this deposit, which may be explained perhaps on the supposition that the animals fell into the fissure while the opening was hghtly covered with snow, the young being the most frequently entrapped. Among the chief localities where this grand Fourth Glacial and Postglacial fauna have been dis- covered are the following: * Ponuic, H.: “Vorliufige Mittheilungen iiber das Plistocen, insbesondere Thiirin- gens,” Sitzungsber. Niederrhein. Ges. Bonn, pp. 2-15. Mar. 3, 1884. 88 NEHRING, A.: “Ubersicht iiber vierundzwanzig mitteleuropiische Quartir-Faunen,” Zeitschr. deutsch. geol. Ges., pp. 468-509. Jahrg. 1880. 8) BouLE, M., and CuHaAuyrt, G.: “Sur lexistence dune faune d’animaux arctiques dans la Charente 4 l’époque quaternaire,’ C. R. Acad. Sci. Paris, Vol. XXVIII, pp. 1188- 1190. 1899. ene OSBORN. REVIBW OF THE PLEISTOCENE 301 Localities Culture Zones KESSLERLOCH Cave, near Thaingen (Fig. 9, 37) ScHWEIZERSBILD Cave. near Schaffhausen (57) Seattered deposits in Thuringia, in northern Germany, Saalfeld. Gera. Jena, Leipzig (29, 30. 31) Wiurzpure, Bavaria, Loess deposits (33) SwaBia and FRANCONIA, cave deposits VGKLINSHOFEN, Alsace (28) Magdalenian Stage Magdalenian Stage Aurignacian and Solu- trean Stages Solutrean Stage Aurignacian and Solu- trean Stages MonrMaAvrin Cave (Haute-Garonne), Upper levels (35) Magdalenian Stage CHATEAU NEUF-SUR-CHARENTE (Charente) (36) The large mammals of the period are more fully known than in any previous Pleistocene stage both through paleontological researches, which date back to the beginnings of the science of vertebrate paleontology in Germany and France, and through the extraordinarily accurate carvings and drawings in the caverns of Dordogne and northern Spain. These drawings have been reproduced with remarkable fidelity, chiefly by Breuil.°° The chief elements in the larger mammal fauna were as fol- lows: ForREST AND MEADOW FAUNA Moose, Alces palmatus Persian deer, Cervus maral Red deer, Capreolus capreolus Roedeer, Cervus elaphus Wild cattle. Bos primigenius Bison. Bison priscus Forest horse. Hquus caballus Bear. Ursus speleus Lion, Felis leo spelea Brown bear, Ursus arctos Wild boar, Sus scrofa TUNDRA FAUNA Woolly rhinoceros, Diceros antiqui- tatis Woolly elephant, Elephas primigenius Musk ox, Ovibos moschatus teindeer. Rangifer tarandus Schwervzershild Cave (Fig. 9, 37).—The Magdalenian, or rein- deer man apparently arrived in the Schaffhausen region long after the maximum Fourth gla- ciation, during the Biihl advance, the period of deposition of the “Upper Rodent” layer with its cold Arctic and Steppe fauna. The deposits of the Schweizers- bild cave as recorded by Neh- ring*? present the fullest succes- sion and extend over a very long period of time as exhibited in the Aretic fox. Vulpes lagopus Elasmothere, Hlasmotherium sibiri- cum following layers: Neolithic : 5. Gray culture layer, forest fauna. % See publications of Abbé Henri Breuil, Marcellin Boule, Le Comte Begouen, P. #1 NEHRING, A.: Bourrinet, L. Capitan. Emile Cartailhac, Lalanne, Lartet and Christy, Déchelette, Hugo Obermaier, Péyrony, Reinach. Others are in preparation. “ibersicht itiber vierundzwanzig mitteleuropiiische Quartir-Faunen.” Zeitschr. Deutsch. Geol. Ges., pp. 468-509. - “Die kleineren Wirbeltiere vom Schweizersbild bei Schaffhausen.” allg. schweiz. Ges. gesim. Naturwiss., Vol. xxxv, pp. 41-77. 1880. Denkschr. 1896. 302 ANNALS NEW YORK ACADEMY OF SCIENCES STEPPE FAUNA Paleolithic, Magdalenian : Saiga antelope, Antelope saiga 4. Upper Breccia, or “Upper Ro- Asiatie wild ass, Hqguus hemionus dent” layer, steppe fauna. Wild horse, EZ. przewalskii ? sp. 3. Yellow culture layer, palzolithie Jerboa, Alactaga jaculus “Reindeer Age,” steppe fauna. bo . Lower Breccia, or “Lower Ro- dent” layer, animal remains ALPINE FAUNA and traces of man, tundra Ibex, lbex priscus fauna. Chamois, Rupicapra 1. Diluvial layer. No fossils. Desert horse, H. caballus celticus (2) Of these the “Lower Rodent” layer contains a pure arctic tundra fauna, such as the vole, hare, fox, the reindeer, the ptarmigan. There are no traces of man. In the layer above these the early steppe animals begin to appear, the hamsters and picas. (3) Then in the “Yellow Cul- ture’ layer there is an assemblage of pure steppe forms, susliks, dwarf picas and wild horses, all pointing to the absence of forests; but at the top of this layer the first squirrel (Sciwrus) appears as the harbinger of forests. (4) In the “Upper Rodent” layer the steppe fauna begins to be intermingled with an increasing number of forest types, such as squir- rels, dormice, and the pine marten. (5) Finally we reach the “Gray Culture” layer, composed of the modern forest dwellers, such as the squir- tel, the beaver, the pine marten, the stag, the roe, the wild boar, the brown bear. The “Lower Rodent” layer is contemporaneous with the Mousterian culture, while the “Upper Rodent” layer belongs to Magda- lenian times. The uppermost “Gray Culture” layer with its forest fauna belongs either to the closing Paleolithic or to Neolithic times. Kesslerloch Cave.—Similar conclusions result from the study of the geologic conditions surrounding the Kesslerloch Cave of Thaingen in Switzerland (Fig. 9, 37). This famous cave les on the edge of a moderately wide valley, traversed by a brook.®? In this sheltered, well- watered, hilly region, woods flourished and harbored the forest animals, at the same time that the glaciers retreating southward left damp and stony areas, closely followed by a tundra fauna. The woolly rhinoceros and the mammoth persisted longer here than in other parts of Hurope. As analyzed by Niiesch, we discover here mammals distinctive of the tun- dras, of the steppes, of the modern Alps (marmot, chamois, ibex), of the meadow-forests (bison, urus), and finally of the modern forest type (lion, wolf, brown bear, pine marten, squirrel, wild boar, and stag). These mammal zones undoubtedly correspond with the passing or evolution of ®2 NUmscH, J.: “Das Kesslerloch, eine Hoéhle aus paladolithischer Zeit. Neue Grabungen und Funde.” Neue Denkschr. allg. schweiz. Ges. gesam. Naturwiss., Vol. XXXIX, Pt. 2, pp. 1-72. 1904. ; OSBORN, REVIEW OF THE PLEISTOCENE 303 several human culture stages (perhaps the Aurignacian, Solutrian, and -Magdalenian). While the tundra fauna was pushing southward into the heart of Switzerland, it had already vanished from central Germany, Belgium and France, where it had been superseded by a steppe, or even a meadow-forest fauna. The human artifacts show that these deposits are contemporaneous with those of Schweizersbild, both belonging to Magdalenian times. A hearth, with ashes and coals, and many charred bones of old and young mammals, including the woolly rhinoceros, have _been found here. The human remains show that a race of pigmies dwelt here smaller even than the small men of Schweizersbild, their height being estimated at 120cm. (4 feet).°* The horse of Kesslerloch shows many resemblances to the Przewalsky horse of the high steppes of Cen- tral Asia.** It is characteristic of these faunas that among species still living are mingled remains of the great extinct mammals of the times. Another feature is that occasionally the Steppe, Tundra and Forest faunas are found either nearly pure or entirely distinct and separate as in the lower deposits of Thiede near Braunschweig, above cited: More often as in Schweizersbild and Kesslerloch they are successive or superposed upon each other. Beside the cavern deposits are those in the loess. Thus in the “Upper Loess” near Wiirzburg, Bavaria, Nehring®® has recorded both a Tundra and a Steppe fauna, including beside the still living types the woolly rhinoceros, the mammoth, the urus and the bison. MIGRATIONS OF THE LARGE MAMMALS OF THE FOURTH GLACIAL AND POSTGLACIAL PERIOD Over the greater part of the Iberian Peninsula the stag (Cervus elaphus) took the place of the reindeer. There is no trace of the en- trance of the Steppe Fauna at any period into Spain or Portugal. The Pyrenees also presented a barrier to the greater part of the tundra fauna, yet the Norwegian lemming (Myodes lemmus) penetrated Por- tugal to the vicinity of Lisbon. The cold fauna (2. primigenius, E. antiquitatis, U. speleus, F. spelea) is not represented in Portugal, but EH. primigenius has been discovered in two localities on the extreme northern coast of Spain, in the Province of Santander bordering the Bay of Biscay. D. antiquitatis also occurs in the same _ province. 2% NUESCH : op. cit., p. 21. % STUDER, T.: “Die Knochenreste aus der Héhle zum Kesslerloch bei Thayngen,’’ Neue Denkschr. allg. schweiz. Ges. gesam. Naturwiss., Vol. XXXIX, Pt. 2, pp. 73-112. 1904. % NEHRING, A.: “Ubersicht tiber vierundzwanzig mitteleuropiische Quartir-Faunen,” Zeitsehr. Deutsch. Geol. Ges., pp. 468-509. 1880. 304 ANNALS NEW YORK ACADEMY OF SCIENCES Rangifer tarandus is found in the cavern of Serinya south of the Pyre- nees (Torralba). It also has been recorded recently (Obermaier) in the cavern of Altamira, near Santander. The Alpine chamois (Rupi- capra) occurs south of the Pyrenees and the ibex is traced to Gibraltar. Thus Harlé’® concludes it is certain that the “cold fauna” of France spread along the northern coast of Spain flanking the Pyrenees into Catalonia, including the mammoth, reindeer, chamois, woolly rhinoceros, and spreading as far west as Santander. ‘This is also the range of the Cro-Magnon race of men. Mammoth.—The woolly mammoth (#. primigenius) now reaches the height of its evolution and specialization. As preserved in the frozen tundras of northern Siberia and as represented in very numerous draw- ings and engravings by the Upper Paleolithic artists, it is the most com- pletely known of all fossil mammalia. Its proportions as shown in the accompanying figure, which represents the information gathered from all sources, are entirely different from those of either the Indian or African elephant. ‘The head is very high and surmounted by a great mass of hair and wool. Behind this is a sharp depression separating the back of the head from the great dorsal hump. The hinder portion of the back falls away very rapidly and the tail is short. ‘The overcoat of long hair nearly reaches the ground, and beneath this is a warm under- coating of wool. As described by Salensky®’ from the wonderfully complete specimen discovered in 1901 on the banks of the Beresowka River in northeastern Siberia, this animal developed characters which absolutely exclude the possibility of its ancestry or relationship to the existing Indian elephants. The hind foot was four-toed, or tetradactyl, and not five-toed as in the living forms. The head was larger as compared with the length of the body than in recent elephants, a character which stands in close connec- tion with the enormous development of the tusks; these were distin- guished by their spiral form, the points directed inward. The ears were very small and covered with hair. The tail was relatively shorter than in the existing elephants and was provided with a tassel of long, bristly hair at the end. The color of the hair was yellowish brown, varying from hght brown to pure brown, and a coat of woolly hair, 2 to 214 cm. in length, covered the whole body. Interspersed with these were a large number of longer and thicker hairs which formed mane-like patches on 86 HARLE, Epouarp: ‘‘Les Mammiféres et oiseaux quaternaires connus jusqu’ici en Portugal. Memoire suivi d'une liste générale de ceux de la Péninsule Ibérique.’”’” Com- munic. du Service Géol. du Portugal, Tom. viii, pp. 22-85, pll. I-V. Lisbon. 1910. °*T SALENSKY, W.: Uber die Hauptresultate der Erforschung des im Jahre 1901 am Ufer der Beresowka entdeckten miinnlichen Mammutcadavers,”’ C. R. Séa. Six. Congr. Internat. Zo6l., pp. 67-86. Berne, 1904. OSBORN, REVIEW OF THE PLEISTOCENE 305 the cheeks, on the chin, on the shoulders, flanks, abdomen, ete. A broad fringe of this long hair extended along the sides of the body as depicted mn the paleolithic sketches from the Combarelles Cave discovered by Capitan and Breuil in 1901. Especially interesting is the food found in the stomach and mouth, which consists of a meadow flora such as characterizes this region of Siberia at the present day, thus appearing to disprove the theory that the climate was milder than that now prevailing. Nor does it appear that it was more frigid, because there are few repre- Pic. 18.—The hairy mammoth (ELlephas primigenius) and Paleolithic man (Homo neanderthalensis ) Restored by Charles R. Knight under the direction of the author, 1914. Original in the American Museum of Natural History, New York City. sentatives of tundra vegetation. Grasses (Graminew) and sedges (Cy- peracew) predominate. There were also wild thyme (Thymus), beans of the wild oxytropis (Oxytropis compestris), seeds of the alpine poppy (Papaver), and the boreal variety of the upright crowfoot (Ranunculus acer), all still found in this region. Woolly Rhinoceros—TVhe woolly rhinoceros (D. antiquitatis, D. tich- orhinus) is distinctly a cold-weather, or tundra form and the invariable companion of the mammoth. Like D. mercku it has no front, or cut- ting teeth hence it has been improperly considered as related to this spe- cies, but it really belongs to the modern African group of Diceros (Ate- 306 ANNALS NHW YORK ACADEMY OF SCIENCES ° lodus), which is distinguished by a very elongate front horn (Fig. 19) and a small posterior -horn, as in the existing white rhinoceros (D, sumus) of Africa. The resemblance of the cave drawings of the Pleisto- cene animal to the living form is very close indeed except as regards the heavy coat of hair, which, as in the mammoth, extends far below the body. The hair of the face, of a golden brown color with an under- covering of wool, is preserved in the St. Petersburg Museum. Through a discovery (1911) at Starunia®® in Hast Galicia this animal is now Fic. 19.—The woolly rhinoceros (Diceres antiquitatis) Restored by Charles R. Knight under the direction of the author, 1914. Original in the American Museum of Natural History, New York City. completely known to us except the tail. The remains of the woolly rhinoceros were found at a depth of 13.6m., including the head, the left fore leg and the skin of the left side of the body, all with the muscula- ture but lacking the hair. The Starunia specimen exhibits a broad, truncated upper lip, small, oblique eyes, ears long, narrow and pointed, a long nasal horn with oval base and shorter frontal horn, a short neck, 9 NIBZABITOWSKI, H. L.: ““Die Uberreste des in Starunia in einer Hrdwachsgrube mit Haut und Weichteilen gefundenen Rhinoceros antiquitatis Blum. (tichorhinus FWisch.), Vorlaufige Mitteilung.’’ Bull. Acad. Sci. Cracovie, Ser. B, pp. 240-267. April, 1911. OSBORN. REVIEW OF THE PLEISTOCENE 307 on the back of which is a small hump quite independent of the skeleton. The larger hump on the shoulders is formed by the long vertebral spines. The legs are comparatively short. ‘The skin is smooth. Niezabitowski observes that D. antiquitatis resembles D. simus most closely, having in common the elongate head with prominent orbits, the truncated upper lip, the hump on the neck, and the short legs; it differs from D. simus in the somewhat narrower muzzle, small, pointed ears and the presence of a thick coating of hair. Fic. 20.—Rhinoceros skulls Skulls of the Pleistocene ‘‘woolly rhinoceros,” Diceros antiquitatis of Hurasia (above), and of the recent African ‘‘white rhinoceros,” Diceros simus (below). In the American Museum of Natural History. 2 Like D. simus, D. antiquiiatis was a plains dweller living on grass and small herbs. The woolly rhinoceros was confined more closely to the edges of the great ice sheets than the mammoth; that is, 1t did not migrate so far to the south, stopping at the Alps, while the mammoth wandered into Italy as far south as Rome. Llasmotheres—The elasmothere (Hlasmotherium sibericum) was an- other companion of the mammoth which ranged over eastern Europe, Germany, and southern Siberia. It was probably a steppe dweller. In Pleistocene times it is reported as occurring as far south as Sicily.°? It 28 BranpT, J. F.: “Mittheilungen iiber die Gattung EHlasmotherium, besonders den Schiidelbau derselben.” Mem. Acad. Imper. Sci. Pétersbourg, Ser. VII, Vol. XXVI, No. 6. St. Vetersburg, 1878. and Gavupry and Bove‘. “Matériaux pour l’Histoire des Temps Quaternaires.” Fasc. 31. L’Blasmotherium. Paris, 1888. 308 ANNALS NEW YORK ACADEMY OF SCIENCES. is a gigantic animal, distinguished from all the European Pleistocene rhinoceroses by the entire absence of the anterior horn and the pos- session of an enormous horn situated on the forehead between the eyes; also by the elaborate foldings of its dental enamel, to which the name Elasmotherium refers. Its hypsodont and folded teeth were especially adapted to a grassy diet, and Gaudry connects its appearance in Europe with the extensive deforestation accompanying the Steppe and Tundra periods of mammalian life. It apparently wandered into EKurope from central Asia and never became very abundant. The elasmothere is pos- sibly descended?’ from the Aceratherium of the Upper Miocene of Ep- pelsheim which has perfectly smooth, pointed nasals, and the rudiment of a horn between the eyes. Horses of the Pleistocene-—The horse was distributed all over the northern hemisphere in Pleistocene times in the Glacial, Interglacial, and Postglacial Epochs. In America no Postglacial horses are found. In Europe horses were apparently abundant in Postglacial times and two of the natural breeds appear to have given origin to two of the modern domesticated types. The horses of the long warm Second Interglacial Stage were remarkable for their great size (#. siissenborn- ensis, #. mosbachensis) which exceeded that of the largest recent breeds (Pohlig, 1907).*°* According to Pohlig the horses were at all times ac- companied by the wild asses (H#. hemionus) but this we are inclined to believe was a special feature of the dry and cold steppe periods in which we should expect to find asses similar to the dzeggetai of Asia of present time. The existing wild ass, or kiang, of the Asiatic steppes certainly appears in early Postglacial times at Wildscheuer, Thaingen, NKessler- loch, and Schweizersbild associated with the late Aurignacian Palzo- lithic culture. Reference of the ancient Pleistocene horses to #. caballus is certainly erroneous. The connection of these ancient Pleistocene horses with the modern species and races requires further investigation. We should expect to discover in Europe horses of three different habi- tats or life zones, namely, of the dry African-Asiatic plains, of the Eurasiatic forests and meadows, of the tundras and steppes. Such an- ticipation appears to be verified through the new lines of study instituted by Ewart?’ since 1904. Following more or less closely the work of pre- vious students of the Equide he has shown that the different wild breeds of horses have evolved in three kinds of environment. Thus we discover 10 OsBorN, OH. F.: “Frontal Horn on Aceratherium incisirum. Relation of the type to Hlasmotherium.” Science, N. S., Vol. IX, No. 214, pp. 161-162. Feb., 1899. 1 PoHLIG, H.: Hiszeit und Urgeschichte des Menschen. Leipzig, 1907. 102 Hwart, J. Cossar: “The Multiple Origin of Horses and Ponies.’’ Trans. High- land. Agric. Soc. of Scotland, pp. 1-39. 1904. OSBORN, REVIEW OF THE PLEISTOCENE 309 horses adapted to: (1) forests and upland valleys; (2) high, dry, cold steppes: (3) warm deserts and plateaus. In these three chief habitats the horses may be respectively known as the “forest horse,” the “steppe horse,” and the “desert horse.” Hach has its distinctive coloring, tooth structure, and proportions of the skull, body and limbs, in adaptation to its peculiar mode of life. The forest horse is relatively a large, clumsy animal. The face is arched, as in the modern draught horse. The limbs are short, the front cannon bone (Mtc. II1) being short and stout, the length only 514 times the width. The tail is depressed in contrast with that of the desert horse. According to Ewart this type of horse (EH. robustus) occurs at Solutre and in the Neolithic deposits of Ilford (Essex), and Kent. In Aurignacean times Solutré was the site of a great open air Paleolithic camp. ‘Toussaint enumerates fragments of ‘at least 100,000 horses, which mingled with other bones of the chase formed a sort of rampart around the camp. The majority of these horses belonged to the stout- headed, short-limbed forest, or Norse type, measuring 54 inches (13.2 hands) at the withers, the size of the existing pony.'?* The large joints and hoofs were especially adapted to the low-lying marshy ground in the vicinity of forests, and the long teeth and powerful jaws were adapted to feeding during parts of the year on coarse grasses, roots and other hard substances. There is no evidence that the men of the Aurignacean age either bred or reared these animals. The majority of the remains are those of horses from five to seven years of age. This type of horse is engraved on the walls of the cave of Combarelle, where the drawings are chiefly of old Magdalenian age and the pure forest type of horse is most frequently represented. ‘There is also a small, fine-headed type suggest- ing the desert horse, and a larger, long-armed type suggesting the wild ass. The desert horse is the Pleistocene animal identified by Richard Owen as an ass (Ff. asinus fossilis), but considered to be a horse by Ewart and named by him #. gracilis. This is a small animal, not over 12.2 hands in height, slender-limbed, with long, slender front cannon bones (Mte. ITI), the length being 714 times the width. The head is small, the face fine and narrow, with a straight profile only slightly deflected upon the cranium. ‘The internal cusp (protocone) of the upper molars is short. Remains of an animal of this type are found in the Pliocene of Italy (smal!, slender-limbed varieties of H. stenonis) and France, and in the Pleistocene of France and northern Africa. It agrees, so far as known, with the existing Celtic pony type (H. caballus celticus), a variety of 102 WWARL: Op. cit. 210 ANNALS NEW YORK ACADEMY OF SCIENCES horse distinguished by small, fine head, large eyes, slender limbs, five lumbar vertebre, now found in more or less pure form in the outlying islands and on the coast of western Europe. This animal is believed to be a northern, hardy, thick-coated relative of the pure desert type, better known as the Arabian, which gave rise to the modern thoroughbred. Perfect representations of this type of horse are found in the engravings and mural paintings of the Magdalenian artists in the caverns of Font de Gaume, Combarelle, and Grotte de la Mairie. A possible contributory to the desert breed of the Pleistocene and of the modern domesticated horses is the animal of the /. swvalensis type of the Upper Pliocene in the Siwaliks of India. This animal is tall, with long, fairly slender limbs, long neck, well elevated tail, long face, which is strongly deflected on the cranium with a slightly convex profile and broad brow, and teeth with a narrow protocone. Bears.—The Postglacial bears (Ursus speleus) are found in greater abundance than the lions. They include a gigantic and a smaller va- riety. The former (Ursus spelwus) nearly equalled the larger recent bears in size and were more thick-set and of heavier proportions; the front paws especially were of tremendous size. During a long period the cave bears undoubtedly haunted the caverns undisturbed by Paleeo- lithic man and developed certain peculiarities of structure in these haunts: thus the claw-bearing phalanges are feebly developed, indicating that the claws had partly lost their prehensile function; the anterior premolar teeth are practically vestigial: the cusps of the grinding teeth are blunted in a way which is indicative of an omnivorous diet. It would. appear, therefore, that the large herbivorous mammals and even primi- tive man found no very formidable enemy in the cave bear. While the other and smaller races were contemporary, there are certain indications that the smaller race (Ursus sub-speleus) was geologically older, being found at Mosbach during the Second Interglacial Stage. Both races became extinct during Postglacial times without leaving descendants. . The ancestor (Ursus arctos priscus) of the brown bear of Europe, by some believed to be a descendant of the etruscan bear (Ursus etruscus) of the First or Norfolkian Interglacial Stage, is also occasionally found in the Pleistocene cave deposits. It is not so large as the cave bear and while it has been mistakenly identified with the American grizzly (U. horribilis) in reality it has closer affinities to the European brown bear (Ursus arctos). The badger (Meles tarus) also probably originated in west-central Asia, since the only other species known are confined to Asia. The two OSBORN, REVIEW OF THE PLEISTOCENE 211 o extinct Lower Pliocene species are found in Maragha, Persia (M. polaki, M. Maraghanus) .*%* TRANSITION TO THE EUROPEAN FOREST STAGE This transition is believed to have begun late in Postglacial times, toward the end of the Magdalenian culture period. Evidence that the mammoth fauna lingered late both in the Dordogne region of central France and to the north is found in the abundant representation of the mammoths in the very latest paintings and engravings by the Magda- lenian artists. MIGRATION OF THE TUNDRA FAUNA The backward or northward migration of the Tundra fauna is be- lieved to have occurred in the following manner.*® As the glacial caps retreated they left barren stretches behind them and the valleys and plateaus now free from ice became tundras where swamps alternated with patches of polar willows and stunted fir trees, while other areas were covered merely with low, scrubby birches or reindeer moss and lichens. As these climatic conditions shifted northward before the re- treat of the great Scandinavian glaciers the Tundra fauna followed. It was a slow change that drove the Tundra mammals toward the dry re- gions of the east to make room for the forests and their faunas advancing from the south. It is clear that the north and east were the only direc- tions of retreat for the damp climate and the spread of the woodlands. RETREAT OF THE STEPPE FAUNA As long ago as 1890 Nehring’ held that the Steppe period of central Europe was partly in Postglacial times. This opinion was supported by Woldrich (1896),1° and has been abundantly confirmed by Harlé’s ob- servations in southern France and by the recent researches of Koken and Schmidt. Steppe conditions of climate appear probable from the exten- sive depositions of the “Newer Loess” in Postglacial times (Koken, 1% ScHarrr, R. F.: The History of the European Fauna, p. 44. London, 1899. 16 StupErR, T.: “Die Tierreste aus den pleistocenen Ablagerungen des Schweizers- bildes bei Schaffhausen.” Neue Denkschr. allg. schweiz. Ges. Gesam. Naturwiss., Vol. XXXV, pp. 1-38. 1896. 106 NEHRING, A.: Uber Tundren und Steppen der Jetzt- und Vorzeit, mit besonderer Beriicksichtigung ihrer Fauna, pp. 81-166. Berlin, 1890. 107 WoLDRICH, J. N.: “Ueber die Gliederung der anthropologischen Formationsgruppe Mitteleuropas.” Sitzber. kgl. béhm. Ges., math. naturwiss. Class., 1896. Ref. Matiegka in Centralblatt Anthrop., pp. 142-143. 1896. 312 ANNALS NEW YORE ACADEMY OF SCIENCES Schmidt). On the other hand, Kobelt*®® and Scharff*°? agree in think- ing that the presence of Steppe mammals affords inadequate proof of the steppe character of the country in Inter- and Postglacial times. The de- posits of the “Newer Loess” in Postglacial times point to a dry steppe period because according to the theory of Richthofen, which is now gen- erally aecepted, the loess owes its origin to wind-borne dust and sand acting under the influence of a dry climate either in summer or winter. The Steppe Fauna in deposits at several points is shown to have hn- gered longer than the Tundra fauna. As regards the lingering of the Steppe Fauna it is indicated in the succession of the three rodents char- acteristic of the Tundra, Steppe and Forest conditions respectively, namely : Forest climate and conditions, the squirrel (Sciurus vulgaris) Steppe climate and conditions, the jerboa (Alactaga jaculus) Tundra climate and conditions, the banded lemming (Myodes torquatus) The absence of fossil plants in the deposits of the steppe period is due to the unfavorable conditions for the preservation of plant remains. Small stretches of woodland were probably confined to the banks of rivers, to favorable mountain slopes, ete. The flora was probably hke that of eastern Eurasia or southwestern Siberia to-day. In their migrations such animals as the jerboa which were unable to swim presumably crossed the rivers while frozen over. Saiga.—Of the Steppe fauna (fully described on p. 248) the saiga antelope (Saiga tartarica) has at the present time retreated to the steppes of eastern Hurope and western Siberia. This animal is represented in the carvings and engravings of Upper Paleolithic or late Magdalenian times in the Dordogne region of France. Its fossil remains have been found in thirteen localities in southwestern France in association with a cold steppe fauna. In the same region have been found remains of the musk ox (Ovibos). SURVIVAL OF FOREST AND MEADOW FAUNA The final retreat of the cold faunas of the tundras and steppes occurred during the late stages of the Upper Paleolithic Magdalenian culture. The most advanced Magdalenian art continues to represent the woolly mammoth in the cavern of Font de Gaume (Dordogne) and elsewhere, but in the very latest Magdalenian culture stages it would appear that the mammoth and woolly rhinoceros were becoming rare. This final 108 KOBELT, W.: Die Verbreitung der Tierwelt. Gemissigte Zone. Leipzig, 1902. 109 Scuarrr, R. F.: The History of the European Fauna. London, 1899. a OSBORN, REVIEW OF THE PLEISTOCENE 313 Magdalenian culture, which is correlated with the Gschnitz advance (Schmidt, op. cit., p. 270), is later than the Steppe period of the “Upper Rodent” layer, which is correlated with the preceding Bihl advance. At the same time the Cré-Magnon, or Aurignacian type of Homo sa- piens, which we believe to be the artistic race of the Reindeer period, dis- appears or becomes greatly reduced in numbers and new brachycephalic and dolichocephalic races of men enter Europe. Azilian-Tardenoisian, Final Upper Paleolithic Culture-—This is re- garded as the closing culture of Upper Paleolithic times. It is be- heved to be associated with the newly arriving broad-headed Fwrfooz- Grenelle race. Although this point is not positively determined this race is first found at Ofnet in Bavaria. It is readily distinguished from the preceding Magdalenian culture by the degeneration of the stone in- dustry into microlithic and other types and by the entire disappearance of art m all its forms. The Azilian culture is essentially Paleolithic although it embodies only its last degenerate stages. While the perfec- tion of the older crafts was lost forever the Neolithic arts of polishing stone, making pottery, cultivating land and domesticating animals are as yet utterly unknown. The Azilian is the age of the stag for there is no longer any trace of the reindeer or other Tundra forms. The bone imple- ments are now made of the horns of the stag. The Tardenoisian culture, supposed by some to be distinct from the Azilian, is characterized by flint microliths of unusual fineness, but it appears that the Azilian and Tarde- noisian cultures are contemporary (Obermaier, 1912). There were two or more human races in Europe in these pre-Neolithic times including brachycephalic and dolichocephalic types which are found commingled at Furfooz. In the meantime Paleolithic races were ad- vyancing in the north along the shores of the Baltic and preceding the Campignian culture, which is the first of the Neolithic arrivals in the Baltic region. Forest Fauna.—The spread and multiplication of the Eurasiatic For- est Fauna thus occurred before the close of Paleolithic times. Following the retreat of the glaciers and the disappearance of steppe conditions of climate there came a gradual subsidence of the coasts of northern Europe and with it a more humid climate favorable to reforestation. Besides the common squirrel (Sciwrus vulgaris) which is the herald of forest condi- tions all over the northern hemisphere, there appear in larger numbers the entire Forest Fauna which we have traced from its beginnings in early Pleistocene times and which we regard as having been resident in favorable localities throughout the entire epoch. With the Tundra and Steppe Faunas disappear also the wolverine (Gulo luscus) and the lion S34! ANNALS NEW YORK ACADEMY OF SCIENCES 2 (Felis leo spelea), which are never found in western Europe after the Pleistocene although the lion lingered until a late period in eastern Kurope. The Alpine Fauna, which is mainly of central Asiatic rather than of northern relationships, retreats to the higher levels of the Alps and the Pyrenees. ‘Thus there remained in the forests, in the plains and in the lower mountain regions of Hurope the direct descendants of the Hura- siatic Forest and Meadow Fauna of the Pleistocene. It is noteworthy that no new mammals appear in Europe at this time except those intro- duced by man. The fauna of early Neolithic times is directly sequent upon that of late Paleolithic times. This fauna has been discovered in the Swiss lake dwellings? (Fig. 9, 38-40) at Moosseedorf, Wauwy]l, Robenhausen, Concise, etc. In the peat bogs of Hassleben (41), etc., in the travertines of Jena, Langensalza (42), etc.,11t have been found the following mammals: Forrest AND MEADOW Bison bonasus, the European bison, still surviving in Lithuania. Bos primigenius, collateral ancestor of the long-horned larger existing cattle of western Europe. The “urus,’ of Cesar’s text. Surviving in Germany until the sixteenth century. Bos longifrons, the “Celtic short-horn,” the probable ancestor of the small breeds of British short-horned and hornless cattle. Cervus elaphus, the red deer or stag. Cervus capreolus, the roe deer. Alces machlis, the elk or moose. Rangifer tarandus, the reindeer, surviving in central Europe until the twelfth century. Cervus dama, the fallow deer. Sus scrofa ferus, the wild boar. Sus scrofa palustris, the turf pig. Equus caballus celticus, the Celtic pony, representative of the “plateau” type. ; Equus caballus typicus, the Norse, or “forest” horse. Castor fiber, the beaver. Sciurus vulgaris, the common squirrel. Lepus timidus, the European hare. Lepus variabilis, the arctic hare, in Ireland and the north. Mus sylwaticus, the field mouse. Arcitomys marmotta, the marmot of the alpine fauna. Ursus arctos, the brown bear. Meles taxus, the badger. Ho RUTIMEYER, L.: “Die Fauna der Pfahlbauten der Schweiz,’ Neue Denkschr. allg. schweiz. Gesell. gesam. Naturwiss., Vol. XIX. Ziirich, 1862. 11 PoHLiG, H.: “Vorliinfige Mittheilungen iiber das Plistoeen, insbesondere Thiirin- gens,” Sitzungsber. Niederrhein. Ges. Bonn, pp. 2-15. Mar. 3, 1884. OSBORN, REVIEW OF THE PLEISTOCENE 315 Mustela maries, the pine marten, also the weasel, pole cat, the ermine, ete. Lutra vulgaris, the otter. Gulo luscus, the wolverine. Canis lupus, the wolf. Canis vulpes, the fox. Felis catus, the wild cat. ALPINE Capra ibex, the ibex of the mountain or alpine fauna. Rupicapra tragus, the chamois of the mountain fauna. There is evidence of the “plateau” or “Celtic” horse in the Neolithic deposits of Essex and of Switzerland (La Téne) ; it was widely distrib- uted in Europe and Asia in prehistoric times.’ It is beyond the purpose of this volume to trace the history of domes- tication. The dog (Canis familiaris), a descendant of the wolf (Canis lupus), first appears in western Europe late in Upper Paleolithic times.*** The Neolithic immigrants, or men of the New Stone Age, pos- sessed or brought with them cattle, sheep, goats, pigs, horses and dogs. Appreciating the value of domestication, they certainly captured and domesticated three indigenous Huropean species, namely, the Celtic short- horn cattle, the forest horse (H#. caballus typicus) and the Celtic horse (#. caballus celticus). The wild ox (Bos primigenius) was hunted but not domesticated. The domestic ox (Bos taurus) shows many points of resemblance to the Urus, but is not directly descended from it, but rather from the Bos trochoceros type of the Pleistocene of Italy. Riitimeyer has made an exhaustive study of this subject,‘‘* tracing the origin of the various types of domesticated cattle. M2 WWART, J. C.: Op. cit., 1907. 13. TROUESSART considers the diminutive wolf of India, Canis pallipes, as the prin- cipal if not the sole source of all our races of domestic dog. This species of wolf, with the exception of the Sumatran wolf, Canis sumatrensis, is also more closely related thin any other to the dingo of Australia. “L’Origine préhistorique du chien domestique,” Revue des Idées, pp. 388-411. June 15, 1911. 4 RUTIMpyeER, L.: “Die Fauna der Pfahlbauten der Schweiz,’ Neue Denkschr. allg. schweiz. Gesell. gesam. Naturwiss., Vol. XIX. Zjiirich, 1862. ret cantatas ae % PUBLICATIONS OF THE NEW YORK ACADEMY OF SCIENCES - (Lyceum or Naturat History, 1817-1876) The publications of the Academy consist of two series, viz. : (1) The Annals (octayo series), established in 1823, contain the sci- entific contributions and reports of researches, together with the records of meetings and similar matter. A volume of the Annals coincides in general with the calendar year and is seld at the uniform price of three dollars per volume. 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